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Fishes associated with mesophotic coral ecosystems (MCEs) of the La Parguera shelf-edge were surveyed between 2007 and 2011 using mixed-gas rebreather diving. Fishes were identified and counted within belt transects and roving surveys at 30, 40, 50, 60 and 70 m depth. Vertical transects from 70 to 30 m depth helped determine depth distribution ranges. One hundred and three species were identified at MCEs (40–70 m), with high abundances and species richness, though both varied greatly among transects. Most species at MCEs were common inhabitants of shallow reefs, but some were restricted to mesophotic depths. An additional 15 species were added to those previously classified as indicator species of mesophotic areas in Puerto Rico. The MCE fish assemblage was distinct from shallow areas (30 m), with taxonomic composition, abundance and the proportion of trophic guilds varying with increasing depth. The dominant trophic guild within MCEs was the zooplanktivores, while herbivores dominated shallow reefs. Both herbivores and zooplanktivores responded strongly, and oppositely, to depth. The few herbivores associated with deep MCEs are small-bodied species. The largest changes within the mesophotic fish community along the depth gradient occurred at 60 m, similar to that reported for algae and corals, and seem to represent both a response to reduced light and variations in herbivory. The presence of commercially important fishes at MCEs, many considered to be threatened by fishing pressure in shallow areas, suggests that MCEs are important for the conservation of these species. This study represents the first quantitative in situ observations and descriptions of fishes inhabiting MCEs at depths of 50–70 m in Puerto Rico and highlights the role of MCEs as valuable habitats for reef fishes. The composition and distribution of the MCEs fish community should be incorporated when planning for the spatial management of coral reef resources.
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Coral Reefs
Journal of the International Society for
Reef Studies
ISSN 0722-4028
Volume 33
Number 2
Coral Reefs (2014) 33:313-328
DOI 10.1007/s00338-014-1125-6
Fishes associated with mesophotic coral
ecosystems in La Parguera, Puerto Rico
I.Bejarano, R.S.Appeldoorn &
M.Nemeth
1 23
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REPORT
Fishes associated with mesophotic coral ecosystems
in La Parguera, Puerto Rico
I. Bejarano R. S. Appeldoorn M. Nemeth
Received: 12 April 2013 / Accepted: 20 January 2014 / Published online: 13 February 2014
ÓSpringer-Verlag Berlin Heidelberg 2014
Abstract Fishes associated with mesophotic coral eco-
systems (MCEs) of the La Parguera shelf-edge were sur-
veyed between 2007 and 2011 using mixed-gas rebreather
diving. Fishes were identified and counted within belt
transects and roving surveys at 30, 40, 50, 60 and 70 m
depth. Vertical transects from 70 to 30 m depth helped
determine depth distribution ranges. One hundred and three
species were identified at MCEs (40–70 m), with high
abundances and species richness, though both varied
greatly among transects. Most species at MCEs were
common inhabitants of shallow reefs, but some were
restricted to mesophotic depths. An additional 15 species
were added to those previously classified as indicator
species of mesophotic areas in Puerto Rico. The MCE fish
assemblage was distinct from shallow areas (30 m), with
taxonomic composition, abundance and the proportion of
trophic guilds varying with increasing depth. The dominant
trophic guild within MCEs was the zooplanktivores, while
herbivores dominated shallow reefs. Both herbivores and
zooplanktivores responded strongly, and oppositely, to
depth. The few herbivores associated with deep MCEs are
small-bodied species. The largest changes within the
mesophotic fish community along the depth gradient
occurred at 60 m, similar to that reported for algae and
corals, and seem to represent both a response to reduced
light and variations in herbivory. The presence of com-
mercially important fishes at MCEs, many considered to be
threatened by fishing pressure in shallow areas, suggests
that MCEs are important for the conservation of these
species. This study represents the first quantitative in situ
observations and descriptions of fishes inhabiting MCEs at
depths of 50–70 m in Puerto Rico and highlights the role of
MCEs as valuable habitats for reef fishes. The composition
and distribution of the MCEs fish community should be
incorporated when planning for the spatial management of
coral reef resources.
Keywords Fish assemblage Mesophotic coral
ecosystems (MCEs) Depth Rebreather Trophic guilds
Herbivores
Introduction
Coral reef fishes have been extensively studied in many
parts of the tropics, and the composition and ecology of
reef fish communities are well characterized for the upper
30 m, largely because of accessibility provided by con-
ventional scuba diving (Thresher and Colin 1986; Itziko-
witz et al. 1991; Pyle 2000). However, coral ecosystems
can extend to depths of 100 m or more, with large gradi-
ents occurring in key physical parameters such as light,
temperature and vertical relief that are expected to have a
significant impact on overall fish diversity and community
composition (Hinderstein et al. 2010). Additionally, these
mesophotic coral ecosystems (MCEs), i.e., between 40 and
150 m, are generally near coastal margins and are thus
potentially impacted by anthropogenic activities, particu-
larly fishing. As a consequence, documenting the compo-
sition and variability of fishes within MCEs is important
not only for characterizing the full biodiversity of reef
fishes but also for understanding how species and com-
munities respond to natural and anthropogenic change.
Communicated by Biology Editor Dr. Glenn Almany
I. Bejarano (&)R. S. Appeldoorn M. Nemeth
Department of Marine Sciences, University of Puerto Rico,
P. O. Box 9000, Mayagu
¨ez, PR 00681, USA
e-mail: ivonnebeja@gmail.com
123
Coral Reefs (2014) 33:313–328
DOI 10.1007/s00338-014-1125-6
Author's personal copy
At present, research on fishes associated with MCEs is
very limited. Previous studies have shown that MCE fish
assemblages differ in taxonomic structure and abundance
from that of shallower reefs because MCEs include several
species commonly found at shallow reefs but also deep
species restricted to mesophotic depths. In addition, some
shallow species occur in lower numbers within MCEs
while others occur in higher numbers, compared to shallow
reefs (Colin 1974,1976; Feitoza et al. 2005; Brokovich
et al. 2008; Garcı
´a-Sais 2010). Mesophotic coral ecosys-
tems represent the lower distribution of many shallow
species (Colin 1974,1976; Thresher and Colin 1986;
Brokovich et al. 2008; Garcı
´a-Sais 2010) and include en-
demics (species with limited geographic extent) (Pyle
2000; Pyle et al. 2008; Brokovich et al. 2008) and species
restricted to deep areas (Feitoza et al. 2005; Brokovich
et al. 2008). Additionally, MCEs sometimes harbor large
commercially important species threatened by overfishing
at shallower depths (Garcı
´a-Sais et al. 2004; Feitoza et al.
2005). Although management of MCE fish communities is
important for maintaining healthy fisheries and local and
regional biodiversity (Riegl and Piller 2003), a better
understanding of the fishes at MCEs is critical, and only
possible after more robust data (quantitative and qualita-
tive) are obtained, and MCE fish distributions are described
in detail and discussed.
Past work at mesophotic depths has been primarily
conducted using submersibles or remotely operated vehi-
cles (ROVs) (Colin 1974; Nelson and Appeldoorn 1985;
Thresher and Colin 1986), which has provided general
qualitative descriptions but limited quantitative informa-
tion. Due to the extreme geomorphology of slope envi-
ronments, diver-based survey techniques can provide more
quantitative data, but for work in MCEs, this requires the
use of trimix rebreather diving. This technique provides
reasonable bottom time at mesophotic depths to perform
detailed fish censuses, take photographs, collect cryptic
species, or effectively approach fishes within cracks or
crevices. An added advantage is that fishes are less evasive
to the presence of divers because no bubbles are released
with closed-circuit diving. Although this technology has
proven to be ideal for researching mesophotic assemblages,
it is still currently underutilized.
Thus far, MCE research in Puerto Rico and USVI has
focused on environments at depths of 40–50 m. Only
limited information exists on deeper MCEs at depths of
50–70 m. Preliminary observations of the MCE fish com-
munities off the shelf-edge of La Parguera, Southwest
Puerto Rico, were made using scuba in the 1970s by PL
Colin (pers. comm.), with deeper observations obtained
using the Johnson-Sea-Link II submersible to survey
deepwater fish habitats and abundance at depths ranging
from about 100–450 m (Nelson and Appeldoorn 1985).
However, in the last 6 years, a detailed study of the MCEs
off La Parguera has been undertaken under the auspices of
the Coral Reef Ecosystem Studies (CRES) program of the
U.S. National Oceanic and Administration (NOAA). This
program has accessed the MCEs off La Parguera using a
combination of remotely operated vehicle (ROV) transects
for gross characterization and rebreather diving for quan-
titative assessments. Sherman et al. (2010) provided a
detailed description of the geomorphology, biota and flora
of these ecosystems, while a number of new species
descriptions (Ballantine et al. 2009,2011; Ballantine and
Ruı
´z2010,2011; Corgosinho and Schizas 2013; Pesic et al.
2012; Petrescu et al. 2012) have increased our under-
standing of their biodiversity. As part of this larger study,
the main objective of this paper is to present the first
quantitative characterization of the La Parguera MCE fish
assemblage between 30 and 70 m, to describe the changes
in the fish community structure with increasing depth and
to compare and contrast shallow and mesophotic assem-
blages along the insular slope. Here, shallow fish assem-
blages are considered to be those associated with reefs
located along the insular slope/shelf-edge above mesoph-
otic depths, i.e., at 30 m or shallower. This study increases
our understanding of reef fish ecology, highlights the role
of MCEs as valuable habitats for reef fishes and represents
some of the first in situ observations and descriptions of
MCEs deeper than 50 m in Puerto Rico. Results emphasize
the importance of incorporating the composition and dis-
tribution of the MCE fish community when planning for
the spatial management of coral reef resources.
Materials and methods
Study area
The insular shelf of La Parguera is a broad carbonate
platform that extends approximately 10 km offshore, has
an average depth of 20 m and supports an extensive
development of coral reefs, seagrass beds and mangrove
forests. There are distinct cross-shelf gradients unrelated to
depth, but rather to proximity to land (e.g., turbidity, Be-
jarano and Appeldoorn 2013), and there is a marked dis-
tinction between fish assemblages off the shelf-edge and
inshore reefs (e.g., Aguilar-Perera and Appeldoorn 2008;
Nemeth and Appeldoorn 2009). Both the entire shelf and
the shelf-edge have a karst topography (Morelock et al.
1977). The insular slope consists of an upper portion that
extends from 20 to 160 m depth, followed by the basal
slope beneath (Sherman et al. 2010). The shelf breaks at
20–35 m depth and supports a marginal barrier reef with
several narrow channels through which sand is transported
over the shelf-edge (Morelock et al. 1977). Deep buttresses
314 Coral Reefs (2014) 33:313–328
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formed during preexisting sea levels lower than present are
common between 45 and 65 m depth, and a prominent
terrace occurs at a depth of approximately 80 and 90 m.
Below 90 m, a steep wall drops precipitously down to
160 m (Sherman et al. 2010). Coral ecosystems on the
shelf-edge slope of La Parguera extend to waters deeper
than 100 m, but most benthic MCEs are located above
90 m. This zone is characterized by eastward-facing slopes
having a gentler gradient and lower relief than westward-
facing slopes, as a result of geomorphologic changes
related to the prevailing wave exposure over recent geo-
logic time (Sherman et al. 2010). Because active down-
slope sediment transport negatively affects the growth of
benthic organisms, the better-developed MCEs are located
in association with topographic highs, which are patchily
distributed along the insular slope.
Data collection
The distribution of MCEs along the shelf-edge slope of La
Parguera, southwest Puerto Rico, was determined by cou-
pling exploratory diving and ROV surveys with high-res-
olution multibeam bathymetry available from Battista and
Stecher (2006). Potential study sites were then selected
from areas where the multibeam imagery indicated the
presence of complex topography between 40 and 70 m
depth, which suggested the presence of MCEs. Sites were
visited to corroborate MCE development, and six MCE
study sites were chosen for study (Fig. 1). In order to
develop an overall description of the MCE fish assemblage,
these sites were spread evenly over southwest and south-
east facing slopes. Data were collected between November
2007 and December 2011 using closed-circuit rebreather
trimix technical diving. Each site was sampled with a
minimum of four replicates taken at each of five depth
intervals: 30, 40, 50, 60 and 70 m. Fish species saturation
curves showed that few species were added after the fourth
transect at all sites. Fishes were identified and counted
during 15 min along 10 93 m modified belt transects
(30 m
2
) (Brock 1954). The first species recorded were
those that were evasive at the presence of divers (e.g.,
snappers) followed by the more territorial species. In
addition, roving surveys were made simultaneously with
transect surveys, which consisted of recording the fishery
target species outside belt transects. Six vertical roving
transects were performed at each study site, from 70 to
30 m to help determine the maximum depth distribution of
the species in the area. Observations from three exploratory
dives to deeper depths (up to 91 m) were also considered
when determining species depth range. Species unidenti-
fied in the field were photographed and eventually col-
lected using quinaldine and dip nets, to identify them later
in the laboratory. Identifications followed Nelson (2006).
All species were assigned to one of six trophic guilds:
planktivore, herbivore, piscivore, omnivore, mobile inver-
tebrate feeder or sessile invertebrate feeder, following
Randall (1967) or fishbase.org. Depending on its depth
distribution, each species was classified as shallow (when
present at B30 m, or previously reported in the area
occurring at B30 m, even if are now rare due to overfish-
ing) or deep (when restricted to C40 m). Chromis insolata
(sunshine fish), Prognathodes aculeatus (longsnout but-
terflyfish) and Sparisoma atomarium (greenblotch parrot-
fish) were exceptions in that they were considered deep
species although they occasionally occurred in more shal-
low areas.
Data analysis
Multivariate analyses were used to examine the variation in
fish species composition and abundance along a depth
gradient from 30 to 70 m, using nonmetric multidimen-
sional scaling (NMS) ordination analysis in PC-ORD ver-
sion 5.0 (McCune and Mefford 2006), while a
nonparametric multivariate test for group differences
(multiresponse permutation procedure, MRPP) was used to
evaluate compositional differences of fish assemblages
along the depth gradient and between each depth (pairwise
comparisons) (McCune and Mefford 2006). Both ordination
and MRPP analyses used the Sorensen distance measure on
log(x?1) transformed mean fish densities (number of
fishes/transect) of all observed species. This transformation
was useful for the analyses of numerical abundance because
it rescaled disproportionally large and small values, thereby
reducing the range in values. Joint plots were displayed on
the ordination to explore the relationship of trophic guilds
responses to ordination axes. Species most frequent and
abundant at shallow reefs (30 m) and at mesophotic depths
were identified using indicator species analysis (ISA) in PC-
ORD (McCune and Mefford 2006) in order to allow inter-
pretation of single-species responses to depth. To lend more
weight to our results, data were compared with previous
shallow fish data collected in the area at 20 m depth, as part
of a CRES program supported by the NOAA. These data
were collected with the same methodology used in our
research but with larger transects (100 m
2
) (Nemeth and
Appeldoorn 2009). Comparisons with our data were pos-
sible after standardizing densities per m
2
, and calculating
sighting frequencies of the species, and proportion of tro-
phic guilds and species richness in 30 m
2
.
Results
The MCEs off La Parguera between 40 and 70 m depth
possess a fish fauna composed of at least 103 species from
Coral Reefs (2014) 33:313–328 315
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31 families: 82 species were identified within transects and
37 within roving surveys (Tables 1,2). The mean abun-
dance across all species was 56 ind 30 m
-2
(range 5–260
ind 30 m
-2
). The mean species richness per transect was 9
(range 4–16). Species richness at mesophotic depths was
similar among depths, but fish abundance decreased with
depth (Fig. 2a, b).
Most species within MCE (76 %) were common
inhabitants of coral reefs above 40 m, or were previously
common but are now rare. These fishes were classified as
‘shallow’’ but extended their lower distribution to
mesophotic depths (Tables 1,2). Abundance and frequency
of occurrence of most shallow species decreased rapidly
with depth (Fig. 3a). The largest group of shallow species
(34, 43 %) does not occur below 60 m (Tables 1,2). For
example, Stegastes partitus (bicolor damselfish) was the
most common species at 30 m (present in 85 % of the
transects with a mean of 4.2 ind 30 m
-2
±3.7) and was
frequently seen in the upper MCE (in 67 % of the transects
at 40 m and 31 % at 50 m), but rarely reached deeper
zones (Fig. 3a). Other common species (present in [50 %
of the transects at 30 m) following this trend were
Acanthurus bahianus (surgeonfish) and Sparisoma auro-
frenatum (redband parrotfish) (Fig. 3a). In contrast, another
11 shallow species followed an opposite trend as they were
more abundant at depth (Fig. 3b). This group included
large predators historically targeted by fishermen in the
area such as Lutjanus jocu (dog snapper) and L. cyan-
opterus (cubera snapper), Mycteroperca bonaci (black
grouper), and Carcharhinus perezii (reef shark), as well as
Clepticus parrae (creole wrasse) (Tables 1,2). There was a
small group of fishes (9 species) that were common and
abundant along the entire depth range (30–70 m) (Fig. 3c).
Halichoeres garnoti (yellowhead wrasse), Coryphopterus
personatus (masked goby) and Cephalopholis cruentata
(graysby) were present in the majority of the transects
([50 %) at any depth, showing some variability among
depths but without a trend. C. personatus was the most
abundant species between 30 and 60 m, but at 70 m, its
frequency of occurrence and abundance declined (Table 1;
Fig. 1 Map of the mesophotic
study sites location along the
insular platform margin (shelf-
edge) south of La Parguera,
Puerto Rico
316 Coral Reefs (2014) 33:313–328
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Table 1 Fishes recorded at six locations at mesophotic depths along the shelf-edge of La Parguera, southwest Puerto Rico
Species Common name Depth
ranges
Max.
depth
Depth
class
30 m 40 m 50 m
% F MD SD % F MD SD % F MD SD
Acanthurus bahianus Ocean surgeonfish 30–56 56 Sh 51.5 1.2 2.0 16.7 0.5 2.0 10.3 0.1 0.4
Acanthurus chirurgus Doctorfish 30–60 60 Sh 27.3 0.9 1.7 0.0 0.0 0.0 17.2 0.2 0.5
Acanthurus coeruleus Blue tang 30–60 61 Sh 15.2 0.2 0.5 4.2 0.0 0.2 3.4 0.1 0.6
Anisotremus virginicus Porkfish 30–61 61 Sh 15.2 0.2 0.4 4.2 0.1 0.4 6.9 0.1 0.4
Apogon lachneri Whitestar cardinalfish 50–82 106 Dp 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.3 1.2
Aulostomus maculatus Trumpetfish 30–50 50 Sh 6.1 0.1 0.2 0.0 0.0 0.0 3.4 0.0 0.2
Balistes vetula Queen triggerfish 30–71 275 Sh 9.1 0.2 0.8 8.3 0.1 0.3 3.4 0.0 0.2
Bodianus rufus Spanish hogfish 30–60 70 Sh 6.1 0.1 0.2 0.0 0.0 0.0 3.4 0.0 0.2
Calamus penna Sheepshead porgy 61 87 Sh 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0
Canthigaster rostrata Sharpnose puffer 30–70 160 Sh 18.2 0.3 0.7 4.2 0.0 0.2 24.1 0.4 0.9
Caranx lugubris Black jack 52–91 364 Sh 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0
Caranx ruber Bar jack 30–61 91 Sh 12.1 0.9 3.4 0.0 0.0 0.0 6.9 0.2 0.9
Centropyge argi Cherubfish 40–80 106 Dp 0.0 0.0 0.0 8.3 0.2 0.6 0.0 0.0 0.0
Cephalopholis cruentata Graysby 30–80 170 Sh 63.6 1.2 1.2 50.0 0.7 0.9 69.0 1.0 0.9
Cephalopholis fulva Coney 30–70 216 Sh 6.1 0.1 0.2 4.2 0.0 0.2 0.0 0.0 0.0
Chaetodon capistratus Foureye butterflyfish 30–74 91 Sh 36.4 0.7 1.1 20.8 0.3 0.6 37.9 0.6 0.8
Chaetodon ocellatus Spotfin butterflyfish 70 70 Sh 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0
Chaetodon sedentarius Reef butterflyfish 30–70 105 Dp 3.0 0.0 0.2 4.2 0.0 0.2 0.0 0.0 0.0
Chromis cyanea Blue chromis 30–70 70 Sh 75.8 3.7 4.5 50.0 1.4 2.5 62.1 1.7 2.2
Chromis insolata Sunshinefish 30–80 152 Dp 12.1 0.7 2.3 58.3 2.4 3.4 86.2 15.7 18.1
Chromis multilineata Brown chromis 30–50 91 Sh 3.0 0.1 0.3 0.0 0.0 0.0 3.4 0.1 0.7
Chromis scotti Purple reeffish 30–80 120 Dp 0.0 0.0 0.0 8.3 0.1 0.3 3.4 0.3 1.5
Clepticus parrae Creole wrasse 30–80 190 Sh 33.3 4.1 7.8 16.7 4.0 11.7 13.8 1.4 5.8
Coryphopterus glaucofraenum Bridled goby 40–61 61 Sh 0.0 0.0 0.0 8.3 0.1 0.3 6.9 0.2 0.7
Coryphopterus lipernes Peppermint goby 30–60 60 Sh 24.2 0.6 1.4 4.2 0.1 0.4 6.9 0.2 0.8
Coryphopterus personatus Masked goby 30–70 70 Sh 63.6 27.2 68.4 75.0 49.3 67.6 62.1 28.2 43.0
Elacatinus evelynae Sharknose goby 30–74 74 Sh 15.2 0.3 0.9 25.0 0.3 0.6 20.7 0.3 0.7
Epinephelus guttatus Red hind 30–64 100 Sh 3.0 0.0 0.2 8.3 0.1 0.3 0.0 0.0 0.0
Gramma loreto Fairy basslet 30–70 70 Sh 33.3 0.8 1.4 33.3 0.5 0.8 17.2 0.6 1.9
Gramma sp. Basslet 60–64 130 Dp 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0
Haemulon flavolineatum French grunt 30–70 70 Sh 39.4 0.5 0.8 12.5 0.1 0.3 10.3 0.1 0.3
Haemulon plumieri White grunt 30–70 70 Sh 9.1 0.1 0.3 4.2 0.0 0.2 3.4 0.0 0.2
Haemulon sciurus Bluestriped grunt 30–60 60 Sh 18.2 0.2 0.4 8.3 0.1 0.3 10.3 0.1 0.4
Haemulon striatum Striped grunt 70–91 210 Dp 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0
Halichoeres cyanocephalus Yellowcheek wrasse 52–70 91 Dp 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0
Halichoeres garnoti Yellowhead wrasse 30–80 90 Sh 69.7 1.2 1.3 87.5 2.3 1.7 79.3 1.9 1.4
Holacanthus ciliaris Queen angelfish 30–61 82 Sh 6.1 0.1 0.2 4.2 0.0 0.2 10.3 0.1 0.4
Holacanthus tricolor Rock beauty 60–70 135 Sh 6.1 0.1 0.2 0.0 0.0 0.0 0.0 0.0 0.0
Holocentrus adscensionis Squirrelfish 30–53 190 Sh 39.4 0.6 0.8 0.0 0.0 0.0 6.9 0.1 0.3
Holocentrus rufus Longspine squirrelfish 30–70 216 Sh 18.2 0.3 0.7 25.0 0.3 0.7 34.5 0.4 0.7
Hypoplectrus chlorurus Yellowtail hamlet 30–74 74 Sh 6.1 0.2 1.1 4.2 0.0 0.2 3.4 0.0 0.2
Hypoplectrus puella Barred hamlet 50–70 70 Sh 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.1 0.3
Hypoplectrus sp. Hamlet 60–70 70 Sh 6.1 0.1 0.2 0.0 0.0 0.0 0.0 0.0 0.0
Liopropoma carmabi Candy basslet 61 70 Dp 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0
Liopropoma mowbrayi Cave bass 40–70 130 Dp 0.0 0.0 0.0 8.3 0.1 0.4 27.6 0.5 0.9
Liopropoma rubre Peppermint bass 50–70 70 Dp 0.0 0.0 0.0 0.0 0.0 0.0 6.9 0.1 0.3
Liopropoma sp. Bass 60–70 70 Dp 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0
Coral Reefs (2014) 33:313–328 317
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Table 1 continued
Species Common name Depth
ranges
Max.
depth
Depth
class
30 m 40 m 50 m
% F MD SD % F MD SD % F MD SD
Lutjanus analis Mutton snapper 48–91 170 Sh 3.0 0.2 1.0 0.0 0.0 0.0 0.0 0.0 0.0
Lutjanus apodus Schoolmaster 29–70 70 Sh 12.1 0.1 0.3 12.5 0.2 0.7 10.3 0.1 0.3
Lutjanus buccanella Blackfin snapper 52–91 273 Dp 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0
Lutjanus mahogoni Mahogany snapper 30–60 100 Sh 6.1 0.1 0.5 0.0 0.0 0.0 6.9 0.1 0.3
Mulloidichthys martinicus Yellow goatfish 30–36 110 Sh 3.0 0.1 0.3 4.2 0.0 0.2 0.0 0.0 0.0
Muraena retifera Reticulate moray 65–70 76 Sh 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0
Myripristis jacobus Blackbar jacobus 30–62 130 Sh 48.5 1.2 1.7 0.0 0.0 0.0 3.4 0.0 0.2
Neoniphon marianus Longjaw squirrelfish 30–70 91 Sh 48.5 0.8 1.1 25.0 0.3 0.4 37.9 0.6 1.0
Ocyurus chrysurus Yellowtail snapper 30–70 180 Sh 24.2 0.3 0.7 12.5 0.4 1.4 13.8 0.2 0.7
Ophioblennius macclurei Redlip blenny 30–70 11 Sh 3.0 0.1 0.3 0.0 0.0 0.0 0.0 0.0 0.0
Paranthias furcifer Creole fish 50–70 190 Sh 0.0 0.0 0.0 0.0 0.0 0.0 3.4 0.1 0.4
Pomacanthus arcuatus Gray angelfish 40–73 91 Sh 0.0 0.0 0.0 4.2 0.0 0.2 3.4 0.0 0.2
Pomacanthus paru French angelfish 30–70 100 Sh 3.0 0.0 0.2 0.0 0.0 0.0 3.4 0.1 0.4
Prognathodes aculeatus Longsnout butterflyfish 30–79 145 Dp 9.1 0.1 0.3 0.0 0.0 0.0 13.8 0.2 0.6
Pterois volitans Lionfish 30–70 55 Sh 12.1 0.1 0.3 16.7 0.2 0.5 3.4 0.1 0.6
Rhinesomus triqueter Smooth trunkfish 30–41 50 Sh 6.1 0.1 0.2 4.2 0.0 0.2 0.0 0.0 0.0
Rypticus saponaceus Greater soapfish 30–70 140 Sh 6.1 0.1 0.4 8.3 0.1 0.3 3.4 0.0 0.2
Scarus iseri Striped parrotfish 30–60 53 Sh 18.2 0.5 1.5 0.0 0.0 0.0 0.0 0.0 0.0
Scarus taeniopterus Princess parrotfish 30–70 53 Sh 66.7 2.6 2.8 29.2 0.8 1.4 27.6 0.6 1.2
Serranus annularis Orangeback bass 39–82 70 Dp 0.0 0.0 0.0 4.2 0.0 0.2 0.0 0.0 0.0
Serranus luciopercanus Crosshatch basslet 70 190 Dp 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0
Serranus sp. Bass 40 130 Dp 0.0 0.0 0.0 4.2 0.1 0.4 0.0 0.0 0.0
Serranus tabacarius Tobaccofish 40–70 190 Dp 0.0 0.0 0.0 8.3 0.1 0.3 10.3 0.3 1.1
Serranus tigrinus Harlequin bass 30–42 53 Sh 6.1 0.1 0.4 20.8 0.2 0.4 0.0 0.0 0.0
Serranus tortugarum Chalk bass 40–70 396 Dp 0.0 0.0 0.0 16.7 0.5 1.5 3.4 0.1 0.4
Sparisoma atomarium Greenblotch parrotfish 40–73 106 Dp 0.0 0.0 0.0 12.5 0.3 0.9 24.1 0.5 0.9
Sparisoma aurofrenatum Redband parrotfish 30–70 53 Sh 57.6 1.1 1.3 12.5 0.1 0.3 24.1 0.4 0.7
Sparisoma viride Stoplight parrotfish 30–60 50 Sh 30.3 0.5 0.9 0.0 0.0 0.0 3.4 0.0 0.2
Sphyraena barracuda Great barracuda 30–73 110 Sh 3.0 0.0 0.2 4.2 0.0 0.2 0.0 0.0 0.0
Stegastes leucostictus Beaugregory 30–55 11 Sh 48.5 0.8 0.9 12.5 0.3 1.0 6.9 0.1 0.3
Stegastes partitus Bicolor damselfish 30–70 116 Sh 84.8 4.2 3.7 66.7 2.6 2.6 31.0 1.1 2.3
Stegastes variabilis Cocoa damselfish 60 190 Sh 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0
Synodus intermedius Sand diver 50 320 Sh 0.0 0.0 0.0 0.0 0.0 0.0 3.4 0.0 0.2
Thalassoma bifasciatum Bluehead wrasse 30–53 53 Sh 48.5 1.9 3.0 12.5 0.6 1.6 3.4 0.0 0.2
Xanthichthys ringens Sargassum triggerfish 40–70 190 Dp 0.0 0.0 0.0 20.8 0.3 0.5 31.0 0.6 0.9
Species Common name Depth
ranges
Max.
depth
Depth
class
60 m 70 m
% F MD SD % F MD SD
Acanthurus bahianus Ocean surgeonfish 30–56 56 Sh 0.0 0.0 0.0 0.0 0.0 0.0
Acanthurus chirurgus Doctorfish 30–60 60 Sh 3.3 0.0 0.2 0.0 0.0 0.0
Acanthurus coeruleus Blue tang 30–60 61 Sh 3.3 0.0 0.2 0.0 0.0 0.0
Anisotremus virginicus Porkfish 30–61 61 Sh 10.0 0.1 0.4 0.0 0.0 0.0
Apogon lachneri Whitestar cardinalfish 50–82 106 Dp 0.0 0.0 0.0 0.0 0.0 0.0
Aulostomus maculatus Trumpetfish 30–50 50 Sh 0.0 0.0 0.0 0.0 0.0 0.0
Balistes vetula Queen triggerfish 30–71 275 Sh 0.0 0.0 0.0 0.0 0.0 0.0
Bodianus rufus Spanish hogfish 30–60 70 Sh 3.3 0.0 0.2 0.0 0.0 0.0
Calamus penna Sheepshead porgy 61 87 Sh 3.3 0.0 0.2 0.0 0.0 0.0
318 Coral Reefs (2014) 33:313–328
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Table 1 continued
Species Common name Depth
ranges
Max.
depth
Depth
class
60 m 70 m
% F MD SD % F MD SD
Canthigaster rostrata Sharpnose puffer 30–70 160 Sh 6.7 0.1 0.3 7.7 0.1 0.4
Caranx lugubris Black jack 52–91 364 Sh 3.3 0.0 0.2 3.8 0.0 0.2
Caranx ruber Bar jack 30–61 91 Sh 0.0 0.0 0.0 0.0 0.0 0.0
Centropyge argi Cherubfish 40–80 106 Dp 13.3 0.2 0.8 38.5 0.6 0.9
Cephalopholis cruentata Graysby 30–80 170 Sh 50.0 0.6 0.8 53.8 0.9 1.1
Cephalopholis fulva Coney 30–70 216 Sh 3.3 0.0 0.2 3.8 0.0 0.2
Chaetodon capistratus Foureye butterflyfish 30–74 91 Sh 23.3 0.4 0.8 11.5 0.1 0.3
Chaetodon ocellatus Spotfin butterflyfish 70 70 Sh 0.0 0.0 0.0 3.8 0.0 0.2
Chaetodon sedentarius Reef butterflyfish 30–70 105 Dp 3.3 0.1 0.4 3.8 0.1 0.4
Chromis cyanea Blue chromis 30–70 70 Sh 26.7 0.7 1.9 15.4 0.5 1.2
Chromis insolata Sunshinefish 30–80 152 Dp 96.7 15.0 16.2 100.0 11.5 9.4
Chromis multilineata Brown chromis 30–50 91 Sh 0.0 0.0 0.0 0.0 0.0 0.0
Chromis scotti Purple reeffish 30–80 120 Dp 26.7 0.6 1.3 30.8 1.8 4.6
Clepticus parrae Creole wrasse 30–80 190 Sh 10.0 0.4 1.9 3.8 1.2 5.9
Coryphopterus glaucofraenum Bridled goby 40–61 61 Sh 6.7 0.2 0.9 0.0 0.0 0.0
Coryphopterus lipernes Peppermint goby 30–60 60 Sh 3.3 0.1 0.5 0.0 0.0 0.0
Coryphopterus personatus Masked goby 30–70 70 Sh 73.3 30.1 30.2 26.9 6.0 11.7
Elacatinus evelynae Sharknose goby 30–74 74 Sh 16.7 0.4 0.9 3.8 0.0 0.2
Epinephelus guttatus Red hind 30–64 100 Sh 3.3 0.0 0.2 0.0 0.0 0.0
Gramma loreto Fairy basslet 30–70 70 Sh 20.0 0.4 0.8 7.7 0.1 0.3
Gramma sp. Basslet 60–64 130 Dp 3.3 0.0 0.2 0.0 0.0 0.0
Haemulon flavolineatum French grunt 30–70 70 Sh 6.7 0.1 0.3 7.7 0.1 0.3
Haemulon plumieri White grunt 30–70 70 Sh 3.3 0.0 0.2 3.8 0.0 0.2
Haemulon sciurus Bluestriped grunt 30–60 60 Sh 3.3 0.0 0.2 0.0 0.0 0.0
Haemulon striatum Striped grunt 70–91 210 Dp 0.0 0.0 0.0 11.5 1.7 6.2
Halichoeres cyanocephalus Yellowcheek wrasse 52–70 91 Dp 0.0 0.0 0.0 11.5 0.1 0.3
Halichoeres garnoti Yellowhead wrasse 30–80 90 Sh 66.7 2.7 3.3 80.8 1.8 1.5
Holacanthus ciliaris Queen angelfish 30–61 82 Sh 10.0 0.1 0.3 0.0 0.0 0.0
Holacanthus tricolor Rock beauty 60–70 135 Sh 3.3 0.0 0.2 3.8 0.0 0.2
Holocentrus adscensionis Squirrelfish 30–53 190 Sh 0.0 0.0 0.0 0.0 0.0 0.0
Holocentrus rufus Longspine squirrelfish 30–70 216 Sh 20.0 0.2 0.5 15.4 0.2 0.4
Hypoplectrus chlorurus Yellowtail hamlet 30–74 74 Sh 0.0 0.0 0.0 3.8 0.0 0.2
Hypoplectrus puella Barred hamlet 50–70 70 Sh 0.0 0.0 0.0 3.8 0.0 0.2
Hypoplectrus sp. Hamlet 60–70 70 Sh 6.7 0.1 0.3 3.8 0.0 0.2
Liopropoma carmabi Candy basslet 61 70 Dp 6.7 0.1 0.4 0.0 0.0 0.0
Liopropoma mowbrayi Cave bass 40–70 130 Dp 43.3 0.7 1.1 50.0 0.7 0.9
Liopropoma rubre Peppermint bass 50–70 70 Dp 3.3 0.0 0.2 3.8 0.0 0.2
Liopropoma sp. Bass 60–70 70 Dp 3.3 0.1 0.5 3.8 0.0 0.2
Lutjanus analis Mutton snapper 48–91 170 Sh 3.3 0.0 0.2 0.0 0.0 0.0
Lutjanus apodus Schoolmaster 29–70 70 Sh 10.0 0.1 0.3 0.0 0.0 0.0
Lutjanus buccanella Blackfin snapper 52–91 273 Dp 30.0 0.3 0.5 57.7 0.8 1.1
Lutjanus mahogoni Mahogany snapper 30–60 100 Sh 16.7 0.2 0.5 0.0 0.0 0.0
Mulloidichthys martinicus Yellow goatfish 30–36 110 Sh 0.0 0.0 0.0 0.0 0.0 0.0
Muraena retifera Reticulate moray 65–70 76 Sh 0.0 0.0 0.0 3.8 0.0 0.2
Myripristis jacobus Blackbar jacobus 30–62 130 Sh 26.7 0.6 1.2 0.0 0.0 0.0
Neoniphon marianus Longjaw squirrelfish 30–70 91 Sh 50.0 0.7 0.8 34.6 0.5 0.9
Ocyurus chrysurus Yellowtail snapper 30–70 180 Sh 6.7 0.1 0.3 3.8 0.1 0.4
Coral Reefs (2014) 33:313–328 319
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Fig. 3c). The rest of the shallow species (25) were rare or
varied without a pattern in relation to depth. Finally, 25
species associated with MCEs were considered to be
‘deep’’ species because they were only found at or below
40 m, or were scarce at shallow depths and markedly
increased in frequency of occurrence at mesophotic depths
(Tables 1,2; Figs. 3d, 4). Of these, the most common and
abundant was C. insolata, present in 87 % of all transects
between 40 and 70 m and representing on average 23 % of
the individuals per transect. Frequency of occurrence and
relative abundance (%) of this species increased with
depth. Though less abundant, Liopropoma mowbrayi (cave
basslet), Lutjanus buccanella (blackfin snapper), P. acule-
atus and S. atomarium increased in frequency and density
with depth as well. Xanthichthys ringens (sargassum trig-
gerfish) was also common within MCEs up to 60 m. This
group included rare species such as Serranus chionaraia
(snow basslet), S. luciopercanus (crosshatch basslet),
Lipogramma klayi (bicolor basslet) and Phaeoptyx pig-
mentaria (dusky cardinalfish).
The ISA recognized 21 species as mainly responsible for
distribution differences with depth. Thirteen species were
characteristic of shallow habitats and eight of the
mesophotic assemblage (Table 3).
The main trophic guilds within MCEs were zooplank-
tivore, mobile invertebrate feeder and piscivore; however,
some herbivores, sessile invertebrate feeders, and omni-
vores were present at all depths (Figs. 5,6). Zooplankti-
vores were the most diverse and abundant group (36 % of
all species and 52 % of the fishes within 30 m
2
transects),
followed by mobile invertebrate feeders (25 and 20 %,
respectively). Piscivores were diverse and abundant, and
Table 1 continued
Species Common name Depth
ranges
Max.
depth
Depth
class
60 m 70 m
% F MD SD % F MD SD
Ophioblennius macclurei Redlip blenny 30–70 11 Sh 3.3 0.0 0.2 3.8 0.1 0.6
Paranthias furcifer Creole fish 50–70 190 Sh 3.3 0.0 0.2 15.4 0.2 0.5
Pomacanthus arcuatus Gray angelfish 40–73 91 Sh 6.7 0.1 0.3 11.5 0.2 0.5
Pomacanthus paru French angelfish 30–70 100 Sh 0.0 0.0 0.0 3.8 0.1 0.4
Prognathodes aculeatus Longsnout butterflyfish 30–79 145 Dp 16.7 0.2 0.4 50.0 0.7 0.8
Pterois volitans Lionfish 30–70 55 Sh 13.3 0.2 0.5 3.8 0.0 0.2
Rhinesomus triqueter Smooth trunkfish 30–41 50 Sh 0.0 0.0 0.0 0.0 0.0 0.0
Rypticus saponaceus Greater soapfish 30–70 140 Sh 6.7 0.1 0.3 3.8 0.0 0.2
Scarus iseri Striped parrotfish 30–60 53 Sh 3.3 0.0 0.2 0.0 0.0 0.0
Scarus taeniopterus Princess parrotfish 30–70 53 Sh 30.0 0.7 1.2 19.2 0.3 0.7
Serranus annularis Orangeback bass 39–82 70 Dp 0.0 0.0 0.0 3.8 0.0 0.2
Serranus luciopercanus Crosshatch basslet 70 190 Dp 0.0 0.0 0.0 3.8 0.0 0.2
Serranus sp. Bass 40 130 Dp 0.0 0.0 0.0 0.0 0.0 0.0
Serranus tabacarius Tobaccofish 40–70 190 Dp 3.3 0.0 0.2 3.8 0.0 0.2
Serranus tigrinus Harlequin bass 30–42 53 Sh 0.0 0.0 0.0 0.0 0.0 0.0
Serranus tortugarum Chalk bass 40–70 396 Dp 20.0 1.0 2.8 11.5 0.5 1.5
Sparisoma atomarium Greenblotch parrotfish 40–73 106 Dp 20.0 0.2 0.4 15.4 0.3 1.2
Sparisoma aurofrenatum Redband parrotfish 30–70 53 Sh 13.3 0.2 0.5 3.8 0.0 0.2
Sparisoma viride Stoplight parrotfish 30–60 50 Sh 6.7 0.1 0.3 0.0 0.0 0.0
Sphyraena barracuda Great barracuda 30–73 110 Sh 0.0 0.0 0.0 3.8 0.0 0.2
Stegastes leucostictus Beaugregory 30–55 11 Sh 0.0 0.0 0.0 0.0 0.0 0.0
Stegastes partitus Bicolor damselfish 30–70 116 Sh 10.0 0.1 0.3 3.8 0.2 0.8
Stegastes variabilis Cocoa damselfish 60 190 Sh 3.3 0.0 0.2 0.0 0.0 0.0
Synodus intermedius Sand diver 50 320 Sh 0.0 0.0 0.0 0.0 0.0 0.0
Thalassoma bifasciatum Bluehead wrasse 30–53 53 Sh 0.0 0.0 0.0 0.0 0.0 0.0
Xanthichthys ringens Sargassum triggerfish 40–70 190 Dp 30.0 0.5 0.9 19.2 0.3 0.8
Depth ranges (m) are based on minimum and maximum depths recorded in vertical and horizontal transects, roving surveys and exploratory dives
in the area to depths up to 90 m. Max. depth (m) is the maximum depth recorded here or reported by Colin (1974,1976); Nelson and Appeldoorn
(1985); Garcı
´a-Sais et al. (2007); Garcı
´a-Sais (2010), or www.fishbase.org. Depth classification (Depth class, Sh: shallow, D: deep). Sighting
frequency (% F) and mean density (MD) are from standard horizontal visual census transects along a depth gradient from 30 to 70 m. SD is the
standard deviation of the mean density
320 Coral Reefs (2014) 33:313–328
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their proportion increased with depth (Fig. 6). However,
this group was better characterized using data from roving
surveys.
The distribution of trophic guilds varied with depth
(Table 4). The most prevalent trophic guild at 30 m
(shallow depth) was herbivores, representing 31 % of the
species and 36 % of the fishes per transect. Mobile inver-
tebrate feeders and zooplanktivores were also abundant (22
and 21 % of the fishes per transect, respectively), with the
former more diverse (30 % of the species) than the latter
Table 2 Fishes recorded at six locations at mesophotic depths along the shelf-edge of La Parguera, southwest Puerto Rico
Species Common name Depth ranges Max. depth Depth class 40 m 50 m 60 m 70 m
%F %F %F %F
Anisotremus surinamensis Black margate 30–67 20 Sh 4004
Apogon sp. Cardinalfish 42–60 273 Dp 0040
Balistes vetula Queen triggerfish 30–71 275 Sh 8448
Canthidermis sufflamen Ocean triggerfish 41–48 60 Sh 0400
Caranx latus Horse-eye jack 73 140 Sh 4004
Caranx lugubris Black jack 52–91 364 Sh 28 4 8 28
Caranx ruber Bar jack 30–61 91 Sh 0040
Carcharhinus perezii Shark 48–91 65 Sh 12 7 8 12
Cephalopholis cruentata Graysby 30–80 170 Sh 12 0 0 12
Cephalopholis fulva Coney 30–70 216 Sh 0 11 0 0
Dasyatis americana Southern stingray 55 53 Sh 0400
Elagatis bipinnulata Rainbow runner 41–52 150 Sh 0400
Epinephelus guttatus Red hind 30–64 100 Sh 0400
Epinephelus itajara Jewfish 55 100 Sh 0400
Ginglymostoma cirratum Nurse shark 58 130 Sh 0040
Halichoeres cyanocephalus Yellowcheek 52–70 91 Dp 0400
Lachnolaimus maximus Hogfish 30–67 35 Sh 4444
Lipogramma klayi Bicolor basslet 80 145 Dp 0000
Lutjanus analis Mutton snapper 48–91 170 Sh 24 11 8 24
Lutjanus apodus Schoolmaster 29–70 63 Sh 4 26 16 4
Lutjanus buccanella Blackfin snapper 52–91 273 Dp 48 7 12 48
Lutjanus cyanopterus Cubera snapper 40–73 55 Sh 24 7 4 24
Lutjanus jocu Dog snapper 40–74 70 Sh 56 44 56 56
Lutjanus mahogoni Mahogany snapper 30–60 100 Sh 0 11 4 0
Mycteroperca bonaci Black grouper 41–91 50 Sh 12 11 12 12
Mycteroperca venenosa Yellowfin grouper 41–45 137 Sh 0000
Ocyurus chrysurus Yellowtail snapper 30–70 180 Sh 8 26 4 8
Phaeoptyx pigmentaria Dusky cardinalfish 70 50 Dp 4004
Pterois volitans Lionfish 30–70 55 Sh 0440
Scarus guacamaia Rainbow parrotfish 71 25 Sh 4004
Scarus taenopterus Princess parrotfish 30–70 53 Sh 0400
Scomberomorus sp. Mackerel 29–50 55 Sh 0400
Serranus chionaraia Snow basslet 60 90 Dp 0000
Serranus luciopercanus Crosshatch basslet 70 190 Dp 4004
Sphyraena barracuda Great barracuda 30–73 110 Sh 12 4 4 12
Sphyrna sp. Hammerhead 61 80 Sh 0040
Trachinotus falcatus Permit 91 130 Sh 0000
Depth ranges (m) are based on minimum and maximum depths recorded in vertical and horizontal transects, roving surveys and exploratory dives
in the area to depths up to 90 m. Max. depth (m) is the maximum depth recorded here or reported by Garcı
´a-Sais et al. (2007), Garcı
´a-Sais
(2010), Nelson and Appeldoorn (1985), Colin (1974,1976), or www.fishbase.org. Depth classification (Depth class, Sh: shallow, D: deep).
Sighting frequency (% F) and mean density (MD) are from roving surveys along a depth gradient from 30 to 70 m. SD is the standard deviation
of the mean density
Coral Reefs (2014) 33:313–328 321
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(16 %). Omnivores, sessile invertebrate feeders, and pi-
scivores were present with few species and low densities.
However, the dominance of herbivores rapidly declined
with depth (Fig. 5), except for S. atomarium and Centro-
pyge argi (cherubfish). From 30 to 40 m depth, overall
species richness and densities decreased sharply (Fig. 2a,
b), mostly due to the rapid reduction in the number and
densities of herbivores when moving from shallow to
mesophotic depths (Fig. 5). The number of species of all
trophic groups diminished in this depth range, except for
zooplanktivores, which instead increased in species num-
ber and densities (Table 3; Fig. 5). As a result, a domi-
nance shift occurred at 40 m from herbivores to
zooplanktivores, a trend which became stronger with depth
as herbivores continuously decreased in species richness
and density while zooplanktivores increased (Fig. 5). The
other trophic groups remained relatively similar among all
depths.
Data from roving surveys unmasked a large represen-
tation of piscivores within MCEs, which was underesti-
mated using only transect data. Roving surveys provided
information on another 22 species that were in the area but
not within transects, and most of them were piscivores
(Table 2). Lutjanus apodus (schoolmaster) was the most
abundant fish in roving surveys at 40 and 50 m depth, but
its abundance and frequency of occurrence decreased from
60 to 70 m. L. jocu, on the contrary, became more frequent
and abundant until 60 m depth, where it was dominant in
number. The upper limit of L. buccanella was 60 m, and its
abundance increased with depth, becoming the dominant
piscivore at 70 m. In addition, roving surveys detected the
presence of species now rarely seen shallower in La
3
6
9
12
15
18
20 30 40 50 60 70 80
Number of Species
Depth (m)
a
0
40
80
120
160
20 30 40 50 60 70 80
Fish density
Depth (m)
b
Fig. 2 Changes in fish community structure along a depth gradient from 30 to 70 m. aMean species richness; bmean fish density, with standard
error bars, per 30 m
2
across six reefs in La Parguera shelf-edge, southwest Puerto Rico
0
10
20
30
40
50
60
70
80
90
20 40 60 80
Sighting frequency (%)
Depth (m)
average trend surgeonfish
redband parrotfish bicolor damselfish
a
0
10
20
30
20 30 40 50 60 70 80
Sighting frequency (%)
Depth (m)
large predators average trend
creole wrasse
b
0
10
20
30
40
50
60
70
80
90
20 40 60 80
Sighting frequency (%)
Depth (m)
average trend graysby
yellowhead wrasse masked goby
c
0
10
20
30
40
50
60
70
80
90
100
20 30 40 50 60 70 80
Sighting frequency (%)
Depth (m)
average trend sunshinefish
cave basslet blackfin snapper
longsnout butterflyfish greenblotch parrotfish
d
Fig. 3 Species sighting
frequency (%) along a depth
gradient from 30 to 70 m, across
six reefs in La Parguera shelf-
edge, southwest Puerto Rico.
Shallow species are those
present at B30 m, or previously
reported in the area occurring at
these depths, even if are now
rare due to overfishing.
aShallow species that
decreased in frequency with
depth, bshallow species that
increased in frequency with
depth, cshallow species that
were frequent along the entire
depth range. dDeep species
restricted to C40 m, except for
&C. insolata (sunshine fish), P.
aculeatus (longsnout
butterflyfish) and S. atomarium
(greenblotch parrotfish), which
occasionally occur shallower.
‘Average trend’’ is the averaged
sighting frequency for each
group
322 Coral Reefs (2014) 33:313–328
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Parguera, such as Epinephelus itajara (goliath grouper), M.
bonaci,M. venenosa (yellowfin grouper), Lutjanus cyan-
opterus,Scarus guacamaya (rainbow parrotfish), sharks
(e.g., C. perezii,Sphyrna sp., Ginglymostoma cirratum)
and rare small species (basslets, cardinalfishes).
Fish communities showed greater structure along depths
gradients than expected by chance (Monte Carlo procedure,
p=0.004) (Fig. 7a), with depth strongly associated with
two independent ordination axes (orthogonality =93 %)
that explained 90 % of the fish assemblage variance. Thus,
shallow water sites were located in the lower right portion
of Fig. 7a, while deep water sites were located to the upper
left. Sites with intermediate depths were located between
these extremes. The overlay of joint plots with trophic
guild responses to ordination axes showed herbivores and
zooplanktivores strongly associated with axis 1 (r=0.82,
s=0.65, and r=0.51, s=0.35, respectively) and axis 2
(r=0.47, s=0.30, and r=0.15, s=0.05), and there-
fore with depth (Fig. 7b, c). Significant variation in species
composition and abundance for the entire fish assemblage
was demonstrated among depths (MRPP, p\0.0001).
Fishes from 30 and 70 m depth were significantly different
from all others (40, 50, 60 m) (MRPP, all p\0.01,
Table 5). Fish assemblages at 70 m were dissimilar from
all other mesophotic assemblages (p\0.005) (Fig. 7a).
Discussion
The MCE fish assemblage between 40 and 70 m off La
Parguera was diverse and unique because its taxonomic
composition and abundance differed from that of shallower
areas and varied with increasing depth. Results from NMS
ordination and MRPP analysis showed that the shallow
assemblage at 30 m differed from all mesophotic assem-
blages. Thirteen species were more abundant at shallow
depths than expected by chance, and 8 species were char-
acteristic of MCEs (Table 3).
A wide variety of fishes inhabit MCEs off La Parguera,
including species common on shallow reefs (e.g., acan-
thurids), species scarce at shallow but common at
mesophotic depths (e.g., S. atomarium) and species
restricted to deep areas (e.g., L. buccanella). Overall, the
presence of ‘‘shallow’’ and ‘‘deep’’ species at mesophotic
depths has been reported for other areas, e.g., Jamaica,
Belize, Bahamas (Colin 1974,1976), the Marshall Islands
(Thresher and Colin 1986), the Red Sea (Brokovich et al.
2008), and Bajo de Sico and Isla Desecheo, Puerto Rico
(Garcı
´a-Sais et al. 2007; Garcı
´a-Sais 2010). However,
some differences were noted. For example, juveniles of
both L. buccanella and Haemulon striatum (striped grunt)
were part of the deep species assemblage in our study
(reaching up to 70 m depth); at MCEs in Jamaica and
Belize, these juveniles occurred shallower and only adults
were seen between 50 and 100 m depth (Colin 1974).
Mesophotic coral ecosystems are considered to be exten-
sions of shallower coral reefs; therefore, the presence of
shallow reef fishes at mesophotic depths is possible if there
is biological and physical connectivity between these
ecosystems (Hinderstein et al. 2010). In our study, 25
species were classified as deep (Tables 1,2), an increase in
15 species as indicators of mesophotic assemblages in
Puerto Rico over those previously described by Garcı
´a-Sais
0
25
50
75
100
20 30 40 50 60 70
Relative density (%)
Depth (m)
Deep species Shallow species
Fig. 4 Proportion of shallow species and deep species (see Fig. 3)
per depth, along a gradient from 20 to 70 m pooled across six reefs in
La Parguera shelf-edge, southwest Puerto Rico. Data from 20 m were
taken from the CRES program (see ‘Materials and methods’’ )
Table 3 Indicator species analysis with Monte Carlo test of signifi-
cance (pvalues) indicating primary characteristic species of shallow
and mesophotic habitats pooled across six reefs in La Parguera shelf-
edge, southwest Puerto Rico
Species Depth group (m) pvalue
Acanthurus bahianus 30 0.001
Acanthurus chirurgus 30 0.019
Acanthurus coeruleus 30 0.054
Chromis cyanea 30 0.010
Coryphopterus lipernes 30 0.007
Haemulon flavolineatum 30 0.009
Scarus iseri 30 0.023
Scarus taeniopterus 30 \0.001
Sparisoma aurofrenatum 30 \0.001
Sparisoma viride 30 0.001
Stegastes leucostictus 30 \0.001
Stegastes partitus 30 0.004
Thalassoma bifasciatum 30 0.002
Serranus tigrinus 40 0.019
Chromis insolata 60 0.025
Liopropoma mowbrayi 60 0.036
Centropige argi 70 0.045
Chromis scotti 70 0.013
Haemulon striatum 70 0.027
Lutjanus buccanella 70 \0.001
Prognatodes aculeatus 70 0.012
Coral Reefs (2014) 33:313–328 323
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(2010); six of them were found in this study because they
live deeper than 50 m. Thus, similar to other localities
(Kahng et al. 2010; McClain and Barry 2010; Rooney et al.
2010), MCEs in La Parguera are valuable habitats con-
taining a high diversity of species ecologically connected
to shallower areas. Habitats with high diversity of species
usually support high genetic variability, functional
redundancy and resilience (Walker and Salt 2006; Ives and
Carpenter 2007). Therefore, to really understand the
dynamics of reef fishes and manage their use and conser-
vation, a landscape approach that includes mesophotic
populations should be considered.
0
25
50
75
100
20 30 40 50 60 70
Relative density
Depth (m)
a
0
25
50
75
100
20 30 40 50 60 70
Relative number of species
Depth (m)
b
Fig. 5 Mean trophic guild proportions observed in 30 m
2
transects
distributed along a 20–70 m depth gradient, pooled across six reefs in
La Parguera shelf-edge, southwest Puerto Rico. Data from 20 m were
taken from the CRES program (see ‘Materials and methods’’).
Zooplanktivore (black), herbivore (white), mobile invertebrate feeder
(diagonal stripes), omnivore (dark gray), piscivore (black dots) and
sessile invertebrate feeder (light gray). aRelative density of each
trophic guild per 30 m
2
.bRelative species richness of each trophic
guild per 30 m
2
0.0
0.5
1.0
1.5
2.0
2.5
3.0
40 50 60 70
Number of species
De
p
th (m)
Fig. 6 Distribution of the mean species richness of trophic guilds in
roving surveys in a 40–70 m depth gradient across six reefs in La
Parguera shelf-edge, southwest Puerto Rico. Piscivore (black dots),
all other trophic guilds (gray)
Table 4 Relative species richness (% R) and density (% D) of tro-
phic guilds per 30 m
2
, along a depth gradient from 30 to 70 m pooled
across six reefs in La Parguera shelf-edge, southwest Puerto Rico
30 m 40 m 50 m 60 m 70 m
%
R
%
D
%
R
%
D
%
R
%
D
%
R
%
D
%
R
%
D
Z21163231563560415836
H3631 28211316 611 610
MI 22 30 23 27 16 24 20 24 19 25
O11965354659
P 8 10 8 11 8 13 6 12 6 19
SI35344736 611
Zzooplanktivore, MI mobile invertebrate feeder, Hherbivore,
Oomnivore, Ppiscivore, SI sessile invertebrate feeder
Fig. 7 a Nonmetric multidimensional scaling (NMS) plot based on
Bray–Curtis similarities of fish assemblages at six reefs in La
Parguera shelf-edge, southwest Puerto Rico, in a depth gradient from
30 to 70 m. NMS with herbivore (b) and zooplanktivore (c) density
distributions (circles size is proportional to density value), and trophic
guilds responses to ordination axes. H, herbivore; Z2, zooplanktivore
excluding dominant masked goby (C. personatus); O, omnivore; P,
piscivore
324 Coral Reefs (2014) 33:313–328
123
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The proportion of shallow species rapidly decreased
with increasing depth, as a result of changes in abundances
and in species composition, and this trend was evident at
even shallower depths. Comparison of these results to data
collected at 20 m shows that the decline in abundance of
shallow species within mesophotic depths is a continuation
of a trend that starts at shallower depths (Fig. 4). At the
species level, those shallow species that declined the most
in sighting frequency with depth (e.g., S. partitus, acan-
thurids and S. aurofrenatum) were the same species that
had the highest sighting frequencies at 20 m depth
(C97 %). Even though decreases in the relative abundance
of shallow species with depth were gradual, the greatest
decline occurred from 60 to 70 m, where a high species
turnover occurs. Thirty-four of 79 shallow species did not
occur at 70 m, so that this was also the depth over which
deep species become dominant within the mesophotic fish
assemblage (Fig. 4). Deep species were not recorded in
studies at 20 m, except for relatively infrequent sightings
of S. atomarium and P. aculeatus. Despite the disappear-
ance of shallow species with increasing depth, overall
species richness and mean densities were conserved over a
broad depth range due to the appearance of deep species.
However, at 70 m, both species richness and especially
density decreased, and the fishes at 70 m formed a distinct
mesophotic assemblage. Most deep species were solitary,
and shallow species that reached deeper zones were present
in far lower densities.
Sharp changes in the fish assemblage between 60 and
70 m depth were consistent with variations in the compo-
sition and abundance of the mesophotic coral and algal
assemblages at these areas (Sherman et al. 2010). Coralline
algae are more abundant at deeper zones of MCEs in La
Parguera, especially below 60 m where they are the dom-
inant algal taxa (Sherman et al. 2010). In contrast, abun-
dance of non-calcareous algae abruptly declines between
60 and 70 m, and those that do extend into deeper depths
often undergo morphological change. For example, the
low-light adapted alga Lobophora variegata (Runcie et al.
2008) has a foliose form in the upper 60 m in La Parguera,
but below that depth, it is only found in encrusting form (H
Ruı
´z and DL Ballantine pers comm). In corals, the domi-
nant species (68 % of overall coral cover) at 70 m were
Agaricia undata, where it forms large plate-like colonies,
but it was rare (2.5 %) at shallower depths (Sherman et al.
2010). Significant shifts in assemblages at 60–70 m depth
are reported in different countries and oceans (e.g.,
Thresher and Colin 1986; Rooney et al. 2010; Bridge et al.
2011); therefore, this may be a common transition depth
for mesophotic communities across a wide geographical
area and in different ecological settings. Nevertheless, this
particular depth limit will depend on local physical and/or
biological parameters, such as, light penetration. The low-
light conditions that result from steep decreases in light
irradiance and spectral quality with depth can be utilized
by only those few autotrophic organisms that can adapt to
increase their photosynthetic efficiency (Lesser et al.
2010). To maximize light capture, some algae can increase
their surface area by lateral spreading, growing in flattened
morphologies (Hanisak and Blair 1988; Aponte and Ball-
antine 2001). However, despite the presence of low-light
adapted algae below 60 m, light levels are probably too
low below this depth off La Parguera to maintain sufficient
production to support benthic herbivores, and therefore
changes in the food chain are expected. On the other hand,
nutrients and particulate matter that may be imported from
rich deep waters to mesophotic depths by upwelling and
internal waves (Leichter et al. 1996,2003; Leichter and
Genovese 2006) may provide the main source of energy in
mesophotic habitats through plankton production (Lesser
2006), and therefore benefit planktivores and filter feeders.
Additionally, changes in temperature, sedimentation, geo-
morphology, competition and predation also influence the
ecology of MCEs (Kahng et al. 2010). Thus, the resources
needed by most common shallow species in La Parguera
shelf-edge were in sufficient supply up to 60 m, but below
this depth, a different biota becomes dominant. Lastly,
changes in the benthic environment could be linked to
changes in the fish assemblage at this depth. Although this
study did not survey habitat variability among depths,
personal observations, as well as data from Sherman et al.
(2010), suggest little differences in habitat availability in
the range of 60–70 m. Nevertheless, marked habitat
changes occur deeper: at the formation of the terrace, at
80 m and at 90 m, where the vertical wall starts.
As a consequence of the above transition, the trophic
structure of the fish assemblage at MCE depths differed
from that of shallower reefs. Zooplanktivores were domi-
nant on MCEs, representing more than half of the fishes
within the 30 m
2
transects. At shallow reefs, zooplankti-
vores only represented 19 % of the fishes and the pre-
dominant guild was herbivores, a group that was scarce at
MCEs (Fig. 5) and that represented as low as 6 % of the
Table 5 Multi-response permutation procedures (MRPP)
T 30m 40m 50m 60m 70m
30 m 23.3 25.5 26.5 26.9
40 m 20.9 24.1 24.8
50m–––21.4 23.3
60m––––23.6
70m–––––
Pairwise comparisons between depths, from 30 to 70 m, of fish
assemblages from six reefs in La Parguera shelf-edge, southwest
Puerto Rico. Tvalues. Bold values correspond to significant differ-
ences (p\0.005)
Coral Reefs (2014) 33:313–328 325
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fishes within transects at 70 m depth. Herbivores (e.g.,
acanthurids), mostly reached the upper (40–50 m) but not
the deeper (60–70 m) portion of MCEs, and their densities
decreased markedly with depth. This relative scarcity of
herbivorous fishes is characteristic of MCEs (e.g., Gil-
martin 1960; Van den Hoek et al. 1978; Liddell and Ohl-
horst 1988; Garcı
´a-Sais et al. 2008; Brokovich et al. 2008,
2010). At the same time, while algae are less diverse and
grow slower in MCEs (Brokovich et al. 2010), they
maintain high abundances (Sherman et al. 2010) due in part
to low grazing pressure (Van den Hoek et al. 1978; Liddell
and Ohlhorst 1988; Morrison 1988; Leichter et al. 2008;
Brokovich et al. 2010). Several hypotheses have been
postulated to explain the low representation of herbivores
in MCEs, e.g., low algal productivity, nutritional value,
palatability or digestibility. While changes in algal nutri-
tional characteristics with depth are still unproven (Cle-
ments et al. 2009), some of the genera occurring in high
abundances at MCEs, such as Lobophora or Halimaeda,
although edible (Colin 1978), are known to be less palat-
able to most herbivorous fishes (Duffy and Hay 1990). In
addition, large grazers concentrate in zones with high rates
of algal turf production (Russ 2003); therefore, the low
number of these fishes within MCEs may be related with
the slower growth and turnover of the algae at deep depths.
Some herbivorous species had their maximum occurrence
at the deepest depths surveyed, but these were only small-
bodied herbivores, such as the S. atomarium and C. argi,
which were the most abundant at 70 m. These fishes may
have found an ecological niche at these depths where the
low productivity of algae on which they feed might be
sufficient for their energetic demand given their small size.
By grazing deep in MCEs, these small fishes do not need to
compete with larger grazers for food.
Does the decline of herbivorous fishes with depth affect
the algal community within MCEs? In this study, herbivore
relative abundance decreased exponentially with depth
(R
2
=0.97, Fig. 8). This trend is opposite to that of turf
and coralline algae, which increase from 20 to 60 m depth
and 46 to 76 m, respectively (H Ruı
´z and DL Ballantine
pers comm). Nevertheless, coralline algae were relatively
abundant in 20 m reefs. The latter results are in agreement
with the relative-dominance model (Russ 2003) developed
for shallow reefs (20 m), which states that high grazing
leads to a reduction in turf and an increase in coralline
algae (Fig. 8). Nemeth (pers. obs.) has reported a signifi-
cant correlation between herbivore density and % cover of
crustose coralline algae on the shallow forereefs of La
Parguera. This suggests that fish grazing pressure at shal-
low areas of the shelf-edge of La Parguera is at least in part
shaping the algal community. This pattern is lost with
increasing depth as both the % cover of coralline algae and
turf increase as herbivores decline. Between 60 and 70 m,
this pattern changes again; at 70 m depth, where the min-
imum herbivore density occurs, turf abundance becomes
low but coralline algae are dominant, similar to shallow
depths where grazing pressure is high. This lack of corre-
lation of herbivore density with changes in the algal
community with depth suggests that factors other than
grazing pressure (e.g., light availability) are strongly
influencing algal communities at these deeper depths.
Coralline algae are low-light adapted species (Runcie et al.
2008) and constituted the dominant algae group between
60 and 120 m depth in Bahamas (Aponte and Ballantine
2001). Thus, grazing pressure seems to partially influence
algal distribution, but changes in algal community structure
and productivity with depth (Ballantine and Ruı
´z2011)
might also influence the distribution of herbivorous fishes
(Nemeth and Appeldoorn 2009).
Both herbivores and zooplanktivores were strongly
associated with depth, demonstrating major and opposite
trajectories in composition and abundance along the depth
gradient (Fig. 7b, c). The reduction in herbivores with
depth was only compensated by increases in zooplankti-
vores, and not by increases across the other trophic guilds.
Therefore, plankton seems to become the main source of
energy for mesophotic fishes (Kahng et al. 2010) and
replaces benthic primary production as the base of the food
chain. These results are consistent with descriptions of fish
assemblages of MCEs from other localities (Thresher and
Colin 1986; Feitoza et al. 2005; Garcı
´a-Sais 2010).
The greater extent of piscivores within MCEs, as evi-
denced in the roving surveys and the change in composi-
tion from medium-size species (e.g., L. apodus) shallow to
larger-size species deeper (e.g., L. jocu and sharks), sug-
gests that MCEs in La Parguera are relatively healthy
ecosystems that still sustain a balance of functional groups
that includes top predators. Top-down control in the fish
population exerted by apex predators acts to reduce the
dominance of a few species, allowing a more diverse
0
5
10
15
20
25
20 30 40 50 60 70
Mean abundance
Depth (m)
Fig. 8 Mean cover (%) of coralline algae (black), and turf (gray), and
relative density (%) of herbivore fish (white) per depth, along a
gradient from 20 to 70 m at six reefs in La Parguera shelf-edge,
southwest Puerto Rico. Algal and fish data from 20 m were taken
from the CRES program (see ‘Materials and methods’’ )
326 Coral Reefs (2014) 33:313–328
123
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ichthyofauna. Some apex predators (e.g., sharks) are
thought to visit shallow areas to feed, especially at night, so
their ecological role may not be limited to mesophotic
areas but also may extend into shallow areas. While this
may contribute to the higher diversity observed at shallow
reefs (Fig. 2a), multiple additional factors determine this
community parameter. For example, rugosity is strongly
correlated with both fish abundance and diversity, and the
greater small- to medium-scale habitat complexity pro-
vided by coral colonies growing in several different, three-
dimensional shapes at shallow depths is significantly
greater than that provided by the flattened morphology of
mesophotic colonies. The integrity of functional groups is a
crucial component of ecosystem stability and resilience
(Bellwood et al. 2004). Thus, these MCEs may be more
resistant to invasion by Pterois volitans (lionfish) than
MCEs in the Bahamas (Lesser and Slattery 2011) and may
potentially contribute to shallow reef resilience. Currently
(2012), lionfish are not common at mesophotic depths in La
Parguera, and understanding the role of a healthy predator
guild on the potential for lionfish colonization would be an
interesting area of future research.
Mesophotic coral ecosystems in La Parguera are subject
to a lower impact from fisheries than shallow reefs due to the
difficulty of targeting a steep narrow area with prevailing
onshore winds and currents, and because the bulk of the deep
commercial fishery targets the Mona Passage. In shallow
areas, large-size snappers and groupers have declined in
abundance as they have long been the main fishery targets in
Puerto Rico (Matos-Caraballo 2004). The presence of these
commercial species within MCEs suggests that these habi-
tats play a key refuge role and therefore are essential for the
conservation of these threatened species. Research and
monitoring of MCE fish assemblages are critical to enhance
our knowledge of species composition, accurately assess the
stocks of important commercial species, and to compare
ecological processes with shallow reefs.
Acknowledgments We thank deep divers Milton Carlo, Hector J.
Ruı
´z and Clark Sherman for assisting in field sampling, and D.
Mateos-Molina for his continuous help and support. This work was
supported by the National Oceanic and Atmospheric Administration’s
Center for Sponsored Coastal Ocean Research (NOAA/CSCOR)
(Grant No. NA06NOS4780190), through the Caribbean Coral Reef
Institute (CCRI). This project used the facilities of the University of
Puerto Rico, and the University of North Carolina—Wilmington’s
Undersea Research Center oversaw the training of rebreather divers.
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... Declines in herbivorous fish density with depth agree with previous findings (e.g. Brokovich et al. 2010;Bejarano et al. 2014), where decreases in herbivores with depth can occur due to reductions in food source and declines in coral cover (Ferreira and Goncalves 2006;Rotjan and Lewis 2006;Nemeth and Appeldoorn 2009). ...
... This may cause fish to preferentially feed at shallower depths, where reductions in surgeonfishes and parrotfishes across small depth gradients (3-15 m) have been correlated with changes in light penetration and algal productivity (Nemeth and Appeldoorn 2009). Macroalgal production, which also decreases with depth (Bejarano et al. 2014), is likely to influence the large decrease in browser biomass and density observed here, where macroalgae is the dominant food source of this group. The change in herbivore composition across depth could also be driven by ontogenetic habitat shifts, where previous work in the Caribbean found that striped parrotfish move from shallower to deeper reefs as they mature from juveniles to adults (Cocheret de la Morinière et al. 2002), supporting our findings of larger body lengths of scrapers at increased depth. ...
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Herbivorous fish are key to maintaining a balance between coral and algae on reefs, where reefs with greater herbivore biomass often show lower algal cover. For reefs worldwide, algal turf cover is expanding and is increasingly used as an indicator of disturbance. Water depth affects reef fish composition; thus, it may be expected that herbivory could also differ by depth. We examined relationships between algal turf cover and biomass (g m−2), density (# m−2) and size (cm) of herbivore groups (grazers, browsers and scrapers) across shallow (< 6 m), mid (6–18 m) and deep (18–30 m) coral reefs in the Main Hawaiian Islands. We find that across all depth classes, algal turf cover decreased with increasing grazer and scraper density, with steeper relationships observed at mid and deep reefs than in shallow reefs. In contrast, algal turf cover slightly increased with increasing grazer and browser biomass at deep reefs. Considering fish size, algal turf cover increased with larger grazer and scrapers at mid and deep reefs. The results indicate that herbivorous fish density, rather than biomass, is a better indicator of reductions in algal turf cover and resulting coral-algal balance on Hawaiian reefs, where smaller fish exert greater top-down control on cover than larger fish. Despite significant differences in herbivorous fish compositions, length-frequency distributions and fishing intensities across depth, algal turf cover remains similar across depths. Increases in fishing would have a disproportionately negative impact in deep than shallow reefs due to a lower overall fish density, where grazing functions in deep reefs are maintained by significantly fewer and smaller grazers and browsers, and larger scrapers, than in shallow reefs. Developing an understanding of patterns of algal turf herbivory by depth is important to understanding the spatial scale at which herbivory and regime shifts operate.
... En Bermudas también encontraron gran cantidad de estómagos con presas muy digeridas, infiriéndose que posiblemente la riqueza de presas del pez león podría ser mucho más amplia que la registrada (Eddy et al., 2016). Además, la menor riqueza de presas en el PNNCP puede ser consecuencia a que en ambientes mesofóticos hay menor riqueza de especies con respecto a los someros, como lo registraron en Puerto Rico (Bejarano et al., 2014), Curazao (Pinheiro et al., 2016) y una revisión global de publicaciones entre 1968 y 2010 2019; Bustos-Montes et al., 2020), which relates to the following: (a) captures are mainly made in shallow sectors, which decreases the average size and weight over time (Frazer et al., 2012;Henly, 2017); and (b) there could be ontogenetic migrations of shallow ecosystems to deep ones (Barbour et al., 2010;Biggs and Olden, 2011;Brightman-Claydon et al., 2012). ...
... In Bermuda, a big number of stomachs with very digested prey were found, inferring that possibly lionfish prey richness could be much broader than the registered one (Eddy et al., 2016). Besides, the lower richness of preys in the CPNNP may be a consequence of the lower richness of species in mesophotic environments than in shallow ones, as was recorded in Puerto Rico (Bejarano et al., 2014), Curaçao (Pinheiro et al., 2016), and a global review of publications between 1968 and 2010 (Pyle et al., 2019). In addition, lower diversity and biomass have been found in the deeper strata in Bermuda (Stefanoudis et al., 2019a). ...
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Lionfish was studied in the mesophotic environment. 237 specimens were observed, most in the upper mesophotic zone. Males were larger than females and differences between sexes in growth models were found. Sixteen prey items were identified, the most important being the teleost families Acanthuridae and Monacanthidae, and the crustacean Penaeidae. The mean of δ13C was -17.08 ± 0.36 ‰ and δ15N was 8.68 ± 0.46 ‰, with no differences between sexes. Lionfish occupies a less extensive isotopic niche in mesophotic environment than in shallow sectors; there is an isotopic niche overlap between sexes. Likewise, lionfish has specialized trophic habits. All specimens were mature and in females regression phase predominated. In females, condition factor (CF), gonadosomatic index (GSI) and hepatosomatic index (HSI) increased with gonadal development, reaching peak in the active spawning phase, and decreasing in regression. Males had a condition factor similar to spawning females, but IGS and IHS were lower
... Prior to protection, the historical fishing pressure at Middleton Reef was extremely low due to its remote location. Our work suggests that patterns in abundance and biomass of the mid-trophic level predators (e.g., spotcheek emperor, comet grouper) positively associated with depth may be more closely attributed to predator fishes in this region being naturally more abundant at mesophotic depths-potentially preying on the abundant small planktivorous species commonly aggregated near the steeper flanks of these atoll-like seamounts [77][78][79]. ...
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Predatory fishes on coral reefs continue to decline globally despite playing key roles in ecosystem functioning. Remote atolls and platform reefs provide potential refugia for predator populations, but quantitative information on their spatial distribution is required to establish accurate baselines for ongoing monitoring and conservation management. Current knowledge of predatory fish populations has been derived from targeted shallow diver-based surveys (<15 m). However, the spatial distribution and extent of predatory fishes on outer mesophotic shelf environments has remained under described. Middleton Reef is a remote, high-latitude, oceanic platform reef that is located within a no-take area in the Lord Howe Marine Park off eastern Australia. Here we used baited remote underwater stereo video to sample predatory fishes across lagoon and outer shelf habitats from depths 0–100 m, extending knowledge on use of mesophotic depths and habitats. Many predatory fish demonstrated clear depth and habitat associations over this depth range. Carcharhinid sharks and Carangid fishes were the most abundant predators sampled on Middleton Reef, with five predatory fishes accounting for over 90% of the predator fish biomass. Notably, Galapagos shark ( Carcharhinus galapagensis ) and the protected black rockcod ( Epinephelus daemelii ) dominated the predator fish assemblage. A higher richness of predator fish species was sampled on reef areas north and south of the lagoon. The more exposed southern aspect of the reef supported a different suite of predator fish across mesophotic habitats relative to the assemblage recorded in the north and lagoonal habitats, a pattern potentially driven by differences in hard coral cover. Biomass of predatory fishes in the more sheltered north habitats was twice that of other areas, predominantly driven by high abundances of Galapagos shark. This work adds to the growing body of literature highlighting the conservation value of isolated oceanic reefs and the need to ensure that lagoon, shallow and mesophotic habitats in these systems are adequately protected, as they support vulnerable ecologically and economically important predator fish assemblages.
... Those used CCR observations to assess changes in taxonomic and ecological composition in reef-fish assemblages down to 100 m depth at Bermuda (Pinheiro et al., 2016;Goodbody-Gringley et al., 2019;Stefanoudis et al., 2019), and 130 m at Curacao (Pinheiro et al., 2016). CCRs also have been used to characterize changes in species presence, diversity and ecological groupings of shallow and upper-mesophotic reef fishes at Puerto Rico (Bejarano et al., 2010(Bejarano et al., , 2014Garcia-Sais, 2010) and record depth ranges of fishes down to 130 m in the Bahamas (Pinheiro et al., 2019a) and conduct exploratory work to midmesophotic depths at San Andres Island (Chasqui et al., 2020). In the central and western Pacific CCRs are commonly used to record quantitative depth-distribution data throughout most of the mesophotic zone of reefs (e.g., Fukunaga et al., 2017;Coleman et al., 2018;Pinheiro et al., 2019b) and also to collect specimens of new species of deep-reef fishes (e.g., Pyle, 2000;Pinheiro et al., 2019b;Shepherd et al., 2020;Tea et al., 2020). ...
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Understanding the diversity and ecology of deep-reef fishes is challenging. Due to intensive and widely dispersed sampling, the Greater Caribbean (GC) fauna of species found on shallow reefs is much better characterized than the fauna of deep-reef species restricted to mesophotic (40–130 m) and rariphotic (130–300 m) depths. Our knowledge about deep-reef fishes is based on ship-board sampling and the recent use of rebreather diving, remotely operated vehicles (ROVs), baited remote underwater videos, and crewed submersibles. Submersible research on GC deep-reef fishes began in the 1960s and has flourished over the last decade through research by the Smithsonian Institution’s Deep Reef Observation Project (DROP). Here we quantify the contribution of submersible research, particularly the surge by DROP, to our understanding of the diversity of the deep-reef fish fauna of the GC. We compared shallow- and deep-reef fish faunas of three GC sites subjected to DROP research to faunas of three sites without such research. DROP increased the size of the deep faunas at three islands ∼9-fold, and they have deep-reef faunas ∼2–4 times the size of those of the other three sites. Those deep-reef faunas have high proportions of small cryptobenthic fishes, which also represent a major component of shallow faunas. That research increased the rate of discovery (collection) of new species of deep-reef fishes ∼6-fold and accounts for 31% of the deep-reef species first discovered within the GC. Substantial numbers of new species at each of the three DROP islands were not found at the other two. This indicates that other parts of the GC likely harbor many undetected deep-reef fishes, and that the size of the deep-reef fauna of the GC is significantly underestimated. These results show that small research submersibles are versatile, highly productive tools for deep-reef studies. They allow long-duration dives at any depth, while offering unparalleled views of their surroundings to study the ecology of deep-reef fishes (e.g., DROP’s definition of the rariphotic assemblage from fish depth distributions). Submersibles can efficiently collect reef fishes of a broad range of taxa, ecotypes and sizes, leading to a more comprehensive understanding of the regional GC deep-reef fish fauna.
... Hamlets are found on coral reefs at depths ranging from just a meter in sheltered areas down to at least 90 meters (Bejarano et al. 2014;Anderson et al. 2015), with all species commonly encountered in shallow habitats. Adults are occasionally seen in seagrass beds (where juveniles are common), as well as mangrove roots, wrecks, rocky reefs, and sandy rubble areas fringing reefs. ...
Article
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The hamlets (Hypoplectrus spp., Perciformes: Serranidae) constitute a distinctive model system for the study of a variety of ecological and evolutionary processes including the evolution and maintenance of simultaneous hermaphroditism and egg trading, sex allocation, sexual selection, social-trap, mimicry, dispersal, speciation, and adaptive radiation. Addressing such fundamental and complex processes requires a good knowledge of the taxonomy and natural history of the hamlets. Here, we review the taxonomy of the hamlets, from early ichthyological studies to the most recent species description in 2018. We report a total of 72 different binomial names for Hypoplectrus, synonymized or invalidated down to 17 unambiguously recognized species today. In addition, we redescribe Hypoplectrus affinis (Poey, 1861) as a valid species. In Bocas del Toro (Panama), this hamlet is distinct from eight sympatric congeners in terms of colour pattern, body size and behaviour. Whole-genome analysis and spawning observations indicate that it is genetically distinct from sympatric congeners and reproductively isolated through assortative pairing. Based on the colour pattern we detail in its redescription, live-fish photographs, videos, and earlier reports, H. affinis occurs in Panama, Nicaragua, Mexico, the Florida Keys, Cuba, Grand Cayman, Jamaica, the Dominican Republic, Los Roques (Venezuela), Bonaire, and Tobago. We conclude with a discussion of pending taxonomic issues in this group and the species status of the hamlets in general.
... MCEs have been described as transition zones between shallow and deepreef fish assemblages (Brokovich et al., 2008;Weijerman et al., 2019), and represent a frontier for research as demonstrated by the continuing discovery of new and conspicuous reef fish species (Pyle, 1996;Pyle et al., 2008;Rocha et al., 2017;Arango et al., 2019). Despite increasing interest in MCEs during the last decade (Lindfield et al., 2016;Abesamis et al., 2020), the effective management of this habitat remains problematic due to limited knowledge about the community structure and habitat associations of mesophotic fish assemblages (Dennis and Bright, 1988;Hinderstein et al., 2010;Bejarano et al., 2014;Pinheiro et al., 2016). ...
Article
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Mesophotic reefs (30–150 m) occur in the tropics and subtropics at depths beyond most scientific diving, thereby making conventional surveys challenging. Towed cameras, submersibles, and mixed-gas divers were used to survey the mesophotic reef fish assemblages and benthic substrates of the Au‘au Channel, between the Hawaiian Islands of Maui and Lāna‘i. Non-parametric multivariate analysis: Non-metric Multidimensional Scaling (NMDS), Hierarchical Cluster Analysis (HCA), Multi-Response Permutation Procedure (MRPP), and Indicator Species Analysis (ISA) were used to determine the association of mesophotic reef fish species with benthic substrates and depth. Between 53 and 115-m depths, 82 species and 10 genera of fish were observed together with 10 types of benthic substrate. Eight species of fish ( Apolemichthys arcuatus , Centropyge potteri, Chaetodon kleinii, Chromis leucura, Chromis verater, Forcipiger sp., Naso hexacanthus , and Parupeneus multifasciatus ) were positively associated with increasing depth, Leptoseris sp. coral cover, and hard-bottom cover, and one species ( Oxycheilinus bimaculatus ) of fish was positively associated with increasing Halimeda sp. algae cover. Fish assemblages associated with rubble were not significantly different from those associated with sand, Montipora coral beds and Leptoseris coral beds, but were distinct from fish assemblages associated with hard bottom. The patterns in the data suggested two depth assemblages, one “upper mesophotic” between 53 and 95 m and the other deeper, possibly part of a “lower mesophotic” assemblage between 96 and 115 m at the edge of the rariphotic and bottomfish complex.
... At the extreme, one medium sized Mycetophyllia aliciae colony at HW 50 m suffered full mortality within 6 months, while a ∼20-cm wide colony of Agaricia grahamae at HW 70 m underwent rapid and total mortality in 2 months after showing only a slow progression and partial mortality during the preceding 9 months. Invasive lionfish were not present within mesophotic depths at the time of the transect study and were still uncommon 2 years later (Bejarano et al., 2014). ...
Article
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There is limited information on the intra-annual variability of mesophotic coral ecosystems (MCEs), worldwide. The benthic communities, measured as % cover, of two geomorphologically different mesophotic sites (El Hoyo and Hole-in-the-Wall) were examined during 2009–2010 in southwest Puerto Rico. Depths sampled were 50 and 70 m. At each site/depth combination, two permanent transects, measuring 10-m long by 40-cm wide, were surveyed by successive photoquadrants, 0.24 m ² in area. Scleractinian corals, octocorals, macroalgae, crustose coralline algae (CCA), sponges and unconsolidated sediment were the main components along the transects. Significant community differences were observed both among sites and among depths. Differences among sites were greater at 50 m than at 70 m. The El Hoyo site at 50 m was the most divergent, and this was due to a lower coral and sponge cover and a higher algal cover ( Amphiroa spp., Peyssonnelia iridescens , turf) relative to the other site/depth combinations. As a consequence, the differences in community structure with depth were larger at El Hoyo than at Hole-in-the-Wall. The communities at 70 m were distinguished from those at 50 m by the greater proportion of the corals Agaricia undata, Madracis pharensis and CCA, and a reduced cover of the cyanobacterium Schizothrix . Temporal variation in the benthic assemblages was documented throughout the year. For both mesophotic sites, the magnitude of change at 50 m was significantly greater than at 70 m. For both depths, the magnitude of change at El Hoyo was significantly greater than at Hole-in-the-Wall. All assemblages experienced almost the same temporal patterns, despite the differences in species composition across sites and depths. Changes in temporal patterns are driven by an increase in the percent cover of the macroalgae Dictyota spp., and a decrease in the percent cover of non-colonized substrata (sand, pavement or rubble). Relatively rapid, intra-annual changes are dictated by the negative correlation between cyclic Dictyota spp. cover and open substrata cover. Other observed mechanisms for rapid community changes in the photoquadrants were diseases and collapses of substrata along with their associated fauna indicating that small-scale disturbance processes may play an important role within MCEs.
... However, in the current research, the water column's zooplankton community showed a greater abundance than the mesophotic community. This particular difference from the study made in Honduras, could be explained because coral reefs are highly biodiverse and most of its associated species, like fishes, corals, among others, feed on plankton (Lesser et al., 2009;Bejarano et al., 2014;Kahng et al., 2014), likely leading to a decrease in the zooplankton abundance. Besides, plankton associated with reefs had exclusive species and some species that modify their behaviour to live in these ecosystems (Heidelberg et al., 2004). ...
Article
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Mesophotic coral ecosystems shelter unique communities, but have not been studied enough due to the high cost for the available technologies. The zooplankton have become the primary food resource of the polyps at these environments, due to the low photosynthesis rate of its zooxanthellae, . Therefore, the purpose of this work was to study the zooplankton community associated to the MCEs in Bajo Frijol, in the Corales de Profundidad National Natural Park, and compare its composition with the zooplankton community from shallower parts of the water column. Three samples were taken, filtering 24 L of seawater (45 µm mesh size) at each station with a device designed to collect zooplankton right on top of the reef substrate. The taxonomic composition, density and relative abundance were obtained. A resemblance analysis was performed, complemented with a cluster, an MDS and a modified Kandoorp test. The analysis showed clear differences between the water column samples from those taken close to the reef. It also showed the separation of the community at the seamount into two large groups: north and center-south, both with exclusive species.
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Coral reef fishes often exhibit specific or restricted depth distributions, but the factors (biotic or abiotic) that influence patterns of depth use are largely unknown. Given inherent biological gradients with depth (i.e. light, nutrients, habitat, temperature), it is expected that fishes may exploit certain depths within their environment to seek out more favourable conditions. This study used baited remote underwater video (BRUV) systems to document variation in the taxonomic and functional (trophic and size) structure of a fish assemblage along a shallow to upper-mesophotic depth gradient (13–71 m) at a submerged, offshore shoal in the northern Great Barrier Reef. BRUVs were deployed during two separate time periods (February and August 2017), to separately examine patterns of depth use. Both the relative abundance and diversity of reef fishes declined with depth, and there were pronounced differences in the taxonomic and functional structure of the fish assemblage across the depth gradient. In shallow habitats (< 30 m), the fish assemblage was dominated by herbivores, detritivores, planktivores and sessile invertivores, whereas the fish assemblage in deeper habitats (> 30 m) was dominated by piscivores and mobile invertivores. Depth and habitat type were also strong predictors for important fisheries species such as coral trout (Plectropomus spp.), emperors (Lethrinus spp.) and trevallies (Carangid spp.). We found limited evidence of temporal changes in depth and habitat use by fishes (including fisheries target species), although recorded temperatures were 4 °C higher in February 2017 compared to August 2017.
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Mesophotic coral ecosystems (MCE) are located between 30 and 150 m deep and shelter unique communities. However, they have been poorly studied due to the high cost of the available sampling technologies. Zooplankton are key organisms for the maintenance of the coral colonies as their primary food source because the zooxanthellae in these habitats have a low photosynthetic rate due to the low light incidence. The purpose of this work was to characterize the zooplankton community associated with MCEs. We took 15 samples (three samples per station) by filtering 24 L of seawater (45 µm mesh size) in five stations at Bajo Fríjol with a device designed to collect zooplankton right just above the reef substrate. We determined the taxonomic composition, density, and relative abundance, and we categorized the species according to their trophic level. Crustaceans (especially nauplii and copepods), tintinnids, and foraminifers were the most abundant while tintinnids and radiolarians had the higher species richness. Herbivores dominated in both composition and abundance as usually happens in other pelagic communities. We present here the first reports of planktonic radiolarians and tintinnids from oceanic zones and MCEs in the Colombian Caribbean and the first global list of zooplankton species associated with MCEs.
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Five new species of the damselfish genus Chromis 2 (Perciformes 3 : Labroidei 4 : Pomacentridae 5 ) are described from specimens collected from deep (>60 m) coral-reef habitat in the western Pacific by divers using mixed-gas closed-circuit rebreather gear. Two of the five new species (C. abyssus and C. circumaurea) are each described from specimens taken at a single locality within the Caroline Islands (Palau and Yap, respectively); one (C. degruyi) is described from specimens collected or observed throughout the Caroline Islands, and two (C. brevirostris and C. earina) are described from specimens collected from several localities throughout the Caroline Islands, Fiji, and Vanuatu. All five species can easily be distinguished from other known Chromis, and from each other, on the basis of color and morphology. These new species represent the first five scientific names prospectively registered in the official ICZN ZooBank registry 6 . Moreover, the electronic online edition of this document has been specially formatted with many embedded links to additional resources available online via the internet to enhance access to taxonomically-relevant information, and as a demonstration of the utility of international standards for biodiversity informatics.
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Recent collections of the benthic macro- and meiofauna associated with the mesophotic coral ecosystems of Puerto Rico have revealed two new Cumacea from the family Nannastacidae. A new genus Cumellana and two new species, Cumellana caribbica and Cumella alexandrinae are described herein. The new genus Cumellana can be distinguished from the other genera of the family Nannastacidae by having a long antennule and pereopod 2 with short terminal setae, equal in length.
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Examining the relationship between habitat characteristics and utilization patterns by herbivorous fishes on coral reefs will add to our understanding of the factors that influence the abundance and distribution of this important group. The abundances of parrotfishes (Scaridae) and surgeonfishes (Acanthuridae) on fore-reef habitats were sampled along an inshore-offshore gradient to provide for within reef and crossshelf comparisons in relation to the environmental parameters of depth and topographic relief. Temporally replicated visual surveys were conducted along permanent belt transects (100m2) at three depth intervals (3, 10, 15 m) to obtain data on fish species density and lengths, which were used to calculate biomass. The roving herbivorous fish assemblage was dominated by three species of parrotfishes (Scarus iseri, Sparisoma aurofrenatum and S. viride) and three surgeonfishes (Acanthurus bahianus, A. chirurgus and A. coerulus). Overall the biomass of both families was highest at 3m compared to 10 or 15m (p<0.05). However, the relative decrease in biomass across depths for both families was greatest at inshore reefs where water transparency is lowest. The mean biomass for both families differed between inner and mid-shelf reefs at 10 and 15 m (p<0.05) but not at 3m. Fish biomass was correlated to reef topographic relief at 3m for parrotfishes (p<0.05) and at all three depth intervals for surgeonfishes (p<0.05). Overall patterns of herbivore biomass across the shelf reflect differences in light penetration, suggesting that fish may be responding to algal productivity. Thus, within fore-reef habitats along a cross-shelf gradient water transparency and topographic relief may interact to structure biomass patterns.
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The heart of the PC-ORD system is a group of Fortran programs for multivariate analysis on the MS-DOS family of microcomputers. Analytical features include ordinations (detrended correspondence analysis, Bray-Curtis ordination, principal components analysis, and reciprocal averaging), descriptive statistics, and diversity indices. -from Author
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Describes the distribution of Algae, Corals and Gorgonians in relation to depth, light attenuation, water movement and grazing pressure in the fringing coral reef of Curaçao.
Article
Tropical marine communities from shallow-water (<30 m) carbonate environments are often dominated by hermatypic scleractinian corals with lesser amounts of crustose coralline algae and endolithic demosponges. Living cover is typically high (80-100%). Along the north-central coast of Jamaica and at many western Atlantic sites, communities existing below 55 m inhabit a vertical to overhanging wall of reef limestone, the deep fore reef, which extends to approximately 130 m. At 60 m the community resembles that of shallower water, although sclerac-tinians are less abundant and encrusting and erect demosponges are much more abundant. Coralline algae and macroalgae are also important space occupants at 60 m and living cover approaches 65%. Encrusting sponges and coralline, filamentous, and macroalgae predominate in the middle region of the deep fore reef. A low-diversity assemblage occupying 40% of the substratum and dominated by diminutive encrusting and endolithic? demosponges and largely endolithic filamentous algae occurs from 100-130 m, the lower limit of the deep fore reef. Community structure and zonation on the shallower reefs is controlled by a number of biotic and abiotic factors, most notably predationlgrazing, light intensity, and turbulence. On the deep fore reef, grazing and turbulence are greatly reduced. While reduced in intensity, light continues to exert a strong influence on community bathymetric zonation. Sedimentation also exerts an important control on the spatial distribution of the deep fore-reef biota with the most diverse assemblages flourishing in areas protected from sediment. Despite a regime of reduced disturbance in deep water, community diversity remains relatively constant to a depth of 90-100 m. Copyright © 1988, The Society of Economic Paleontologists and Mineralogists.