Deep-Sea Research I 48 (2001) 2185–2194
Depth distribution of algal species on the deep insular fore reef
at Lee Stocking Island, Bahamas
Nilda E. Aponte, David L. Ballantine*
. 00681, Puerto Rico
Department of Marine Sciences, University of Puerto Rico, PO Box 9013, Mayaguez
Received 13 June 2000; received in revised form 8 December 2000; accepted 16 February 2001
Abstract
Deep-water benthic algal composition and cover were studied with a submersible on the deep fore reef of
Lee Stocking Island, Bahamas, from 45 to 150 m. Algal cover decreased from 57% to 16% over this depth
range. Although there was substantial overlap in depth distributions, different species or groups of species
dominated benthic cover at different depths. Lobophora and Halimeda copiosa co-dominated the fore reef
from 45 to 60 m. A Corallinales/Peyssonnelia group was abundant from 60 to 120 m. The Corallinales/
Peyssonnelia group shared dominance with Ostreobium between 90 and 120 m. Ostreobium was the only
alga observed below 150 m and remained abundant below 200 m. Movement of sand down the fore reef is
recognized as having substantial influence on algal cover. # 2001 Elsevier Science Ltd. All rights reserved.
Keywords: Atlantic; Bahamas; Coral reef; Deep-water algae; Submersible
1. Introduction
Submersibles have been utilized for the last two decades to collect and examine algal
populations in water deeper than can be accessed with SCUBA. In the tropical western Atlantic
region, this has resulted in the characterization of species new to science (including Eiseman and
Moe, 1981; Eiseman and Norris, 1981; Eiseman and Earle, 1983; Ballantine and Wynne, 1988;
Norris and Olsen, 1991; Bucher and Norris, 1992; Ballantine and Norris, 1994; Ballantine and
Aponte, 1996) and new algal depth and range extensions (Eiseman and Blair, 1982; Hanisak et al.,
1987; Hanisak and Blair, 1988; Blair and Norris, 1988; Norris and Olsen, 1991; Wynne and
Schneider, 1996). Submersible work in the region also led to discovery of the deepest known
*Corresponding author. Tel.: +787-899-5783; fax: +1-787-899-2630.
E-mail address: d [email protected] (D.L. Ballantine).
0967-0637/01/$ - see front matter # 2001 Elsevier Science Ltd. All rights reserved.
PII: S 0 9 6 7 - 0 6 3 7 ( 0 1 ) 0 0 0 1 1 - 5
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habitat for an attached algal species (Littler et al., 1985). The latter was a crustose coralline
rhodophyte collected at a depth of 268 m off the Island of San Salvador, Bahamas.
Algae are well known to be important components of coral reef communities. They provide (i)
structural framework (Adey, 1976, 1978; Wray, 1977); (ii) physical consolidation (Borowitzka
et al., 1974); (iii) high species diversity (Van Den Hoek et al., 1975; Wanders, 1976; Conner and
Adey, 1977; Littler and Littler, 1994); (iv) food (Borowitzka, 1981; Carpenter, 1981; Hay, 1984);
(v) contribution to reef sediment (Wray, 1977; Bach, 1979); (vi) primary productivity (Vooren,
1981; Hawkins and Lewis, 1982; Morrissey, 1985); and (vii) nitrogen-fixation (Wiebe et al., 1975;
Mague and Holm-Hansen, 1975). Although substantially less information is available on deepwater macroalgae, their productivity, abundance and potential importance have been reviewed by
Jensen et al. (1985) and Littler et al. (1985, 1986).
Littler et al. (1986) pointed out that much of the literature resulting from submersible research
on coral reefs includes only incidental observations of algal presence. To date there has been only
a single study specifically concerned with algal distribution in deep fore reef habitats. That study,
by Littler et al. (1986), was conducted from 81 to 268 m on a seamount off San Salvador Island,
Bahamas. The present paper provides a quantitative description of the distribution and species
composition of the macroscopically visible benthic marine algae across the entire depth range of
deep fore reef habitats, from 45 to 150 m, at Lee Stocking Island, Exuma Cays, Bahamas.
Fig. 1. Location of submersible transects at Lee Stocking Island, Bahamas.
N.E. Aponte, D.L. Ballantine / Deep-Sea Research I 48 (2001) 2185–2194
2187
2. Study area
Lee Stocking Island is part of the Exuma Cays in the southeast of the Bahamas Archipelago,
located approximately 450 km southeast of Miami, Florida. The study area (Fig. 1), is located at
the insular shelf break, east of Lee Stocking Island (Transect AA: 238470 4600 N and 768050 8500 W;
Transect BA: 238460 7800 N and 768050 0000 W). The fringing reef system that surrounds the island
gradually slopes to a depth of approximately 30 m, where the slope precipitously increases. Below
approximately 40 m the reef slope ranges from roughly 458 to nearly vertical, with occasional
terraces, steps and overhangs.
3. Materials and methods
Observations and sampling utilized the two-manned submersible Nekton-Gamma leased by the
Caribbean Marine Research Center (CMRC) of the NOAA National Underwater Research
Program (NURP). The submersible was towed to site and launched from the R/V Bahama
Hunter. The DSV Nekton-Gamma, with operating depths to 300 m, was equipped with an
hydraulically manipulated mechanical arm, which allowed sampling of small to medium-sized
dislodged pieces of substratum (3–10 kg). Collected samples were brought to the surface in an
externally mounted bag or basket, or were held in the manipulator claw for the duration of the
dive. The submersible was also equipped with internal still (35 mm) and video cameras plus an
external light system and strobe.
Submersible dives were made on two previously established transects (AA, BA) on the insular
shelf edge (Fig. 1). The transects were oriented perpendicularly to the shelf and extended from
depths of 30–300 m. Results of quantitative sampling are limited to the latter transect, although
data from both transects was used for depth distributions of algal species. Transect BA was
sampled three times between September 1994 and May 1995, at intervals of four months. Species
distribution is based on collection, visual observations, annotated video transects and quantitative
photo sampling (below).
Quantitative sampling (photographic) was conducted at 15 m intervals between 45 and 150 m.
At each depth increment the submersible worked perpendicular to transect line BA and
approximately 1.0 m from the substratum. Sample scale and right angle orientation were provided
by a 0.75 m long steel rod calibrated at 2 cm intervals. The rod was held by the hydraulic arm and
positioned just over the photographed surfaces. Eighteen color transparencies were taken at each
sampling depth.
Quantification utilized the ‘photo quadrat’ method of Littler and Littler (1985). Twelve
transparencies representative of each depth increment were randomly selected. The slides were
projected on a screen that contained a fixed array of 150 randomly marked points in a delimited
rectangular area. Identity of the algal component or substratum type at each point was recorded.
Non-algal organisms were lumped in a single category, and small filamentous algal-forms were
collectively designated as turf. It was not always possible to differentiate between Peyssonnelia and
coralline red algal crusts in the photographs, hence they are combined here as a single functional
form group. Percent cover values were expressed as the number of points touching each species,
divided by the total of points in the quadrat. For each depth, 2–4 randomly selected
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transparencies were scored twice or by different personnel as well as at different times to assure
reproducibility. Reproducibility was generally within 5–7%. Significance of difference between
means of percentage cover between adjacent depths was determined by one-way analysis of
variance following arcsine transformation of the square root of cover. Statistical difference was
P40:05. Algal species authority designations are according to Brummitt and Powell (1992).
4. Results
The depth distribution of macroalgae and the sand distribution at transect BA is summarized in
Figs. 2 and 3. There was substantial variability between species composition and algal percent
cover within quadrats sampled at the same depth over all of the depths sampled. The high
coefficients of variation (CV=Standard Deviation/Mean), frequently over 1.0 for algal species
cover, is indicative of patchy distributions (Table 1). Nevertheless, total algal cover was highest at
the two shallowest quantified depths, 50% at 45 m and 57% at 60 m. The dominant species in
terms of cover at 45 m was Lobophora variegata (J.V.Lamour.) Womersley, 23%; followed by
Halimeda spp., 10%; and turf, 13%. Corallinales/Peyssonnelia had a cover of 3% at this depth
(Corallinales are non-articulated crustose calcified Rhodophyta species). At 60 m, Lobophora and
Halimeda spp. had essentially equal covers (15% and 14%). While several Halimeda species
(including H. discoidea Decne. and H. incrassata (J.Ellis) J.V.Lamour.) were lumped together in
Fig. 2. Depth distributions of commonly identified algal species on the deep fore reef of Lee Stocking Island.
N.E. Aponte, D.L. Ballantine / Deep-Sea Research I 48 (2001) 2185–2194
2189
Fig. 3. Percent cover as a function of depth of common deep-water algae on the fore reef of Lee Stocking Island. Note
that scale for percent cover of algal species and algal groups differs from scale for sand cover. Stars between adjacent
depths indicate significant difference in percent cover (P50:05).
the scoring, H. copiosa Goreau and E.A.Graham was by far the most abundant species. Within
the Halimeda depth range, it was common to see areas in which H. copiosa cascaded over vertical
wall faces, with localized cover approaching 100%. Corallinales/Peyssonnelia accounted for 14%
and turf 13% cover at 60 m. From 45 m to a depth of approximately 70 m, other regularly present
macroscopic algae included the chlorophytes Udotea sp., Rhipocephalus phoenix (J.Ellis and Sol.)
.
Kutz.,
Avrainvillea spp., and Penicillus dumetosus (J.V.Lamour.) Blainv.
Lobophora variegata (ochrophyta) was a dominant algal element from 30 to 45 m and was
conspicuous to 70 m (Fig. 4). At its maximum depth limit, the Lobophora zone ended abruptly
with no evident transitional zone. Lobophora was observed chiefly on steps or terraces and upper
parts of overhangs, more rarely on vertical walls. At localized areas, cover of L. variegata
approached 90%. The zone of Lobophora dominated substratum was also rich in turf algae and
corallines, the latter forming an under-story, not quantifiable with the methods utilized. The
functional form group, turf algae, included numerous small, filamentous red and green forms that
could not be accurately identified to specific taxonomic category from examination of the
transparencies. The turf assemblage is, nevertheless, an important component of the algal
composition to a depth of 75 m, with average abundance ranging between 13% and 15%.
Although the composition varied with specific site, Microdictyon marinum (Bory) Silva,
Cladophora sp. and Dictyota spp. were among the more common members.
The deep-water genus Verdigellas, represented chiefly by V. peltata D.L.Ballant. and J.N.
Norris (Chlorophyta), was never abundant but was highly conspicuous because of its bright green
coloration (Fig. 5). It was present at depths between 60 and 120 m, and its cover rarely exceeded
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Table 1
Mean cover (Av.) and coefficients of variation (CV) of dominant algal groups plus sand at the deep fore reef at Lee
Stocking Island, Bahamas
Depth (m)
45
60
Lobophora
Av.
CV
23.3
0.75
14.9
1.36
0.5
1.58
Halimeda spp.
Av.
CV
10.4
0.55
14.0
0.80
6.1
1.04
0.0
Turf
Av.
CV
13.4
1.02
12.8
0.67
14.5
0.70
0.9
1.91
0.3
1.90
0.0
3.2
1.43
13.5
0.80
12.7
1.07
17.2
0.61
16.7
0.49
6.8
0.76
2.3
1.17
1.4
1.44
Ostreobium
Av.
CV
0.2
2.15
2.5
1.54
14.3
0.52
14.5
0.52
12.6
0.68
13.3
0.56
14.1
0.51
Verdigellas
Av.
CV
1.7
1.04
2.1
1.17
1.3
1.65
0.5
3.2
0.1
2.30
0.0
28.3
0.69
36.15
0.76
56.6
0.28
62.1
0.25
77.2
0.17
79.6
0.14
Corallinales/Peyssonnelia
Av.
CV
Sand
Av.
SD
31.1
0.69
75
90
105
120
135
Fig. 4. Fore reef at Lee Stocking Island, 60 m depth; benthic cover is dominated by Lobophora variegata.
150
80.5
0.12
N.E. Aponte, D.L. Ballantine / Deep-Sea Research I 48 (2001) 2185–2194
2191
2% although it averaged 3.6% during a single sampling period at 75 m. In general, the percent
coverage by Verdigellas did not vary significantly throughout its depth range. Verdigellas shows
its maximum abundance at depths of 60–75 m.
Total algal cover fell to 38% at 75 m and to 34% at 90 m. Crustose Corallinales/Peyssonnelia
provided the dominant algal cover, constituting 13% and 17% at these depths, respectively. Turf
algae made up 15% at 75 m although this component fell to less than 1% at 90 m. Ostreobium
quekettii Bornet and Flahault, a filamentous endolithic chlorophyte that is recognizable as a light
green cast on exposed rock surfaces was first observed to be abundant (14%) at 90 m. Ostreobium
may have been obscured by the greater cover of algae and corals in shallow water and may be
more abundant at shallower depths than indicated here. Halimeda spp. were not observed deeper
than 90 m
At 105 m, total algal cover was 32% with Corallinales/Peyssonnelia (Fig. 5) representing 17%
and Ostreobium making up 15%. At depths from 120 to 150 m, the total algal coverage was 15–
20%, being almost entirely represented by Ostreobium growing on the lower faces of the sand
covered rock ledges and on non-occupied exposed surfaces (Fig. 6). Ostreobium was present
Fig. 5. Fore reef at Lee Stocking Island, 105 m depth; benthic cover is dominated by Corallinales (asterisk) and
Peyssonnelia sp. (arrow). Verdigellas peltata is also present (arrowhead).
Fig. 6. Fore reef at Lee Stocking Island, 200 m depth; bottom is mostly covered by sand; however, Ostreobium is
apparent on vertical rock face (arrow).
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throughout the range of depths studied and was observed at 200 m. It was obvious, however, only
at depths below 75 m. Its percent cover ranged from 13% to 15% at depths between 90 and 150 m.
5. Discussion
In general, a decline in algal percent cover (from 57% to 16%) was observed through the depth
range studied. Sand cover along the shelf fore reef increased with depth, from 31% cover at 45 m
to 81% cover at 150 m. A continual and substantial transport of sand down the fore reef is a
normal feature of the habitat. The sand movement and sediment accumulation on the fore reef
undoubtedly have a large effect on the living cover of all benthic species due to periodic burying.
The dynamics of the movement and the degree of burying and unburying of benthic algae are
unknown; however, it is felt that burying substantially influences algal cover at any portion of the
fore reef. At depths below 90 m the effect of burying was evident particularly on low angled ledges
and overhangs. The coralline algae and the endolithic algae found at those depths were confined
mostly to the vertical walls of the lower faces of the rims. Steneck (1997) studied the effects of
sediment burial on a coralline alga in the same vicinity and demonstrated survival after at least
100 days. Dill (1991) has commented on movement and downwelling of ooid-sand sediments in
the vicinity of Lee Stocking Island. Downwelling was evident by the abundance of seagrass leaves
and Lobophora thalli, which were commonly observed drifting loosely at depths in excess of
150 m. Liddell and Ohlhorst (1988) indicated that sedimentation was an important controlling
factor on deep Jamaican fore-reef biota. Where substrata are low-angled, biota are subjected to
sediment burial.
Littler et al. (1986) recognized four algal zonal assemblages between 81 and 268 m at San
Salvador, Bahamas. These assemblages were: (1) Lobophora (81–90 m); (2) Halimeda (90–130 m);
(3) Peyssonnelia (130–189 m); and (4) crustose corallines (189–268 m). Our results share some
similarity to these overall patterns, but there are exceptions. At Lee Stocking Island, the zones are
seen as overlapping entities. Lobophora and Halimeda copiosa co-dominated the fore-reef from 45
to 60 m, with Lobophora being more abundant at the shallower end of the range and Halimeda
being more abundant at the deeper, remaining in abundance to 75 m. At a depth of 60 m, the
Corallinales/Peyssonnelia group was also abundant and remained abundant to a depth of 120 m.
The Corallinales/Peyssonnelia group shared dominance with Ostreobium between 90 and 120 m.
Ostreobium was the only alga observed below 150 m and remained abundant below 200 m. All of
the depth distribution limits of recognizable algal groups at Lee Stocking Island were shallower
than their San Salvador counterparts. This may be a function of relatively greater turbidity due to
sediment spilling over the reef and also sediment cover at Lee Stocking as opposed to the
presumably less terrestrially influenced seamount at San Salvador.
Aside from recognition that a coralline alga is the deepest occurring species (Littler et al., 1985),
the importance of Corallinales and Peyssonneliaceae as components of deep benthic fore reefs has
been indicated in other studies. Agegian and Abbott (1985) also reported abundant crustose
coralline red algae on deep fore reefs of south Pacific atolls. Dullo et al. (1990, p. 265), based on
submersible research of deep fore reef environments in the Red Sea, indicated that ‘‘corallinaceans
and peyssonneliaceans represented the dominant biomass of the foralgal crusts’’ (to 110 m).
N.E. Aponte, D.L. Ballantine / Deep-Sea Research I 48 (2001) 2185–2194
2193
Liddell and Ohlhorst (1988) reported a maximum cover of coralline algae (presumably including
Peyssonneliaceae) of 14–19% at 60–90 m in Jamaica.
Liddell and Ohlhorst (1988) considered factors responsible for changes with depth in reef biota.
They attributed overall distribution to a combination of factors, including: (i) decrease in light
irradiance and change in spectral qualities with depth, (ii) decrease in biotic and physical
disturbance with depth, and (iii) increase in burial by sediment with depth. We would agree that
these factors apply to the present study.
Acknowledgements
This study was partially funded by the Caribbean Marine Research Center of the National
Oceanographic and Atmospheric Administration-National Undersea Research Program, which
made available ship time aboard the R/V Bahama Hunter and dives with the research submersible
DSV Nekton Gamma. We wish to express our appreciation to the submersible and tender crew,
particularly Mr. Robert Wicklund Jr., Ray Brockway and Don and Sally Tondro, as well as to the
administrative staff of Lee Stocking Island. Ms. Bonnie Bower Dennis assisted in drafting the
figures. Dr. Paul Yoshioka provided valuable discussion which improved the manuscript.
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