Reproduction of the commercial sea cucumber Holothuria scabra (Echinodermata: Holothuroidea) in the Solomon Islands

Over a 3-year period (1996-1998), reproduction of the commercial sea cucumber Holothuria scabra (Jaeger, 1833) was investigated in the Solomon Islands to determine the spawning pattern and whether gametogenesis is continuous or seasonal. The gonad consisted of a single cohort of tubules that developed uniformly. Macroscopic examination of the gonads revealed that mature gametes were present throughout the year. Individuals with gonads at different stages of maturity were present in most samples. Partly spawned gonads were prevalent in females, whereas mature gonads were prevalent in males. The time at which the peak gonad index was recorded differed among years. Although gametogenesis was continuous, with a potential for prolonged gamete release, a period of enhanced spawning occurred during the dry season, from September to December. Maximum gonad indices were reached prior to and during this period of enhanced spawning. Histology revealed that gametogenesis reinitiated in partly spawned gonads, resulting in the presence of gametes at different stages of development in the gonad. The uniform growth of gonad tubules indicated that H. scabra does not conform to the progressive tubule recruitment model described for holothurian oogenesis. Continuous reproduction in H. scabra and prolonged availability of mature gametes would facilitate use of this species for aquaculture.     

Reproductive cycles of tropical sea cucumbers (Echinodermata: Holothuroidea) in southern Taiwan

Reproductive cycles of Afrocucumis africana (Dendrochirotida), Actinopyga echinites, Holothuria leucospilota, H. cinerascens, H. difficilis (Aspidochirotida), Synaptamaculata, Opheodesoma grisea, Patinapta taiwaniensis and Polycheira rufescens (Apodida), representing three orders of intertidal holothurians in southern Taiwan, were determined by gonad index and histological examination from March 1990 through September 1991. All nine species of holothurians have annual spawning periods lasting 2 to 4 mo in spring or summer. The early spawning of H. cinerascens and P. rufescens (suspension feeders) suggests that their feeding mode is related to food resources and may influence the reproductive period. In the direct-developing dendrochirotid (suspension feeder), gonad development occurs in cool months, and gametes are released at the beginning of warm months. In indirect-developing aspidochirotid and apodid (deposit feeder), gamete release occurs in late spring or summer and appears to be correlated with summer phytoplankton growth. The holothurians in the present study display spawning periodicity and duration similar to those of temperate species. Four species of holothurians lengthened spawning season with decreased latitude.     

L-Lactate dehydrogenase in the longitudinal muscle of the sea cucumber Sclerodactyla briareus (Echinodermata: Holothuroidea)

The infaunal holothurian Sclerodactyla briareus (Thyone briareus) is able to tolerate exposure to hypoxic conditions for over 2 days. Since the in vitro anaerobic degradation of glucose-U-C¹⁴ by longitudinal muscle preparations leads to an accumulation of labeled lactic acid, it is apparent that lactate dehydrogenase plays a key role during anoxia. Disc electrophoresis resolved one major band of lactate dehydrogenase (LDH) activity in longitudinal muscle extracts. Muscle extracts of s. briareus oxidized L-lactate only, as is the case with most other echinoderms. The apparent Michaelis constants for lactate oxidation and pyruvate reduction are dissimilar from values reported for other marine invertebrates. The LDH activity in this holothurian is higher than the activities reported for other marine invertebrates with the exception of some active, large arthropods. Substrate inhibition of LDH maximal activity occurs at substrate concentrations (pyruvate) exceeding 10 mM, and then to only a minor extent. The characteristics of lactate dehydrogenase from the longitudinal muscle of S. briareus support the hypothesis that it functions to maintain glycolytic flux during prolonged periods of exposure to anoxic or hypoxic conditions.     

Reproductive cycle of the sea urchin Holopneustes purpurascens (Temnopleuridae: Echinodermata)

This study documents the annual reproductive cycle of the Temnopleurid echinoid Holopneustes purpurascens near Sydney. H. purpurascens reached sexual maturity between 11 and 26 mm diameter and was gonochoristic. The sequence of oogenesis was similar to previously described echinoids. However, the proportion of eggs at each stage showed no temporal pattern in abundance, suggesting that vitellogenesis is continuous throughout the year. Spermatogenesis was also similar to other echinoids and showed a significant increase in the proportion of partially spawned testes in spring, followed by a similar increase in the proportion of partially spawned testes in late spring and summer. Gonad indices showed a peak in weight from late winter to mid-spring (9.10ǃ.47 for females; 6.28ǂ.43 for males; mean-SE), then a decline for both sexes. Although spawning was variable over time, there was a peak in the proportion of spawning from late winter to early summer that correlated with the observed peaks in gonad indices and gametogenic cycles. The demographic implications of this reproductive pattern are that new recruits can enter the population through synchronous spawning during the breeding period, and smaller haphazard spawning events at other times of the year.     

Spawning behaviors of northeast Pacific sea cucumbers (Holothuroidea: Echinodermata)

Spawning behaviors observed from May 1981 through August 1983 in the laboratory and field are described and compared for twelve species of northeast Pacific holothuroids collected from waters of the San Juan Archipelago, Washington. For many species, elevation of the anterior end, stretching of the introvert, tentacle waving, and swaying are behaviors commonly associated with gamete release. Less common activities include waving of introvert podia, waves of body wall contractions, errant males, aggregations, and pseudocopulation. Positively buoyant eggs of four species [Psolus chitonoides Clark, Cucumaria miniata (Brandt), C. piperata (Stimpson), and C. fallax (Ludwig)] were packaged in pellets, ropes, or strands and released passively; a female Molpadia intermedia (Ludwig) discharged negatively buoyant eggs in a single explosive burst. Two externally brooding sea cucumbers [C. lubrica (Clark) and C. pseudocurata Deichmann] produced strands of dilution-resistant sperm that were carried by currents to nearby females. Three species commonly sympatric [P. chitonoides, C. miniata, and Eupentacta quinquesemita (Selenka)] in high-current rocky habitats have overlapping spring breeding seasons, but spawn at different hours of the day.Please address all requests for reprints to Dr. McEuen at the Harbor Branch Oceanographic Institution     

Reproduction and growth of Holothuria atra (Echinodermata: Holothuroidea) at two contrasting sites in southern Taiwan

Reproductive periods and growth of two populations of Holothuria atra Jaeger distinctly different in body size at two sites of southern Taiwan were determined. Individuals examined in the present study were collected between march 1990 and February 1992. At Nanwan (21° 57N, 120° 45E), large individuals (351 to 1400 g wet wt) spawned from June to September. At Wanlitung (22°N, 120° 42E), a small proportion of frequently dividing individuals (<190 g wet wt) have mature gonads in May, June and September, but histological examination revealed no sign of spawning. Sexual recruits, defined as small individuals <5 g wet wt without sign of regeneration, were not found at either site during this 2-yr study. After the peak of fission at Wanlitung, 40% of the population showed signs of external regeneration. At Nanwan, small individuals transferred from Wanlitung grew from 6 g (n=6) to 166±8 g within 8 mo, and from 48±4 g (n=50) to 324±16 g within 1 yr, with a 6.8-fold biomass increase in 1 yr. At Wanlitung, the monthly average body weight of H. atra is between 33 and 62 g, apparently due to frequent fission, and the biomass increased only 2.9-fold in 1 yr. In southern Taiwan, sexual reproduction of H. atra occurs in large individuals. Asexual reproduction, occurring in small individuals, is the chief mechanism for population maintenance and increase, but it may decrease sexual reproductive potential.     

Reproductive biology of the commercial sea cucumber Holothuria fuscogilva in the Solomon Islands

 Reproduction of Holothuria fuscogilva (Selenka, 1867) in the Solomon Islands was investigated over a 4 yr period (1994 to 1998) by macroscopic and microscopic examination of the gonad tubules, the gonad index (GI) method, histological examination of gametogenesis, and spawning-induction trials. The gonad consisted of numerous tubules that dominated the coelom of gravid specimens. New tubules appeared in March, and grew in size and extent of branching until they reached their maximum size and maturity in August. Spawning occurred from August to October, with the majority of gametes released during October, although it was only partial in many individuals. After spawning, the tubules appeared wrinkled and resorbed into the gonad basis. A five-stage gonad maturity scale based on the macroscopic appearance of the gonad tubules corresponded with discrete stages of gametogenesis identified by histology. Gametogenesis was initiated in mid-March, with oogenesis and spermatogenesis occurring in parallel, followed by the growing stage (May to July) which was marked by active gamete development. Successful induction of spawning during the breeding period corroborated the GI and histological data. The uniform growth of gonad tubules indicated that H. fuscogilva in the Solomon Islands does not conform to the progressive tubule recruitment model described for other holothurians. An important application of this study is that the appearance of gonad tubules, removed by biopsy, can be used to determine the gonad condition of wild adults or captive broodstock. Received: 28 June 1999 / Accepted: 8 March 2000     

Investigations on brood protection inPsolus dubiosus (Echinodermata: Holothuroidea) from Antarctica in spring and autumn

Brood protection in the holothurianPsolus dubiosus Ludwig and Heding, 1935 was investigated in the Antarctic zone of permanent packice, an environment with an extreme seasonality in planktonic activity. Yearly reproductive cycles on the continental shelf of this area have been poorly investigated. Brooded juveniles of this species were found for the first time between the retracted tentacles of adult females at two different localities in the southeastern Weddell Sea, sampled in spring (October 1986) and autumn (February 1984). Mean lengths of the juveniles of each brooding female showed significant differences within each sample. However, there was no detectable difference in size of the juveniles between the two seasons. The results are discussed as an uncoupling of the dynamics of the early life history of these benthic invertebrates in space and time from processes in the euphotic layer of the Southern Ocean. The species investigated follows the general trend of some Antarctic invertebrates towards a non-pelagic development.     

Reproduction of the commercial sea cucumber Holothuria whitmaei [Holothuroidea: Aspidochirotida] in the Indian and Pacific Ocean regions of Australia

Seasonal trends in the gonad index (GI) of two widely separated populations of black teatfish, Holothuria whitmaei (formerly included in Holothuria nobilis), were investigated between Pacific (Great Barrier Reef) and Indian Ocean (Ningaloo Reef) coral reefs of Australia. Reproductive activity followed a similar annual trend, with the GI of both populations peaking typically between April and June. Macroscopic and histological analysis of Ningaloo Reef specimens revealed that large germinal tubules, positioned centrally on the gonad basis, progressed through four maturity stages: growing (II); mature (III); partly spawned (IV); and spent (V). Growing tubules dominated the central region of the gonad basis between January and March, followed by an increase in the number of mature tubules throughout the GI peak (April–June). The progressive appearance of partly spawned and spent tubules between June and October suggests that spawning in H. whitmaei continues intermittently over an extended period throughout the austral winter. The examination of the gonad structure of sexually mature male and female specimens identified five tubule size classes in total (C₁ to C₅), each of differing physical and gametogenic status. In females, smaller C₁ tubules located at the anterior edge of the gonad basis contained pre- to early vitellogenic oocytes. Larger C₂ and C₃ tubule cohorts, positioned centrally on the gonad basis, contained mid- to late-stage vitellogenic oocytes. Smaller C₄ and C₅ tubules, located at the posterior edge, contained only relict oocytes. Similar physical and gametogenic differences were apparent between tubule cohorts in male specimens. We propose that these results, together with evidence of incomplete gonad resorption over the austral summer, indicate that gonad development in H. whitmaei conforms to the predictions of the Tubule Recruitment Model (TRM). The TRM is reported rarely among tropical aspidochirotes, and results presented here (1) provide the first direct evidence of this model in H. whitmaei, and (2) confirm that this species is one of the few winter-spawning tropical invertebrates.     

Functional effects of the hadal sea cucumber <Emphasis Type=Italic>Elpidia atakama</Emphasis> (Echinodermata: Holothuroidea,Elasipodida) reflect small-scale patterns of resource availability

Holothuroidea represent the dominant benthic megafauna in hadal trenches (~6,000–11,000 m), but little is known about their behaviour and functional role at such depths. Using a time-lapse camera at 8,074 m in the Peru–Chile Trench (SE Pacific Ocean), we provide the first in situ observations of locomotory activity for the elasipodid holothurian Elpidia atakama Belyaev in Shirshov Inst Oceanol 92:326–367, (1971). Time-lapse sequences reveal ‘run and mill’ behaviour whereby bouts of feeding activity are interspersed by periods of locomotion. Over the total observation period (20 h 25 min), we observed a mean (±SD) locomotion speed of 7.0 ± 5.7 BL h⁻¹, but this increased to 10.9 ± 7.2 BL h⁻¹ during active relocation and reduced to 4.8 ± 2.9 BL h⁻¹ during feeding. These observations show E. atakama translocates and processes sediment at rates comparable to shallower species despite extreme hydrostatic pressure and remoteness from surface-derived food.     

Development and metamorphosis of two psolid sea cucumbers,Psolus chitonoides andPsolidium bullatum,with a review of reproductive patterns in the family Psolidae (Holothuroidea: Echinodermata)

Reproduction and development of two psolid sea cucumbers in waters of the San Juan Archipelago, Washington, were studied from April 1981 through August 1984. Spawning inPsolus chitonoides Clark andPsolidium bullatum Ohshima in the laboratory and in the field waters of the San Juan Archipelago, Washington, occurred from February through may, with heaviest spawning in April and May. The egg ofPsolus chitonoides was bright red in color and 627µm in diameter;Psolidium bullatum released eggs that were golden yellow in color and 330µm in diameter. Maximum measured fecundities (spawned eggs) forPsolus chitonoides andPsolidium bullatum were 34 700 and 3 074, respectively. Developmental schedules were similar in both species. Early cleavages were equal, radial, and holoblastic; slightly subequal cleavage was common inPsolus chitonoides. The gastrula elongated to form a uniformly ciliated larva and the vestibule first appeared as a ventral crescent-shaped depression, after which the larva decreased in length and formed three posterior ridges that encircled the early doliolaria. Just prior to settlement, there was loss of cilia between the three ridges and on the posterior end of the early doliolaria, leaving three ciliary rings that are characteristic of the dendrochirote doliolaria larva. Vertical swimming ensued, and five primary tentacles protruded through the vestibule, with occasional attachment to the substratum using the primary tentacles. Upon settlement, two primary podia emerged and all external ciliation was lost except among tentacular papillae and on the end of tube feet. Timing of appearance and of loss of ciliary rings suggest that the rings function in vertical swimming and substratum selection behaviors. Metamorphosis involved the following simultaneous events: (1) resorption of the preoral lobe, (2) formation of large, dorsal, calcareous ossicles, (3) flattening of the trivium, and (4) proliferation of ventral tube feet on Ambulacral Radii B and E to form the sole. Reproduction in the family Psolidae is reviewed. The large proportion of brooders in this family appears to be related to small size, the possession of protective armor and sole, and geographic distribution (mainly in Antarctica). Lecithotrophic doliolaria larvae with three ciliary rings are produced by large northern psolids.     

Structural and functional peculiarities of the digestive system of Cucumaria frondosa (Echinodermata: Holothuroidea)

The fine structure of the intestinal epithelium of Cucumaria frondosa has been examined. The digestive cells possessed some specific ultrastructural peculiarities which testify to their participation in the transport of substances and in the processes of intracellular and cavitary digestion. The main units of the intestinal epithelium appear to be multifunctional digestive cells in which the phenomenon of compartmentalization among the organelles' groups, intended to carry out various functions, occurs. The heterogeneity of the structure of populations of the digestive cells is explained. Both primitive and progressive features of the structure of the digestive tract of C. frondosa are indicated.     

Reproduction of the intraovarian brooding apodid Leptosynapta clarki (Echinodermata: Holothuroidea) in British Columbia

Reproduction in the viviparous apodid sea cucumber Leptosynapta clarki was examined in Grappler Inlet, Bamfield, Vancouver Island, Canada from July 1989 through July 1991. The reproductive cycle of L. clarki is annual with gonad growth during the summer months. Male sea cucumbers spawn during November and December and internally fertilize, through as yet undetermined means, the oocytes within the ovary. Embryonic development is rapid, so that within 2 wk fertilized eggs reach the pentactula stage. Pentactulae have an open mouth and anus and actively move in the ovarian tubules feeding on droplets and fluid derived from the resorption of unfertilized eggs. Histological examination of gametogenesis shows similar development to that found in non-brooding holothurians. However, the protandric sex change observed in some male L. clarki after spawning and changes to the overian wall structure prior to fertilization are previously undescribed in sea cucumbers.     

Fission and its effect on population structure of Holothuria atra (Echinodermata: Holothuroidea) in Taiwan

Populations of Holothuria atra Jaeger occurring at Wanlitung and Nanwan were sampled monthly at spring tide from March 1990 to February 1992 in southern Taiwan. At Wanlitung, small individuals with evidence of fission and regeneration were abundant in shallow tidepools on the wide reef flats (density: 98±40 ( ± SD) 100 m^-2, body weight: 6 to 182 g). At Nanwan, large individuals with no evidence of fission were sparse in deep tidepools and narrow flats [density: 0.24±0.07 ( ± SD) 100 m^{- 2}, body weight: 351 to 1400 g]. At Wanlitung, fission occurred throughout the year with an average monthly fission frequency of 4.5%, peaking at 18% in September 1990 and 16% in August 1991. Fission was accomplished by revolving, twisting, and stretching of the body, resulting in two unequal fragments. The minimal weight of 3007 individuals measured during 1 yr at Wanlitung was 6 g. All ten individuals weighing 6 to 9 g ( ± SD = 6.7 ± 1.1) 100 m^{- 2} showed evidence of regeneration from fission products. Nine large individuals (>500 g) transferred from Nanwan to Wanlitung and 50 small individuals (<100 g) transferred from Wanlitung to Nanwan in February 1991 showed no evidence of fission during the following year. Fission occurs in small individuals living in shallow tidepools, suggesting that fission probably is triggered by a stressful environment resulting from solar radiation at the unusually low water level of spring tide. Fission of H. atra at Wanlitung results in a population consisting of small individuals.     

Reproductive variability over a four-year period in the sea urchin Evechinus chloroticus (Echinoidea: Echinodermata) from differing habitats in New Zealand

 The endemic New Zealand echinoid, Evechinus chloroticus (Valenciennes), was sampled approximately monthly from September 1990 to October 1994 at three sites in Tory Channel, Marlborough Sounds, New Zealand. These channel sites (outer, mid and inner) were up to 20 km from the open ocean and differed in their shore type, exposure to wave action and macrophyte abundance. E. chloroticus showed an annual reproductive cycle at each site, with gametogenesis commencing in the late austral winter and spawning in summer. Maximum gonad indices (reproductive potential) varied spatially, with the outer site generally having higher maximum gonad indices than the inner site, and temporally, with maximum gonad indices occurring earlier in 3 of the 4 seasons at the outer site than the inner site. The mid and inner sites showed much greater variation in maximum gonad indices (range 15.83 to 26.99% and 11.87 to 20.90%, respectively) than the outer site (range 19.31 to 22.95%). Reproductive output (weight of gametes released per gram echinoid) also varied, with the different sites showing significantly different outputs in the different years. A regression of maximum reproductive potential against reproductive output was significant ( p < 0.001), and had a positive slope with an r ² of 0.79. While, the initiation of gametogenesis was relatively synchronous between sites and years, and is possibly cued by increasing daylength, it progressed at different rates among populations. Spawning did show spatial and temporal variability, occurring near the time of highest sea-surface temperatures (∼15 °C). The observed variations in reproductive cycle may be related to small-scale variability in diet and environmental conditions. Furthermore, asynchronous spawning, variable spawning duration, and variable reproductive output are likely to strongly influence annual recruitment variability in E. chloroticus, with different larval subpopulations contributing unequally in different years. The ecological consequences of this, both for the ability of E. chloroticus to propagate itself in space and time and for the management of the developing E. chloroticus fishery in New Zealand, are discussed. Received: 9 December 1998 / Accepted: 13 June 2000     

Reproductive cycle,larval development,juvenile growth and population dynamics of Patiriella pseudoexigua (Echinodermata: Asteroidea) in Taiwan

A field study was conducted at Wanlitung, southern Taiwan, in 1986–1089, to determine the reproductive cycle, development mode, growth rate and population dynamics of the small seastar Patiriella pseudoexigua (Dartnall), which occurs in highly stressful and disturbed intertidal pools in this area. An inverse relationship between gonad index and pyloric-caccum index was only recorded immediately prior to spawning. A short, well-synchronized seasonal spawning occurs in October. When reared at 25 °C, lecithotrophic larvae develop directly, lack a bipinnaria stage, and metamorphose completely on the seventh day after fertilization. The growth curves of field juveniles are linear, those of laboratory-reared juveniles are sigmoid. Juveniles appear in tide pools in spring-early summer of each year. Adults spawn mainly in late fall, enabling spawning to occur in time for the larvae to benefit from the environmentally favorable winter season. Populations in high-tidal pools decrease in later summer, but remain more stable in lowtidal pools and lagoons.     

Reproductive strategies of two deep-sea gastropod species from the Porcupine Seabight (Northeast Atlantic)

Two species of small gastropods (<6 mm in length), Amphissa acutecostata (Philippi, 1844) and Gymnobela subaraneosa (Dautzenberg and Fischer, 1896), widely distributed in the northeast Atlantic, were found in large numbers in the Porcupine Seabight (Northeast Atlantic). Except for some aspects of taxonomy and distribution, as well as some data on larval development, the biology of these species is unknown. This study describes basic aspects of the life-history strategies of both species. Histological studies showed that oocyte and sperm development in both species was similar to the gametogenetic patterns observed in other deep-sea gastropods. In females, oogonia proliferated in the germinal epithelium and developed into previtellogenic oocytes (30–40 m), which grew into vitellogenic primary oocytes. Vitellogenic oocytes were covered by a thin layer of follicle cells involved in the vitellogenic processes. The maximum size for mature oocytes was 99.06 m for A. acutecostata and 114.82 m for G. subaraneosa. In A. acutecostata most of the volume of the ovary was occupied by previtellogenic and early vitellogenic oocytes, whereas in G. subaraneosa most of the volume was filled by large vitellogenic oocytes. Both species showed quasi-continuous production of oocytes. The oocyte size-frequency diagrams suggested a continuous release of a small number of oocytes throughout the year for A. acutecostata, and asynchronous periodic spawning events for G. subaraneosa. Gonad development and gametogenesis could be strongly affected by presence of parasites in one of the populations of G. subaraneosa.Communicated by J.P. Thorpe, Port Erin     

Bacterial diversity of the sediments transiting through the gut of Holothuria scabra (Holothuroidea; Echinodermata)

This work analyzes bacterial diversity of sediments transiting through the gut of Holothuria scabra which is an important bioturbator in tropical shallow waters. This edible holothurian species has a social and economic importance for coastal populations in many developing countries. Bacterial biodiversity was analyzed by sequencing the 16S rRNA of bacterial cultures and clones. DAPI and FISH methods were used to determine and compare the number of bacteria found in the various gut compartments. A total of 116 phylotypes belonging to the γ-Proteobacteria (60.5 %), α-Proteobacteria (24.5 %), Bacteroidetes (6 %), Actinobacteria (2.75 %), Fusobacteria (1.75 %), Firmicutes (1.75 %), Cyanobacteria (1.75 %) and δ-Proteobacteria (1 %) were identified. The number of bacteria is significantly greater (1.5×) in the foregut than in the ambient sediments. The number of bacteria significantly decreases in the midgut and remains stable until defecation. Some γ-Proteobacteria, especially Vibrio, are less affected by digestion than other bacterial taxa. The season has an impact on the bacterial diversity found in the sediments transiting through the gut: in the dry season, γ-Proteobacteria are the most abundant taxon, while α-Proteobacteria dominate in the rainy season. Vibrio is the most frequent genus with some well-known opportunistic pathogens like V. harveyi, V. alginolyticus and V. proteolyticus. Findings show that sediment-associated microbial communities are significantly modified by H. scabra during their transit through the gut which supports the view that holothurians play a substantial role in the structuring of bacterial communities at the sediment–seawater interface.     

Reproductive biology of the common antarctic crinoid Promachocrinus kerguelensis (Echinodermata: Crinoidea)

Histological examinations of the gonads of Promachocrinus kerguelensis Carpenter, 1888 collected in November 1984 from McMurdo Sound, Antarctica, revealed ripe ovaries containing three distinct size classes of oocytes: 20 to 90, 91 to 150 and 151 to 180 m diameter as well as ova. Testes were ripe, and filled with mature sperms. There was marked inter-pinnule and inter-individual synchrony of gametogenesis and spawning is probably synchronous, at least within the sub-population sampled. The number of eggs per pinnule is allometrically related to body size. Body weight and arm length are positively correlated with total egg number. Annual fecundity was estimated to be 29 000 eggs (full-grown oocytes and ova) for a crinoid weighing 53.6 g wet weight with an arm length of 16.3 cm. The eggs float. Female genital pinnules contain more lipid and energy than male pinnules. Reproductive output and effort (23.4 to 65.2 kJ and 26 to 38%, respectively) increase with body size and are similar to values reported for other echinoderms with lecithotrophic larvae. The greatest overall energetic contribution is represented by the arms and genital pinnules. Spawning in P. kerguelensis probably occurs in November and December within McMurdo Sound. If development is slow, as in all other antarctic echinoderms studied to date, then settlement most likely occurs 2 to 3 mo later, after the short summer period of high phytoplankton productivity. High fecundity, a pelagic mode of reproduction, and the high probability of a slow rate of development may contribute to the wide geographic distribution of this common antarctic crinoid.