Species Description: 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.
Potentially Misidentified Species: Panulirus argus closely resembles a related species, the smoothtail spiny lobster, P. laevicauda, whose range is sympatric. The two are easily distinguished based on two 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.
Regional Occurrence: 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).
IRL Distribution: 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.
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).
Reproduction: 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 24 ºC 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).
Embryology: 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 available (Marx and Herrnkind 1986).
Temperature: 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 - 28 ºC (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).
Salinity: Postlarvae do not tolerate salinity below 19 parts per thousand (ppt) (Witham et al 1968).
Trophic Mode: 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 extract prey.
Competitors: Competition among 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.
Predators: 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 1980).
Habitats: 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 five 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 long 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 1980).
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 al 1975).
Activity Time: Juveniles and adults are primarily nocturnal, with juveniles being somewhat more nomadic (Marx and Herrnkind 1986).
Special Status: Commercially important.
Fisheries Importance: Commercial Fishery
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 million. Most 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 five-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 23rd in commercial value to IRL counties, and 44th 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%) counties.
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 2001).
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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