LIFE HISTORY AND POPULATION BIOLOGY
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 female's 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 female's 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
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