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The Caribbean spiny lobster, Panulirus argus (Latreille, 1804), is an important resource in the Western Atlantic Ocean, Caribbean Sea, and Gulf of Mexico. A nemertean egg predator of P. argus, Carcinonemertes conanobrieniSimpson, Ambrosio & Baeza, 2017, was recorded in Florida in 2017 and Colombia in 2018. This study sought to investigate the presence of C. conanobrieni in P. argus in Saint Kitts and Nevis, eastern Caribbean Sea. Nemerteans were observed in the brood masses of P. argus. Molecular techniques confirmed its presence in the P. argus population on Saint Kitts. Egg, juvenile, and adult stages of C. conanobrieni and of an undescribed Carcinonemertes sp. were found in the brood masses of 27 of 31 (87%) ovigerous P. argus. Differences in the shape of the nemertean egg masses (string versus spherical arrangement) were confirmed as two distinct species of Carcinonemertes Coe, 1902 using molecular analyses. The juvenile stage of the undescribed Carcinonemertes sp. was detected in the gills of 72 of 320 (23%) of the P. argus individuals from Saint Kitts. The prevalence of branchial nemertean infestation was not significantly different among male and female P. argus. Female lobsters with ovaries in early stages of gonadal maturation were more likely to have branchial nemerteans relative to those with ovaries in late stages of maturation, suggestive of nemertean migration from gills to egg mass once the host oviposits. This is the first report of C. conanobrieni in the Caribbean spiny lobster from the eastern Caribbean, which is approximately 1,300 miles from Florida and 1,000 miles from Colombia. This is also the first report of an undescribed nemertean worm in P. argus. The high prevalence of Carcinonemertes spp. in P. argus raises concern for the extent to which these egg predators may be impacting P. argus in Saint Kitts, an important fishery on the island.

The review explores the current diversity of methods used to determine the nutritional condition of crustaceans and to provide evidence that standardization is needed to make results comparable among the increasing number of studies in this field. The advantages and disadvantages of inexpensive and wide-ranging indices are discussed with respect to the blood refractive index (BRI) and the effectiveness of the total weight/carapace length relationship based on their application to the spiny lobster Panulirus argus (Latreille, 1804). Also reviewed are the environmental factors that impact metabolism based on the nutritional condition concept and the nutritional condition itself. Ocean acidification, temperature, salinity, dissolved oxygen, and noise have been the most studied factors impacting nutritional condition in crustaceans, usually under controlled (laboratory) conditions. Methodological aspect could be nevertheless one of the most important factors affecting nutritional condition estimations. More than twenty methods have been applied to determine nutritional variations in crustaceans, all then following different techniques, procedures, and instrumentation. These differences make results almost impossible to compare. BRI determined by a hand-held refractometer, usually inexpensive, is a good option to standardize studies so that the results from different species and different regions can be comparable; other methodologies could nevertheless be used in combination. Laboratory studies, which typically use few individuals that are exposed to extreme and usually non-natural conditions, show limited results. Emphasis should be placed on field studies that integrate the effects of several environmental factors.

There is little information on the early life history of the spotted spiny lobster Panulirus guttatus (Latreille, 1804), an obligate reef resident, despite its growing importance as a fishery resource in the Caribbean and as a significant predator. We cultured newly-hatched P. guttatus larvae (phyllosomata) in the laboratory for the first time, and the growth, survival, and morphological descriptions are reported through 324 days after hatch (DAH). Phyllosomata were cultured at 25 °C in a flow-through seawater system within a series of custom 80 l plankton-kreisel tanks and provided with ongrown Artemia and mussel gonad. Mean body length (BL) of phyllosomata was 1.70 mm (N = 10) at hatch and increased linearly to 22.20 mm at 226 DAH (N = 3). Morphological characters from a total of 164 sampled phyllosomata were ascribed to nine distinct developmental stages (stages I-IX), and described and illustrated. Although no final stage phyllosomata (stage X) were obtained, the BL in the final stage was extrapolated at 39.6 mm using a Gompertz function, expressing the relationship between phyllosoma stages and BL. The total duration of phyllosomata for P. guttatus was estimated at 410 d, with the 5th and 95th percentiles at 334 and 526 d, respectively. Our data suggest that P. guttatus has a larger body size in the final larval stage and a substantial pelagic larval duration compared with other related panulirid lobsters in its geographic range. The described morphological and biological attributes associated with the early-life history for this species can inform future studies, and add value to models of distribution and population connectivity.

The Caribbean spiny lobster Panulirus argus (Latreille, 1804) is an important species in shallow-water coral reefs and target of the most lucrative fishery in the Caribbean Sea. We explored historical demography in P. argus inferred using single nucleotide polymorphisms (SNPs). We expected an increase in population size of P. argus from Florida, USA starting ~18,000-24,000 years ago, after the Last Glacial Maximum, when ice sheets started to retreat and sub-tropical/tropical shallow coastal waters warmed up. A total of 10 lobsters were collected from shallow reefs in the Florida Keys, Florida, USA. One microgram of gDNA extracted from each specimen was used for RAD library construction using established protocols. A panel of 1643 SNPs obtained after interrogation of RAD-tags was used to calculate a site frequency spectrum (SFS). The observed SFS for the Florida population of P. argus exhibited a non-normal distribution peaking at singleton SNPs. The expected SFS in a total of six different candidate demographic models with dissimilar population size changes through time (i.e., standard neutral, exponential growth, bottleneck, bottleneck + growth, two epochs, and three epochs) were numerically computed in the software A i and a model selection approach was implemented to test which expected model(s) best fitted the empirical SFS. The model selection approach indicated that the bottleneck + growth model most closely matched the observed SFS; P. argus experienced a population decline at about 1.9 (0.75-5.7) mya, to then recover and growth exponentially until present time. In disagreement with expectations, population expansion started much earlier than ~18,000-24,000 years ago. Fisheries and conservation studies are expected to profit from the evaluation of genomic and population variability in this species using demographic models, as shown here. Studies exploring population connectivity and locality-specific demographic history of P. argus are underway. © 2019 The Author(s) 2019. Published by Oxford University Press on behalf of The Crustacean Society. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com. La langosta espinosa del Caribe Panulirus argus (Latreille, 1804) es una especie importante en los arrecifes de coral de aguas poco profundas y objetivo de la pesquería más lucrativa del Mar Caribe. Exploramos la demografía histórica en P. argus inferida mediante polimorfismos de un solo nucleótido (SNP). Esperábamos un aumento en el tamaño de la población de P. argus de Florida, EE. UU., Comenzando hace ~ 18.000-24.000 años, después del Último Máximo Glacial, cuando las capas de hielo comenzaron a retirarse y las aguas costeras poco profundas subtropicales / tropicales se calentaron. Se recolectaron un total de 10 langostas de arrecifes poco profundos en los Cayos de Florida, Florida, EE. UU. Se utilizó un microgramo de ADNg extraído de cada muestra para la construcción de la biblioteca RAD utilizando protocolos establecidos. Se utilizó un panel de 1643 SNP obtenidos después de la interrogación de etiquetas RAD para calcular un espectro de frecuencia de sitio (SFS). El SFS observado para la población de Florida de P. argus exhibió una distribución no normal con un pico en los SNP únicos. El SFS esperado en un total de seis modelos demográficos candidatos diferentes con cambios de tamaño de población diferentes a lo largo del tiempo (es decir, estándar neutral, crecimiento exponencial, cuello de botella, cuello de botella + crecimiento, dos épocas y tres épocas) se calcularon numéricamente en el software A i y Se implementó un enfoque de selección de modelos para probar qué modelo (s) esperado (s) se ajustaba mejor al SFS empírico. El enfoque de selección del modelo indicó que el modelo de cuello de botella + crecimiento coincidía más estrechamente con el SFS observado; P. argus experimentó una disminución de la población de aproximadamente 1,9 (0,75-5,7) millones de años, para luego recuperarse y crecer exponencialmente hasta la actualidad. En desacuerdo con las expectativas, la expansión de la población comenzó mucho antes que hace ~ 18.000-24.000 años. Se espera que los estudios de pesca y conservación se beneficien de la evaluación de la variabilidad genómica y poblacional de esta especie utilizando modelos demográficos, como se muestra aquí. Se están realizando estudios que exploran la conectividad de la población y la historia demográfica específica de la localidad de P. argus. Department of Biological Sciences, 132 Long Hall, Clemson University, Clemson, SC, United States Smithsonian Marine Station at Fort Pierce, 701 Seaway Drive, Fort Pierce, FL, United States Departamento de Biología Marina, Facultad de Ciencias Del Mar, Universidad Católica Del Norte, Coquimbo,Larrondo, Chile Laboratorio de Análisis Genómico (LAGEN), Escuela de Ciencias Biológicas, Universidad Nacional, Heredia, Costa Rica Laboratorio de Bioespeleología y Carcinología, División de Desarrollo Sustentable, Universidad de Quintana Roo, Quintana Roo, Mexico Universidad Nacional, Costa Rica Escuela de Ciencias Biológicas

7 páginas, 2 figuras , 3 tablas, 1 apéndice con 2 tablas., Most fisheries involving spiny lobsters of the genus Palinurus have been overexploited during the last decades, so there is a raising concern about management decisions for these valuable resources. A total of 13 microsatellite DNA loci recently developed in Palinurus elephas were assayed in order to assess genetic diversity levels in every known species of the genus. Microsatellite markers gave amplifications and showed polymorphism in all species, with gene diversity values varying from 0.650 6 0.077 SD (Palinurus barbarae) to 0.792 6 0.051 SD (Palinurus elephas). Most importantly, when depth distribution was taken into account, shallower water species consistently showed larger historical effective population sizes than their deeper-water counterparts. This could explain why deeper-water species are more sensitive to overfishing, and would indicate that overexploitation may have a larger impact on their longterm genetic diversity., This work was supported by a pre-doctoral fellowship awarded by the Autonomous Government of Catalonia to FP (2006FIC-00082). Research was funded by projects CGL2006-13423 and CTM2007-66635 from the Ministerio de Educacio´n y Ciencia. The authors are part of the research group 2009SGR-636, 2009SGR-655 and 2009SGR-1364 of the Generalitat de Catalunya. FP acknowledges an EU-Synthesys grant (GBTAF- 4474).

Structure of the mouthparts and foregut of Panulirus ornatus phyllosomata (stages I-VI hatchery reared, IX-X wild caught) were examined using scanning electron microscopy and histology to gain an understanding of ingestive and digestive processing mechanisms, to identify potential shifts in diet during development, and to suggest appropriate physical characteristics for the development of a suitable formulated diet for commercial aquaculture of this species such as size, texture, and buoyancy. Mouthpart and foregut structure indicates that P. ornatus phyllosomata are capable of ingesting zooplankton of any hardness during this life history stage and are only limited by their ability to capture and manipulate prey as the mandibular molars are well developed to masticate prey further. Mouthpart morphology changes little during development, however, the disposition of the mouthparts and size of the mouth aperture increases with each successive stage of development, suggesting a greater capacity to m...

The ability to culture larval lobsters is of paramount importance to the commercial development of effective aquaculture methods. Recently, we developed two separate laboratory culturing strategies that yielded the complete larval development from egg to puerulus (post-larva) for the commercially important and transpacific Pronghorn spiny lobster, Panulirus penicillatus (Olivier, 1791). Individual phyllosomal culture of 10 newly hatched animals was carried out in a static seawater system. Two of the 10 phyllosomata held at 24.5-26.0°C metamorphosed after 22 molts to the puerulus stage at 256 and 294 days respectively (final body lengths = 30.80 mm and 32.00 mm). Mass culture of 500 newly hatched phyllosomata was also carried out in two specialized acrylic flow-through seawater tanks. Of the 500 larval animals, 215 were randomly sampled and morphologically staged (10 distinct stages were observed and documented as well as two sub-stages). Seven phyllosomata that were mass cultured metamorphosed to...

15 pages,10 figures, 2 tables Puerulus and early juvenile stages of the spiny lobster Palinurus mauritanicus Gruvel, 1911 from the western Mediterranean are described and illustrated, and their morphological characteristics are compared with those described in other palinurids. This constitutes the first detailed morphological description of a puerulus of the genus Palinurus. The puerulus of P. mauritanicus presents a carapace already with some spines characteristic of the adult phase, which allows to differentiate it from the congeneric species Palinurus elephas (Fabricius, 1787) co-occurring in most of its distribution area. The puerulus of P. mauritanicus can be differentiated from those described for pueruli of the genera Panulirus and Jasus by the presence of very long, flagellated, and setose exopods on maxillipeds 2 and 3. The puerulus of P. mauritanicus resembles those described for Projasus by the presence of the crista dentata on the third maxilliped. The morphology and differential colouration of the early juvenile with respect to the adult are described. We wish to thank all participants in the cruises NERIT-2 on board R/V ‘‘García del Cid’’ and MEDITS_ES_2004 and 2005 on board R/V ‘‘Cornide de Saavedra’’, as well as Ferran Palero for helpful comments and discussion.

Stage IV phyllosoma larvae of the packhorse lobster, Jasus verreauxi (H. Milne Edwards, 1851), were observed while feeding to determine the processing ability and involvement of mouthparts. Phyllosomata were presented live brine shrimp, Artemia salina (Linnaeus, 1758), mussel flesh, Perna canaliculus (Gmelin, 1791), and jellyfish, Aurelia sp., and were given chemical or tactile stimulation to induce a feeding response. Larvae were observed tearing their food with maxillipeds 2 and 3, before maceration by maxillipeds 2 and maxillae 1, mastication by mandibles and labrum, and subsequent ingestion of finer particles via a suction action induced by foregut contractions. Phyllosomata became entangled in food material and did not feed when provided soft tissue, such as jellyfish and mussel gonad. Maxilliped action increased when larvae were provided with harder prey, which indicated greater energetic investment required to process the food. Our observations suggest that phyllosomata are capable of processing fleshier prey items than Artemia, which is traditionally used as food in artificial culture. An ideal food source for phyllosomata should not readily break up and foul the water, should be free of microbes, and should be energetically economical to capture and masticate.

The reproductive biology of Panulirus guttatus from northern Quintana Roo, Mexico, was studied as part of a large research program on the biology and ecology of this poorly studied species. Lobsters were caught with baited traps deployed on the coral reef throughout 17 months (May 1988-September 1989). A total of 870 lobsters (558 males and 312 females) were obtained. The strongly biased sex ratio may have been an artifact of the sampling method, but differential movements of males and females related to reproductive activity were not excluded as possible causes. Ovigerous females (40% of total females, i.e., 125) occurred throughout the year, but were more prevalent during the spring of 1988, and the spring and summer of 1989. The size range of ovigerous females was 40.0-73.5 mm in carapace length (CL). Brood size was determined for the 125 ovigerous females plus an additional 41 sampled during 1987. After excluding the data of females with eggs about to hatch (of smaller sizes than eggs in previous stages), seasonal log brood size-log CL relationships were examined. No significant differences were found either seasonally or interannually, indicating that the average brood size of females of equal CL remained fairly similar throughout the study period. Females between 55 and 65 mm CL had the highest indices of reproductive potential, and accounted for 54% of the total egg production of the population. These results were compared with others obtained from populations of P. guttatus in Florida and other parts of the Caribbean.

The gonadosomatic index (GSI), ratio of area of the lumen to that of the seminiferous tubules, and sperm density were analyzed against body size and season in male spiny lobsters, Panulirus japonicus, sampled off Oshima Island, Tokyo, Japan. For functionally mature individuals (≥54 mm carapace length), regressions of GSI and area of the lumen against body size were significant only before the spawning season. GSI showed no clear seasonal changes. Germinal cells at all stages of spermatogenesis were found in the testis, and sperm were present in the vas deferens throughout the year. However, seasonal changes in the 3 reproductive indices indicate that P. japonicus has a distinct seasonal pattern for spermatogenesis and storage of sperm. The ratio of the area of the lumen to that of the seminiferous tubules was highest before the spawning season. Sperm density during and 1 month after the spawning season (from June to September), however, was significantly lower than in other months.

Specimens of the puerulus stage of the rock lobster Jasus edwardsii were collected as they settled from the plankton. They were raised in a closed marine aquarium system. The molt cases were collected with each ecdysis during growth. The development of the Cuticular Articulated Peg (CAP) organs on the walking legs of sequential molts from the same animals was studied using scanning electron microscopy. The number of CAP sensilla in the organ increased with each molt. The size of individual CAP sensilla and the distance between them increased during growth, with the result that the length of the organ also increased. Sensilla could be added or lost at each molt. The number of sensilla in each organ increased steadily, because the rate of addition was greater than the rate of loss. Most new sensilla were added to the organ proximally and most losses occurred distally. New proximal sensilla appeared fully formed. As the animals grew, the distal sensilla became gradually smaller until finally they were lost by a process of resorbtion. This is the first example of development of a crustacean proprioceptor by ordered, sequential turnover of receptor elements. Since each sensillum is associated with a single neuron, the pattern of development has implications for the way in which the sensory nervous system associated with the receptors develops.

Reproduction of the tropical spiny lobsters Panulirus inflatus and P. gracilis was studied in Guerrero, Mexican Pacific coast, from February 1979 to May 1980, from samples with bottomset nets and from commercial divers' catches. Lobsters caught with nets were tagged, and some females were recaptured up to 4 times during the study period. The smallest ovigerous females measured 49.4 mm (P. gracilis) and 54.7 mm (P. inflatus) carapace length (CL). Reproductive females of both species were found throughout the year, but analysis of the percentage of reproductive females by size class showed that small lobsters ( 70 mm CL). Females of both species > 70 mm CL were capable of producing at least 3 or 4 broods per year, those from 60-70 mm CL up to 3, and smaller mature lobsters 1 or 2. In addition, females between 65 and 80 mm CL had the highest index of reproductive potential, due to their highest numbers in the population and their number of broods per year. Fishery implications of these findings are discussed.

The gross morphology and fine structure of the mouthparts and foregut of the rock lobster Jasus edwardsii were studied in its late larval and early juvenile stages. The stage-XI phyllosoma has third maxillipeds with specialized setae, second maxillipeds and first maxillae with strong setae, mandibles with well-developed molar and canine-like processes and teeth, and a foregut with well-developed grooves and brushes but no gastric teeth. The first-molt juvenile has third and second maxillipeds and first maxillae with rows or tufts of numerous strong setae, wellcalcified mandibles with strong molar and incisor processes, and a foregut with well-developed ossicles, grooves, brushes, and strong gastric teeth. In contrast, the puerulus has maxillipeds and maxillae with only small numbers of setae, uncalcified mandibles with a rudimentary incisor, and a foregut with a small number of ossicles and setae, and rudimentary lateral teeth. These observations suggest that the late-stage phyllosoma of J. edwardsii feeds primarily on large, soft materials and the early postpuerulus on hard, as well as soft, materials. In contrast, the puerulus probably does not feed during some or all of its existence, and any feeding is on small, soft materials. Use of nutrients stored in the digestive gland is one possible way that the puerulus could meet its energy requirements.

Relationships between relative growth of various body parts, gonadal maturation, and functional maturity were examined in male and female Panulirus japonicus by regression techniques and histological study of the ovary, testis, and vas deferens. Relative growth of the first pleopod and the first and second pereiopods against the carapace length (CL) is described best by two power regressions for both sexes, except for the male first and second pereiopods, which are described by three power regressions. Sexual differences in relative growth were observed when size exceeded 28-mm CL for the pleopod, and 52- and 54-mm CL for the first and second pereiopods, respectively. In all sizes less than these CLs, regressions between males and females were not significantly different from each other. For female pleopods, an inflection point was observed at 43-mm CL, whereas it occurred at 52- and 54-mm CL in male pereiopods. Females with immature ovaries often were less than 45-mm CL and their gonadosomatic indices were less than 1%, whereas all females greater than and including 45-mm CL had developing or ripe ovaries. In males, sperm were present in the testis and vas deferens of all individuals examined histologically (≧37-mm CL). In individuals greater than and including 54-mm CL, ratios of the diameter of the middle vas deferens to that of the posterior vas deferens were almost constant, and high sperm density was observed in the middle vas deferens, whereas in some individuals less than 54-mm CL sperm were absent in a part of the middle vas deferens and the ratio was low. These histologically observed values correspond well with the inflection points of the relative growth regressions for female pleopods (43-mm CL) and those in male pereiopods (52- and 54-mm CL). This indicates that the inflection points are closely related to gonadal maturation in females and functional maturity in males.

The species of the rare spiny lobster genus Palinustus A. Milne Edwards, 1880, are reviewed, based on a total of 54 specimens. Although the characteristics used previously to separate the species are rather variable, all 4 named species are nevertheless considered to be valid; with P. truncatus A. Milne Edwards, 1880, restricted to the western Atlantic, P. mossambicus Barnard, 1926, known only with certainty from the eastern coast of Africa, while both P. waguensis Kubo, 1963, and P. unicornutus Berry, 1979, are widely distributed in the Indo-West-Pacific. A new species, P. holthuisi, is found in Taiwan and Japan. The distinguishing characteristics of all the species are described in detail, together with line drawings and photographs. The different species have distinct coloration and color photographs of live or fresh specimens are provided where possible. A key to the species is given for their identification.

This study describes the sensory structures on the antennae and antennules of the puerulus and postpuerulus stage juveniles of the western rock lobster Panulirus cygnus. Both stages have short and long spines, which probably function in protection and in contact-detection. Both also possess setae structurally similar to those shown to be near-field vibration receptors in other crustaceans. Both have aesthetasc setae on the antennules and petaloid setae on the antennae, with both types of setae appearing to be chemosensory. The puerulus stage has an almost continuous linear series of pinnate setae that overlie each other and also the segmental boundaries along the antenna in such a way that the series is well positioned to act as a detector for vibration of the whole antenna. This arrangement is unique to the puerulus stage, which is also noted for its 40-60-km directional swim from the offshore waters to the coast. We suggest that the complex receptor system formed by the antennae and the pinnate setal system plays a role in navigation by allowing the animal to orient to vibrational clues associated with the coast.

Pueruli of spiny lobsters, Panulirus argus, observed at Manchoniel Bay, Carriacou, Grenada, swam only at night. Swimming was in a linear, straightforward manner within a few centimeters of the surface, at speeds up to 10 cm/s. Swimming, sinking, approach, and retreat postures are described. Pueruli did not detect motionless objects prior to contact, but reacted to turbulence caused by waves or current acting upon objects. Pueruli in the laboratory usually swam only within 3 h after sunset; swimming behavior was the same as in the field and confirmed the inability to detect motionless objects prior to contact. Individuals with more than 2 days of pigment development never swam. During daylight, pueruli buried themselves in sand or hid in algae or crevices. Pueruli seldom exhibited any attraction to light. Reliability of recruit monitoring with artificial floating habitats is discussed.