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Potentially Misidentified Species:
Harengula jaguana co-occurs with several clupeids throughout much of
its range. The congeners Harengula clupeola and Harengula
humeralis occur alongside H. jaguana in the IRL, as do the
Spanish Sardine, Sardinella aurita, and the Atlantic thread herring,
Opisthonema oglinum. Another co-occurring clupeid group, the
menhaden (Brevortia spp.), are considerably larger than scaled
sardines, reaching a standard length of 25 cm.
The body depth of the Spanish sardine is substantially less than that of
any other Harengula species, and the long thread-like dorsal
filament present on the thread herring is sufficient to allow
differentiation. The presence of a small toothed hypomaxilla between the
hind tip of the premaxilla and the expanded blade of the maxilla also
distinguishes the genus Harengula from co-ocurring clupeid genera
(FAO 1985).
The scales of H. jaguanaare less deciduous (shed less easily) than
those of H. humeralis (FAO 1985), and the lack of a dark shoulder
spot aids in discriminating H. humeralis and . clupeola from
H. jaguana. Scaled sardines also lack the diffuse reddish-orange
'ear spot'at the edge of the opercle that is typical of H. humeralis
(Robbins et. al 1986).
II. HABITAT AND DISTRIBUTION
Regional Occurrence:
The genus Harengula is a New World clupeid group found on both the
Atlantic and Pacific coasts. Harengula jaguana occurs in the
western Atlantic from New Jersey and Bermuda southward throughout Florida
and the Gulf of Mexico (abundant), in the Caribbean and down to southern
Brazil (FAO 1985).
IRL Distribution:
Scaled sardines can be found throughout the IRL system.
III. LIFE HISTORY AND POPULATION BIOLOGY
Age, Size, Lifespan:
Carpenter (2002) notes a SL of 12 cm as typical for the species, with the
largest specimens growing to 22 cm.
Scaled sardines are a fast-growing, short-lived species (FMRI undated).
Scale ring count analyses by Martinez and Houde (1975) suggested a lifespan
of three years for the species. More refined analysis involving the
enumeration of otolith daily rings buy Pierce et al. (2001), however,
indicated that scaled sardines on both coasts of Florida live for a maximum
of just one year. The authors report rapid growth rates exceeding 0.5
mm/day.
Abundance:
Harengula jaguana is abundant in nearshore estuaries and bays,
particularly from spring through autumn (Carpenter 2002).
Acoustic-geostatistical population assessments conducted by Paramo and Roa
(2003) in the northern Colombian Caribbean revealed that H. jaguana
were 80% more abundant as the Atlantic thread herring (Opisthonema
oglinum). Interestingly, O. oglinum, was a species under
commercial exploitation in the region while H. jaguana was not.
Thayer et al. (1999) record that the sardines Harengula jaguana and
Sardinella aurita were the 15th most numerous fish species collected
in Florida bay by trawl in 1984-1985, and the 6th most abundant species
collected a decade later. In the 1994-1996 trawl survey, sardines were
collected at a mean abundance of approximately 25 individuals per hectare.
Although its commercial importance in Florida is minimal (see below), a
Gulf of Mexico survey of eggs and larvae conducted by Houde (1977)
indicated a potential fishery yield of 41-92 million kg/year.
Reproduction:
Most individuals are mature at age-1 (80-130 mm), and all by age-2
(Carpenter 2002). Garcia-Abad et al. (1999) estimated that females in the
southern Gulf of Mexico reached maturity at around 117 mm.
Scaled sardines spawn offshore, typically 5-20 km from shore (Higgs and
Fuman 1998). Carpenter (2002) suggests spawning occurs at night from
January to September, with peak activity occurring from April to August
depending on location. Garcia-Abad et al. (1999) found that reproduction
persisted somewhat longer in the southern Gulf of Mexico, from February
through October and at depths of 18-36 m.
Martinez and Houde (1975) published details suggesting a serial spawning
behavior for this species. The authors found fish maturing at 78-85 mm SL
as well as continuous ripening of oocytes during a protracted spawning
season from February to August. Multiple spawning peaks have been noted by
other authors as well (Modde and Ross 1981).
Fecundity is estimated to range from 5,500 to more than 52,000 eggs per
individual, representing a relative fecundity of 323 to 807 eggs/g body
mass. Scaled sardine eggs are pelagic, transparent, and spherical,
averaging 1.55-1.85 mm in diameter (Carpenter 2002).
Embryology:
Pierce et al. (2001) examined combined Florida commercial purse-seine and
cast-net length-frequency data sets to find multiple juvenile recruitment
peaks consistent with the serial spawning reported by Martinez and Houde
(1975). Peaks occurred between June and November 1993 in Tampa Bay and
between June and August 1992 off of West Palm Beach. The authors note that
length frequency data suggests near-continuous, year-round spawning,
particularly in Tampa Bay. They further suggest that the earliest recruits
of the year could reach maturity to become spawners in the fall of the same
year. Garcia-Abad et al. (1999) also noted two distinct recruitment periods
in their study, one occurring March-June and a second occurring
August-November.
Whereas spawning occurs well offshore, most of the larval development
stages take place in nearshore and inshore waters (Shaw and Drullinger
1990, Hettler and Barker 1993). Vasquez-Yeomans (2000) reports that H.
jaguana eggs and larvae were present in samples collected from the
(Bahia de la Ascension estuary) in the western Caribbean from
April through October, and then again in December. Larvae occur offshore,
but their distribution patterns over time suggest shoreward movement (Houde
1977, Ruple 1984, Vasquez-Yeomans 2000).
IV. PHYSICAL TOLERANCES
Temperature:
Harengula jaguana is found from the tropics to warm temperate waters
as far north as New Jersey. The northern distributional limit for the
species is likely to be temperature-based.
Salinity:
Harengula jaguana is a euryhaline species encountered in habitats
ranging from coastal marine areas to mesohaline estuaries to hypersaline
lagoons (Carpenter 2002).
Paperno et al. (2000) found that in the upper IRL system H. jaguana
was most prevalent in the vicinity of Ponce de Leon Inlet, occurring in
substantially lesser abundance in less saline portions of the estuary.
Other investigators have also indicated that among the clupeids, H.
jaguana and Opisthonema oglinum are two species typically
associated with higher salinities (Richards et al. 1974, Hoese and Moore
1977, Higgs and Fuiman 1998).
Dissolved Oxygen:
LC50 tests conducted by Goodman and Campbell (2007) to determine lethal
levels of hypoxia for seven Gulf of Mexico fish species and two crustacean
species indicated that Harengula jaguana was the most
hypoxia-sensitive of all the species tested. Oxygen concentrations of 2.17
mg/L were low enough to kill a substantial number of test animals in less
than four hours.
V. COMMUNITY ECOLOGY
Trophic Mode:
Adult scaled sardines are planktivores, feeding on a variety of prey items
including copepods, mysids, amphipods, isopods, ostracods, insect larvae,
and small molluscs (FMRI undated).
Competitors:
Likely competitors with adult scaled sardines include other planktivorous
fish species (e.g., anchovies), but it is unlikely that dietary resources
are often a limiting factor.
Predators:
Harengula jaguana represent an important prey resource for a wide
variety of piscivoroue predators, including mackerels, gag grouper,
bluefish, crevalle jack, tunas, and sharks (FMRI undated, Hoffmayer and
Parsons 2003).
The species is also an important prey item for sea birds and wading birds,
as well as bottlenose dolphin (Sogard et al. 1989, FMRI undated).
Habitats:
Scaled sardines are marine (especially coastal) and estuarine schooling
pelagic and demersal fish commonly found over sand and mud bottoms and on
and around seagrass meadows (Carr and Adams 1973, Williams 1985, Sogard et
al.1989, Carpenter 2002).
In coastal areas, individuals typically remain within the 40-m isobath, but
are rarely collected from as deep as 500 m (Pierce et al. 2001).
Activity Time:
Although the species is often encountered by day, Sogard et al. (1989)
listed Harengula jaguana among the nocturnally active fish species
on seagrass-covered shallow banks in Florida Bay. Animals increased in
abundance at dusk with high catch rates continuing for a few hours
thereafter and then declining through the night. The authors note, however,
that activity patterns were influenced by tidal patterns as well as diel
cycles. In addition, the authors pointed out that nocturnal activity seen
in this study may be related to vertical migration in crustacean prey that
reside on the bottom by day but ascend into the water column at night.
VI. SPECIAL STATUS
Special Status:
None
Economic/Ecological Importance:
Scaled sardines represent an important link in estuarine and coastal food
webs (Garcia-Abad et al. 1999). They are abundant consumers of zooplankton
as well as an important prey species for a variety of piscivorous fish,
birds, and marine mammals (Carr and Adams 1973, Pierce et al. 2001).
A small Harengula jaguana baitfish industry exists in south Florida,
and the species is caught as part of the mixed-species 'trashfish' fishery
as well. It is marketed fresh for human consumption primarily outside of
the US and it is canned in Cuba and Venezuela.
Total statewide landings in 1999 were nearly 140,000 kg, with 61% of the
total taken from the Gulf coast and almost all of the landings attributable
to the commercial fishery. Annual landings are highly variable, dropping
by as much as 60% in two years (FMRI undated). Such variability is typical
of several 'trashfish' fishery species.
VII.
REFERENCES
Carr WES, and CA Adams. 1973. Food habits of juvenile marine fishes
occupying sea-grass beds in the estuarine zone near Crystal River, Florida.
Transactions of the American Fisheries Society 102:511-540.
Carpenter KE (ed.). 2002. The living marine resources of the Western
Central Atlantic. Volume 2: Bony fishes part 1 (Acipenseridae to
Grammatidae). FAO Species Identification Guide for Fishery Purposes and
American Society of Ichthyologists and Herpetologists Special Publication
No. 5. pp. 601-1374.
Florida Marine Research Institute. Undated. Scaled Sardines (Harengula
jaguana). Species account for scaled sardines in Florida. FMRI Stock
Assessment Group. 3 p.
Garcia-Abad MC, Tapia-Garcia M, Yanez-Arancibia, and P. Sanchez-Gil. 1999.
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Goodman LR and JG Campbell. 2007. Lethal levels of hypoxia for gulf coast
estuarine animals Marine Biology 152:37-42.
Hettler WF and DL Barker. 1993. Distribution and abundance of larval fishes
at two North Carolina inlets. Estuarine and Coastal Shelf Science
37:161-179.
Higgs DM and LA Fuiman. 1998. Associations between sensory development and
ecology in three species of clupeoid fish. Copeia 1998:133-144.
Hoese HD and RH Moore. 1977. Fishes of the Gulf of Mexico. Texas,
Louisiana, and Adjacent Waters. Texas A&M University Press, College Station
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Hoffmayer ER and GR Parsons. 2003. Food Habits of Three Shark Species from
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Houde ED. 1977. Abundance and potential yield of the scaled sardine,
Harengula jaguana, and aspects of its early life history in the
eastern Gulf of Mexico. Fishery Bulletin 75:613-628.
Martinez S, and ED Houde. 1975. Fecundity, sexual maturation, and spawning
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Paperno R, Milleand KJ, and E Kadison. 2001. Patterns in species
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Paramo J and R Roa. 2003. Acoustic-geostatistical assessment and
habitat-abundance relations of small pelagic fish from the Colombian
Caribbean. Fisheries Research 60:309-319.
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scaled herring, Harengula jaguana, from Florida waters, as indicated
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Richards WJ, Miller RV, and ED Houde. 1974 Egg and larval development of
the Atlantic threadfin herring, Opisthonema oglinum. Fishery
Bulletin 72:1123-1136.
Robins CR, Ray GC, and J Douglas. 1986. A Field Guide to Atlantic Coast
Fishes. The Peterson Field Guide Series. Houghton Mifflin Co., Boston. 354
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Ruple DL. 1984. Occurrence of larval fishes in the surf zone of a northern
Gulf of Mexico barrier island. Estuarine, Coastal and Shelf Science
18:191-208.
Shaw RF and DL Drullinger. 1990. Early-life- history profiles, seasonal
abundance, and distribution of four species of clupeid larvae from the
northern Gulf of Mexico, 1982 and 1983. NOAA Technical Report NMFS 88.
Sogard SM, Powell GVN, and JG Holmquist.1989. Utilization by fishes of
shallow, seagrass-covered banks in Florida Bay: 2. Diel and tidal patterns.
Environmental Biology of Fishes 24:81-92.
Thayer GW, Powell AB, and DE Hoss. 1999. Composition of Larval, Juvenile,
and Small Adult Fishes Relative to Changes in Environmental Conditions in
Florida Bay. Estuaries 22:518-533.
Vasquez-Yeomans L. 2000. Seasonal variation of ichthyoplankton in a western
Caribbean bay system. Environmental Biology of Fishes 58: 379-392.
Whitehead PJP. 1985. Clupeoid fishes of the world. An annotated and
illustrated catalogue of the herrings, sardines, pilchards, sprats,
anchovies and wolf-herrings. Part 1 - Chirocentridae, Clupeidae and
Pristigasteridae. FAO Fisheries Synopsis No. 125, Volume 7. 303 p.
Report by:
J. Masterson, Smithsonian Marine Station
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