Size and temperature-dependent variations in intermolt duration and size increment at molt of Northern Shrimp, Pandalus borealis

Growth of Pandalus borealis post-larval stages was measured in relation to size and temperature. Growth characteristics, including intermolt period (IP), molt increment (MI) in size and mass, and tissue allocation in juvenile, male, and female shrimp, were evaluated at 2, 5, and 8°C, the temperature range where this species is generally found in the Northwest Atlantic. Significant variations in growth were associated with temperature and shrimp size. IP (days) increased significantly with shrimp size and was inversely related to temperature. Size (cephalothorax length in mm) and temperature effects were best described by IP = 10^{(0.67 log(CL) − 0.06 T − 1.34)}. The pronounced effect of temperature on IP while MI_S changed little indicated that the main influence of temperature on growth rate of P. borealis was through IP. Specific growth rate (SGR_S) decreased rapidly with size to near zero values in females. Overall, juveniles were much more sensitive to temperature variations than adults, suggesting that temperatures encountered during the juvenile stage will largely influence the growth trajectory of the population.     

Latitudinal and bathymetric trends in body size of the deep-sea gastropod<Emphasis Type="Italic"> Troschelia berniciensis</Emphasis> (King)

Body size is a fundamental topic in ecology with important implications for community structure and biodiversity. Although there are numerous studies addressing patterns of geographic variation of body size in deep-sea benthos, results are conflicting. Thus geographic patterns of body size in deep-sea organisms remain poorly described. We analysed depth and latitudinal trends of body size in a species of gastropod, Troschelia berniciensis (King, 1846) from the eastern North Atlantic. We tested the hypotheses that (1) body size increases linearly with latitude and (2) body size increases with depth. Results partially supported the predictions. Firstly, there was a significant linear increase of body size with latitude, but this trend was weak. Secondly, body size decreased with depth. Environmental gradients that cause large-scale patterns of body-size variation in surface environments would have little effect on communities living at great depths. Latitudinal and depth clines may be produced by independent mechanisms that operate on different scales of time and space.Communicated by J.P. Thorpe, Port Erin     

Area-specific temporal changes of species composition and species-specific range shifts in rocky-shore mollusks associated with warming Kuroshio Current

Increases of low-latitude species in various sea areas and poleward shifts of ranges of many species have been reported and linked to warming environments. To examine the generality of these trends for mollusks, we conducted 7 quadrat surveys during 1978–2006 on Pacific rocky shores in Japan (26.6–45.0°N). Results showed that the dominance of southern species increased on 11 of the 15 shores in a southern, warming sea area but on only 1 of the 6 shores in a northern area with no warming trend. The latitudes of species ranges increased on average but showed large interspecific variations which showed a weak, positive correlation with interspecific taxonomic distance. The differences of these results from the previously reported trends are discussed in relation to the unique current patterns in our study area and the phylogenetic constraints of species-specific responses to a warming environment.     

Life history and production of <Emphasis Type="Italic">Corophium urdaibaiense</Emphasis> (Crustacea: Amphipoda) in the Urdaibai estuary (NE Spain)

The population of species Corophium urdaibaiense has been studied for a year in the Urdaibai estuary (NE Spain). The new species is characterised by its preference for muddy sand sediments (15% of mud) and the long body size (with a maximum length of 10.24 mm). The maximum abundance occurs in November (11,600 ind m⁻²); it almost disappears at the end of spring. The juveniles represent 79% of the population and the recruitment is concentrated within the October–December period. The breeding period begins in May, when the temperature is above 15°C, and it ceases in November, when the temperature is below 15°C. Bivoltinism is probable, taking into account the latitude (43°), temperature range (12–25°C) and breeding period (May and August–October), but the data are not sufficiently consistent to confirm the hypothesis. The production ranges from 2.93 to 5.85 g m⁻² year⁻¹, considering one and two generations per year, respectively. Some aspects concerning the food availability, predation and intraspecific competition could have an influence on the dynamics of the population and are discussed.     

Latitudinal patterns of diversity and abundance in North Atlantic intertidal boulder-fields

In order to study taxon richness, biodiversity and abundance patterns in the North Atlantic from temperate latitudes through Arctic to high Arctic latitudes, we recorded the faunas (at ELWS level) colonizing 20 cobbles from three sites at each of seven boulder-field localities (south-west England, 50°N; Wales, 51°N; west Scotland, 56°N; Iceland, 64°N; Tromsø, 70°N; Svalbard, 77°N, 79°N). Inverse correlations were found between latitude and all measures of richness (species, orders, and phyla numbers) and biodiversity (S–W, P). However, these correlations were driven mostly by the consistently impoverished Arctic sites; an even cline of decreasing diversity from south to north did not exist. Multidimensional scaling revealed two communities, temperate–subarctic species-rich and high arctic species-poor. Evenness as measured by Pielou’s index was similar across all latitudes. Abundance data exhibited a similar trend to the species richness and diversity data with a significant negative correlation with latitude, but when Arctic data were excluded the correlation vanished.     

Tracking habitat and resource use for the jumbo squid Dosidicus gigas: a stable isotope analysis in the Northern Humboldt Current System

To determine the habitat and resource use of Dosidicus gigas in the Northern Humboldt Current System, we analysed carbon and nitrogen stable isotopes of 234 individuals collected during 2008–2010. Large variations in mantle stable isotope ratios were recorded, with values ranging from −19.1 to −15.1 ‰ (δ¹³C) and from 7.4 to 20.5 ‰ (δ¹⁵N). Most of the variation was explained by latitude, followed by distance to shelf break for carbon and by squid size for nitrogen. Latitudinal variations with increasing values from north to south were also found in zooplankton samples and were related to changes in isotope baseline values probably due to oxygen minimum zones that occur off Peru. This similar latitudinal trend in both zooplankton and D. gigas samples reveals that D. gigas is a relatively resident species at the scale of its isotopic turnover rate (i.e. a few weeks), even if this is not necessarily the case at the scale of its life. A small but significant size effect on δ¹³C values suggests that jumbo squid perform offshore–onshore ontogenic migration, with juveniles distributed offshore. For nitrogen, the high inter-individual variability observed with mantle length indicates that D. gigas can prey on a high variety of resources at any stage of their life cycle. This large-scale study off the coast of Peru provides further evidence that D. gigas have the capability to explore a wide range of habitats and resources at any stage of their life.     

Multi-year observations on the gametogenic ecology of the Antarctic seastar <Emphasis Type="Italic">Odontaster validus</Emphasis>

This study reports the first multi-year observations on the reproductive patterns for an Antarctic predator/scavenger, Odontaster validus (Koehler 1912). Seastars were collected monthly from a shallow site (15–20 m depth) near the British Antarctic Survey (BAS) Rothera Research Station (Adelaide Island, 67°34′S 68°08′W) from July 1997 to January 2001. Reproductive condition, oocyte size frequencies and spermatogenesis were examined in at least ten seastars each month using histological and image analysis techniques. Gonad indices (GI) and pyloric caeca indices (PI) were also examined in the same samples. Female and male GIs varied seasonally, in parallel with a reduction in the proportion of large oocytes and mature sperm in the gonad in August to mid-October following winter spawning. Despite there being remarkable consistency in the timing of spawning from year to year, differences in the reproductive condition of individuals were apparent. Patterns in the digestive tissues also varied with season, peaking in December and reaching a minimum in February in two of the three study years. This weaker annual pattern may partly reflect the varied diet of this predator/scavenger species, which is not directly dependant on the timing and magnitude of the annual phytoplankton bloom. Pooled oocyte size distributions and residual analysis suggested that oogenesis progressed over 18–24 months, with the largest of the two size classes (maximum diameter = 183 μm) being spawned annually. This pattern of oocyte growth and spawning was previously reported in the early 1960s for an O. validus population from McMurdo Sound, which lies south of Rothera by 10° latitude. The extremely catholic diet of this predator/scavenger suggests the reproductive patterns of the seastar will be less susceptible to changes in food supply compared to polar suspension feeders or deposit feeders. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Thermal effects on swimming activity and habitat choice in juvenile Pacific cod (Gadus macrocephalus)

The behavioral responses of fishes to temperature variation have received less attention than physiological responses, despite their direct implications for predator–prey dynamics in aquatic ecosystems. In this paper, we describe the temperature dependence of swimming performance and behavioral characteristics of juvenile Pacific cod (Gadus macrocephalus; 75–125 mm total length). Maximum swimming speeds increased with temperature and body size. Routine swimming speeds of Pacific cod in small groups of similarly sized fish (N = 6) increased with body size and were 34 % faster at 9 °C than at 2 °C. The response to temperature was opposite that previously described for juvenile walleye pollock (Theragra chalcogramma), reflecting species-specific differences in behavioral responses. In a separate experiment, we demonstrated the effect of temperature on habitat selection of juvenile Pacific cod: Use of an artificial eelgrass patch in a 5-m-long laboratory tank was significantly greater at 9 °C than at 2 °C. These results illustrate that temperature affects a range of behavioral traits that play important roles in determining the frequency and outcomes of predator–prey interactions.     

Energy content and fecundity of capelin (<Emphasis Type="Italic">Mallotus villosus</Emphasis>) along a 1,500-km latitudinal gradient

Capelin is a planktivorous key fish species in many subarctic ecosystems representing the link between lower trophic levels and apex predators. Little is known, however, of capelin in Greenland waters, but it has been suggested that size and growth increases due to difference in food availability and temperature conditions along a 1,500 km south–north gradient on the west coast. It is presently unknown how the qualitative state of capelin energy content is affected along this gradient. Based on 2007 and 2008 samples, we show that energy content increases with capelin length in both spawning and non-spawning fish and that it varies with latitude in spawning fish along West Greenland (60–71°N). Combining our results on energy content with information on capelin growth along the same latitudinal climate gradient demonstrates that less and lower-quality food is available to capelin predators in the south than in the north.     

Archival tagging of subadult and adult common thresher sharks (Alopias vulpinus) off the coast of southern California

The common thresher shark (Alopias vulpinus) is a secondary target species of the California drift gillnet fishery (CA-DGN) and supports a growing recreational fishery in California waters. This study used archival tags to examine the movement patterns and habitat preferences of common threshers of the size range captured in the CA-DGN (>120 cm fork length). Depth and temperature-logging archival tags were deployed on 57 subadult and adult common threshers in the Southern California Bight. Tags from five individuals (8.8%) were recovered, and 154 days of data were successfully obtained from four of these. By night, shark movements were primarily limited to waters above the thermocline, which ranged in depth from 15 to 20 m. Sharks were significantly deeper by day, and daytime vertical distribution consisted of two distinct modes: a ‘shallow mode’ (wherein sharks occupied only the upper 20 m of the water column) and a ‘deep mode’ (characterized by frequent vertical excursions below the thermocline). This modal switch is interpreted as relating to regional differences in abundance of surface-oriented prey and prey in deeper water. Maximum dive depth was 320 m, greatest dive duration was 712 min, minimum temperature experienced during a dive was 9.1°C, and dive descent rate was significantly greater than ascent rate. Sharks inhabited waters corresponding to a sea surface temperature range of 16 to 21°C. The nocturnal depth distribution of common threshers has implications for management of drift gillnet deployment depths in the CA-DGN.     

Reproductive traits of pioneer gastropod species colonizing deep-sea hydrothermal vents after an eruption

The colonization dynamics and life histories of pioneer species determine early succession at nascent hydrothermal vents, and their reproductive ecology may provide insight into their dispersal and population connectivity. Studies on the reproductive traits of two pioneer gastropod species, Ctenopelta porifera and Lepetodrilus tevnianus, began within a year after an eruption on the East Pacific Rise (EPR) that eliminated vent communities near 9°50′N from late 2005/early 2006. Standard histology was used to examine gamete release, instantaneous female fecundity, and time to maturation. Both species exhibited two-component oocyte size–frequency distributions indicating quasi-continuous reproduction with high fecundity. In samples collected in December 2006, both C. porifera and L. tevnianus individuals were reproductively mature. The smallest reproducing C. porifera were 4.2 mm (males) and 5.4 mm (females) in shell length, whereas reproductive L. tevnianus were smaller (2.3 and 2.4 mm in males and females, respectively). Most C. porifera were large (>6.0 mm) compared to their size at metamorphosis and reproductively mature. In contrast, most L. tevnianus were small (<1.0 mm) and immature. Reproductive traits of the two species are consistent with opportunistic colonization, but are also similar to those of other Lepetodrilus species and peltospirids at vents and do not fully explain why these particular species were the dominant pioneers. Their larvae were probably in high supply immediately after the eruption, due to oceanographic transport processes from remote source populations.     

Population dynamics of naturalised Manila clams <Emphasis Type="Italic">Ruditapes philippinarum</Emphasis> in British coastal waters

The Manila clam Ruditapes philippinarum was introduced to Poole Harbour (lat 50°N) on the south coast of England in 1988 as a novel species for aquaculture. Contrary to expectations, this species naturalised. We report on individual growth patterns, recruitment, mortality and production within this population. On the intertidal mudflats the abundance of clams (>5 mm in length) varied seasonally between 18 and 56 individuals m⁻². There appear to be two recruitment events per year and there were 6 year classes in the population. A mid-summer decline in abundance was partly due to increased mortality but probably also a result of down-shore migration in response to high water temperatures and the development of anoxic conditions. A winter fishery removes c 75% of clams of fishable size (maximum shell length ≥40 mm) and c 20% of the annual production. The fishery depresses the maximum age and size attained by the clams but appears to be sustainable. Clam mortality due to factors other than fishing is highest in late-winter to early spring. The growth of the clams is intermediate in comparison with many published studies but remarkably good given their intertidal position. As on the coasts of the Adriatic Sea, where the clam is also non-native, the Manila clam has thrived in a shallow, eutrophic, lagoon-like system on the English coast. While the Poole Harbour population is currently Europe’s most northerly reported self-sustaining, naturalised population, given forecasts of increasing air and sea temperatures it might be expected that this species will eventually spread to more sites around the coasts of Northern Europe with associated economic and ecological consequences.     

Growth parameters of deep-water decapod crustaceans in the Northwestern Mediterranean Sea: a comparative approach

 Relative and absolute growth were studied in 17 species of deep-water decapod crustaceans, spanning nine families of six different infra-orders, in the Northwestern Mediterranean Sea. The overall maximum abundance of these species lay between 200 m and 750 m (i.e. upper- and mid-slope species). Relative and absolute growth rates were compared by contrasting the slopes of the size–weight relationships for the different species and calculating the von Bertalanffy growth-equation parameters asymptotic length (L _∞ ) and growth rate (k). The size–weight relationships differed significantly as function of the species' life habits. The results revealed a significant decrease in weight relative to size in mesopelagic species (which carry out diel vertical migrations), an almost isometric relationship between size and weight in the less mobile nektobenthic species, and a significant increase in weight relative to size in strictly benthic species. The mean allometric coefficient for each group increased significantly from mesopelagic to benthic species. However, no general trend was observed in the growth-performance index, Φ (an index used to compare absolute growth rates between species, as a function of habit and depth of maximum abundance for all species combined), suggesting that the deep-water decapod crustaceans studied have similar absolute growth rates. Nevertheless, comparison of growth-parameter and growth-performance index values within families did reveal differences. Mesopelagic species of the families Sergestoidae and Pasiphaeidae showed slightly increased growth rates with increasing depth of distribution. Nektobenthic species of the genus Plesionika followed a trend opposite to that shown by mesopelagic species, with a higher growth rate for the shallowest-dwelling species (P. heterocarpus) than the deepest-dwelling species (P. acanthonotus). Taking growth as one of the major components of an organism's energy budget, the growth rates for the decapod crustacean species in this study were significantly lower than those reported in the literature for shallow-water penaeid crustacean species (which are distributed in higher-temperature habitats than deep-water Mediterranean crustaceans) and higher than those reported for mesopelagic myctophid fish species. Hence, the well-defined growth trends shown by deep-water decapod crustacean species in the Northwestern Mediterranean Sea, compared to the less well-defined trends in the other taxa, is discussed in the framework of the overall dynamics of their ecosystem. Received: 25 May 1998 / Accepted: 27 September 1999     

Physiological tolerance predicts species composition at different scales in a barnacle guild

This study examined how the species composition of an intertidal barnacle guild varied according to physical gradients in the environment at small scales governed by microclimates, medium scales of wave exposure and large scales of latitude. Barnacle distributions at small and medium scales were sampled in Ireland between 51°29′ and 52°44′N and 6°50′ and 10°08′W. Sampling on European shores spanned ~18° latitude from 37°05′ to 55°16′N. Barnacle surveys mainly took place in 2003–2004. An index of wave fetch was calculated along the wave exposure gradient using a digital coastline-based model that was supported by a biological exposure scale. A ‘dryness’ index was defined according to mean monthly wind speed, fetch along the average wind direction and mean monthly air or sea surface temperatures for 2 years (January 2001–December 2002) which is the period when the most recent adults in the barnacle community would have settled and grown to adulthood. The proportion of the dry-loving barnacle Chthamalus montagui Southward increased within the barnacle guild at all scales as the habitat became warmer and drier. Barnacle densities were high in all habitats, mean densities ranged from a minimum of 4.16 cm⁻² on moderately exposed shores to a maximum of 6.27 cm⁻² in sunlit or south-facing microclimates. Percentage cover of barnacles across the gradient of latitudes was usually >70%. The results suggest that the distribution and abundance of interacting barnacle species on European coasts is strongly controlled by abiotic factors, most likely temperature and desiccation.     

Biodiversity of coastal polyclad flatworm assemblages in the wider Caribbean

Polyclads are mobile predators and possibly an important functional component of hard substrate marine environments globally. To understand the natural patterns of spatial differences, polyclad assemblages were sampled in seven coastal regions across the wider Caribbean spanning 15° latitude and 24° longitude between May and June 2005 and May and September 2006. In total, 67 species in 28 genera and 17 families were recorded from 62 sites. Only two species were found in all surveyed regions, Melloplana ferruginea and Pseudoceros bicolor. Conversely, 41 species were restricted to one or two sites, and 34 species were represented by one or two individuals. The distribution and abundance of species varied between the two suborders. Cotyleans were most species rich and had a higher number of species of restricted range, while Acotylea showed a higher proportion of rare species; however, two species were highly abundant comprising over half of the individuals counted. In most habitats, polyclads were rare, but in some intertidal habitats two species, Styloplanocera fasciata and Boninia divae were densely aggregated and dominant members of the benthic epifauna. Alpha diversity was variable but showed no evidence of a relationship with latitude, longitude or depth. Beta diversity increased with the number of habitats sampled and was highest for cotyleans. Highest gamma diversity was recorded in Jamaica and the US Virgin Islands and was not significantly correlated to alpha diversity. Overall assemblages from the seven regions were similar, revealing faunal homogeneity across the wider region. Reef assemblages were distinct from other habitats dominated by species of the Cotylea. Reefs from Panama and the US Virgin Islands were the most species rich. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Temporal, spatial, and body size effects on growth rates of loggerhead sea turtles (Caretta caretta) in the Northwest Atlantic

In response to a call from the US National Research Council for research programs to combine their data to improve sea turtle population assessments, we analyzed somatic growth data for Northwest Atlantic (NWA) loggerhead sea turtles (Caretta caretta) from 10 research programs. We assessed growth dynamics over wide ranges of geography (9–33°N latitude), time (1978–2012), and body size (35.4–103.3 cm carapace length). Generalized additive models revealed significant spatial and temporal variation in growth rates and a significant decline in growth rates with increasing body size. Growth was more rapid in waters south of the USA (<24°N) than in USA waters. Growth dynamics in southern waters in the NWA need more study because sample size was small. Within USA waters, the significant spatial effect in growth rates of immature loggerheads did not exhibit a consistent latitudinal trend. Growth rates declined significantly from 1997 through 2007 and then leveled off or increased. During this same interval, annual nest counts in Florida declined by 43 % (Witherington et al. in Ecol Appl 19:30–54, 2009) before rebounding. Whether these simultaneous declines reflect responses in productivity to a common environmental change should be explored to determine whether somatic growth rates can help interpret population trends based on annual counts of nests or nesting females. Because of the significant spatial and temporal variation in growth rates, population models of NWA loggerheads should avoid employing growth data from restricted spatial or temporal coverage to calculate demographic metrics such as age at sexual maturity.     

Growth of the scyllarid lobsters Ibacus peronii and I. chacei

The growth rates of the morphologically similar scyllarid lobsters Ibacus peronii (Leach, 1815) and I. chacei (Brown and Holthuis, 1998) are described using data from a tag/recapture study and from tagged lobsters kept in captivity. Within particular size classes, we found no differences in moult increments between male and female I. peronii nor between male and female I. chacei. Small individuals of both species always had larger moult increments than larger individuals. For I. peronii, females moulted more frequently than males, and smaller size classes moulted more frequently than larger size classes. Female I. peronii therefore grew more quickly than males and reached their estimated size at sexual maturity (51 mm carapace length) after ∼2 yr. Moulting of I. peronii was seasonal, with most lobsters (96.3%) moulting between October and January. We found no differences in growth rates of I. peronii at two locations along the east coast of Australia: Coffs Harbour in New South Wales (30°18′S; 153°08′E), and Lakes Entrance in Victoria (37°53′S; 148°00′E). For I. chacei, we found no differences in the frequency of moulting between males and females and, because we also found no differences in the moult increments between males and females, the growth rates of both sexes were the same. Received: 14 August 1999 / Accepted: 20 January 2000     

Isotopic evidence of distinct feeding ecologies and movement patterns in two migratory predators (yellowfin tuna and swordfish) of the western Indian Ocean

Ecologists primarily use δ¹⁵N values to estimate the trophic level of organisms, while δ¹³C, and even recently δ¹⁵N, are utilized to delineate feeding habitats. However, many factors can influence the stable isotopic composition of consumers, e.g. age, starvation or isotopic signature of primary producers. Such sources of variability make the interpretation of stable isotope data rather complex. To examine these potential sources of variability, muscle tissues of yellowfin tuna (Thunnus albacares) and swordfish (Xiphias gladius) of various body lengths were sampled between 2001 and 2004 in the western Indian Ocean during different seasons and along a latitudinal gradient (23°S to 5°N). Body length and latitude effects on δ¹⁵N and δ¹³C were investigated using linear models. Both latitude and body length significantly affect the stable isotope values of the studied species but variations were much more pronounced for δ¹⁵N. We explain the latitudinal effect by differences in nitrogen dynamics existing at the base of the food web and propagating along the food chain up to top predators. This spatial pattern suggests that yellowfin and swordfish populations exhibit a relatively unexpected resident behaviour at the temporal scale of their muscle tissue turnover. The body length effect is significant for both species but this effect is more pronounced in swordfish as a consequence of their different feeding strategies, reflecting specific physiological abilities. Swordfish adults are able to reach very deep water and have access to a larger size range of prey than yellowfin tuna. In contrast, yellowfin juveniles and adults spend most of their time in the surface waters and large yellowfin tuna continue to prey on small organisms. Consequently, nitrogen isotopic signatures of swordfish tissues are higher than those of yellowfin tuna and provide evidence for different trophic levels between these species. Thus, in contrast to δ¹³C, δ¹⁵N analyses of tropical Indian Ocean marine predators allow the investigation of complex vertical and spatial segregation, both within and between species, even in the case of highly opportunistic feeding behaviours. The linear models developed in this study allow us to make predictions of δ¹⁵N values and to correct for any body length or latitude differences in future food web studies.     

Reproduction patterns of four Antarctic octocorals in the Weddell Sea: an inter-specific,shape, and latitudinal comparison

The reproductive patterns of four Antarctic gorgonian species have been investigated. Two of them, Dasystenella acanthina and Thouarella sp., present the bottle-brush-shape type; the other two, Fannyella rossii and Fannyella spinosa, are fan-shaped. Two different reproductive patterns have been observed in D. acanthina and Thouarella sp., which point to two size classes in the frequency distribution of oocytes. This feature indicates a reproduction cycle with overlapping generations, being each of them of more than one year, probably with seasonal spawning. F. rossii and F. spinosa show only one size class of oocytes, which could point towards an annual reproductive cycle. The presence of larvae in the gastrovascular cavities in both Fanyella species and Thoaurella sp. is a sign of a possible larvae release during austral summer. The number of oocytes per polyp ranges from 1.1 ± 0.10 SE to 1.5 ± 0.06 SE, and the size ranges from 50 to 1200 μm. The number of spermatic cysts ranges from 2.6 ± 0.19 SE to 5.0 ± 0.21SE, and their sizes range from 50 to 800 μm. These values are comparable to the reproductive trends and features found in gorgonians from other latitudes, which correspond in some cases with similar morphotypes. These coincident strategies could be related to morphological similarities rather than with latitude or water temperature. The results of this study indicate that octocoral morphology may play a crucial role in determining the reproductive output of these organisms.