Observations on Octopus joubini: Some aspects of reproductive biology and growth

Observations were made on the brooding behavior of 10 laboratory reared female Octopus joubini, including egg laying, care of the eggs, feeding during brooding, and survival of the female from the onset of egg laying. Data concerning the numbers of eggs laid by each female and hatching (duration, frequency, correlation of duration to water temperature, etc.) are given. The newly hatched octopuses (whose ages were determined within 24 h) were fed Uca spp. to determine their growth rate from hatching to 4 months of age. In addition, some specimens were presented with a variety of foods that were easily accessible in the laboratory or field in an effort to find a food suitable for mass culture.Contribution No. 1830 from the University of Miami, Rosenstiel School of Marine and Atmospheric Science.     

Effects of embryonic exposure to conspecific chemical cues on hatching and larval traits in the common frog <Emphasis Type="Italic">(Rana temporaria)</Emphasis>

Summary. Recent studies indicate that amphibian eggs are capable of hatching plasticity in response to chemical cues released by predators feeding on conspecific eggs or larvae. However, information is scarce on the relative importance of predator and conspecific cues in such a process. In particular, no attempt has been made to compare the effects of embryonic exposures to chemical cues indicative of a predation risk for eggs and larvae, although both life stages can co-occur in natural habitats. In this context, common frog embryos (Rana temporaria) were raised until hatching in the presence of crushed conspecific extracts from eggs and tadpoles to assess their respective influences on some hatching and larval traits. While a significant delay in hatching time was observed in embryos exposed to chemical cues from tadpole extract, this life-history shift appeared unaffected by embryonic exposure to egg extract. Hatchlings derived from eggs incubated in the presence of both conspecific extracts showed a significantly greater weight than unexposed controls. However, such an effect was no longer apparent 15, 30 and 50 days after hatching, suggesting that embryonic exposure to chemical cues from damaged conspecific eggs and tadpoles has no influence on larval growth. Lastly, morphological measurements performed on hatchlings and older tadpoles (15, 30 and 50 days old) revealed no significant effect of embryonic treatments on the shape of body and tail.     

Hatching with the enemy: Daphnia diapausing eggs hatch in the presence of fish kairomones

Summary. Infochemicals are known to play a key role in mediating predator-prey interactions, both in aquatic and terrestrial communities. However, state-dependent variation may exist in how effectively individuals can use this information, depending on genotype, life stage and experience. For our study, we used the predator-prey model system fish-waterflea Daphnia magna Straus (Cladocera, Daphniidae). Adult Daphnia use fish-derived infochemicals, so-called kairomones, as indicators of predation risk, and exhibit a spectrum of morphological, behavioural and life-history responses to the presence of fish kairomones. Here, we investigate whether diapausing eggs, an embryonic resting stage in the life cycle of D. magna, also use fish kairomones and tune their hatching to the risk of fish predation, as reported for diapausing stages of dinoflagellates. In two laboratory experiments, we studied hatching proportion and time until hatching of D. magna diapausing eggs in the absence and presence of fish kairomones. D. magna families differed significantly in their response to the presence of fish kairomones; some families reduced hatching proportion, whereas others increased it. Our results imply genotype-dependent differences in the hatching reactions to fish kairomones as observed for other traits in adult Daphnia.     

Reproduction of the invasive slipper limpet, Crepidula fornicata, in the Bay of Brest, France

The reproduction of Crepidula fornicata was studied in the Bay of Brest in order to characterise the first step of the reproductive cycle of this invasive species. The survey was carried out from 2000 to 2003 and different parameters were measured, namely, the percentage of the different sexual stages, the straight length of the shell and the percentage of brooding females using a survey of the embryonic development and the fecundity. The juvenile frequency increases generally from mid-June or mid-August, depending on the year. In 2001 and 2003, a first peak was observed as early as May, but it was followed by a rapid disappearance of the individuals. The sex-ratio female/male increased from 0.22 to 0.46 between 2001 and 2003. The sex change between intermediates and females took place mainly in summer and was well marked in 2001 and 2003. The survey of the embryonic development in the egg capsules brooded by the females provided an annual phenology of the laying and hatching processes. The laying period extends from February to September with three to four major periods of egg-laying per year and corresponding hatching periods about 1 month later. Each female lays two to four times per year on average. The first egg-laying concerned fewer females than subsequent ones, except in 2003, and exhibited a higher fecundity. The annual mean of the number of eggs for each stage was not significantly different, thus indicating no significant mortality rate during embryonic development. For the C. fornicata population in the Bay of Brest, several reproductive characteristics tend to highlight its invasive capacity: (1) a long reproductive period, (2) reproduction in a ‘multi-trials’ process equivalent to a spreading out of the risks and (3) a relatively high fecundity.     

Brooding behaviour of the northern sea star Leptasterias polaris

The northern sea star Leptasterias polaris (Müller and Troschel) has a unique mode of brooding its young. The mother curves her arms along the plane of the disc, the whole individual assuming a flattened plate-like shape. In this form she covers the larvae, which are attached to the substratum beneath her. The central disc is not raised to form a brood chamber, as in related species, and there is no tendency to hide while brooding. In the St. Lawrence Estuary, brooding L. polaris are common from February through May and decline in numbers during June and early July. The genus Leptasterias originates in the Arctic and probably all species brood their young.This paper is dedicated to François Dorion who lost his life during benthic studies in the St. Lawrence Estuary. His enthusiasm in studying marine organisms will always be an inspiration to usContribution to the program of GIROQ (Groupe Interuniversitaire de Recherches Océanographiques du Québec)     

Study of the life history of Capitella capitata (Polychaeta: Capitellidae) in Amakusa,South Japan including a comparison with other geographical regions

The life history of Capitella capitata (Fabricius), occurring in Amakusa, South Japan, was studied in detail through field investigations and laboratory experiments. The important life-history characteristics of this species in the present study area are described and compared with those reported in previous studies from various geographic regions. Though some life-history characteristics (extended breeding season throughout almost the whole year, parental brooding habit, and early maturation) are common to all localities, the reproductive mode, larval developmental pattern and morphology are very distinct among the worms from various localities. C. capitata showing these considerable geographical variations should not be treated as a single species. It is suggested that C. capitata, originally defined on the basis of morphological features of adult worms, needs to be subdivided into several species, subspecies, or biological races.     

Brooding behaviour and reproductive success in two species of free-living simultaneous hermaphrodites

Although polyclads are amongst the most structurally simple of the triploblastic metazoans, they adopt a wide range of reproductive strategies. Parental care behaviour in this group is yet to be quantified for any species. We assessed the significance of brooding behaviour to the reproductive success of two free-living marine flatworms. Echinoplana celerrima and Stylochus pygmaeus were collected from the field and placed in pairs in containers of filtered seawater where they laid batches of eggs. Both parents were then removed from half of the containers and the brooding behaviour and hatching success of eggs were quantified. There were interspecific differences in brooding behaviour. Egg masses were covered by one E. celerrima parent for 12 ± 2% of time, whereas egg masses of S. pygmaeus were covered by one or both parents simultaneously for 85 ± 8% of time. Egg batches were abandoned by both species immediately prior to the onset of hatching (10–12 days). Hatching success was generally high (~90%) and brooding did not enhance the hatching success of eggs. We assessed the significance of parental care to hatching success of E. celerrima egg masses in the presence of three potential egg predators; in the presence of other organisms. E. celerrima devoted less time to brooding; however, hatching success was not affected. The amount of time spent brooding eggs differed greatly between the two polyclad species but was not essential to their reproductive success under benign conditions. Parental care may be of adaptive value under more stressful environmental conditions commonly experienced in estuarine environments such as lowered salinity, increased hypoxia or turbidity. Covering egg batches may play an additional role of advertising sexual status and a willingness to care for eggs.     

Apparent resource competition among embryos in the brood pouch of a male pipefish

Embryo success was studied in the paternally brooding pipefish Syngnathus typhle. During brooding, which lasts about a month, males provide embryos in their brood pouch with nutrients and oxygen via a placenta-like structure. Egg size depends on female size. In aquaria, males were mated with differently sized females to give the following treatments: M, mixed-egg-size broods of approximately half large and half small eggs; L, single-egg-size broods of large eggs; S, single-egg-size broods of small eggs; and F, field mated males. All males were kept in aquaria for a full brooding period. For each egg-size category, the number of newborn was compared with the number of eggs the male initially fertilized in his brood pouch. Within mixed-egg-size broods, a higher proportion of large eggs survived and large eggs resulted in heavier newborn than small eggs. Indeed, small eggs from a mixed-egg-size brood had significantly lower relative success (proportion of embryos surviving to birth) than those from a brood entirely composed of small eggs. The implication is that embryos compete for resources within the brood pouch, and that competitive success depends on egg size. Given that females produce eggs corresponding in size to their body size, and that females are known to compete indirectly for access to mates (i.e., the sex-roles are reversed), this intrabrood competition could be seen as an extension of female-female competition, but alternative explanations are discussed. Received: 28 April 1995/Accepted after revision: 28 October 1995     

Life-history variation in three coexisting cheilostome bryozoan species of the genus Stylopoma in Panamá

Life-history variation was studied in three sympatric species of Stylopoma Levinsen [S. spongites (Pallas), S. projecta Canu and Bassler and Stylopoma n. sp. 15] in Panamá. Bryozoan colonies were collected from 27 reefs along 300 km of the Caribbean coast of Panamá. The distribution and abundance of each species were very patchy, but with broad overlap in occurrence among localities and depths. Nevertheless, species differed considerably in colony size, size at first reproduction and numbers of brooding larvae; implying that interspecific differences in life-history traits may contribute to their coexistence. To examine closely this variation in life-history patterns, we grew, in a common garden experiment, the offspring of the two most common species that were obtained from parent colonies collected from several reefs. There were highly significant differences in growth rates and the timing and extent of sexual reproduction which corresponded well to patterns observed in individuals from the field. Other factors, including size of larvae, extent of secondary zooidal calcification and numbers of avicularia were also correlated with differences in life histories. Despite this additional complexity, however, ecological consequences of trade-offs in life history among modular species such as Stylopoma spp. appear very similar to those among unitary species.     

Brooding and development of <Emphasis Type="Italic">Anasterias minuta</Emphasis> (Asteroidea: Forcipulata) in Patagonia,Argentina

Brooding, embryonic and larval development, and the influence of environmental and biological factors in tidepool habitats were studied in the sea star, Anasterias minuta, at various sites along ~220 km of the Patagonian coast. This species has a benthic, lecithotrophic development that includes eight distinct developmental stages. A larval organ, the connection cord, is developed from a small preoral lobe at early stages of development and becomes larger and thinner at advanced stages. Fecundity and average egg size increased with female body size. The regression of log egg number to log sea-star size and weight at different sites had a slope significantly less than 3.0, resulting in negative allometry and indicating that brood capacity was limited in large females. Development was generally synchronous among sites, but varied within each brood at advanced stages, with more developed brooded larvae located at the periphery of the brood mass. Brooding was synchronous among various populations at different years and spatial scales, and extended over a period of 8 months. The highest proportion of brooding females occurred during May and June (austral winter). Juveniles were released mainly during September. The likelihood of finding brooding sea stars decreased with increasing sea water temperature, tidal height, and wave exposure, and increased with increasing body size. Both body size of brooding females and brooding rate were higher in the infralittoral fringe than at midlittoral levels. A revision of the current model of brooding behavior and development among forcipulate sea stars is given.     

The effect of maternal body size on embryo survivorship in the broods of pregnant male pipefish

The occurrence of male pregnancy in the family Syngnathidae (seahorses, pipefishes, and sea dragons) provides an exceptionally fertile system in which to investigate issues related to the evolution of parental care. Here, we take advantage of this unique reproductive system to study the influence of maternal body size on embryo survivorship in the brood pouches of pregnant males of the broad-nosed pipefish, Syngnathus typhle. Males were mated with either two large females, two small females, a large then a small female, or a small then a large female. Our results show that offspring survivorship depends on an interaction between female body size and the number of eggs transferred by the female. Eggs of larger females deposited in large numbers are more likely to result in viable offspring than eggs of smaller females laid in large numbers. However, when females deposited smaller numbers of eggs, the eggs from smaller females were more likely to produce viable offspring compared to those from larger females. We found no evidence that this result was based on mating order, the relative sizes of competing females, or egg characteristics such as dry weight of eggs. Additionally, male body size did not significantly influence the survivorship of offspring during brooding. Our results suggest that the factors underlying offspring survivorship in pipefish may be more complex than previously believed, with multiple factors interacting to determine the fitness of individual offspring within the broods of pregnant males.     

Effects of sublethal application of Deepwater Horizon oil to bird eggs on embryonic heart and metabolic rate

Following large crude oil spills, oil from feathers of brooding birds and oiled nesting material can transfer to eggs, resulting in reduced embryonic viability for heavily oiled eggs. Eggs may also be subjected to trace or light oiling, but functional teratogenic effects from sublethal crude oil exposure have not been examined. We assessed whether sublethal application of weathered Deepwater Horizon crude oil to the eggshell surface alters heart rate and metabolic rate in Zebra Finch (Taeniopygia guttata) embryos. We first determined sublethal applications with a dosing experiment. Embryo viability for eggs exposed to 5 μL or more of crude oil decreased significantly. We conducted a second experiment to measure heart rate and metabolic rate (CO₂ production) 5 and 9 d after 1 sublethal application of crude oil to eggshells on day 3 of incubation. One application of 1.0 or 2.5 µL of crude oil reduced embryonic heart rate and metabolic rate on day 12 of incubation. Using unfertilized eggs, we measured the transfer of polycyclic aromatic hydrocarbons (PAHs) from the eggshell surface to egg contents 9 d after a single application of sublethal crude oil. Our results suggest avian eggs externally exposed to small amounts of crude oil may exhibit protracted embryonic development and impaired postnatal cardiac performance.     

Growth and reproduction in the Antarctic brooding bivalve Adacnarca nitens (Philobryidae) from the Ross Sea

We present information on the reproductive biology, population structure, and growth of the brooding Antarctic bivalve Adacnarca nitens Pelseneer 1903, from the Ross Sea, Antarctica. Individuals ranging from 0.85 to 6.00 mm were found attached to a hydrozoan colony. This species shows low fecundity and large egg size, common to other brooding species. The minimum size at which oogenesis was detected was 2.3 mm and the minimum size at which brooding was evident was 3.9 mm. Embryos of a full range of developmental stages were brooded simultaneously in females. The population showed a log–normal distribution and results suggest non-periodic reproduction with continuous embryonic development. The reproductive traits of A. nitens are discussed in the context of circum-Antarctic species distribution and limitations to dispersal in brooding benthic invertebrates.     

Pelagic larval development in the brooding Antarctic brachiopod Liothyrella uva

Larvae of the Antarctic brachiopod Liothyrella uva (Broderip, 1833) were released by brooding females between 10 and 15 January 1992. They were collected and cultured for the ensuing 45 d. At release they were at the gastrula stage, and were recognisable as the usual brachiopod three-lobed larva from around 18 d after release. At 22 d post-release, larvae began to congregate under shell fragments placed in culture vessels as potential settlement substrata. However, the larvae showed no signs of settlement which, combined with the absence of many of the attributes of mature larvae seen in previous studies, was taken as evidence that these larvae had not reached fully competent stages. Previous work had shown that L. uva spawns in October, indicating that the developmental period lasts for a minimum of between 115 and 160 d. The release and subsequent development of larvae also shows that what was previously thought to be a brooding species combines a long brooding period with a free-swimming phase. If this is true of other brachiopod species, then there may need to be a recategorization of some species from brooding to combined brooding and free-swimming developmental types. Even though the development period in L. uva was likely to have been underestimated because a fully competent stage was not reached, a comparison with development rates for temperate species indicated a slowing of between 8 and 70 times. This dramatic slowing of development rate in brachiopods now joins similar data previously reported for echinoderms and nemerteans, suggesting that this is a very widespred, perhaps universal, attribute of Antarctic marine invertebrates.     

Sexual reproduction and settlement of the coral reef spongeChalinula sp. from the Red Sea

Characteristics of the sexual reproduction and larval settlement of the haplosclerid spongeChalinula sp., which inhabits the shallow waters (1 to 6 m) of Eilat, Red Sea, were investigated from September 1985 through to November 1987. This species was found to be a simultaneous hermaphroditic brooder, hence gonochorism is not the rule in the order Haplosclerida. Brooding always takes place in special brooding chambers. While the oocytes in the brooding chambers are among the largest known in sponges (355±37µm), the spermatic cysts distributed in the choanosome are among the smallest known for this phylum (average 26±7µm).Chalinula sp. breeds throughout the year and in experiments most larvae (74%) settled within 1 to 8 h post-release, generally within 4.5 h. Metamorphosis from larval shape to a sessile sponge lasts 1 to 6 h. Thus, larvae had a short swimming period, settled fast, and metamorphosed rapidly (within 1 to 6 h). The large size of the larvae may contribute to their ability to rapidly reorganize their body shape into that of a sessile sponge. In addition, the existence of already differentiated choanocyte chambers in the larvae, facilitates fast construction of the water filtration system in the newly settled sponges. The reproductive and larval characteristics ofChalinula sp. enable the larvae to settle on any vacant space in the reef, which may explain its abundance in the Red Sea.     

Egg survival,embryonic development,and larval characteristics of northern shrimp (<Emphasis Type="Italic">Pandalus borealis</Emphasis>) females subject to different temperature and feeding conditions

Laboratory experiments on ovigerous females of northern shrimp (Pandalus borealis) were used to assess the effects of temperature and food ration on female condition during incubation and examine how combined effects of temperature and female condition influenced egg survival, embryonic development, and larval characteristics. Ovigerous females were maintained at 2°C, 5°C, and 8°C and fed on a low (three times/week; 2–2.7% W/W) or high ration (five times/week at satiation). The increase in temperature accelerated the developmental time of the eggs but their survival at 8°C was reduced. Conversion efficiency of yolk reserves in developing embryos was significantly reduced at elevated temperatures and larvae hatching at 2°C and 5°C were significantly larger and heavier than those hatching at 8°C. The experimental design did not result in any effect of food ration on the energetic condition of females or on egg characteristics and their biochemical composition. However, lower energy reserves were observed for females held at 8°C.     

Factors affecting the allocation of paternal care in waterbugs (Belostoma flumineum Say)

Summary Male giant waterbugs (Belostoma flumineum Say) brood eggs oviposited on their dorsa by conspecific females. Laboratory observations indicate that viable egg pads are occasionally discarded before hatching. Theory predicts that such behavior should occur only if the costs incurred by brooding exceed the benefits of hatching the egg pad. We studied the effects of egg pad size, time invested in brooding, and egg viability upon the continuation of paternal care in the giant waterbug. We found that smaller egg pads are less likely to hatch than larger ones, and males appear to be less likely to discard egg pads as temporal investment increases. However, the inviability of eggs did not appear to affect the probability of an egg pad being discarded. Males of this species appear to have evolved a decision-making process involving the continuation of paternal care. Offprint requests to: K.C. Kruse     

Sexual reproduction in solitary corals: Overlapping oogenic and brooding cycles,and benthic planulas in Balanophyllia elegans

The temperate solitary coral Balanophyllia elegans Verrill, 1864 was collected monthly between January 1977 and August 1978 off the California coast. B. elegans reproduces only sexually, is gonochoric, and broods its embryos. Males are ripe only in late summer, but oocytes and embryos (ca. 40 per female) are found throughout the year. The presence of oocytes and embryos throughout the year may be the result of very long, overlapping oocyte growth and embryonic developmental times. Large, crawling, benthic planulas are released mainly in the winter. The skeletal design of the corallum of B. elegans accommodates the brooding of unusually large embryos; this design may be indicative of brooding large embryos in other species of corals, particularly dendrophyllids.