Apical organs in echinoderm larvae: insights into larval evolution in the Ambulacraria

The anatomy and cellular organization of serotonergic neurons in the echinoderm apical organ exhibits class-specific features in dipleurula-type (auricularia, bipinnaria) and pluteus-type (ophiopluteus, echinopluteus) larvae. The apical organ forms in association with anterior ciliary structures. Apical organs in dipleurula-type larvae are more similar to each other than to those in either of the pluteus forms. In asteroid bipinnaria and holothuroid auricularia the apical organ spans ciliary band sectors that traverse the anterior-most end of the larvae. The asteroid apical organ also has prominent bilateral ganglia that connect with an apical network of neurites. The simple apical organ of the auricularia is similar to that in the hemichordate tornaria larva. Apical organs in pluteus forms differ markedly. The echinopluteus apical organ is a single structure on the oral hood between the larval arms comprised of two groups of cells joined by a commissure and its cell bodies do not reside in the ciliary band. Ophioplutei have a pair of lateral ganglia associated with the ciliary band of larval arms that may be the ophiuroid apical organ. Comparative anatomy of the serotonergic nervous systems in the dipleurula-type larvae of the Ambulacraria (Echinodermata+Hemichordata) suggests that the apical organ of this deuterostome clade originated as a simple bilaterally symmetric nerve plexus spanning ciliary band sectors at the anterior end of the larva. From this structure, the apical organ has been independently modified in association with the evolution of class-specific larval forms.     

Development of the nervous system in the brittle star Amphipholis kochii

There are several studies of neural development in various echinoderms, but few on ophiuroids, which develop indirectly via the production of pluteus larvae, as do echinoids. To determine the extent of similarity of neuroanatomy and neural development in the ophiuroids with other echinoderm larvae, we investigated the development of the nervous system in the brittle star Amphipholis kochii (Echinodermata: Ophiuroidea) by immunohistochemistry. Immunoreactive cells first appeared bilaterally in the animal pole at the late gastrula stage, and there was little migration of the neural precursors during A. kochii ontogeny, as is also the case in echinoids and holothuroids. On the other hand, neural specification in the presumptive ciliary band near the base of the arms does occur in ophiuroid larvae and is a feature they share with echinoids and ophiuroids. The ophiopluteus larval nervous system is similar to that of auricularia larvae on the whole, including the lack of a fine network of neurites in the epidermis and the presence of neural connections across the oral epidermis. Ophioplutei possess a pair of bilateral apical organs that differ from those of echinoid echinoplutei in terms of relative position. They also possess coiled cilia, which may possess a sensory function, but in the same location as the serotonergic apical ganglia. These coiled cilia are thought to be a derived structure in pluteus-like larvae. Our results suggest that the neural specification in the animal plate in ophiuroids, holothuroids, and echinoids is a plesiomorphic feature of the Ambulacraria, whereas neural specification at the base of the larval arms may be a more derived state restricted to pluteus-like larvae.     

Development of the sand dollar <Emphasis Type="Italic">Scaphechinus mirabilis</Emphasis>

Under laboratory conditions, the development of larvae of the sand dollar Scaphechinus mirabilis (Agassiz) took 28.5–29 days from fertilization to settling and the end of metamorphosis at a temperature 20°C and salinity 32.2–32.6‰ The cleavage divisions were completed in 12 hours after fertilization (AF) by the release of free swimming ciliary blastula from the egg membrane. The larvae attained pluteus I stage with one pair of arms at an age of 40 hours. In 4.5–5 days the pluteus II stage with three pairs of arms, and in 9 days the pluteus III stage with four pairs of arms were formed. On the 20–21st day AF the larvae developed a vestibule, in which the adult skeleton rudiments, spicules, plates and pedicellariae were formed on the 26–27th day AF. The size of the larvae at an age of 22.5 days was 1064.3 ± 44.7 μm. The settling of larvae was recorded on the 28–29th day of development. Most of the larvae completed their metamorphosis in 4.5–5 hours after settling.     

Peptidergic and serotonergic immunoreactivity in the metamorphosing ophiopluteus of Ophiactis resiliens (Echinodermata, Ophiuroidea)

Abstract. Antibodies against the echinoderm-specific neuropeptide S1 and against 5HT were used to examine the fate of the larval nervous system during metamorphosis in the ophiuroid Ophiactis resiliens . In contrast to most echinoderms, the onset of peptidergic and serotonergic expression was delayed to the advanced ophiopluteus stage, in particular for 5HT. In advanced ophioplutei, peptidergic immunoreactivity was located in simple fibres associated with the ciliated bands, a stomach nerve ring, and cells along the antero-lateral arms. 5HT immunoreactivity was concentrated in 2 oral ganglia in the adoral projections, located at the posterior rim of the mouth. Clusters of 5HT-positive cells were also found along the antero-lateral arms. The ophiopluteus lacked a serotonergic (or peptidergic) anterior ganglion. In echinoids, holothuroids, and crinoids, anterior ganglia are thought to have a sensory role in settlement and metamorphosis. Given that ophioplutei metamorphose in the plankton and that larval structures degenerate before settlement, the absence of apical ganglia correlates with the lack of a functional role for larval structures in substrate selection and settlement. Although most of the larval nervous system degenerated during metamorphosis, the adoral projections and associated oral ganglia appeared to be incorporated into the juvenile mouth, suggesting a potential role for larval neurons in contributing to oral neuronal structures in the adult. S1-positive neurons and fibres in the rudiment developed de novo and in parallel with development of the epineural canal. This structure gives rise to the primordia of the adult circumoral nerve ring and radial nerves, indicating that differentiation of the adult nervous system begins in the early stages of metamorphosis.     

菜粉蝶幼虫、蛹和成虫的雌雄形态鉴别

利用体视显微镜观察、测量、记录菜粉蝶不同虫态雌雄个体间的形态特征差异。雌雄菜粉蝶幼虫、蛹和成虫均存在显著的雌雄二型特征,3龄之后雄性幼虫的第6腹节有一对肾型黑斑,雌性幼虫无此斑。雄蛹第9腹节有生殖孔和一条短纵裂缝,裂缝两侧有凹凸不平的半圆形瘤状突起;雌蛹腹部第8、9腹节分别有生殖孔和产卵孔,且两孔之间亦有一条纵裂缝。雌蝶前翅背面有一对上下排列的黑圆斑,其腹部末端外生殖器为圆筒型;雄蝶前翅背面一对黑斑相对较小,颜色较浅,尤其下斑为浅灰色,腹部末端具钳状外生殖器。本研究揭示了菜粉蝶幼虫、蛹及成虫不同虫态的雌雄二型差异,比较研究并提出了便于快速鉴别雌雄个体的典型特征和识别方法。     

The ectosymbiont crab Dissodactylus mellitae-sand dollar Mellita isometra relationship

The presence of the ectosymbiont Dissodactylus mellitae on Mellita isometra was studied at Tybee Island in Georgia. The number of crabs found on sand dollars, stage of maturity, sex, numbers of eggs, and size of eggs produced by crabs were noted. The test diameter of sand dollars, and the number and diameter of eggs produced were also noted. Infestation rates of crabs on sand dollars varied over time. The maximum number of crabs found on a sand dollar was 10. The three types of population dispersion, random, uniform, and clumped, were observed for crabs on sand dollars throughout the sampling period. Clumped or gregarious settling was observed when juvenile crabs were abundant, uniform and random distribution when mature crabs were abundant. Female crabs were significantly larger than male crabs, with carapace width of the largest mature female crab being 4.6 mm and the largest male 3.1 mm. Crabs produced between 80 and 300 eggs from 0.188 to 0.291 mm in diameter. Bigger crabs produced significantly more eggs than smaller crabs. Sand dollar sizes were 50-110 mm, with a mode between 60 and 70 mm. The effect of crab burden on egg production in the sand dollar was time-dependent. The presence of crabs on sand dollars correlated with total egg production of sand dollars in May, the peak of the spawning season, with sand dollars carrying one or two crabs having a lower egg production than those without crabs. Overall, variation in egg size was attributed to variation among females followed by variation between seasons and crab burden.     

Spawning behavior and early life history of the sharpnose puffer,Canthigaster rivulata,in the aquarium

Specimens ofCanthigaster rivulata (Temminck et Schlegel) were collected from Kominato and Hayama, central Japan, from May, 1985 to October, 1986. On the basis of the gonadosomatic index, gonadal histology and results of artificial fertilization of these specimens, the spawning season is considered to extend from late June to mid-September. The specimens exhibited the following dimorphic differences associated with sex: 1) The male is larger than the female. 2) Ventral side of the body is brownish orange in the male with vermiculated or reticulated patterns of bright violet, while it is white in the female. 3) The male has a well-developed skin fold along the mid-dorsal and mid-ventral lines, which is greatly elevated during courtship; whereas the female’s skin folds are not or slightly developed and conspicuous only during courtship. In an aquarium with the water temperatures of 22 to 26°C, a pair of fish spawned every four days late in the morning for three consecutive months. Courtship and spawning occurred in a pair. The male swam in front of the female, and elevated the skin folds both dorsally and ventrally, fully spreading the unpaired fins, with the ventral side of the body flashing bright blue and the dorsal side turning dark. Both fish swam in a circular fashion, elevating the skin folds. The male followed the female nudging her abdomen with his snout. Both fish turned upward, and released gametes. The eggs are spherical, 0.53–0.73 mm in diameter, demersal, adhesive, transparent, and pale yellowish orange in color, and contain a cross-shaped or asteroid cluster of oil globules. The egg membrane was thick and consisted of about 14 concentric layers. The incubation period ranged from 73.5 hours at 28.2–28.5°C to 145.0 hours at 22.1–22.4°C. The newly hatched larvae were 1.38–1.98 mm in total length (TL) with 84-11-13 = 19–21 myomeres. The yolk was absorbed when the larvae attained 1.49–2.22 mm TL, three days after hatching. The larvae were fed on oyster larvae, blue mussel larvae, sea-urchin larvae and rotifers, but all of them died in 16 days. During the embryonic and early larval stages, the only pigment cells that appeared on the body were the black chromatophores.     

Larval feeding structure plasticity during pre-feeding stages of echinoids: Not all species respond to the same cues

Much of the work on phenotypic plasticity has focused on inducible defenses. As a result, little is known about induced phenotypes that improve the acquisition of resources (i.e. inducible offenses). Feeding larvae of several marine invertebrate species, gastropods and echinoderms, have inducible offenses, and produce larger feeding structures when given less food. To better understand inducible offenses, I investigated when in development sea urchin and sand dollar larvae can first alter their feeding morphology in response to different concentrations of food. Food induced feeding structure changes in both sea urchin and sand dollar larvae before larvae were able to ingest food. This suggests that the nervous system and a regulator gene, orthopedia, play a mechanistic role. In addition, larvae of the two species, Strongylocentrotus purpuratus and Dendraster excentricus, responded to different cues. Pre-feeding larvae of both species developed relatively shorter arms when given algal cells (i.e. chemical and physical stimuli), whereas only pre-feeding larvae of D. excentricus developed shorter arms when exposed to algal exudates (i.e. chemical stimuli). Larvae of neither species responded morphologically to the presence of polystyrene beads (i.e. physical stimuli).     

Thyroid hormones determine developmental mode in sand dollars (Echinodermata: Echinoidea)

Evolutionary transitions in larval nutritional mode have occurred on numerous occasions independently in many marine invertebrate phyla. Although the evolutionary transition from feeding to nonfeeding development has received considerable attention through both experimental and theoretical studies, mechanisms underlying the change in life history remain poorly understood. Facultative feeding larvae (larvae that can feed but will complete metamorphosis without food) presumably represent an intermediate developmental mode between obligate feeding and nonfeeding. Here we show that an obligatorily feeding larva can be transformed into a facultative feeding larva when exposed to the thyroid hormone thyroxine. We report that larvae of the subtropical sand dollar Leodia sexiesperforata (Echinodermata: Echinoidea) completed metamorphosis without exogenous food when treated with thyroxine, whereas the starved controls (no thyroxine added) did not. Leodia sexiesperforata juveniles from the thyroxine treatment were viable after metamorphosis but were significantly smaller and contained less energy than sibling juveniles reared with exogenous food. In a second starvation experiment, using an L. sexiesperforata female whose eggs were substantially larger than in the first experiment (202+/-5 vs. 187+/-5 microm), a small percentage of starved L. sexiesperforata larvae completed metamorphosis in the absence of food. Still, thyroxine-treated larvae in this experiment completed metamorphosis faster and in much higher numbers than in the starved controls. Furthermore, starved larvae of the sand dollar Mellita tenuis, which developed from much smaller eggs (100+/-2 microm), did not complete metamorphosis either with or without excess thyroxine. Based on these data, and from recent experiments with other echinoids, we hypothesize that thyroxine plays a major role in echinoderm metamorphosis and the evolution of life history transitions in this group. We discuss our results in the context of current life history models for marine invertebrates, emphasizing the role of egg size, juvenile size, and endogenous hormone production for the evolution of nonfeeding larval development.     

A gastric-brooding asteroid, Smilasterias multipara

The gastric-brooding asterinid sea star, Smilasterias multipara, broods from late August to early November in the shallow sublittoral zone of southeastern Australia. We observed males and females spawning in the laboratory. They shed gametes through gonopores on the sides of the arms. The eggs were orange, about 1.0 mm in diameter, and heavier than seawater. They were externally fertilized by sperm, and placed into the stomach of the female by the tube feet. Twenty-four hours after fertilization, the first cleavage occurred. Cleavage was equal, total, and radial. Development via a non-feeding lecithotrophic brachiolaria was direct, there being no planktrotrophic bipinnaria or brachiolaria larva. Embryos developed, through wrinkled blastula and gastrula stages, into brachiolariae with arms. All of the surfaces of the brachiolaria were covered by cilia. At metamorphosis, a starfish rudiment appeared on the posterior portion of the larval body, while the anterior portion of the larval body was absorbed. Two months after fertilization, metamorphosis was complete. After metamorphosis, juveniles in the stomach grew six pairs of tube feet in each arm. Juveniles, 3 mm in diameter, emerged from the mouth of the mother in early November. Developmental evidence suggests that this asteroid has evolved mechanisms for the protection of larvae and juveniles from gastric digestion.     

Direct development of the brittle star Amphiodia occidentalis (Ophiuroidea, Amphiuridae) from the northeastern Pacific Ocean

Abstract. The highly modified development of the brittle star Amphiodia occidentalis is described from post-fertilization to the juvenile stage. Fertilized eggs are negatively buoyant, ∼190 μm in diameter, surrounded by a thick hyaline layer and a tough fertilization envelope. After gastrulation, embryos flatten into a bilaterally symmetrical disk with a U-shaped ridge surrounding an indented stomodeum on the oral surface. Internally, a ring of ∼22 calcitic ossicles grows at the edge of the disk. Vestigial ophiopluteal structures such as a ciliated band, paired larval spicules, or larval arms are not expressed during development. Although the fertilization envelope disintegrates on day 3, developmental stages remain immotile for five more days until they move with podia. At hatching, five hydrocoelic lobes are evident on the left side of the post-gastrula, and these migrate clockwise around the stomodeum, establishing pentamerous radial symmetry. Central and radial plates originate on the right side and migrate to a dorsocentral location as pentamerous symmetry is established. Development of the juvenile oral skeletal frame follows closely that described by Hendler (1978) for Amphioplus abditus except that A. occidentalis did not form buccal scales. The juvenile mouth opened by day 12. Fifty-five days after fertilization, juveniles had not added their first arm segments, although the first lateral arm plates had appeared. Developmental stages identical to those described here have been found in plankton tows taken in Oregon usually after storms that bring high waves. The unusual development of this species probably occurs in both benthic and pelagic environments.     

An ophiuroid trackway from the Lower Devonian Hunsrück Slate, Germany

A new trace fossil, Arcichnus saltatus , from the Lower Devonian Hunsrück Slate, Germany, is interpreted as the trackway of a protasterid ophiuroid, Taeniaster. The trackway consists of a series of horseshoe-shaped impressions, produced by a pair of arms, a feature characteristic of some ophiuroid trackways. This new trace fossil demonstrates that protasterids lived both infaunally and epifaunally.     

Geometrical resolution limits and detection mechanisms in the oral cavity

The objective of this work is to gain more insight into the processes of oral perception of food texture. Particularly, the limits for detectable thickness differences of objects, which are evaluated in the human mouth, are investigated. In a sensory study small, flexible circular disks (diameter in mm range) of varying thickness (in microm range) and material properties are evaluated between tongue and palate in human subjects. The thicker sample is identified in pair comparison tests. Experimental evidence suggests the existence of one detection process (attempt to align tongue and palate and the disk between them) to which the tongue-palate system reacts in two different ways: (1) by bending the disk (thickness below 125 microm, Young's modulus of 480 MPa) and (2) by impressing the disk into the tongue (thickness above approximately 200 microm, Young's modulus of 480 MPa), whereas the first reaction is necessarily followed by the second if the first one fails. For both ranges, differences in thickness of 25 microm can be detected. The two reaction processes cover isolated ranges and leave an insecure detection range in between them, for which neither one of the processes applies. Since deformation and load distribution on the disk are supposed to play a major role in the first detection process (the loads exerted on the disk in order to bend it are compared), we formulate a mathematical model to quantify these mechanical effects. The model is employed to identify parameter constellations (thickness, material properties) for which the insecure range is omitted or the range is enlarged. Theoretical findings are confirmed by further experiments. Their results are consistent with the characteristics and functioning of the mechanoreceptors in-mouth.     

Lecithotrophic development and metamorphosis in the Indo-West Pacific brittle star Ophiomastix venosa (Echinodermata: Ophiuroidea)

Summary

The larval development of the ophiocomid ophiuroid Ophiomastix venosais described using SEM. The gastrula transforms into a uniformly ciliated early larva which progressively changes into a lecithotrophic late premetamorphic larva with a continuous bilateral ciliated band. This stage is short-lived and equivalent to a highly reduced ophiopluteus. Comparisons between O. venosa and other ophiuroid species whose development has been investigated suggest that, whatever the developmental mode (lecithotrophic or planktotrophic), a pluteus stage always occurs in ophiuroids with planktonic development. Two metamorphic stages were identified, the late metamorphic larva differing from the early one by the closure of the larval mouth. The appearance of the permanent mouth marks the end of the metamorphosis. The postlarva still possesses remnants of larval features. The transformation of the reduced ophiopluteus into a barrel-shaped metamorphic larva with transverse ciliated bands, a vitellaria larva, is followed. The possible occurrence of a unique type of metamorphic larva in non-brooding ophiuroids is discussed. Verification of this, however, needs further SEM investigations on metamorphic larva from species having “regular” planktotrophic development.     

Development of eggs and larvae of the flounders Rhombosolea tapirina and Ammotretis rostratus (Pisces: Pleuronectidae)

The stages of development of laboratory-reared eggs and larvae of Rhombosolea tapirina and Ammotrelis rostratus are described. They both exhibited á typical pleuronectid pattern of development. At ambient seawater temperature of 11.1–13.8° C R. tapirina eggs hatched 83–93 h after fertilization and larvae metamorphosed c. 65 days later at mean length of 8.83 mm. Hatching of A. rostratus eggs occurred 93–105 h after fertilization and larvae metamorphosed after c. 69 days (mean length 11.21 mm) at 12.7–16.5° C. The two species can be readily separated by their morphology, meristics and pigmentation. In particular, the eggs differ in diameter, only R. tapirina larvae possess a pair of spines in the otic region, and juvenile A. rostratus have only a left pelvic fin.     

The larvae of Diadema setosum (Leske, 1778) (Camarodonta: Diadematidae) from South China Sea

Abstract

Diadema setosum (Leske, 1778) develops from small isolecithal eggs with a diameter of 84 ± 3 μm. Embryonic development took about 6.5–7 h and finished when a blastula left the fertilization envelope and became a larva. At this stage, the first pigment cells had appeared. At 23 h a prism developed; at 44 h a pluteus with one pair of arms had appeared; at 45 h of development plutei had two pairs of arms. The pigment cells colour the pluteus of D. setosum dark red. When 20-day-old larvae were mechanically stimulated, they flared their arms which may be defensive behaviour. During further development, the post oral arms of plutei grew to 1900 μm or more. Metamorphosis took place at about 40–45 days. At this time, five primary ambulacral podia were visible within the larval body. The duration of metamorphosis from the moment of larval settlement until the juvenile sea urchins began to move along the bottom was 40–60 min. The diameter of the test of the newly metamorphosed juvenile sea urchins was about 500 μm.     

Spawning,eggs and larvae of the hawkfish,Cirrhitichthys aureus,in an aquarium

Spawning of the hawkfish,Cirrhitichthys aureus, occurred repeatedly between a female and a male in the aquariums of the Marine Science Museum, Tokai University. The pair was collected from Suruga Bay by scuba diving and maintained for 50 days prior to spawning. In early September, 1979, the first successful spawning took place at 17:40 h after several repetitions of a courtship behavior sequence, which was initiated by the male. Spawning continued daily for 104 days between the two fish. Fertilized eggs were spherical, transparent and pelagic, and measured 0.75–0.78 mm in diameter. Hatching took place 19–22.5 hours after fertilization at 26.2–28.4°C. Newly hatched larvae, measuring 2.23–2.28 mm in total length, had a rather slender body with 12+18 = 30 myotomes and a large ellipsoid yolk sac. The front tip of the yolk sac protruded forward beyond the snout of the larvae. A single oil globule was situated in the front part of the yolk sac. The larvae were maintained for 4 days after fertilization. Larval characteristics ofC. aureus bore a close resemblance to those of other cirrhitid species especially in melanophore pigmentation along the dorsal and ventral surfaces.     

Below-ground host location by <Emphasis Type="Italic">Campsomeriella annulata</Emphasis> (Hymenoptera: Scoliidae), a parasitoid of scarabaeid grubs

Scoliid wasps are ectoparasitoids that attack soil-dwelling scarabaeid larvae, and little is known about their host-searching behavior. In this study we investigated the cues used in host location by Campsomeriella annulata (Fabricius) and examined whether or not these wasps can detect hosts in the soil from the surface. In a dual-choice test with a Y-tube, female wasps were attracted to sand with host odor, sand with host feces, and sand used for rearing the host, the larvae of Anomala rufocuprea Motschulsky (Coleoptera: Scarabaeidae). In a dual-choice test for cues presented at a distance, the wasps did not discriminate between the Y-tube arms with and without cues. In an experimental arena in which host products and a host grub were buried 0.5 cm below the surface the wasps did not respond to the cues from the surface in terms of the burrowing frequencies and antennal tapping rates. Our results indicate that C. annulata searches for the host grubs by using kairomones, residual cuticular substances, or feces deposited in the soil as the grubs move through it, and that wasps cannot perceive the host in the soil from the surface. We discuss how scoliid wasps search for soil-dwelling hosts using cues that are reliable but not highly detectable.     

The development of the enteropneust hemichordate Balanoglossus misakiensis KUWANO

We describe development from fertilization to metamorphosis of the enteropneust hemichordate Balanoglossus misakiensis. This is the first report to describe the complete development of an indirect-developing hemichordate under laboratory conditions. Mature adults were induced to spawn by shifting the temperature of seawater from 23 to 28 degrees C. Eggs (200 microm diameter) were enclosed within a non-mucilaginous membrane, and dispersed readily in seawater. After artificial insemination, a fertilization envelope was elevated from the egg surface beneath the egg membrane; this was followed by the formation of the first and second polar bodies within the envelope. Zygotes cleaved at 20-min intervals to form blastulae, and gastrulation started 9 h after fertilization. Embryos hatched 1 day after fertilization to become typical feeding tornaria larvae. The larvae metamorphosed 7-10 days after fertilization without undergoing the first (Müller) or forth (Krohn) stage of indirect-developing hemichordate development. Larvae that were not fed failed to metamorphose. Juveniles completed adult body formation within a week of settling in sand at the bottom of the culture tube. We discuss heterochronical modifications of B. misakiensis development, and make the case for this species as a potential model organism for the investigation of indirect-developing hemichordates.     

Short-term fluctuation in salinity promotes rapid larval development and metamorphosis in Dendraster excentricus

The effect of constant and fluctuating salinity on larval development and metamorphosis of the sand dollar Dendraster excentricus was investigated in the laboratory. Sand dollar larvae at different stages of development were kept either at 32‰ (controls), exposed to constant low salinity (22‰) throughout development, or exposed to fluctuating salinity (i.e. transferring larvae from 32‰ to 22‰ for 7 days then back to 32‰ for the rest of their development). Larvae exposed to constant low salinity were significantly smaller but developed all larval arms at a slower rate than larvae in all other treatments. Larvae exposed to fluctuating salinity recovered and developed significantly longer larval arms and bigger rudiments than larvae kept at constant low salinity. Larvae exposed to fluctuating salinity produced more juveniles than larvae at constant high salinity (32‰), while those at constant low salinity produced few or no juveniles. Four-arm larvae exposed to fluctuating salinity produced significantly more juveniles than six-arm larvae exposed to the same treatment. Transferring competent 8-arm larvae from 31‰ to 15‰ for 2 days then back to 31‰, induced metamorphosis with juvenile production being significantly higher than for those kept at a constant salinity of 20, 25 and 31‰. This study indicates that a short-term decrease in salinity might induce metamorphosis for this species.