Panulirus argus closely resembles a related species, the
smoothtail spiny lobster, P. laevicauda, whose range is
sympatric. The two are easily distinguished based on 2 characters:
P. laevicauda lacks dorsal grooves on abdomen, but possesses
a series of small white spots that run along the lateral margins
of the abdomen.
The carapace of Panulirus argus is subcylindrical and bears
many strong, forward-facing spines. A pair of rostral horns projects
forward over the stalked eyes. Whip-like antennae are longer than
the body and are studded with small spines and setae. Antennules
are also elongate, extending approximately 2/3 body length. The
abdomen is notched along its lateral margins, but is otherwise smooth.
Each segment of the abdomen has a transverse groove that is disrupted
at the midline. The central telson is flanked by 2 pairs of biramous
uropods. Body color is varied, but is generally a gray or tan base
color mottled with shades of green, red, brown, purple, or black.
The second and sixth segments of the abdomen have large yellow or
white ocelli, with smaller ocelli scattered dorsolaterally along
the abdomen. Legs are striped longitudinally in a dull blue color.
Dactyls of the walking legs are setose. Pleopods are bright orange
and black. The endopodites in female pleopods are well developed
and hook-like, bearing many setae. In juveniles, antennae and pereopods
are banded with white; and a broad white stripe runs the length
of the dorsal midline across both the carapace and abdomen.
Panulirus argus is sexually dimorphic,
with females distinguished from males by differences in the sternum,
legs and genital openings. In males, the sternum is somewhat broader
and lacks the striations that are commonly found in females. The
second pair of walking legs in males is more elongate than the other
legs, and bears longer, curved dactyls. Females have small chelae
on the dactyls of the fifth pair of walking legs. The raised genital
openings of males are located ventrally, on the bases of the fifth
pair of walking legs. In females, the gonopores are set at the bases
of the third pair of walking legs.
II . HABITAT AND DISTRIBUTION
Spiny lobsters occur from North Carolina south to Brazil including
Bermuda, the Gulf of Mexico, West Indies and Caribbean. The northernmost
extent of the range is North Carolina (Williams 1984).
Adults are most prevalent in nearshore and offshore waters, but
are found throughout the Indian River Lagoon in areas where there
is ample shelter. Juveniles commonly occur in seagrasses, mangrove
creeks and oyster reefs.
III. LIFE HISTORY AND POPULATION
Age, Size, Lifespan:
Panulirus argus reaches a maximum length of approximately
45 cm (1.47 feet), but is more commonly found at lengths of approximately
20 cm (7.9 inches). Growth in the first year averages 5 cm (1.5
inches), with growth thereafter averaging approximately 2.5 cm (1
inch) per year. Male and female growth rates are approximately equal
in south Florida nursery areas (Davis and Dodrill 1980). Females
tend to grow somewhat more slowly than males (Little 1972; Olsen
and Koblick 1975) and do not achieve as large a size (Williams 1984).
Spiny lobsters molt an average of 2.5 times
per year, with most molting occurring from March July, or from
December February in Florida (Williams 1984). Growth characteristics
are correlated with age, with molt frequency and increment of growth
at molting tending to decline with age (Aiken 1980). Variability
in food quality and abundance, population density, water temperature,
as well as rates of predation and injury, all affect growth rates
in local populations.
Eldred et al. (1972) estimated average growth
rates in wild populations of Panulirus argus as 5 mm (0.2
inches) per month for the first 9 -10 months post-settlement in
Biscayne Bay, Florida. Tagging studies in the same area showed that
mean growth rates in 40-85 mm (1.6 3.3 inches) lobsters averaged
1.8 mm (0.07 inches) per month, with physical condition having a
significant impact on growth rates. Davis (1981) reported that individuals
having no injuries grew approximately 2.2 mm (0.09 inches) per month,
while those missing legs and antennae grew only 1.3 mm (0.05 inches)
per month. In Florida Bay, however, growth rates for injured and
uninjured lobsters were approximately equal, at 3.3 mm (0.12 inches)
per month (Davis and Dodrill 1980). In Key West, tagged lobsters
49-83 mm (1.9 3.3 inches) in length grew approximately 3.1 mm
(0.12 inches) per month (Little 1972).
Witham et al. (1968) reared pueruli of
Panulirus argus under laboratory conditions over 7 months.
During this time pueruli averaging 6 mm (0.2 inches) carapace length
(CL) grew to an average of 34 mm (1.33 inches), with growth rates
ranging from 3.8 4.2 mm (0.14 0.16 inches) per month. Other
laboratory populations grew considerably more slowly, with Lewis
et al. (1952) and Sweat (1968) reporting monthly growth rates less
than 2 mm (0.07 inches) per month.
Assuming a post-settlement growth rate of
4.0 mm (0.16 inches) per month, Panulirus argus likely
reaches legal harvest size in 20 40 months, depending on location
and degree of injury (Little 1972; Davis and Dodrill 1980).
Panulirus argus reach sexual maturity at lengths of approximately
70 80 mm (2.8 3.2 inches) (Witham et al 1968. Olsen et al 1975,
Davis 1979). Adult male and females often inhabit estuaries, bays,
and lagoons; but spawning typically occurs in nearshore and offshore
reef fringes and other hardbottom areas from late spring through
summer in Florida waters (Davis 1975; Kanciruk and Herrnkind 1976;
Lyons et al. 1981). Annual variation in peak spawning period is
dependent on water temperature. Lyons et al. (1981) reported spawning
begins when water temperatures reach 24º C in deeper reef areas.
In Florida, there is little evidence that spiny lobsters spawn more
than once per year; however, in Bermuda, evidence supports multiple
spawning (Sutcliffe 1952).
Mating follows short courtships that involve
signals being given by both sexes. In copulation, males hold females
sternum to sternum and extrude a spermatophore which is adhered
to the females sternum, and will remain until the time of spawning.
Sperm may be viable for as long as a month (Marx and Herrnkind 1986).
Spawning occurs when females flex the abdomen under the carapace,
spreading the telson and uropods, as well as the pleopods. Eggs
are shed onto the abdomen, while the female scratches at the spermatophore
to liberate sperm and fertilize eggs as they are extruded.
Fecundity varies with body size. Females
71-75 mm (2.8 3 inches) in length carry approximately 230,000
eggs; while females over 100 mm (3.9 inches) carry more than 700,000
eggs (Mota-Alves and Bezerra 1968). Williams (1984) reported females
23 cm (9.1 inches) with 500,000 eggs, 30 cm (11.8 inches) with 1.18
million eggs, and 38 cm (15 inches) with 2.6 million eggs.
Using the Index of Reproductive Potential
(IRP) Lyons et al. (1981) estimated that female Panulirus argus
in the upper Florida Keys measuring 76-85 mm (3.0 3.3 inches)
carapace length (CL) contribute 48% of all egg production in the
population. Females over 85 mm (3.3 inches) CL comprise approximately
20% of the female population, but contribute 41% of egg production.
Females smaller than 76 mm (3.0 inches) comprise approximately 25%
of all females, but contribute 11 % of all eggs.
Minimum spawning size has declined in Florida
Panulirus argus females, perhaps as a response to intense
fishing pressures. In 1922, minimum spawning size of females was
reported to be 76mm (3.0 inches) (Crawford and De Schmidt 1922).
Recent surveys have revealed reproductive females as small as 65
mm 71 mm (2.6 2.8 inches) in south Florida (Warner et al. 1977;
Lyons et al 1981). However, unfished populations in the Dry Tortugas
area show the minimum size in egg-bearing females to be 78 mm (3.1
inches) (Davis 1975). Suggested reasons for this decline in minimum
spawning size have been genetic selection; modified sexual behavior
when large females become rare; and reduced growth rates (Davis
1975; Warner et al. 1977; Lyons et al 1981). Decline in size of
mature females has caused a marked reduction in reproductive potential,
with Lyons et al. (1981) estimating that egg production in Florida
Keys spiny lobsters was only 12% of that expected from a similarly
sized, unfished population (Lyons et al. 1981).
Eggs are bright orange in color and measure approximately 0.5 mm
(0.02 inches) In diameter. Upon being extruded, eggs adhere to hook-like
setae of pleopods located on the underside of the abdomen. They
will remain in place until hatching at approximately 3 weeks. Several
days before hatching occurs, eggs turn a darker brown color. Eggs
hatch as transparent phyllosome larvae and are dispersed into the
water column by repeated flexing of the females abdomen. A second,
smaller spawning may occur in Caribbean and West Indies populations
about 1 week after release of eggs, though there is no evidence
that a second spawning occurs in Florida waters. Following a second
spawning, the ovary is usually spent, and the spermatophore erodes.
Molting typically occurs after spawning. (William 1984).
Phyllosomes are transparent and morphologically
adapted for long planktonic existence being transported on oceanic
currents for 6-12 months before metamorphosis to the postlarval
stage (Lyons et al 1981). Phyllosomes are dorsoventrally flattened
and have a bi-lobed cephalothorax. Appendages are long and setose
to assist in floatation in the water column. Swimming is accomplished
by flexion in the exopodites of the legs (Provenzano 1968). Phyllosomes
vertically migrate on a daily basis, ascending to surface waters
at night, and descending during the day (Sims and Ingle 1967). There
are 11 stages of phyllosome development. During this period, larval
size increases from 2 mm (0.07 inches) total length (TL) at hatching
to approximately 34 mm (1.3 inches) before the metamorphosis to
the postlarval puerulus stage.
Pueruli persist for several weeks, are nonfeeding
and oceanic (Lyons 1980). Like phyllosomes, they are dorsoventrally
flattened and transparent, with no calcification in the carapace.
Pueruli return to coastal waters from offshore, swimming shoreward
at night, and tending to remain within a few centimeters of the
water surface (Lyons 1980). Pueruli enter estuarine habitats throughout
the year, with peaks occurring during new and first-quarter moons
(Sweat 1968; Little and Milano 1980). Peak recruitment varies from
year to year, but the main peak typically occurs in spring, followed
by a lesser peak in the fall (Lyons 1980).
Upon encountering suitable inshore substrates,
pueruli settle to the benthos, typically in vegetated areas of algal
beds, mangrove areas where prop roots are fouled by algae, seagrasses,
small holes, and sand-mud bottoms (Witham et al 1964). Several days
before the molt to the first juvenile stage, they begin to show
signs of pigmentation, turning a red-brown color. Benthic juveniles
are cryptically colored in varying shades, and have banding and
striping that aids in camouflage. Young juveniles tend to be solitary
and behave aggressively toward conspecifics, lashing them with antennae,
or prying at them to dislodge them from refuges (Marx and Herrnkind
1986). However, an ontogenetic shift occurs to more social, gregarious
behavior as body size increases. This shift appears to be at least
partially dependent on the distribution and quality of shelters
available (Marx and Herrnkind 1986).
IV. PHYSICAL TOLERANCES
Panulirus argus are generally found where minimum monthly
water temperatures exceed 20 ºC (68 ºF) (George and Main
1967). Optimal growth occurs in temperatures of 26 - 28ºC (Marx
and Herrnkind 1986). While postlarvae tolerate short-term temperature
declines to 13 ºC (55.4 ºF) (Little and Milano 1980),
they have been found to grow more slowly and experience higher mortalities
when temperatures are sustained below 16 ºC (60.8ºF) (Witham
1974). Rapid decline to water temperatures below 10 ºC (50.0
ºF), especially in molting lobsters, may cause death (Marx
and Herrnkind 1986).
Rapid growth with poor survival has been
often observed at water temperatures exceeding 32 ºC (89.6
ºF) (Witham 1974; Aiken 1980).
Postlarvae do not tolerate salinity below 19 parts per thousand
(ppt) (Witham et al 1968).
V. COMMUNITY ECOLOGY
The diet of phyllosome larvae is not well described; however, those
in culture consume chaetognaths, euphausids, fish larvae, and ctenophores
(Provenzano 1968; Phillips and Sastry 1980). Pueruli are non-feeding
(Lyons 1980). Benthic juveniles and adults are foragers that utilize
chemoreceptive setae lining the antennules and walking legs assist
in locating food sources (Ache and Macmillan 1980).
Panulirus argus are often the dominant
carnivores within their habitats, and likely effect the population
sizes of other benthic species (Berry and Smale 1980). Typical prey
items include a variety of slow-moving or sedentary organisms including
gastropods, bivalves, crustaceans, annelids and echinoderms. Shelled
organisms are manipulated with the legs and positioned near the
mouth, where powerful mandibles can then crush or chip shells to
lobster species in Florida is thought to be of little consequence
as Panulirus argus is the dominant lobster species in Florida. A
congener, P. laevicauda, though sympatric, is relatively scarce
and generally found only in reef habitats (Marx and Herrnkind 1986);
thus, little competition likely occurs.
Panulirus argus larvae are consumed by a variety of pelagic
fish species (Phillips and Sastry 1980). Pueruli are consumed by
fishes as well as by benthic and epibenthic organisms such as crabs
and octopods (Little and Milano 1980). Later juveniles and adults
are armoured against predators both by their spiny exoskeletons
and by their behaviors, with rapid tail-flipping the most commonly
observed escape response. Spiny lobsters also resist predation by
congregating together in shelters and blocking den openings (Lipcius
et al. 1983).
Large predators of juvenile and adult spiny
lobsters include groupers (especially goliath groupers) (Crawford
and De Smidt 1922), sharks, loggerhead turtles, and octopods (Kanciruk
Spiny lobsters are gregarious and migratory, most commonly found
in coastal and shallow continental waters to depths of 90 m (295
feet) or more. The life cycle consists of 5 phases, each of which
is habitat-associated. The oceanic, planktonic stage is characterized
by the phyllosome larva. Phyllosomes inhabit the epipelagic zone
of open ocean waters where temperature and salinity are relatively
constant. Optimum survival occurs when conditions are stable and
nonturbid, with no environmental pollutants. The swimming postlarval
stage is characterized by the puerulus postlarva, which utilizes
a broad range of nearshore and estuarine habitats, but settles primarily
in well-vegetated habitats such as seagrasses meadows and algal
beds. Algae, particularly the red alga Laurenia spp., appear to
be especially important to newly settled postlarvae. The early benthic
stage is characterized by the banded coloration pattern in young
juveniles, which utilize mangrove creeks and vegetated shallow water.
The late juvenile stage occurs in older juveniles, which utilize
seagrasses and oyster reefs for as much as 2 years before migrating
as sub-adults to shallow banks in nearshore waters. The adult stage
is characterized by mature adults, which utilize hardbottom, patch
reefs and coral reefs (Marx and Herrnkind 1986), commonly using
crevices in coral reefs, overhangs, outcroppings, and other hard
substrates for shelter. Adults use softbottom habitats and seagrasses
primarily during migratory periods (Herrnkind et al 1975; Kanciruk
Though adults often inhabit bays and estuarine
habitats, they do not typically spawn there. Spawning occurs offshore
in sheltered areas having low turbidity, low wave action and adequate
larval transport by currents and waves (Kanciruk and Hernnkind 1976).
Lobsters longer than 20 mm aggregate in shelters
within protected bays and high salinity estuaries (Olsen et al.
1975; Davis 1979). Typical shelters include sponges, corals, mangrove
roots, holes, rocky outcrops, and ledges. Davis (1971) reported
juvenile spiny lobsters taking shelter under sea urchins. At approximately
70-80 mm, at the onset of sexual maturity, lobsters begin the migration
to nearshore and offshore reefs. More females than males migrate
offshore, with females tending to migrate into deeper waters in
spring and summer for mating and larval release (Lyons et al. 1981).
Both sexes migrate offshore in fall and winter months as severe
fall storms arrive and water temperatures begin to decrease (Davis
1977; Herrnkind 1982). On occasion, mass migration of spiny lobster
occurs, with lobsters forming single-file lines that stretch long
distances (Kanciruk and Herrnkind 1978; Marx and Herrnkind 1986).
Offshore populations consist primarily of
adults that live communally or singly in crevices of rocks and corals,
with most lobsters showing high site fidelity (Herrnkind et al.
1975). Spiny lobsters are relatively selective when choosing den
sites and show a preference for those that allow complete concealment,
exclude large predators, and contain other lobsters (Herrnkind et
Juveniles and adults are primarily nocturnal, with juveniles being
somewhat more nomadic (Marx and Herrnkind 1986).
VI. SPECIAL STATUS
The statewide commercial catch of spiny lobsters is the second most
valuable Florida shell fishery, second only to shrimp. Since 1970,
the commercial catch has ranged without trend between 4.3 million
7.9 million pounds per year (Muller 2003). Between
1987 - 2001, the commercial harvest of Panulirus argus
in Florida totaled 94.6 million pounds, and was valued at $365.3
landings occur on the Gulf coast and in Monroe and Dade counties
on the East coast. Generally less than 1.5 million pounds of spiny
lobsters are landed outside the vicinity of the Florida Keys each
year (Muller 2003), and this is reflected in the relatively low
annual value of the spiny lobster catch to IRL counties. Commercial
lobstermen using traps account for approximately 80% of the total
catch, with recreational divers harvesting 20%. The
5 county area encompassing the IRL (Volusia, Brevard, Indian River,
St. Lucie and Martin Counties) accounted for 607,890 pound of the
commercial harvest, which had a value of $2.3 million. This ranks
the spiny lobster twenty-third in commercial value to IRL counties,
and forty-fourth in pounds harvested.
Figure 1 below shows the
dollar value of the spiny lobster commercial fishery to IRL counties
by year. The fishery ranged in value from a high of $260,838 in
2001, to a low of $68,113 in 1988. Brevard County accounted for
the largest share of the catch at 53.3%, followed distantly by Martin
(15.3%), St. Lucie (13.3%),Volusia (13.2%) and Indian River(4.9%)
Annual dollar value of the commercial catch of spiny lobster to
the 5-county area of the Indian River Lagoon.
Total spiny lobster dollar value and percentage by county for the
years 1987 - 2001.
Table 1. Total dollar value
of IRL spiny lobster, Panulirus argus, between 1987 - 2001.
Table 2. By-county annual
and cumulative percentages of the spiny lobster harvest for the
Table 3. By-county cumulative
dollar value and percentage of total for the spiny lobster harvest
from 1987 - 2001.
Beginning in 1991, the Florida Fish and Wildlife Conservation Commission
began utilizing mail surveys to estimate the recreational harvests
of spiny lobster by recreational fishers during the 2-day mid-summer
sport season, and the regular season, which opens in early August
and runs through late March. In a typical year, the recreational
harvest of spiny lobsters averages 20 - 22% of the commercial catch
with approximately 2 million pounds of lobster harvested (Eaken
Harvest regulations for spiny
lobsters state the carapace must be longer than 3 inches as measured
from the groove between eyes to the end of carapace. Recreational
harvesting requires a saltwater fishing license. Recreational anglers
must measure lobsters in the water and immediately release undersized
animals and females with eggs. The bag limit in most areas is 6
lobsters per person per day.
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