L. Coston-Clements, L.R. Settle, D. E. Hoss and F.A. Cross
|U.S. DEPARTMENT OF COMMERCE Robert A. Mosbacher,
NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION John A. Knauss, Administrator
NATIONAL MARINE FISHERIES SERVICE William W. Fox, Jr. Assistant Administrator for Fisheries
The National Marine Fisheries Service (NMFS) does not approve,
recommend or endorse any
proprietary product or proprietary material in this publication. No
reference shall be made to
NMFS, nor to this publication furnished by NMFS, in any advertising or
sales promotion which
would indicate or imply that NMFS approves, recommends or endorses any
or proprietary material mentioned herein, or indirectly, the advertised
product to be used or
purchased hecause of this NMFS publication.
This report should be cited as follows:
Coston-Clements, L., L.R. Settle, D.E. Hoss and F.A. Cross. 1991.
Utilization of the Sargassum
habitat by marine invertebrates and vertebrates - a review. NOAA
NMFS-SEFSC-296, 32 p.
Copies may be obtained by writing:
National Technical Information Service 5285 Port Royal Road
Springfield, VA 22161
National Marine Fisheries Service, NOAA
Southeast Fisheries Science Center
101 Pivers Island Road
Beautort, NC 28516-9722
Cover illustration from Teal and Teal, 1975.
Numerous species of brown algae (Class Cyclosporeae: Order Fucales:
Family Fucaceae) of the
genus Sargassum occur throughout the world's tropical and temperate
oceans. The pelagic
complex in the western North Atlantic is comprised primarily of Sargassum
natans and S.
fluitans. Both species are hyponeustonic and fully adapted to a pelagic
existence (Parr, 1939).
Known commonly as gulf-weed, sea holly, or sargassum, they are
characterized by a brushy,
highly branched thallus (stem) with numerous leaf-like blades and
(floats). These floating plants may be up to several meters in length but
are typically much
smaller. See Hoyt (1918), Winge (1923), Parr (1939), Taylor (1960),
Prescott (1968), and Humm
(1979) for detailed descriptions of the various species.
Sargassum floats contain mostly oxygen with some nitrogen and carbon
dioxide, and are
responsible for buoyancy. Oxygen content is dependent on the oxygen
partial pressure of the
surrounding medium and independent of photosynthetic activity (Hurka,
(1950) also found diurnal fluctuations in gas pressure within the floats,
and attributed it to similar
changes in oxygen partial pressure during daylight and darkness.
There is some debate as to whether sargassum found in the Gulf Stream,
Gulf of Mexico,
Caribbean Sea, and the North Atlantic Central Gyre is material detached
from littoral plants in
the Antilles and other tropical regions during storms (Peres, 1982) or
whether the oceanic
components are a separate species group independent of their coastal
littoral relatives. Most
consider the pelagic forms a separate and distinct species group
having evolved from benthic
species (Winge, 1923; Parr, 1939; Friedrich, 1969; Butler et al., 1983;
Stoner and Greening,
1984). Supporting evidence includes the lack of sexual reproduction
characteristic of benthic
forms, the loss of a basal holdfast, and the number of endemic organisms
in the associated
community (10 invertebrates and 2 vertebrates). Benthic forms, such as S.
filipendula, occur in
the open ocean in small quantities, but should be considered flotsam
(Hoyt, 1918; Winge, 1923;
Parr, 1939; Dooley, 1972).
Sargassum circulates between 20 and 40 N latituded and 30 W longitude
and the western edge of the Florida Current/Gulf Stream, with an apparent
center of distribution within the North Atlantic Central Gyre between 28
and 34 N latitude (Fig. 1) (Winge 1923; Ryther, 1956; Dooley, 1972;
Butler et al. 1983). Large quantities also occur on
the continental shelf. Some of this material is cast upon beaches
along the eastern seaboard (Hoyt, 1918; Humm, 1979; Winston, 1982), while
much of it remains
on the shelf or is entrained into the Gulf Stream. Throughout the region,
aggregates into large windrows in response to wind forcing (Winge, 1923;
Faller and Woodcock, 1964) or shear currents along frontal boundaries
Woodcock (1950) demonstrated that sargassum can be downwelled along such
zones; the depth of descent being dependent on the buoyancy of individual
algal clumps and the
magnitude of the Langnuir circulation cell. Clumps that do not sink below a
critical depth (» 100 m) can withstand the hydrostatic pressure and
will ultimately rise to the
surface. There is also, however, a time-at-depth relationship that can
influence the true critical
depth at which buoyancy is lost and the algae sink to the bottom (Johnson
1977). Peres (1982) questioned the return to the surface after being
downwelled. When buoyancy
is lost, sargassum sinks slowly to the bottom (about 2 days to reach 5000
m) and provides a
resource for bottom dwelling consumers (Schoener and Rowe, 1970).
The contribution of sargassum to total primary production in the
western North Atlantic is variable and dependent on the region examined
and on accumulated biomass. Carpenter and Cox (1974) estimated average production across the western
Sargasso Sea at =1.0 mg C m-2d-1, with higher values frequently occuring at
localities on the continental shelf and in the northern Sargasso Sea (above
30 degrees N). Howard and Menzies (1969) found production highest in the
Stream, lowest on the shelf, and the intermediate in the Sargasso Sea. These authors estimate sargassum
probably contributes about
0.5% of the total primary production in the region, but nearly 60% of the
total in the upper 1 m
of the water column (Howard and Menzies, 1969; Carpenter and Cox, 1974;
Peres, 1982). In addition, epiphytic cyanobacteria (Dichothrix and
significantly to overall production and nitrogen fixation within the
community (Carpenter, 1972; Carpenter and Cox, 1974; Phlips and Zeman,
sargassum and its associated blue-green algae
epiphytes are adapted to conditions of strong sunlight, that is,
photosynthesis is not inhibited at
high light intensities (Phlips, et al., 1986).
A survey conducted in the Sargasso Sea from 1933 to 1935 (Parr, 1939)
biomass at 524 to 1642 mg m-2. Stoner's (1983) estimates in the Sargasso
Sea from the years
1977 to 1981 were less than 6% of those earlier estimates. He suggested
that the decline may
have been anthropogenically influenced. Subsequent analysis of Stoner's
concluded that there had been no significant change in the biomass of
sargassum from 1933 to
1981, except in an area northeast of the Antilles. This apparent decline
was attributed to seasonal
variation in sargassum abundance or long-term current shifts, and
apparently not to pollution
(Butler et al. 1983; Butler and Stoner, 1984). Reliable estimates of
total biomass are not
available, however, as no directed statistical study encompassing the
vast range of these species
has been conducted.
Pelagic sargassum supports a diverse community of marine organisms including micro- and macro-epiphytes (Carpenter, 1970; Carpenter and Cox, 1974; Mogelberg et al., 1983), fungi (Winge, 1923; Kohlmeyer, 1971), more than 100 species of invertebrates (Table 1), over 100 species of fishes (Table 2), and four species of sea turtles (Carr, 1987a; Manzella and Williams, 1991). Some inhabitants, unique to the sargassum habitat, have evolved unusual
shapes and coloration affording them the additional advantage of
camouflage among the floating
plants. Others are less specialized and utilize the habitat for foraging
or protection from
predators. Community structure is variable; influenced by season,
geographic location, and algal
"age" (Weis, 1968; Fine, 1970; Butler et al. 1983; Stoner and Greening,
1984). Weis (1968) also
noted differences in epibiont diversity between species of sargassum. An
important factor in the
structure of the community is related to compounds occurring in the
exudate released by the
algae during growth. Tannins produced on the distal growing tips of
sargassum have an
inhibitory effect on colonizing epibionts (Conover and Sieburth, 1964;
Sieburth and Conover,
1965). This antifouling effect lasts a short time and a succession of
bacteria, hydroids, bryozoans,
and blue-green algae rapidly follow (Winge, 1923; Conover and Sieburth,
1964; Ryland, 1974).
Carpenter and Cox (1974) also suggest that low epibiont density within
some areas of the
Sargasso Sea may be nutrient limited rather than limited by the
antibiotic activity of sargassum
exudates. Natural chemical compounds, including phenolic compounds,
produced by algae may
also serve as a deterrent to herbivores (Paul, 1987; Hay and Fenical,
1988; Hay et al., 1988;
For details of community metabolism, respiration, trophic web and
chemistry, we refer the reader
to the works of Culliney (1970), Burns and Teal (1973), Smith et al.
(1973), Johnson and
Braman (1975), Blake and Johnson (1976), Hanson, (1977), Geiselman
(1983), Morris et al.
(1976), and Trapnell et al. (1983).
The invertebrate fauna consists of both sessile and motile forms
(Table 1). Epizoans include
colonial hydroids, encrusting bryozoans, the polychaete Spirorbis,
barnacles, pycnogonids, and
the tunicate Diplosoma. Older plants gradually become heavily encrusted
with these organisms
and ultimately sink. This biomass then gradually disintegrates, providing
for animals in deeper water (Parr, 1939; Weis, 1968; Schoener and Rowe,
1970; Butler et al.
1983). Conspicuous among the motile fauna are decapod crustaceans,
particularly the Portunus
crabs, and shrimps Latreutes and Leander, various molluscs, including the
Litiopa melanostoma, polychaetes, flatworms, and nudibranchs. Fine (1970)
found very high
numbers of portunids in his late summer samples in the Gulf Stream and
Sargasso Sea. Only
Portunus sayi is commonly considered a resident of the community; the
remaining megalopa and
juveniles are transitory and utilize the habitat as a nursery.
Dooley (1972) examined stomach contents of the eight most abundant
fish species yielding
further insight into the invertebrate component of the sargassum
community. These included
hydroids, copepods, phylosoma larvae, shrimp zoea and postlarvae, crabs,
barnacles, tunicates, polychaetes, bivalves, gastropods, and
platyhelminthes. The presence of two
rather enigmatic members of the sargassum fauna were revealed by Morgan
et al. (1985) through stomach content analysis of several large
epipelagic predatory fishes. They
found the large mysis of the penaeoids, Cerataspis monstrosa and
co-occurred with sargassum in the stomachs of surface feeding tuna
(Scombridae) and dolphin
Coryphaena hippurus. Nothing is known about the adult stage or life
history of these rare
Vertebrates - Fishes
There is a well known assemblage of small fishes associated with
sargassum rafts, many of
which serve as forage for commercially or recreationally exploited
species (Table 2). Dooley
(1972) described 54 species from 23 families in the sargassum community
of the Florida Current,
while Bortone et al. (1977) reported 40 species from 15 families in the
eastern Gulf of Mexico.
Only 14 species from 11 families are known from the Sargasso Sea
(Fedoryako, 1980; 1989).
Young jacks (Carangidae) live among the protective branches of sargassum and feed heavily on copepods and larval decapods. Apparently sargassum carries along a resident plankton population capable of sustaining these voracious predators (Yeatman, 1962). Sub-adult jacks range further from the rafts but dart in to feed on shrimp and young fishes living in the sargassum. The filefishes and triggerfishes (Balistidae) are also abundant and feed primarily on hydroids, encrusting bryozoans, and other invertebrates. Another major predator is the voracious sargassumfish, Histrio histrio, which selectively preys upon shrimp and young fish (Adams, 1960).
Large predatory species associated with the sargassum habitat include
(Coryphaenidae), barracudas (Sphyraenidae), mackerels and tunas
(Xiphiidae), and billfishes (Istiophoridae) (Gibbs and Collette, 1959;
Stephens, 1965; Dooley,
1972; Fedoryako, 1980; Carr, 1986; C. Manooch, pers. comm.). It is
believed that dolphin, a
much sought after game and food fish, takes shelter under flatoam
(including sargassum) because
of the enhanced availability of prey (Dooley, 1972). Filefish,
triggerfish, jacks, flyingfish
(Exocoetidae), and puffers (Tetradontidae) are among the species
identified in dolphin stomachs
(Gibbs and Collette, 1959; Dooley, 1972; Manooch et al., 1984). Fragments
of sargassum were
also commonly found. Manooch et al. (1984) stated "The close association
dolphin with fish and invertebrates that form the sargassum community is
Manooch and Mason (1983) also reported finding sargassum fragments in 26%
tuna, Thunnus albacaras, stomachs they examined as well as in 12% of
blackfin tuna, T.
atlanticus. They believed the material was ingested incidentally with
There is less known about the ichthyoplankton associated with the habitat, but it seems likely that the same hydrodynamic mechanisms that drive the formation of sargassum rafts, i.e., convergence of surface water within shear zones or Langmuir cells, will also aggregate surface oriented organisms (Kingsford, 1990). There is some evidence that this is the case for swordfish, Xiphias
gladius, larvae and cobia, Rachycentron canadum, eggs near the Gulf
Stream frontal zone (Hassler and Rainville, 1975; J.J. Govoni, pers.
comm.). In addition to
feeding and shelter, adults of some oceanic pelagic fishes use sargassum
as a spawning substrate
(Dooley, 1972: Peres, 1982) or as a nursery area for larvae and
juveniles. Most notable among
these are the flyingfishes (Winge, 1923; Breder, 1938) which are a major
component of the diet
of large oceanic fishes.
During the pelagic stage, hatchling loggerhead, Caretta caretta,
green, Chelonia mydas,
Kemp's ridley, Lepidochelys kempi, and hawksbill, Eretmochelys imbricata,
sea turtles have
been observed in sargassum off Florida, Georgia, North Carolina, and
Texas (Smith, 1968;
Fletemeyer, 1978; Carr and Meylan, 1980; Carr, 1986; 1987a; Schwartz,
1988; 1989; Manzella
and Williams, 1991; Schwartz, pers. comm.). Hundreds of loggerhead
hatchlings, both dead and
alive, were found in the wrack of sargassum deposited on the shore at
Cocoa Beach, Florida
following a hurricane in September, 1979 (Carr and Meylan, 1980). Stomach
contents of the
dead hatchlings showed that almost all contained sargassum floats and
leafy parts. Schwartz
(1988) reported numerous loggerhead hatchlings captured during commercial
sargassum. This observation constitutes the largest known aggregation of
encountered off the North Carolina coast.
Hatchling turtles are thought to actively seek out frontal zones and
hence sargassum rafts. These
areas are then utilized for forage and protection during the "lost year"
(Carr 1986; 1987a,b).
Witham (1988) suggested an alternative hypothesis for this association.
He noted that it remains
untested as to whether sea turtles actually benefit from their
association with sargassum or
whether they are at increased risk from predation, entanglement, and
The pelagic sargassum habitat of the northwestern Atlantic consists of
both truly pelagic forms
and flotsom detached from coastal regions. While within the neuston it
species of invertebrates and vertebrates a source of food, shelter, and
structure varies with season, location, and algal age.
While the relationship of many species within this habitat is well understood, others remain less well known. This is particularly true for egg and larval stages of fishes, some crustaceans, and juvenile sea turtles. Functional relationships between the animals and the habitat have not been elucidated (e.g., we do not know the effect of the loss of sargassum on fish or sea turtle populations). Because estimates of oceanic biomass of sargassum are variable and inadequate, we also do not have a clear understanding the population dynamics of the sargassum habitat,
i.e., what is the standing crop, the productivity and the effect of
harvesting on living marine
We would like to express our thanks to Drs. J. J. Govoni and F. J.
Schwartz for providing
unpublished data, Drs. W. W. Kirby Smith and R. E. Robbins for assistance
with the invertebrate
taxonomy, and to Dr. C. S. Manooch, III for review of the report.
Dr. J. J. Govoni, National Oceanic and Atmospheric Administration,
National Marine Fisheries
Service, Southeast Fisheries Science Center, Beaufort Laboratory,
Beaufort, North Carolina,
Dr. C. S. Manooch, III, National Oceanic and Atmospheric
Administration, National Marine
Fisheries Service, Southeast Fisheries Science Center, Beaufort
Laboratory, Beaufort, North
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|Aglaeophenia latecarinata||O. hyalina|
|A. minuta||Plumularia catharina|
|A. perpusilla||P. corrugata|
|A. rigida||P. diaphana|
|Aglaeophenoides mammillata||P. floridana|
|Antenella secundaria||P. margaretta|
|Campanularia volubilis||P. megalocephala|
|Cladocryne pelagica||P. obligua|
|Clytia bicophora||P. sargassi|
|C. cylindrica||P. setaceoides|
|C. johnstoni||P. strictocarpa|
|C. longicyatha||Scandia mutabilis|
|C. noliformis||Setularia amplectens|
|C. raridentata||S. brevicyathus|
|C. simplex||S. corcicina|
|Desmocyphus pumilus||S. exigua|
|Eucopella sargassicola||S. gracilis|
|Gemmaria sp.||S. inflata|
|Gonothyraea gracilis||S. mayeri|
|G. integra||S. rathbuni|
|Halecium nanum||P. stookeyi|
|Hebella calcarata||S. turbinata|
|Laomedea sp.||S. versluysi|
|Obelia bicuspidata||Syncoryne mirabilis|
|O. dichotoma||Zanclea costata|
|O. geniculata||Z. gemmosa|
Table 1. Contd.
Table 1. Contd.
Alpheus sp .
Table 1. Contd.
Table 1. Contd.
*List compiled from Winge,1923; Parr, 1939; Adams, 1960; Yeatman, 1962; Weis, 1968; Friedrich, 1969; Fine, 1970; Dooley, 1972; Morris and Mogelberg, 1973; Ryland, 1974; Teal and Teal, 1975; Peres, 1982; Butler et al., 1983; Deason, 1983; Stoner and Greening, 1984; and Morgan et al., 1985.
Table 2. Fishes associated with pelagic Sargassum in the North Atlantic and Gulf of Mexico. * = early life stage present (i.e. egg, larvae or juvenile). Nomenclature follows Robins et al. (1991).
|Carcharhinus falciformes||silky shark|
|C. limbatus||blacktip shark|
|C. longimanus||oceanic whitetip shark|
|Sardinella aurita||Spanish sardine|
|Urophycis earlli||* Carolina hake|
|U. floridana||* southern hake|
|Histrio histrio||* sargassumfish|
|Cypselurus furcatus||spotfin flyingfish|
|C. melanurus||* Atlantic flyingfish|
|Exocoetus obtusirostris||* oceanic-two-wing flyingfish|
|Hirundichthys affinis||* fourwing flyingfish|
|Hyporhamphus unifasciatus||silverstripe halfbeak|
|Parexocoetus brachypterus||* sailfin flyingfish|
|Tylosurus acus||* agujon|
|Fistularia tabacaria||* bluespotted cornetfish|
|Macroramphosus scolopax||longspine snipefish|
|Hippocampus erectus||* lined seahorse|
|H. reidi||* longsnout seahorse|
|Microphis brachyurus||* opposum pipefish|
|Syngnathus floridae||* dusky pipefish|
|S. louisianae||* chain pipefish|
|S. pelagicus||* sargassum pipefish|
|S. springeri||* bull pipefish|
|Epinephelus inermis||* marbled grouper|
|Pristigenys alta||* short bigeye|
|Apogon maculatus||* flamefish|
|Rachycentron canadum||* Cobia|
|Phtheirichthys lineatus||slender suckerfish|
|Caranx bartholopmaei||* Yelow jack|
|C. crysos||* blue runner|
|C. dentex||* white trevally|
|C. hippos||* crevalle jack|
|C. latus||* horse-eye jack|
|C. ruber||* bar jack|
|Chloroscombrus chrysurus||* Atlantic bumper|
|Decapterus macerellus||* mackarel scad|
|D. punctatus||* round scad|
|D. tabl||* redtail scad|
|Elagatis bipinnulata||* rainbow runner|
|Seler crumenophthalmus||* bigeye scad|
|Seriola dumerili||* greater amberjack|
|S. fasciata||* lesser amberjack|
|S. rivoliana||* almaco jack|
|S. zonata||* banded rudderfish|
|Tachurus lathami||* rough scad|
|Coryphaena hippurus||* dolphin|
|Rhomboplites hippurus||* vermilion snapper|
|Lobotes surinamensis||* tripletail|
|Pagrus pagrus||* red porgy|
|Mullus auratus||* read goatfish|
|Pseudopeneus maculatus||* spotted goatfish|
|Upeneus parvus||* dwarf goatfish|
|Kyphosus incisor||* yelow chub|
|K. sectatrix||* Bermuda chub|
|Chaetodon ocellatus||*spotfin butterflyfish|
|C. striatus||* banded butterflyfish|
|Abudefduf saxatilis||* sergeant major|
|A. taurus||* night sergeant|
|Pomacentrus variabilis||* cocoa damselfisf|
|Mugil cephalus||* striped mullet|
|M. curema||* white mullet|
|Sphyraena barracuda||* great barracuda|
|S. borealis||* northern sennet|
|Polydactylus virginicus||* barbu|
|Bodianus pulchellus||* spotfin hogfish|
|Thalassoma bifasciatum||* bluehead|
|Acanthurus randalli||gulf surgeonfish|
|Unidentified||* snake mackerel|
|Acanthocybium solandri||* wahoo|
|Auxis thazard||frigate mackerel|
|Euthynnus alleteratus||little tunny|
|Katsuwonus pelamis||skipjack tuna|
|Scomber japonicus||* chub mackerel|
|Scomberomorus caballa||king mackerel|
|Thunnus albacares||yelowwfin tuna|
|T. atlanticus||blackfin tuna|
|Xiphias gladius||* swordfish|
|Istiophorus platypterus||* sailfish|
|Makaira nigricans||* blue marlin|
|Tetrapturus albidus||* white marlin|
|Cubiceps pauciradiatus||bigeye cigarfish|
|Hyperoglyphe bythites||black driftfish|
|H. perciformes||barrel fish|
|Peprilus triacanthus||* butterfish|
|Psenes cyanophrys||* freckled driftfish|
|Aluterus heudeloti||* dottorel filefish|
|A. monoceros||* unicorn filefish|
|A. schoepfi||* orange filefish|
|A. scriptus||* scrawled filefish|
|Balistes capriscus||* gray triggerfish|
|Cantherhines macrocerus||* whitespotted filefish|
|C. pullus||* orangespotted filefish|
|Canthidermis maculata||* rough triggerfish|
|C. sufflamen||* ocean triggerfish|
|Monacanthus ciliatus||* fringed filefish|
|M. hispidus||* planehead filefish|
|M. setifer||* slender filefish|
|M. tuckeri||* pygmy filefish|
|Xanthichthys ringens||* sargassum triggerfish|
|Chilomycterus antennatus||bridled burrfish|
|C. schoepfi||striped burrfish|
|Diodon holocanthus||* ballonfish|
|D. hystric||* porcupinefish|
|Sphoeroides spp.||* puffers|
List compiled from: Beebe and Vee-Van, 1928; Breder, 1938; Berry,
1959; Caldwell, 1959;
Gibbs and Collette, 1959; Adams, 1960; Berry and Vogele, 1961; Dawson,
1965; Beardsley, 1967; Bohlke and Chaplin, 1968; Randall, 1968; Weis,
1968; Friedrich, 1969;
Fine, 1970; Dooley, 1972; Hassler and Rainville, 1975; Teal and Teal,
1975; Hastings and
Bortone, 1976; Bortone et al., 1977; Fedoryako, 1980; Schwartz et al.,
1982; Manooch and
Hogarth, 1983; Manooch and Mason, 1983; Manooch et al., 1984; Manooch et
al., 1985; Carr,
1986; Fedoryako, 1989; Minerals Management Service, 1990; J. Govoni,
pers. comm.; C.
Manooch, pers. comm.; L. Settle, unpubl. data.