Strobilation,budding and initiation of scyphistoma morphogenesis in the rhizostome Cassiopea andromeda (Cnidaria: Scyphozoa)

Scyphistomae of Cassiopea andromeda Forskål, 1775 containing symbiontic zooxanthellae did not develop medusae at a constant temperature of 20°C, but monodisc strobilation was initiated after transfer of the polyps to 24°C. After release of the ephyrae and regeneration of the hypostome and tentacular region, the recovered polyps either produced vegetative buds or entered a new strobilation phase. Formation of motile, planula-like buds was not found to be indicative of unfavourable environmental conditions. Intensity of budding was positively correlated with available food and with increase of temperature. Budding was negatively correlated with the number of polyps maintained per dish and with the conditioning of the sea water. Under optimal feeding and temperature conditions, polyps could simultaneously produce chains of buds at 2 to 4 budding regions. Settlement and development of buds into scyphistomae was suppressed in pasteurized sea water and in pasteurized sea water containing antibiotics, but polyps developed from buds in the presence of algal material taken from the aquarium, debris or egg shells of Artemia salina, or on glass slides which had been incubated in used A. salina culture medium. Several species of marine bacteria were detected after staining these slides. One, a Gramnegative coccoid rod, which was identified as a nonpathogenic Vibrio species, was isolated, cultivated as a pure strain, and was proved to induce the development of C. andromeda buds into polyps. Millipore filter-plates coated with Vibrio sp. cells grown in suspension culture were ineffectual, but diluted filtrate initiated polyp morphogenesis. The inducing factor is obviously not a constituent of the bacterial cell surface, but is a product of growing Vibrio sp. cells released into the medium. This product was found to be relatively heat-stable and dialyzable. As to the basic mechanism involved in the induction of polyp formation, it is suggested that the inducing factor (s) acts bimodally by inducing pedal disc development and by eliminating a head inhibitor originating from the basal end of the bud. The life history, and various aspects of medusa-formation and of vegetative reproduction in scyphozoans are reviewed and discussed with particular reference to rhizostome species. Special attention has been paid to some reports of larval metamorphosis controlled by marine bacteria.     

Genetic analysis of sexual reproduction in the dendrophylliid coral Balanophyllia elegans

The ahermatypic scleractinian Balanophyllia elegans has served as a model of limited larval dispersal in an aclonal species. However, other species from the same family (Dendrophylliidae) produce larvae asexually, and closely connected polyps of B. elegans, potentially the result of asexual reproduction, are commonly observed in the field. Here, we use a combination of laboratory experiments, controlled crosses, and genetic surveys of field-collected individuals to demonstrate (1) marker allozymes are inherited in accordance with Mendel's laws, (2) polyps that are connected in the field are not genetically identical, and (3) laboratory manipulation of a single polyp can produce a second polyp on the aboral side of the original, but not adjacent to the original. In combination, these results suggest that the larvae of B. elegans result from mictic sexual reproduction, and that connected polyps result not from asexual budding but rather from the fusion of genetically distinct individuals.     

Why does the white tip of stony coral grow so fast without zooxanthellae?

The photosynthesis of zooxanthellae in a coral polyp greatly enchances the calcification rate of a coral. However, the white tip of a coral branch is free of zooxanthellae yet still has a very high calcification rate. Furthermore, the reason for the difference is not clear. In this study, the amount of photopigment, total protein (TP), total organic carbon (TOC), ATP, and lipid in polyps from the white tip and brown stalk of a branch of stony coral were measured. Samples of Acropora hyacinthus and A. formosa were collected from southern Taiwan between 1985 and 1987. The results showed that the ATP concentration in polyps of the white tip was much higher than that in polyps of the brown stalk. Conversely, the amount of TP, TOC and measured lipids in polyps of the brown stalk were all higher than those of the white tip. It was the high concentration of ATP in cells that gave these polyp tips the vitality to sustain the energy requirements of such a rapid calification rate. Facilitated diffusion, due to the high metabolite gradient created by cell activity, could be the major driving force for the transport of photosynthetic product from stalk to tip.     

Intra-colonial variability in light acclimation of zooxanthellae in coral tissues of Pocillopora damicornis

We investigated heterogeneity of light acclimation of photosynthesis in sun- and shade-adapted coenosarc and polyp tissues of Pocillopora damicornis. The zooxanthellar community within P. damicornis colonies at Heron Island is genetically uniform, yet they showed a large degree of plasticity in their photo-physiological acclimation linked to light microclimates characterised by fibre-optic microprobes. Microscale scalar irradiance measurements showed higher absorption in polyp than coenosarc tissues and higher absorption in the more densely pigmented shade-adapted polyps than in sun-adapted polyps. The combination of an O₂ microelectrode with a fibre-optic microprobe (combined sensor diameter 50–100 μm) enabled parallel measurements of O₂ concentration, gross photosynthesis rate and photosystem II (PSII) quantum yield at the coral surface under steady-state conditions as a function of increasing irradiances. Lower O₂ levels at the tissue surface and higher compensation irradiance indicated a higher respiration activity in sun-adapted polyp tissue as compared to shade-adapted polyps. Shade-adapted coenosarc and polyp tissues exhibited lower maxima of relative electron transport rates (rETR_max) (84±15 and 41±10, respectively) than sun-adapted coenosarc and polyp tissues (136±14 and 77±13, respectively). Shade-adapted tissues showed stronger decrease of rETR at high scalar irradiances as compared to sun-adapted tissues. The relationship between the relative PSII electron transport and the rate of gross photosynthesis, as well as O₂ concentration, was non-linear in sun-adapted tissues over the entire irradiance range, whereas for shade-adapted tissues the relationship became non-linear at medium to high scalar irradiances >200 μmol photons m⁻² s⁻¹. This suggests that rETR measurements should be used with caution in corals as a proxy for photosynthesis rates. The apparently high rates of photosynthesis (oxygen evolution rates) suggest that there must be a considerable electron transport rate through the photosystems that is not observed by the rETR measurements. This may be accounted for by vertical heterogeneity of zooxanthellae in the tissue and the operation of an alternative electron pathway such as cyclic electron flow around PSII.     

Polyp and medusa of Dipurena bicircella n. sp. (Hydrozoa: Corynidae) from Northern California

The polyp and medusa of Dipurena bicircella n. sp., a metagenic hydrozoan from Northern California, USA, is described. The arrangement of capitate tentacles of the polyp, in two alternating cycles or circlets, is unique to the genus Dipurena as well as to the family Corynidae. Medusae released in the laboratory attained sexual maturity in 2 weeks. The adult medusa can be distinguished from other medusae of the genus by means of 2 gonad rings, a slight terminal nematocyst cluster on each tentacle, and a relatively small size (about 2 mm in bell height). The polyp was not found in association with sponges, as is the case with some of the other polyps of the genus Dipurena. The medusa is not yet known from the plankton.     

Comparison of telomere length among different life cycle stages of the jellyfish Cassiopea andromeda

The polyp (scyphistoma) of the jellyfish Cassiopea andromeda reproduces asexually repeatedly, while the medusa, the sexually reproducing stage, exhibits a relatively shorter life span. As a first step to understand the mechanism behind the differences in the life spans of the polyp and medusa stages of the jellyfish, we compared the lengths of the telomere region of one targeted chromosome between the polyp and medusa stages using a modified single telomere length assay (STELA). The double-stranded regions of the telomeres were amplified by PCR, and the average length of the PCR products was estimated by densitometry analysis of the gel smear. Chromosomes within cells of the bell region of the medusa were characterized by longer telomeres than those of polyps, asexual propagules, or other regions of the medusa. This is the first study to estimate the telomere lengths of targeted chromosomes in a cnidarian and opens a way to understand the mechanism underlying different life spans of the polyp and medusa stages.     

Growth trajectories of corallites and ages of polyps in massive colonies of reef-building corals of the genus Porites

Massive colonies of the reef-building coral genus Porites were collected at inshore, midshelf and shelf-edge reefs in the central section of the Great Barrier Reef in November 1987. These colonies were comprised of 4 species: P. lobata, P. lutea, P. solida and P. mayeri. X-radiographs made of skeletal slices cut from the skeletons displayed the annual density-banding pattern characteristic of massive corals, and appeared to show corallites within each slice. The average age of the 36 colonies was 41±12 yr (mean±SD). The images of corallites displayed by the X-radiographs were not images of actual corallites, but approximated the position and size of actual corallites. Consequently, X-radiographs provide information about the formation and growth trajectories of corallites, and about the history of the polyps which deposited the corallites. Individual corallites were always normal to the growth surface. The growth surface of the colonies became bumpy when they reached 50 to 80 mm in height and, as a result, corallites took on a fan-shaped arrangement within a bump. New corallites were initiated at the summit of each bump and grew upwards and outwards. Thus, growth of colonies resulted in corallites becoming increasingly displaced from the summit of a bump. The X-radiographs showed that corallite growth becomes occluded at the bottom of valleys between adjacent bumps. Corallite growth then stops and the associated polyps are probably resorbed. Annual density banding showed that the average age of polyps in these colonies was 2 to 3 yr, average life expectancy 5 yr, and that no polyp was likely to be older than 8 yr. Small but significant variations in polyp longevity between corals from different reefs were probably associated with significant differences in bumpiness of growth surfaces. Even in Porites colonies which have been growing for several centuries, polyp longevity is likely to be 5 yr.     

Nucleic acid and protein levels in strobilating polyps of Chrysaora quinquecirrha and Aurelia aurita

Levels of DNA, RNA, and protein were measured in scyphistomae of the scyphozoans Aurelia aurita and Chrysaora quinquecirrha during strobilation. In synchronously developing populations of A. aurita, the amount of DNA per polyp increases 41 to 260% during strobilation, whereas RNA and protein remain relatively constant. The RNA/DNA ratio drops by about 50% during strobilation in both C. quinquecirrha and A. aurita, and protein/DNA decreases by 50% in A. aurita. Specific RNA reserves are probably not accumulated in advance of strobilation, since strobilation is blocked immediately in both species by 1 to 3 g/ml actinomycin D. The size and DNA content of polyps are usually increased by prolonged incubation at temperatures of 12° to 15°C, which facilitates strobilation; however, the RNA/DNA and protein/DNA ratios of such polyps are not significantly increased by the cold conditioning.Contribution No. 472 from the Virginia Institute of Marine Science.     

Derivation of the reduced life cycle of <Emphasis Type="Italic">Thecoscyphus zibrowii</Emphasis> Werner, 1984 (Cnidaria,Scyphozoa)

The species Thecoscyphus zibrowii Werner, 1984 has an exceptional life cycle, which lacks a medusa stage but develops an extraordinary structure (egg sac) for reproduction. Investigation of the life cycle, as well as anatomical and histological studies of the different developmental stages of T. zibrowii were performed to provide evidence for a possible homology of the egg sac with the medusa stage and to determine whether the reduced metagenesis of T. zibrowii is derived from strobilation. The egg sac showed several characteristics, which were compared to those of coronate medusae. The ectodermis of the egg sac had a plate-like appearance and was completely ciliated as is typical for coronate medusae. The number and the location of the gonads were similar to those of coronate medusae. The cnidocysts were significantly larger in the egg sac than in the polyp. A size difference of cnidocysts in the medusa and the polyp stage is known for several Coronatae. Characteristics of egg sac formation were compared to characteristics of strobilation. The formation of the early operculum was similar in T. zibrowii and N. eumedusoides. The constriction of egg sac and strobila occurred in the same mode and the gastric cavities of two egg sacs stayed in contact in a similar fashion to the gastric cavities of the strobila discs. The developmental zones of cnidoblasts of the egg sac and polyp were separated during the formation of the egg sac which showed a similar developmental gradient to a strobila. The existence of all of these consistent characteristics makes it very likely that the egg sac structure was homologous to a medusa. The species T. zibrowii would therefore be derived from a metagenetic ancestor. This species has reduced the medusa generation to the greatest extent within the Nausithoidae and has demonstrated thus far the endpoint of a regressive evolution of the medusa generation.     

Invertebrate introductions in marine habitats: two species of hydromedusae (Cnidaria) native to the Black Sea,Maeotias inexspectata and Blackfordia virginica,invade San Francisco Bay

The hydrozoans Maeotias inexspectata Ostroumoff, 1896 and Blackfordia virginica Mayer, 1910, believed to be native to the Black Sea (i.e. Sarmatic) and resident in a variety of estuarine habitats worldwide, were found as introduced species in the Petaluma River and Napa River, California, in 1992 and 1993. These rivers are mostly-estuarine tributaries that flow into north San Francisco Bay. Both species appeared to be well-established in this brackishwater habitat. Salinities at the collection sites were about 11 during the summer, rising to nearly 20 in the early autumn and falling to near 0 in the winter. Large numbers of all sizes of both species of medusae were observed and collected, indicating that the hydroid stages of the life cycles of the two are also well-established in these rivers. In the Petaluma River, populations of both species were at maximum in late July, with numbers of individuals declining through August and into September; the Napa River was sampled only in October, and at that time only B. virginica was found. Examination of full guts of M. inexspectata and B. virginica medusae revealed that both species had fed nearly exclusively on small crustaceans, principally barnacle nauplii, copepods and their eggs and nauplii, and crab zoea larvae (M. inexspectata only). All the M. inexspectata medusae were males, indicating that the population has probably developed from the introduction of perhaps only a single male polyp or polyp bud. In spite of its inability to reproduce sexually, this population appears to be maintained by the prodigious ability of the polyp to bud and reproduce asexually, and is fully capable of invading additional low-salinity habitats from its present Petaluma River site. Male and female B. virginica medusae were collected in both the Petaluma River and the Napa River, indicating that B. virginica may have been introduced by either the polyp or medusa stage (or both), but that multiple individuals (of both sexes) must have arrived from another port in one or more invasions. As indicated for M. inexspectata, the B. virginica population will also probably seed new populations in San Francisco Bay and elsewhere. Based on its cnidome as well as the morphology of both medusa and polyp, M. inexspectata has been reclassified by moving it from the family Olindiidae, Limnomedusae, to the family Moerisiidae, Anthomedusae.     

Reproductive biology of the antipatharian black coral Antipathes fiordensis in Doubtful Sound, Fiordland, New Zealand

The reproductive biology of Antipathes fiordensis Grange, a species endemic to south-western New Zealand, was followed from April 1994 to May 1995. Ten colonies were individually tagged in Doubtful Sound and sampled on a monthly basis in order to determine their reproductive activity. The fecundity of each of the five tagged female colonies was determined by estimating the total number of polyps per colony from photographs of each colony and by planimetry, the proportion of gravid polyps per colony, and the mean number of oocytes per gravid polyp. In addition, 56 colonies were sampled in March 1995 to estimate the sex ratio, height at sexual maturity, and mean sizes of females and males. A. fiordensis was found to be a dioecious species which, in the absence of gonads in the polyps, has no obvious external morphological differences between the sexes. Broadcast spawning of gametes is the likely mode of reproduction. Gametogenesis began in November 1994 and was highly synchronous within and between colonies, with spawning occurring in March 1995. The sex ratio in adults was 1:1. Colonies reached sexual maturity between the heights of 70 and 105 cm which, based on existing estimates of growth rate, corresponds to a minimum age for sexual maturity of about 31 yr. The largest oocytes measured ranged from 100 to 140 μm in size. Female colonies produced between 1.3 and 16.9 million oocytes, with the larger colonies dominating the reproductive output of the population. Received: 23 June 1997 / Accepted: 1 August 1997     

Reproductive biology of the deep-sea octocoral Drifa glomerata in the Northwest Atlantic

The present study examined the mode and timing of reproduction of poorly understood deep-water octocorals and the environmental factors that may influence their reproductive patterns. Data on reproductive characteristics of the octocoral Drifa glomerata (Alcyonacea: Nephtheidae) collected between 2004 and 2007 at ca. 100–330 m depth off Newfoundland and Labrador (eastern Canada) were compared among years, months and depth ranges. No male gonad was ever observed during the study. The ratio of fertile colonies possessing large pinkish polyps with oocytes/planulae was >50% throughout the year. The number of brooded planula larvae within a single fertile polyp varied between 1 and 10 for a total of approximately 40–3,000 in a whole colony. The size of oocytes and/or planulae was consistently greater in the polyps than in the branchlets, indicating that the development pathway of oocytes to planulae is from the branchlets to the polyps. Although larval production seemed to persist year round, the onset of major planulation events occurred in December–January of two consecutive years, when large mature planulae were released in correlation with the first increase of photoperiod and maximum temperature at 150 m. A second peak in planulation between April and early June followed the phytoplankton bloom. Seasonal trends were more apparent in colonies from <200 m, and the planula index varied among sampling depths and years. Larval release in a live colony under laboratory conditions occurred between January and June 2008, closely following predictions based on field samples.     

Flesh or bone? Quantifying small-scale coral morphology using with-tissue and without-tissue techniques

The quantification of small-scale coral morphology using tissue-free skeletons often requires colonies, or colony sections, to be removed from the reef. This undesirable destruction can be reduced by using a with-tissue technique based on photographic images of living corals. The results of this study indicate that with-tissue multivariate morphometric data contain similar information to skeleton-derived data and can help identify phenotypic plasticity in the massive coral Favia speciosa. With-tissue data can also be used to discern changes with time in individual polyps exposed to new environments.Communicated by O. Kinne, Oldendorf/Luhe     

Ontogeny and allometric plasticity of Dentex dentex (Osteichthyes: Sparidae) in rearing conditions

The ontogeny of Dentex dentex (Linnaeus, 1758) was studied comparatively under extensive and semi-extensive rearing conditions. The exotrophic phase was divided into three distinct developmental stages (a) larval, from feeding onset up to 10 mm total length (TL); (b) metamorphosis, from 10 mm TL up to 24 mm TL; and (c) further juveniles. The majority of all ontogenic changes was expressed during the larval stage and integrated during the metamorphosis stage. The juvenile stage was characterized by developmental stability. The normal development of D. dentex was seriously affected by the rearing conditions applied during the early exotrophic phase. The two differently reared populations followed similar developmental patterns of general morphology, pigmentation and behaviour, but they presented high developmental plasticity where allometric growth was concerned. Rearing conditions influenced both the boundaries of the sequential stages (in 50% of the morphometric ratios) and the allometry coefficient of each morphometric character in the various developmental stages (in 80% of the characters). The results are discussed with respect to the ontogeny of the functional morphology under both ecological and aquaculture considerations. Received: 1 August 1998 / Accepted: 1 September 1999     

Substrate choice and settlement preferences of planula larvae of five Scyphozoa (Cnidaria) from German Bight,North Sea

The settlement behaviour of planula larvae and their development to young polyps was investigated in laboratory experiments in five scyphozoan species [Aurelia aurita (L.), Cyanea capillata (L.), Cyanea lamarckii Péron and Leseur, Chrysaora hysoscella (L.), and Rhizostoma octopus (L.)]. The undersides of settling plates were strongly preferred for settlement. Shells, the only natural substrate type offered, were less attractive than artificial substrates (concrete, machined wood, polyethylene, and glass). The advantages of colonization of substrate undersides for survival and reproduction of polyps are discussed. It is supposed that the increase of artificial substrates in our seas, due to marine litter pollution and submarine building activities, enlarge the areas of distribution of scyphozoan polyps, in coastal as well as in off-shore regions. Subsequent increases in ephyra production by polyps are probably one reason for the increase in mass occurrences of jellyfish recognized worldwide during the last few decades. It is suggested that the early developmental stages in the cnidarian life cycle, the planula larvae, and the polyps, play the key role in the development of jellyfish outbursts.     

Occurrence and detrimental effects of the bivalve-inhabiting hydroid <Emphasis Type="Italic">Eutima japonica</Emphasis> on juveniles of the Japanese scallop <Emphasis Type="Italic">Mizuhopecten yessoensis </Emphasis>in Funka Bay,Japan: relationship to juvenile massive mortality in 2003

In November 2003, we first observed prevalent occurrence of a hydroid, Eutima japonica, on soft body tissues of age zero Japanese scallop (Mizuhopecten yessoensis) juveniles cultured in large areas of Funka Bay, Hokkaido. The occurrence coincided with massive death of juvenile scallops. A major objective was to clarify ecological relationships between the symbionts, and to infer the relationship between symbiosis and the massive mortality. To do this, we investigated distributions of association rates of hydroids with juvenile scallops at 15–34 sites over 3 years (2003–2005), with age one adult scallops at 24 sites in 2003, and with mussels at 13 sites in 2004. We studied seasonal changes in association rates with juvenile scallops, and numbers of polyps per juvenile scallop at three sites from November 2003 to June 2004. We also quantified the hydroid impacts on juvenile scallop shell length growth and triglyceride accumulation in the digestive gland. The association rate of E. japonica polyps with juvenile scallops was high in large areas of Funka Bay in 2003, and overlapped the distribution of mussels bearing polyps. Association rates with age one adult scallops were very low in November 2003, even at the sites where polyps were very common on juvenile scallops. Levels of hydroid occurrence in juvenile scallops varies by year. We found that hydroids presence in juvenile scallops declined drastically in 2004 and 2005. The association rates with juvenile scallops, and numbers of polyps per juvenile scallop declined during winter, until they disappeared completely in the following June. Since polyps were rare in adult scallops, we believe that infection of juvenile scallops was probably initiated from the planulae produced by medusae released from polyps growing on Mytilus spp., especially M. galloprovincialis. Subsequently, the inhabitation spread intraspecifically and interspecifically. In juvenile scallops, inhabitation of polyps reduced shell length growth by 43%, and triglyceride accumulation in digestive glands by 24–47%. Inhabitation of E. japonica on juvenile scallop is best regarded as parasitism, rather than inquilinism or commensalism. Occurrence of polyps was probably not a direct lethal factor for juvenile scallops, because there were some sites where association rates were high, but mortalities were low. Massive mortalities in 2003 may have resulted from simultaneous impacts of heavy polyp load and stresses caused by the way in which the animals were handled (transferred from cages for pre-intermediate culture to cages for intermediate culture), because the massive mortality occurred within a month of the transfer. The presence of polyps in juvenile scallops does not affect the quality of the product in Funka Bay, because market size scallops are hydroid-free.     

Size-dependent reproduction in the Caribbean gorgonian Pseudoplexaura porosa

Reproductive success among clonal taxa is often portrayed as a simple function of clone size, but reproduction in Pseudoplexaura porosa (Houtuyn), a common Caribbean gorgonian, reflects a more complex relationship between size and gamete production. Tagged colonies were sampled at two reefs in the San Blas Islands, Panama during the 1995 spawning season to determine the colony size at first reproduction, size-dependent polyp fecundities, and whole colony gonad production, and to follow the gametogenic cycle. Additional data were obtained during non-spawning months in 1984 and 1985. Of 120 colonies ranging in height from 20 to 250 cm, only colonies taller than 50 cm were reproductive. An average of 4.33 oocytes polyp⁻¹, 560 to 800 μm in diameter, were released during monthly synchronous spawning events from June to September. Oocytes released each month developed from a group of intermediate size oocytes present at the end of the preceding month. Almost all oocytes >560 μm were released during each spawning event. Spermatogenesis was a shorter process than oogenesis. During each spawning month, mature spermaries developed from a group of <125 μm spermaries. Virtually all spermaries >190 μm were released each month, and most of the spermary volume in mature male polyps was generated anew each month of spawning. Among reproductive colonies, polyp fecundity increased with colony height from 50 to 200 cm. Per capita fecundity of >200 cm tall female colonies was lower than for 150 to 200 cm colonies, but whole colony fecundity of large colonies was greater due to the exponential increase in the number of polyps that occurs with increasing colony height. In male colonies, both polyp and colony reproductive output increased with colony height. The large amounts of sperm produced by large male colonies probably contributes to the high rates of in situ fertilization observed in P. porosa. Differences in fecundity as a function of colony size and sex suggest differences in some combination of the cost of reproduction and/or allocation of resources to reproduction. Received: 9 September 1998 / Accepted: 7 June 1999     

Complex reproductive cycle inEucheilota paradoxica (Hydrozoa: Leptomedusae): Medusae,polyps and frustules produced from medusa stage

Eucheilota paradoxica Mayer, 1900 is a small leptomedusa unknown in the Mediterranean Sea until 1977. Since then, it has become endemic and occurs in great abundance in autumn in the bay of Villefranche-sur-mer. Since no sexual stages have ever been collected, either from the Atlantic Ocean or from the Mediterranean Sea, we decided to investigate the still unknown life cycle of this species. Specimens were obtained in the bay of Villefranche-sur-mer from plankton hauls between 0 and 50 m depth during October and November 1986. At temperatures >18°C, the medusa reproduces intensively by asexual medusal budding (one young medusa liberated per two days). This asexual reproduction explains the rapid proliferation of the species and the origin of the large population every autumn. In cultures maintained at temperatures <18°C, two other reproduction modes, not yet reported in a leptomedusa, were observed: (1) At 16° to 17°C, frustules bud along the canals; these frustules are dormant, giving rise to a small polyp when the temperature increases once more above 18°C. (2) At temperatures close to 15°C each medusa, in place of the gonads, differentiate into one or two polyps; these polyps develop progressively, concomitant with gradual regression of the mother medusa.     

Reproduction and colony integration of the sea pen<Emphasis Type="Italic"> Virgularia juncea</Emphasis>

The distribution of fertile polyps, and the functional variations of polyps within sea-pen colonies, were investigated in Virgularia juncea, a species inhabiting shallow waters in the Penghu Islands, west of Taiwan. This gonochoristic species started gametogenesis between July and September, and mature gametes were broadcast in late summer the following year. Fertile pinnae and their constituent polyps were located in the middle region of the rachis; the upper and lower regions were infertile. Within the fertile regions, fecundity per centimeter decreased linearly toward the tips of the colonies. Abundant zooxanthellae were found in the upper part of the rachis; algal densities decreased down the axis of the colonies, and no algae were present in the lower part. New pinnae were generated only at the basal part of the rachis, They moved upward in relative position with growth of the colonies. These young pinnae near the peduncle were small and infertile. With time, they became fertile and started to harbor a few zooxanthellae. The old pinnae at the upper part of the rachis were large and infertile and contained numerous zooxanthellae. The tips of the colonies were often damaged, and the bare axial rods there were often fragile and could be easily broken off. The developmental gradient of pinnae (and their constituent polyps) along the axes of the colonies, coupled with the environmental gradient within a sea-pen colony, facilitates the functional differentiation within the rachis that may be an adaptation for inhabiting the interface between the water column and the sediment.Communicated by P.W. Sammarco, Chauvin