The fractal branching of an arborescent sponge

The fractal properties of specimens of a planar branching sponge Raspailia inaequalis (Porifera, Demospongiae) were determined by analysing digitised photographs. The specimens, collected from a single site in northeastern New Zealand, had a wide range of morphology. Three different fractal methods were used: box counting; a method that gives the scaling of branch length with distance from the base of the fan; and an allometric analysis of the dependence of frontal area on specimen size. All three approaches gave a similar value for the fractal dimension. The conjecture that the specimens have a fractal branching structure is consistent with the results of a Horton analysis of their branching pattern. There is a significant relationship between fractal dimension and number of fingers, which implies that a simple count of the number of fingers is as useful for discriminating between individuals as the more complex fractal analysis. Using this relation, sponges from a site with less water movement are inferred to have a lower fractal dimension. This result is in agreement with the predictions of the Kaandorp model of sponge growth. Received: 13 March 2000 / Accepted: 11 October 2000     

Growth of zooxanthellae in culture with two nitrogen sources

Physiological characteristics of zooxanthellae were examined under nutrient-saturated conditions created by mixing ammonium (¹⁵NH₄) with nitrate (¹⁵NO₃) to give 0.88 mM total nitrogen. Growth rate varied with the form of nitrogen provided. Ammonium alone resulted in the lowest C:N and C:chl-a ratios. Although zooxanthellae took up nitrate in the absence of ammonium, ammonium assimilation was 1.3 times higher than nitrate assimilation. Ammonium strongly inhibited nitrate assimilation. While high-ammonium treatments resulted in the highest ¹⁴C incorporation into intermediate compounds, high nitrate levels resulted in the highest ¹⁴C incorporation into protein, suggesting that the intermediate compounds are produced prior to the subsequent production of protein when ammonium is the dominant N source. The enhanced production of intermediate compounds at the expense of carbon directed to protein synthesis in the presence of ammonium might be analogous to the “host factor” observed in zooxanthellae–host symbioses, since growth rate is depressed due to low production of protein. Received: 16 March 2000 / Accepted: 26 August 2000     

Foraging behavior of juvenile Carcinus maenas (L.) and Cancer pagurus L.

Information concerning the way juvenile crabs choose their diet from a variety of prey types can be useful for a better understanding of community dynamics, as well as for the adequate management of natural resources. Prey size and species selection by juvenile Carcinus maenas (15-35 mm carapace width, CW) and Cancer pagurus (20-40 mm CW) feeding on four bivalves of contrasting shell morphology were investigated. When offered a wide size range of Mytilus edulis, Ostrea edulis, Crassostrea gigas, and Cerastoderma edule presented individually, crabs generally showed evidence of size-selective predation. Cancer pagurus selected larger mussels relative to the size of their chelae (relative prey size, RPS) than did Carcinus maenas of similar and even larger carapace width. However, the RPS of selected O. edulis and Cerastoderma edule were similar for all crabs, suggesting that certain prey features constitute effective barriers even to the powerful chelae of Cancer pagurus. When offered a wide size range of mussels and oysters simultaneously, all crabs consistently selected mussels. When offered O. edulis and Crassostrea gigas, crabs consumed both these oyster species in similar numbers. Carcinus maenas consumed similar numbers of mussels and cockles; Cancer pagurus, however, showed no preference for either prey in the smaller size classes but selected more mussels than cockles as prey increased in size. Although previous studies report that adult Carcinus maenas select prey species according to their profitability (amount of food ingested per unit of handling time, milligrams per second), consumption rates of the size classes of prey selected by juvenile shore crabs did not always parallel prey value. Although variations in crab strength can account for many of the differences between the foraging strategy of juvenile and adult C. maenas, our results suggest that juvenile crabs are less species selective than adults as a result of the restrictions imposed on small individuals that have limited access to larger prey.     

The ichthyofaunal composition of the Elbe Estuary: an analysis in space and time

A stow net (90 m² opening) was used, at regular intervals between July 1989 and June 1993, to collect fish over ca. 4 h on both the flood and ebb tides at five sites distributed at regularly spaced distances throughout the Elbe Estuary. Overall, 53.4% of the species were marine, and some of these were abundant, e.g.Clupea harengus, Pleuronectes flesus, Pomatoschistus minutus, Merlangius merlangus and Sprattus sprattus. However, in terms of number of individuals, such species contributed only 9.1%, compared with 90.0% by anadromous species, which was predominantly due to exceptionally high catches of Osmerus eperlanus. The ichthyofaunal compositions on the flood and ebb tides were similar, even downstream where salinities change markedly during each tidal cycle. Although this is probably due in part to groups of fish being swept first upstream on the flood tide and then back downstream on the ebb tide, several species clearly tended to occupy particular regions of the estuary. Each year the species composition changed sequentially from the most downstream site (max. salinity=ca. 31.4‰) to the most upstream site (max. salinity=ca. 1.5‰), mainly as a result of a sequential decline in the number and abundance of marine species and a progressive increase in the contributions of both anadromous and freshwater species. However, the marine species diagnostic of differences in the compositions among regions varied between years. Thus, for example, the diagnostic species for the downstream sites were C. harengus and M. merlangus in 1990/1991 and S. sprattus in 1992/1993, reflecting differences in the recruitment strengths of these marine species in the two years. The ichthyofaunal composition exhibited pronounced annual cyclical changes in each region and thus occurred irrespective of whether or not the salinity in a region underwent pronounced changes during the year. This cyclicity was attributable to intra-annual variations in the times of recruitment of certain marine species, particularly in the downstream region, and to seasonal migrations of diadromous species into each region, as well as to the movements of freshwater species into the upstream region in winter and early spring. Received: 18 November 1999 / Accepted: 25 October 2000     

Hepatopancreatic endosymbionts in coastal isopods (Crustacea: Isopoda), and their contribution to digestion

Three isopod species (Crustacea: Isopoda), commonly found in the intertidal and supratidal zones of the North American Pacific coast, were studied with respect to symbiotic microbiota in their midgut glands (hepatopancreas). Ligia pallasii (Oniscidea: Ligiidae) contained high numbers of microbial symbionts in its hepatopancreatic caeca. Numbers of endosymbionts were strongly reduced by ingestion of antibiotics. By contrast, the hepatopancreas of Idotea wosnesenskii (Valvifera: Idoteidae) and Gnorimosphaeroma oregonense (Sphaeromatidea: Sphaeromatidae) did not contain any microbiota. Results of feeding experiments suggest that microbial endosymbionts contribute to digestive processes in L. pallasii, the most terrestrial of the three isopods that we studied. The acquisition of digestion-enhancing endosymbionts may have been an important evolutionary step allowing isopods to colonize terrestrial habitats where relatively indigestible leaf litter is the primary food source. By contrast, the ability to digest phenolic compounds was most developed in one of the more marine species, suggesting that this trait may have evolved independently in isopod species that consume a phenolic-rich diet, whether in marine habitats or on land. Received: 28 August 2000 / Accepted: 8 December 2000     

Population structure of yellowfin tuna (Thunnus albacares) in the western Pacific Ocean, inferred from microsatellite loci

Five polymorphic microsatellite loci were examined in 1391 yellowfin tuna (Thunnus albacares) from eight regions of the western (Coral Sea, eastern Australia, Fiji, Indonesia, Philippines and Solomon Islands) and eastern (California and Mexico ) Pacific Ocean. Across all samples, numbers of alleles per locus ranged from 7 to 30 (mean: 17.0), and observed heterozygosities per locus ranged from 0.223 to 0.955 (mean: 0.593). Temporal collections were available for three areas: no significant temporal heterogeneity was observed for the Coral Sea (1991/1992 and 1995/1996 collections) or eastern Australia (1994/1995, 1995/1996, 1996/1997 and 1997/1998), but there was slight but significant heterogeneity at one locus (cmrTa-161) between the two Philippines collections (1994/1995 and 1996/1997). Genotypes generally showed a good fit to Hardy-Weinberg expectations within populations; only cmrTa-208 in the pooled Coral Sea population gave a significant deviation after Bonferroni correction for 40 tests, with a small but significant excess of homozygotes. Four loci showed no evidence of population differentiation following contingency Chi-squared and F_ST analyses. The fifth locus, cmrTa-161, showed small but significant differentiation (F_ST=0.002, P<0.001). This heterogeneity was largely a result of the Philippines 1994/1995 and Fiji collections; there was no correlation with geographic distance. The average F_ST across all five loci was very low (F_ST=0.002), but it was significant (P<0.001). It is unclear whether this low but significant differentiation reflects noise in the dataset, perhaps arising from experimental error, or real population differentiation. The finding of very limited population heterogeneity accords with most of the earlier allozyme and mitochondrial DNA studies of yellowfin tuna in the Pacific Ocean.     

Ecology of subtropical hermit crabs in S.W. Madagascar: cluster structure and function

The Madagascar coast (both) has a higher density and diversity of hermit crabs than is known from any other locality in the western Indian Ocean. Of the 20 species occurring at Anakao (S.W. Madagascar), 11 aggregated into clusters, including all but one of the species above the subtidal zone. The mean number of hermit crabs and species in clusters varied with several spatial parameters and time. Over 80% of the community clustered in certain habitats in particular tidal zones, whilst as low as 3% clustered in others. The highest intensity of clustering with shore zone was coincident with peak numbers of hermit crabs. The initiation and duration of clusters of hermit crabs above the eulittoral was driven by circadian rhythms, whilst those in the eulittoral were governed by tidal state. Clusters above and below the eulittoral were longer in duration, and those in the subtidal were more temporally variable than those above it. Certain pairs of species showed positive correlations of occurrence and (more rarely) of abundance, and all the correlations (of occurrence) of one, Clibanarius eurysternus, were negative. Positive correlations of occurrence were related to the degree of shell-use commonality between species pairs. Eulittoral species clustered with other individuals of approximately similar size and exchanged shells upon cluster disintegration. There was evidence of a dehydration-reduction function to shell clustering in addition to shell-exchange facilitation. The highly variable species-specific strategies of clustering may be important in alleviating both intra- and interspecific competition in assemblages of similar and highly abundant species.     

Feeding, growth and reproduction of sea urchins (Strongylocentrotus droebachiensis) on single and mixed diets of kelp (Laminaria spp.) and the invasive alga Codium fragile ssp. tomentosoides

Since its introduction to Nova Scotia in the early 1990s, the invasive green alga Codium fragile ssp. tomentosoides has spread rapidly to become a dominant and persistent component of seaweed assemblages in the rocky subtidal zone. To examine the potential of sea urchins (Strongylocentrotus droebachiensis) to regulate Codium, and the potential of the alga to support urchin growth and reproduction, we fed urchins Codium and/or kelp (Laminaria spp., a high quality and preferred native food) in the laboratory for 11 months. Urchins showed a strong and active preference for kelp: they consumed more Laminaria than Codium (as dry weight) in single and mixed diet treatments. Urchins fed only Codium consumed 3.5 times more of the alga than those on a mixed diet, but did not increase their feeding rate in winter when kelp consumption was greatest. Laminaria was consumed at the same rate in single and mixed diets, indicating that the presence of an alternative food had no effect on kelp consumption. Survival and growth (change in test diameter) were lowest on the Codium diet, although the differences among diets were not statistically significant. Urchins on the Codium diet showed no gonadal production over the annual reproductive cycle, compared to a marked rise in gonad index on the Laminaria and mixed diets. Our results indicate that Codium is an unattractive, unpalatable and poor quality food, which is not readily consumed by urchins. Thus we predict that urchins at moderate densities will be much more likely to graze kelp than Codium in mixed stands, creating a mosaic of Codium patches and barren areas. At high densities, however, urchins are likely to destructively graze all seaweeds, although dense stands of the invasive alga may prolong the widespread formation of barrens.     

Molecular response of the sponge Suberites domuncula to bacterial infection

The aim of this study was the documentation of the molecular immune response of Suberites domuncula upon bacterial infection. Additionally, the bacteria that are naturally present in the sponge after prolonged aquarium maintenance were characterized. After 6 months of maintenance of S. domuncula in seawater aquaria, only one bacterial 16S rDNA sequence could be recovered, which belongs to the genus Pseudomonas. Concomitantly, morphologically uniform bacteria were found encapsulated in bacteriocytes. These findings indicate that certain bacteria, possibly of the genus Pseudomonas, are able to persist for long periods in host bacteriocytes. Subsequent to performing a previously established infection assay with S. domuncula, a potentially pathogenic Vibrio sp. was isolated from the tissues. Furthermore, the host tissue disintegrated and asexual propagation bodies (gemmules) were formed. In order to gain insights into the molecular events occurring after bacterial infection, the stress-response kinases, p38 protein kinase and JNK protein kinase, were analyzed. It is demonstrated that these two kinases are activated (phosphorylated) upon incubation of the tissue with the bacterial endotoxin lipopolysaccharide (LPS). Moreover, LPS strongly inhibits protein synthesis. It is concluded that there are many functionally different interactions between S. domuncula and bacteria and that the animal possesses mechanisms to differentiate between bacteria and to respond accordingly.     

Genetic, morphological, and chemical divergence in the sponge genus Latrunculia (Porifera: Demospongiae) from New Zealand

We undertook a comprehensive study of Latrunculia in New Zealand to determine the relationship between taxonomic, environmental, and chemical variation within the genus. Sponges were collected from five locations around New Zealand: Three Kings Islands, Tutukaka, Wellington, Kaikoura, and Doubtful Sound. Allozyme electrophoresis at nine polymorphic loci indicated that sponges from each geographic location were genetically distinct, and that they displayed genetic differences of the magnitude usually associated with reproductively isolated species (Nei's D between locations =0.375-2.476). Additionally, the comparisons revealed that the green and brown colour morphs of Latrunculia that are sympatric at Three Kings Islands and Kaikoura are distinct from each other, and that there are two genetic groups within the green sponges in Doubtful Sound. On the basis of genetic data we conclude that there are at least eight species of Latrunculia in New Zealand waters, not one to four as had been previously thought. Morphological comparisons of the eight genetic species based on skeletal characters (i.e. skeletal organisation of the choanosome, spicule composition, size, and geometry) indicated that the eight Latrunculia species fell into only two morphological groups that could be easily diagnosed on the basis of discorhabd type. Within these two primary morphological groups, skeletal characteristics among the eight species largely overlap and are not diagnostic. These findings emphasise the limitations of traditional taxonomic methods based solely on skeletal characters for distinguishing species of Latrunculia. However, multivariate analysis (MANOVA and CDA) based on six measured skeletal variables did reveal significant morphological variation among the species (Pillai's Trace=3.28, F=6.90, P<0.0001), supporting the division of the genus into eight species. Comparisons of chemical extracts from Latrunculia also showed that the amounts of five different bioactive compounds (discorhabdins A, B, C, D, and J) varied predictably among the eight species. This finding suggests that discorhabdin variation within Latrunculia, previously thought to be associated with intra-specific environmental variability, is more likely to reflect differences among species rather than phenotypic plasticity. Our results also highlight the importance of thorough taxonomic studies associated with marine natural products research to understand fully the variation in bioactive properties among individuals. The potential processes underlying the unusually high speciation rates in New Zealand Latrunculia that are indicated in our study are discussed.