Developmental Changes in the Mouthparts of Juvenile Caribbean Spiny Lobster, Panulirus Argus: Implications for Aquaculture

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Notice: ©2008 Elsevier B.V. The final published version of this manuscript is available at http://www.sciencedirect.com/science/journal/00448486. This article may be cited as: Cox, S. L., Jeffs, A. G., & Davis, M. (2008). Developmental changes in the mouthparts of juvenile Caribbean spiny lobster, Panulirus argus: Implications for aquaculture. Aquaculture, 283(1‐4), 168‐174. doi:10.1016/j.aquaculture.2008.07.019

Aquaculture 283 (2008) 168–174

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Aquaculture

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Developmental changes in the mouthparts of juvenile Caribbean spiny lobster, Panulirus argus: Implications for aquaculture

Serena L. Cox a,b,⁎, Andrew G. Jeffs c, Megan Davis a a Aquaculture Division, Harbor Branch Oceanographic Institute at Florida Atlantic University, 5600 US 1 North, Fort Pierce, Florida 34946, USA b National Institute of Water and Atmospheric Research, Private Bag 14901, Kilbirnie, Wellington 6241, New Zealand c Leigh Marine Laboratory, University of Auckland, P.O. Box 349, Warkworth 0941, New Zealand article info abstract

Article history: Light microscopy and video analysis were used to examine the mouthpart morphology and feeding Received 5 May 2008 behaviour of the Caribbean spiny lobster from puerulus (megalopal stage) (5–8 mm carapace length, CL) to Received in revised form 14 July 2008 adult (85 mm CL). Upon settlement the pueruli did not possess fully functional mouthparts, however, Accepted 14 July 2008 efficient feeding appendages appeared in the first instar juvenile (after the first moult from puerulus). From this stage the density, robustness and complexity of setation on the mouthparts, together with the size and Keywords: calcification of the mouthparts increased progressively with the size of the lobster. The changes correlated Mouthparts Spiny lobster well with previously observed ontogenetic changes in natural prey, from small and soft prey in early Panulirus argus juveniles to large and well-armoured molluscs, crustaceans and echinoderms in late juveniles and adults. The Feeding efficiency of shredding and tearing food items, mostly with the second maxillipeds, increased concomitantly Puerulus with lobster size, as did the ability of the mandibles to grind and process food. Based on morphological and Juvenile behavioural observations it is recommended that formulated feeds for first instar juvenile lobsters should be small (1–2 mm in diameter), soft and pulpy in texture to maximise feed intake. For large juvenile lobsters (45–80 mm CL), food items should increase to 5–10 mm in diameter and be of firmer consistency. Differences in feeding behaviour between spiny lobster species suggest that formulated diets developed to be efficient in one species may not be directly transferable to other species. © 2008 Elsevier B.V. All rights reserved.

1. Introduction inefficient handling by lobsters, and simple changes in pellet dimensions were found to reduce waste by as much as 19% (Sheppard The Caribbean spiny lobster, Panulirus argus, is an excellent et al., 2002). candidate for commercial aquaculture because of rapid growth and Investigations of species-specific feeding behaviour and functional straightforward husbandry (Lellis, 1991; Booth and Kittaka, 2000; Jeffs morphology, as well as digestive capabilities, over a range of life- and Davis, 2003). However, the development of a formulated feed for history stages is an effective means of identifying ontogenetic changes this spiny lobster species remains a significant impediment to its in dietary capabilities (Cox and Johnston, 2004). This approach has commercial aquaculture despite some initial research (Jeffs and Davis, been used successfully in a wide range of crustaceans and in many 2003; Williams, 2007). There is an underlying lack of knowledge of the instances leading to advances in the design and method of delivery of feeding capabilities and behaviour of spiny lobsters, including for the formulated feeds for aquaculture. This approach has already been used Caribbean lobster. Recent reviews of the nutrition of spiny lobsters in with considerable success for larval spiny lobsters (Cox and Johnston, culture concluded that understanding the feeding capabilities and the 2003), as well as slipper lobsters (Suthers and Anderson, 1981; corresponding format of formulated feeds are critical for advancing Johnston and Alexander, 1999), blue crabs (McConaugha, 2002), and the development of formulated feeds for spiny lobster aquaculture, by shrimp (Alexander and Hindley, 1985). While the feeding morphology maximising intake and reducing feed wastage (Nelson et al., 2006; and behaviour of adult spiny lobsters has been described (Mikami and Williams, 2007). For example, up to 50% of formulated feed pellets fed Takashima, 1994), it is not well known in the puerulus (megalopal to juveniles of the spiny lobster, Jasus edwardsii, were wasted through stage which is often incorrectly called the “postlarva”—Williamson, 1982) and the subsequent juvenile, which frequently undergo marked ontogenetic changes in habitat and diet (Butler et al., 2006). For example, the puerulus of the Caribbean lobster settles and remains ⁎ Corresponding author. National Institute of Water and Atmospheric Research, within dense inshore macroalgal beds. Once it has molted to the first Private Bag 14901, Kilbirnie, Wellington 6241, New Zealand. Tel.: +64 4 386 0300; fax: +64 4 386 2153. instar juvenile it begins to feed, mostly on small resident crustaceans E-mail address: [email protected] (S.L. Cox). and molluscs, before emerging at N15 mm carapace length (CL) to seek

0044-8486/$ – see front matter © 2008 Elsevier B.V. All rights reserved. doi:10.1016/j.aquaculture.2008.07.019 S.L. Cox et al. / Aquaculture 283 (2008) 168–174 169 shelter in nearby crevices and begin feeding locally on larger prey The oesophagus was not fully formed and the opening into the (Andrée, 1981; Marx and Herrnkind, 1985; Lalana et al., 1987). Larger oesophagus appeared to be covered by a fleshy membrane. The juveniles then move offshore, particularly around reefs, and are more overlying mandibles and maxillae were fleshy, soft structures which mobile, ranging across an array of habitats in search of a much wider did not move, even when physically stimulated. The mandibles were variety of prey (Cox et al., 1997; Briones-Fourzán et al., 2003). asymmetrical and simplified, with an unsegmented, laterally attached Therefore, the aim of this research was to describe the morphology mandibular palp which was partially covered posteriorly by the and functioning of the mouthparts in the Caribbean spiny lobster from paragnaths (Fig. 1A). The left mandible incisor process consisted of an puerulus to young adult to determine the changes in feeding undeveloped single, blunt tooth. The first maxillae were flattened, capabilities that could lead to improvements in the design of with a spinose exopod and epipod, and a rudimentary endopod, formulated feeds for aquaculture. together forming three flat lobes (Fig.1B). The second maxilla gave rise to an elongated scaphognathite-like epipod, divided into proximal and 2. Methods and materials distal endites, and a smaller lobe-like endopod (Fig. 1C). A fringe of setae was present along the distal edges of the exopod and a small tuft 2.1. Animal collection and husbandry of simple spinose setae on the proximal endite. The distal and proximal endites are much reduced in the first maxilliped, with the Approximately 100 newly settled P. argus pueruli were collected endopod fused to the elongated exopod (Fig. 1D). A distal flagellum from 10 Witham collectors (Witham et al., 1968) moored in surface was present on the anterior tip of the exopod. The epipod resembled a waters in the Florida Keys, following the new moon in March and April flattened and membranous plate. Setae were present along the distal 2005. Newly settled pueruli were sorted from more advanced margins of both the exopod and endopod. The second and third developmental stages on the basis of well-established staging criteria maxillipeds were more elongated compared with the maxilla or first (Lewis et al., 1952). The pueruli were transported in seawater to the maxilliped (Fig. 1E and F). The endopods consisted of five segments Harbor Branch Oceanographic Institute at Florida Atlantic University each. The exopod on the second maxilliped was elongated with four in Fort Pierce, Florida, and placed into holding tanks which included a segments, whereas the exopod on the third maxilliped was much selection of natural macroalgal (Caulerpa sp.) and artificial refuges. reduced and only had two segments. Setae were present along the The lobsters were fed frozen brine shrimp, Artemia salina and wild inner margin of the endopod dactyl and carpus of the third maxilliped. food items (i.e., worms, amphipods, and other various small Among the five individuals examined for feeding behaviour, there crustaceans, as well as frozen natural seafood items). Lobsters were was consistency in their mouthpart morphology and movement, held at 14:10 light–dark cycle, at 28±1 °C and salinity maintained at including the absence of any feeding behaviour. Close observations of 30±2‰. Ten transparent pueruli (b5 days after settling) were used for untethered pueruli in the tanks indicated that the lack of feeding mouthpart analysis immediately upon arrival at the laboratory, and behaviour was not a response to the experimental tethering remaining individuals were on-grown in the tanks until they treatment. subsequently reached the required developmental stage of early juveniles (10–15 mm CL), juvenile (15–44 mm CL), late juvenile–adult 3.2. Early juvenile (10–15 mm CL) (45–80 mm CL) for video analysis of feeding behaviour and mouthpart morphology. The relative position of the six pairs of mouthparts (mandibles, maxillae I and II, and maxillipeds I, II and III) as well as the labrum, 2.2. Mouthpart morphology and video analysis paired paragnaths, and the posterioventral fleshy lobe (membranous lobe), remained constant between the puerulus and all subsequent For each size range of lobsters, five individuals were tethered with juvenile stages, including early juveniles. The main changes between cyanoacrylic adhesive to fine wire (0.25 mm diameter), at the mid- the puerulus and first instar juveniles were the development of section of the dorsal surface of the cephalothorax and then placed into additional setae and spines, increased mouthpart robustness (i.e., a glass evaporating dish containing fresh seawater. Experimental calcification), an immediate transition to fully functional mouthparts, temperature was maintained at 28±1 °C and light levels were kept and almost doubling in the overall dimensions of the mouthparts. In constant at 2 lx. Feeding responses and mouthpart movements were the first instar juvenile, the mandibular palps had developed simple induced by placing frozen A. salina or fresh clam tissue (Mercenaria setae on the proximal tips and had become segmented. The incisor mercinaria) within the oral field of the tethered lobster. The process had also become more defined, bearing a more prominent mouthpart movements and function during food capture, manipula- groove in the molar process. The epipod, endopod and exopod of the tion and mastication were observed using an inverted Meiji EM28TR first maxilla all bear setae and spines (Fig. 2A). The distal and proximal compound microscope (Martin Microscope Company, USA) with fibre endites of the epipod of the second maxilla both bear setae. The optic illumination and Sony Digital Handycam, which recorded exopod and the endopod have a fringe of setae along the lateral edges. directly onto DVD for later analysis. After recording was completed, The first maxilliped is similar in structure to the puerulus, however, the five individual lobsters were euthanized by rapid cooling in an ice the distal flagellum is elongated and the interior edge of the endopod slurry and the mouthparts dissected out, photographed and then is flatter and broader. The major difference between the second and drawn. In the following descriptions, the terminology of Paterson third maxilliped in the early juveniles is the development of setation (1968) and Nishida et al. (1990) is used for the mouthparts and that of along the inner margin of the dactyl, propodus, merus and basis Factor (1978) is used to describe the setae. (Fig. 2B and C). Segmentation develops between the basis and coxa of the third maxilliped (Fig. 2C). The exopod is elongated and the distal 3. Results segment also bears setae. Among the five individuals examined, there was consistency in their mouthpart morphology with mouthparts 3.1. Puerulus (b9 mm CL) increasing in proportion to changes in body size. The mouth aperture was approximately 300 μm diameter, however, soft, fleshy food The mandibles, maxillae and maxillipeds of the puerulus were particles ranging from 1–2 mm and whole juvenile A. salina (up to small in size, lacked calcification, and had only a limited number of 2 mm total length) were observed being pushed between the setae. Microscopic examination of the mouthparts and the gut mandibles by the maxillae. Early juvenile lobsters were capable of confirmed that the puerulus stage is non-feeding by the inadequacy efficiently shredding fleshy clam tissue and adult A. salina into small of the mouthparts and the absence of any food in the gut. (1–3 mm) pieces. 170 S.L. Cox et al. / Aquaculture 283 (2008) 168–174

Fig. 1. Mouthparts of P. argus puerulus. (A) Mandibles. Scale 500 μm. (B) First maxilla. Scale 500 μm. (C) Second maxilla. Scale 1 mm (D) First maxilliped. Scale 500 μm. (E) Second maxilliped. Scale 500 μm. (F) Third maxilliped. Scale 1 mm. b, basis; c, carpus; co, coxa; d, dactyl; de, distal endite; en, endite; ep, epipod; ex, exopod; i, ishium; ip, incisor process; m, merus; mdp, mandibular palp; mp, molar process; p, propodus; pa, paragnaths; pe, proximal endite; pr, protopod; sc, scaphognathite.

3.3. Juvenile (15–44 mm CL) margin of the endopod, and on the distal tip of the exopod. The only major differences between the second and third maxilliped in the The mandibles and paragnaths showed little developmental early juveniles and the subsequent juveniles were the development of change compared with the early juvenile stage, however, the anterior pigmentation and slightly increased setation along the margins of the surfaces of the mandibles and the palps had developed pigmentation. five segments of the endopod. Among the five juveniles examined The incisor process has become more pointed and the molar process there was a proportional increase in mouthpart size in relation to body had low tubercles located along the internal edge. The first and second size, and general increase in setation and robustness of mouthparts. maxilla were almost identical to those observed in early juveniles, Juvenile lobsters were capable of shredding fleshy prey items into although there was increased setation around the anterior margins of small pieces (approximately 2–3 mm) prior to ingestion. Whole adult the exopods and epipods, including both the distal and proximal A. salina (up to approximately 5 mm) were observed being ingested, endites. The first maxilliped of juveniles was also similar to the early indicating the maxillae and mandibles are able to shred and process juveniles, however, there was increased setation on the interior the tough external carapace of these crustacean prey. S.L. Cox et al. / Aquaculture 283 (2008) 168–174 171

Fig. 2. P. argus early juvenile mouthparts. (A) First maxilla. (B) Second maxilliped (C) Third maxilliped. All scale bars 1 mm. am, attachment muscle; b, basis; c, carpus; co, coxa; d, dactyl; en, endite; ep, epipod; ex, exopod; i, ishium; m, merus; p, propodus; pr, protopod.

3.4. Late juvenile–adult (45–80 mm CL) larger, more robust and pigmented, compared with earlier developmental stages. The five individuals examined were capable of The asymmetrical mandibles in the late juveniles were more efficiently shredding and tearing large pieces of clam tissue (up to calcified, with tubercles along the interior edge of the molar process 10 mm), whole adult A. salina (up to 6–7 mm), and various other prey and a notably more defined incisor process on the opposing left items (i.e., worms, crustaceans) into smaller pieces (3–5 mm) prior to mandible (Fig. 3A). A deep notch separated this incisor process from ingestion. the molar process. The mandibles were capable of easily crushing the tough chitinous exoskeletons of large adult A. salina and various 3.5. Feeding behaviour similar-sized crustaceans (5–9 mm), once the maxillae and maxillipeds had shredded them into smaller pieces. The mandibular palp Detailed examination of the video recordings revealed that all had developed from two into three segments, with coarse setae juvenile lobster stages, except the puerulus, exhibited similar feeding covering the distal segment. Although larger, the first maxilla is behaviour and consistently utilised the paired mouthparts in a similar similar in structure and degree of setation to the same structures in manner for manipulating food. Food items were speared in a grabbing the earlier developmental stages. The scaphognathite of the second motion using the terminal dactyls of pereiopods 1 and 2, and passed maxillae was larger in the late juveniles, but essentially the same toward maxilliped 3. The third maxillipeds typically grasped food shape. The endopod was enlarged and setose along the anterior and items and manipulated the food inwards, towards the anterior second lateral edges (Fig. 3B). The distal and proximal endites had coarser maxillipeds. When not holding food items, the third maxillipeds setae along the entire length. The first maxilliped of the late juvenile is swept back and forth across the surface of the mouth region, over an larger than in the juvenile, but the general shape and structure arc of approximately 90°, articulating at the segment between the remains similar. The exopods in both the second and third maxillipeds basis and coxa. Pereiopods 1, 2 and 3 were also observed sweeping in were elongated and both had simple and plumose setae (Factor, 1978) this arc, in synchrony with the third maxillipeds. The exopod either (Fig. 3C and D). The five segments of the endopods were considerably moved dependently with the endopod, or it was observed beating 172 S.L. Cox et al. / Aquaculture 283 (2008) 168–174

Fig. 3. P. argus late juvenile mouthparts. (A) Mandibles (B) Second maxilla (C) Second maxilliped (D) Third maxilliped. All scale bars 1 mm. b, basis; c, carpus; co, coxa; d, dactyl; de, distal endite; i, ishium; ip, incisor process; m, merus; mdp, mandibular palp; p, propodus; pa, paragnaths; pe, proximal endite; pr, protopod; sc, scaphognathite.

sporadically with inconsistent frequency when the lobster was chamber with seawater. The dense setae which line the edge of the feeding. The third maxillipeds also played an important grooming second maxillae would, together with the two endites pointing into role, removing debris from the oral region by sweeping the surface of the mouth, appear to help guide food between the mandibles. the mandibles, maxillae, and first and second maxillipeds. The However, the setose nature of these mouthparts makes them well exopods of the second maxillipeds were very active during feeding, suited for filtration or helping to handle small food items such as beating continuously which appeared to facilitate the movement of crumbly invertebrate gonadal tissue. The first maxillae were ventral to food particles toward the mouth. The endopod of the second the mandibles, but obscured by the second maxillae and the maxillipeds received food items from the pereiopods or maxilliped maxillipeds. However, the second maxillae bear setae as well as 3, and when food items were within close proximity to the mouth prominent, robust spines along the medial edge of the exopods, these endopods were observed moving frequently, sweeping back and endopods and epipods. These spines appeared to assist in probing for forth across the surface of the mandibles and first maxillipeds. The food, holding the food items between the mandibles during each bite, endopods were also observed being held together in a medial position, and were also possibly used for pushing unwanted food particles preventing food items from being lost anterioventrally whilst the food anteriorly where they could be expelled. The mandibles operated was being manipulated by the first maxillipeds. The second along a lateral plane, using the pointed incisor process of the left maxillipeds also adjusted the location of the food between the mandible to shear against the calcified, tubercled surface of the right mandibles, assisting in the shredding and maceration of larger food mandible. The cutting and crushing action of the mandibles effectively items. In addition, the endopods of the second maxillipeds were in processed pieces of food into smaller, masticated pieces that were soft close contact with the mandibular palps. The first maxillipeds played a enough to be pushed into the mouth by the maxillae. The mandibular similar role to the second and third by assisting in guiding food items palps were located distally on each mandible, and were only observed between the mandibles with lateromedial movements. The first making small movements, independent of the mandibles. The setae maxillipeds also prevented food particles being lost anterioventrally. on the distal segment of the palps made contact with the food during The second and first maxillae were not visible on the video as they feeding, however, they did not appear to manipulate the food items. It were obscured by the maxillipeds. The second maxillae were is possible they function as a barrier, preventing food from escaping restricted to movements in the lateromedial plane between the first anteriorly. In addition, the setae on the palps may also have a maxillae and first maxillipeds, but allowing the scaphognathite to chemosensory role, assessing food palatability prior to ingestion. The move dorsoventrally and possibly associated with flushing the gill paragnaths or labrum were not observed to move during feeding. It S.L. Cox et al. / Aquaculture 283 (2008) 168–174 173 appears food particles were pushed into the oesophagus by the (Garm, 2004), are well suited to the handling and opening of large maxillae, after shredding, softening and crushing by the mandibles. It prey with strong shells, such as top-shells (Trochidae) and moon snails is possible peristaltic foregut contractions assist in movement of food (Polinices spp.) which can make up a significant proportion of the into the oesophagus and through into the gut. natural diet of these larger lobsters (Cox et al., 1997; Briones-Fourzán Late juvenile lobsters were capable of handling and ingesting et al., 2003; Garm, 2004). larger prey items than the juvenile or early juvenile lobsters. The Considerable effort is currently being directed toward the devel- mouthparts efficiently shredded prey into 5 mm diameter (on opment of appropriate formulated feeds for spiny lobster aquaculture. average) pieces in late juveniles, whereas juvenile lobsters tended However, early key requirements for designing an effective formu- to tear and shred food into smaller (average 3 mm diameter) pieces, lated feed are ensuring the format of the diet maximises food intake, and smaller again in early juvenile lobsters (1–2mmpieces).The and minimises wastage through leaching and fragmentation. For- ability to handle and ingest larger prey items was reflected in the mulated diet format includes parameters such as size, shape, texture processing ability of the mouthparts, due to increasing setation, and hardness, which may also need to be altered to better meet robustness and calcification, as well as the increasing dimensions of developmental changes in a target aquaculture organism. Relatively the mouthparts observed over the sequential stages of juvenile little research to date has addressed this concern specifically in spiny lobster development. lobsters. Sheppard et al. (2002) found that the intake of pellets of a All stages of juvenile lobsters examined were untidy eaters, with a formulated diet by juveniles of the lobster J. edwardsii could be significant portion of shredded food material being lost laterally, increased as much as 19% with small changes in the size of pellets to especially where the food material was crumbly, such as with ripe match the feeding abilities of the lobsters. Furthermore, early gondal material from clams. Larger prey items also tended to tangle in juveniles (25–30 mm CL) had difficulty handling and feeding on the pereiopods and third maxillipeds, particularly in the late juvenile– experimental pellets which resulted in high levels of wastage (up to adult stage. 50%) due to fragmentation of the formulated diet. Our observations of the feeding of P. argus from puerulus to adult 4. Discussion indicated marked morphological and behavioural changes in the food handling and processing ability of the mouthparts. These The puerulus stage of all spiny lobster species is widely considered changes must be considered in the development of a formulated diet to be a non-feeding, transitional stage between pelagic and benthic because size, shape, texture and consistency of the diet need to lifestyles (Nishida et al., 1990; Phillips et al., 2006). The incomplete reflect the physical characteristics and capabilities of different development of the mouthparts, a fleshy obstruction at the entrance juvenile stages. This study indicates that food does not need to to the oesophagus, the absence of food in the gut, and the absence of be provided to the non-feeding puerulus stage of P. argus. However, any feeding behaviour confirms earlier speculation that the puerulus the feeding behaviour and mouthpart structure of early juveniles of P. argus is also non-feeding (Wolfe and Felgenhauer, 1991). In the (10–15 mm CL) suggests that small (1–2 mm) and elongated food subsequent first instar juvenile, the feeding appendages show items that are soft and pulpy would be ideal. For the subsequent significant development, almost doubling in size and setation from juveniles, late juveniles and adults, food items that are increasingly the puerulus. The mouth was only 300 μm in diameter, but larger soft, larger in size, while being firmer, and fleshier will assist in improving fleshy food particles of up to 2 mm in diameter were handled and artificial diet intake. The co-ordinated development of teeth on the consumed. This is entirely consistent with the size and texture of the mandibles and spines/setae on the maxillipeds indicates that P. argus majority of the prey items found in the stomachs of juvenile lobsters in lobsters are able to deal successfully with more substantial food in the wild (Marx and Herrnkind, 1985) and in agreement with the benthic environment (Factor, 1989). Juveniles and late juveniles observations of the mouthparts from other juvenile Palinurids (N15 mm CL) were observed to be capable of processing a wide (Nishida et al., 1990). Beds of the macroalgae, Laurencia spp., into variety of prey, indicating that the size of a formulated feed item may which the lobster pueruli settle and subsequently dwell as early not be important. However, if small fragments were generated when juveniles, contain very large numbers of small invertebrates (mostly larger food items were shredded by the mouthparts, these fragments less than 5 mm long) such as isopods, amphipods, polychaetes, were most frequently not captured and ingested by the lobster. For a gastropods, copepods, ostracods and ophiuroids (Marx and Herrnkind, formulated diet to be effective it will need to avoid this potential 1985). These small crustaceans, worms, molluscs and echinoderms wastage by ensuring an appropriate combination of diet shape and can be readily consumed by early juvenile lobsters because this small consistency to reduce fragmentation. prey can be easily handled and the relatively unprotected soft bodies Observations of the behaviour of juveniles of a range of sizes in the pulled apart in the lobster mouthparts in the same manner as A. salina lobster J. edwardsii feeding on prepared pellets indicated differences were in our experimental observations. In contrast, juvenile lobsters in the handling and processing of food items by the mouthparts when (15–44 mm CL) no longer remain within the macroalgal beds, but compared to our results from P. argus (Sheppard et al., 2002). These occupy crevice shelters in shallow water habitats and forage in the differences are likely to be due to dissimilarities in the ecology and immediate vicinity. The diet of these juvenile lobsters mostly consists diet of juveniles of these two species (Booth, 2006). This being the of gastropods, bivalves, chitons, arthropods and echinoderms (Andrée, case, formulated diets developed to be effective in one species of spiny 1981; Herrera et al., 1991; Cox et al., 1997; Briones-Fourzán et al., lobster may not be directly transferable to other species. 2003; Nizinski, 2007). These prey items are mostly well protected with hardened calcareous exoskeletons. The ability of juvenile lobsters Acknowledgements to overcome the exterior defences of these prey is facilitated by a marked increase in the size, strength, complexity and hardening of the The authors wish to thank the staff at the Florida Fish and Wildlife mouthparts. The increasing ability of progressively larger juvenile Conservation Commission and the Keys Marine Laboratory, as well as lobsters to feed effectively on well-armoured molluscan, arthropod Jerry Corsaut, Harbor Branch Oceanographic Institute at Florida and echinoderm prey has been observed in other species of spiny Atlantic University (HBOI) for the collection of lobsters. Jackie Aronson lobsters (Griffiths and Seiderer, 1980; James and Tong, 1998). The (HBOI volunteer) and Erika McKay (NIWA) kindly prepared illustra- overall trend in the development of the mouthparts is continued with tions of the lobster mouthparts. The draft manuscript benefited from late juvenile to adult lobsters (45–80 mm CL) which forage across a helpful comments from an anonymous referee. This work was funded wide range of habitats in more exposed waters closer to the outer reef by a HBOI Postdoctoral Fellowship. This is HBOI Contribution number margins. The mouthparts of these lobsters, as well as larger adults 1705. 174 S.L. Cox et al. / Aquaculture 283 (2008) 168–174

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