Origin of Antarctic Isopoda (Crustacea,Malacostraca)

An analysis of the horizontal and the vertical zonation of the Antarctic Isopoda, combined with knowledge of the geological history of Antarctica and isopod phylogeny, revealed that the isopod family Serolidae and subfamily Arcturinae are likely to have evolved from ancestors that inhabited a cold-temperate Gondwanian province. Antarctic species of other families, such as the Munnopsidae, Nannoniscidae, Desmosomatidae and Ischnomesidae, are likely to have evolved from deep-sea ancestors. It is deduced that emigration of South Patagonian species into the Southern Ocean, although possible, probably did not occur very often. Evolutionary phenomena such as continental-drift vicariance, radiation of species on the continental shelf of Antarctica, and active migration, including submergence and emergence mechanisms are discussed.     

Peculiarities of the geographical and vertical distribution of marine isopods and the problem of deep-sea fauna origin

Data on isopods processed by the author and the literature available on the subject have made it possible to provide specific lists of isopods for coastal zones of cold and cold-temperate regions in both hemispheres and for isopods from depths of more than 2000 m for the entire World Ocean. A comparative analysis shows both faunas to be very similar with respect to family and genus; in this respect they differ considerably from the shallow-water fauna of warm zones. Comparative-anatomic data indicate that the suborder Flabellifera is the most primitive, the Asellota the most specialized group; the data permit the singling out of the more primitive as well as the more specialized families in each suborder under study. Analysis of vertical distributions of all isopods in the World Ocean suggests that primitive isopods are principally confined to tropical shelves; however, in the shelves of cold and cold-temperate waters, especially in deep seas, a considerable number of phylogenetically recent families is present. Thus, the tropical shallow-water fauna proves to be the most ancient, while the shallow fauna of cold and temperate waters may be regarded as geologically more recent. The deep-sea fauna is considered to be the youngest. The author hypothesizes the probable ascendance of the deep-sea fauna from the Antarctic fauna in the period of increasing cold, as early as the late Cretaceous period. This process was promoted by glaciation in the Antarctic Ocean, and its gradual isostatic dipping together with the shelf.     

Possible role of marine bacteria in providing the creosote-resistance of Limnoria tripunctata

Electron microscopic examination of thin-sectioned Limnoria tripunctata from creosoted and untreated wood substrates from Panama, Florida, and laboratory aquaria, coupled with limnorian behavioral studies revealed that bacteria may contribute to the wood boring, nutrition, and creosote-resistance of these isopods. Ingested along with wood fragments and encased in a peritrophic membrane in the isopod intestine, these bacteria, upon lysis, may provide supplemental nutrition to the nitrogen-poor wood diet of L. tripunctata. Ingested material has never been observed in the digestive diverticula of limnorians and the presence of the peritrophic membrane in the isopod intestine has been correlated with feeding. Isopods from creosoted wood differed from those inhabiting untreated wood in that the former contained relatively larger and more diverse bacterial populations both on their exoskeletons and in their gut contents. These isopods harbored bacteria, which apparently bypassed the peritrophic membrane and lived in association with the isopod intestinal lining. These gut-associated bacteria were lost when laboratory isopods reared on creosoted wood were transferred to untreated wood. Laboratory isopods reared solely on creosoted or untreated wood were exposed to creosoted and untreated wood substrates, both sterilized and unsterilized. Boring and mortality data from these studies indicated that the creosote-reared population had a microbial flora whose activity facilitated isopod colonization of creosoted substrates. We present the hypothesis that creosote hydrocarbons provide nutrition for isopod-associated bacterial populations and that L. tripunctata may benefit from a concomitant bacterial detoxification of the creosote.     

Heteroplasmy in a deep-sea protobranch bivalve suggests an ancient origin of doubly uniparental inheritance of mitochondria in Bivalvia

Most metazoan species have strict maternal inheritance of the mitochondrial genome. In bivalves, a unique inheritance pattern called doubly uniparental inheritance (DUI) occurs in at least seven bivalve families. In this system of mitochondrial inheritance, males inherit and carry mtDNA from both parents, while females only carry mtDNA from the mother. Here, we present evidence of mitochondrial heteroplasmy in deep-sea protobranch bivalves. Divergent 16S rRNA and cytochrome b sequences were obtained within individuals of Ledella ultima. Ledella sublevis also exhibited divergent 16S sequences. Levels of divergence between 16S sequences within individuals were 27 and 15 % for each species, respectively. Ratios of homoplasmic to heteroplasmic individuals were not significantly different from 1:1, in agreement with sex ratios in protobranchs. The results provide the first evidence for mitochondrial heteroplasmy in the protobranchs and suggest DUI might have evolved much earlier in the evolution of the Bivalvia than previously thought.     

Health and population-dependent effects of ocean acidification on the marine isopod Idotea balthica

Three populations of the grazing isopod Idotea balthica were exposed to high CO₂ treatment for a period of 20 days to investigate the effect of ocean acidification (OA) on animal health and immunocompetence. The results of the populations from more saline habitats were comparable and showed a 60–80 % decrease in immune response as a result of the high CO₂ treatment. Analysis of protein carbonyls showed no treatment effect, indicating that short-term OA does not increase oxidative protein damage. Meanwhile, the third tested population from the lower saline Baltic Sea had higher background protein carbonyl levels. Ocean acidification in addition to this resulted in 100 % mortality. The results of this study show that OA reduced immunocompetence of this marine isopod. In addition, populations and individuals in poor health are potentially at greater risk to succumb under OA.     

Mineralization of major lateral teeth in the radula of a deep-sea hydrothermal vent limpet (Gastropoda:Neolepetopsidae)

Radular teeth of a neolepetopsid patellogastropod limpet, Paralepetopsis ferrugivora, from a deep-sea hydrothermal vent site have similar elemental composition (O, Si, P, S, Cl, K and Fe) to radular teeth of the shallow-waters patellogastropod limpets. However, in contrast to shallow waters limpets, the fully mature teeth of P. ferrugivora do not show any crystalline phases. Amorphous silica was found in the cusp of the teeth and amorphous iron oxide in the junction zone and base of the teeth. Ferritin-rich vesicles were observed in cells adjacent to the junction zone of the early mature teeth, suggesting that these vesicles can mediate the delivery of iron to the tooth matrix. The similarity in the elemental composition between the hydrothermal limpet P. ferrugivora and shallow-waters patellogastropod limpets shows that the extreme hydrothermal environment did not alter the elemental composition of the radular teeth in the deep-sea species.Communicated by P. W. Sammarco, Chauvin     

Temperature and pressure tolerance of larvae of Crepidula fornicata suggest thermal limitation of bathymetric range

The extant deep-sea fauna is thought to result from recolonisation of this environment by shallow-water organisms following climate-driven mass extinctions. Planktonic larval tolerance to high pressure is considered an important preadaptation for successful deep-sea invasion. In this study, the pressure and temperature tolerance of a species without any known confamilial deep-sea relative were assessed for the first time. Early- and late-veliger larvae of the shallow-water species Crepidula fornicata were subjected to a temperature/hydrostatic pressure regime from 5 to 25 °C and from 0.1 to 40 MPa. Although early and late veliger survived pressures equivalent to 2,000 m water depth or greater at all temperatures, decreased larval activity indicated significant sublethal temperature and pressure effects. Reduced larval activity of early veliger at low temperatures suggests that the bathymetric range of this species may be thermally constrained. A mechanistic model is proposed to explain the emerging pattern of ontogenetic shifts in pressure tolerance of shallow-water benthic invertebrates.     

Sensory brain areas in three families of deep-sea fish (slickheads, eels and grenadiers): comparison of mesopelagic and demersal species

The sensory brain areas of a sample of <100 deep-sea fish species were studied, and the relative volumes of the olfactory bulb, optic tectum, octavolateral area and gustatory area were determined. In the absence of direct observations on the behaviour of this ichthyofauna these data allow deductions to be made about the kinds of sensory modalities used preferentially in the deep-sea environment. Here, members of three families are compared that have representatives living on or near the sea floor (“demersal”) and in the open water between 200 and 1.000 m (mesopelagic). The findings indicate that both regions present fish with rich and diverse sensory environments. While vision emerges as the dominant sense of the mesopelagic realm, olfaction seems more important on or near the bottom of the sea. However, other sensory modalities supplement these senses in species-specific patterns. Considerations of the phyletic relationships indicate different degrees of pervasiveness. Whilst in slickheads the dominance of vision appears to be a family-related trait, a similar relationship is not found in either eels or grenadiers. By contrast, the common trait in these two families seems to be the greater adaptability to the environment. Published online: 21 August 2002     

Parasitism and ecological relationships among deep-sea benthic fishes

We have studied the metazoan parasite fauna of 52 species of deep-living benthic fishes from depths of 53 to 5000m off the New York Bight (39–49°N; 70–72°W). 17144 parasites were recovered from 1712 fishes. The infestation rate was 80%, with an average of 12.5 worms per host. Percentage occurrence by group among all fishes was Monogenea 12.9%, Digenea 48%, Cestoda 22.1%, Nematoda 54.5%, Acanthocephala 3.8%, and Copepoda 4.5%. Differing composition of the parasite fauna in different fish species reflects differences in diet. Specialized feeders are rather distinct; generalized feeders, which predominate, show overlaps in parasite fauna. In individual species, changes in diet with growth are reflected in changes in the parasite fauna. Infestation rate is directly related to abundance of the free-living fauna; hence, fish from within the submarine canyon are more heavily infested than those living without. Although it contains fewer families and genera than shallow faunas, the deep-sea parasite fauna is not extremely unusual in terms of its abundance, diversity, or host specificity. At the greatest depths, parasite abundance and diversity dramatically decline.     

Effect of temperature on the microdistribution of the isopod Sphaeroma rugicauda from a saltmarsh habitat

The saltmarsh isopod Sphaeroma rugicauda (Leach) is subjected to widespread diurnal and seasonal temperature fluctuations under natural conditions. Laboratory studies on its activity show that there is a relationship between behaviour activity and exposure temperature between 2.5° and 25°C. Although S. rugicauda has no complex metabolic adaptations, this isopod is able to maintain a rate of aerial oxygen consumption which is similar to that in water within the temperature range 5° to 25°C. The responses of S. rugicauda to changes in environmental temperature are discussed in relation to the seasonal microdistribution of the isopod in the salt-marsh habitat.     

Depth-related adaptations, speciation processes and evolution of color in the genus Phyllidiopsis (Mollusca: Nudibranchia)

The nudibranch genus Phyllidiopsis (Phyllidiidae) contains 30 currently recognized species, all of them distributed throughout the tropical Indo-Pacific, eastern Pacific, Northwest Atlantic and Caribbean Sea. Half of the known species of Phyllidiopsis inhabit deep waters, and most of the deep-sea species of the Phyllidiidae belong to this genus. There is no definitive explanation for the high diversity of Phyllidiopsis in the deep-sea or for whether diversity could be related to particular adaptations of this group or to historical events. In light of phylogenetic analysis, several cases of vicariance have been detected in this genus. Apparently two major vicariant events occurred between the tropical Indo-Pacific region and the Atlantic-eastern Pacific area first and subsequently between the eastern Pacific and the Atlantic. Vicariant events could also be involved in producing vertical distributional patterns in a few species of Phyllidiopsis. The scarcity of phyllidiids in the Atlantic Ocean may be explained by historical events, including isolation and subsequent extinction in shallow waters. There is a mimicry species complex in Phyllidiopsis, including several members of a clade that probably acquired this coloration through common ancestry, and also including another unrelated species that probably acquired this coloration through convergent or parallel evolution. There is also a group of white species, lacking any other contrasting colors, that inhabits deep waters. This coloration could constitute an adaptation to the deep-sea environment and not a mimicry complex. In this case, all species acquired this coloration through common ancestry.     

How does burrowing by the isopod <Emphasis Type="Italic">Limnoria agrostisa</Emphasis> (Crustacea: Limnoriidae) affect the leaf canopy of the southern Australian seagrass <Emphasis Type="Italic">Amphibolis griffithii</Emphasis>?

In south-western Australia, the isopod Limnoria agrostisa commonly burrows into leaf clusters and immature shoots of Amphibolis griffithii. The isopod also burrows into the sheath and rhizomes of Posidonia species. In A. griffithii, the isopod consumes new tissue within the sheath, damaging or destroying the meristem. This results in malformation of new leaves or destruction of whole leaf clusters with the potential to reduce the photosynthetic area of a shoot. The isopod has been found in all but one meadow of A. griffithii examined over 1,000 km of the Western Australian coastline. It was present throughout the year and showed little variation in abundance. Young were produced year round, but were more abundant in the summer months. Females, 3.5 mm in length or larger, produced 2–5 young that were brooded within the leaf cluster or base of an immature shoot. Within a meadow, 40–70% of shoots and 10–20% of leaf clusters were damaged by isopods. Seasonal trends were not consistent, but damage appeared to be higher in summer when isopod abundance was higher. Approximately 40% of clusters were destroyed by isopod damage. Isopods attack shoots of all ages, but damage was often located on apical clusters. There was no evidence that isopod damage initiated branching or leaf cluster formation. Estimations of clusters damaged or destroyed may be conservative, as only those clusters remaining on a shoot could be counted, and cluster loss could not be quantified. Examination of defoliated stems on upright shoots and horizontal rhizomes indicated that many were destroyed by isopods. The effect of L. agrostisa although substantial appears to be a feature of healthy seagrasses throughout southern Australia.     

Genetic population structure in the black-spot sea bream (<Emphasis Type="Italic">Pagellus bogaraveo</Emphasis> Brünnich, 1768) from the NE Atlantic

The depletion of shallow-water fish stocks through overexploitation has led to increasing fishing pressure on deep-sea species. Poor knowledge of the biology of commercially valuable deep-water fish has led to the serial depletion of stocks of several species across the world. Data regarding the genetic structure of deep-sea fish populations is important in determining the impact of overfishing on the overall genetic variability of species and can be used to estimate the likelihood of recolonisation of damaged populations through immigration of individuals from distant localities. Here the genetic structure of the commercially fished deep-water species the blackspot sea bream, Pagellus bogaraveo is investigated in the northeastern Atlantic using partial DNA sequencing of mitochondrial cytochrome b (cyt-b) and D-loop regions and genotyping of microsatellite loci. An absence of variation in cyt-b and low genetic variation in D-loop sequences potentially indicate that P. bogaraveo may have undergone a severe bottleneck in the past. Similar bottlenecks have been detected in other Atlantic species of fish and have possibly originated from the last glaciation. P. bogaraveo may have been particularly vulnerable to the effects of low temperature and a fall in sea level because stages of its life history occur in shallow water and coastal sites. However, there are other explanations of low genetic variability in populations of P. bogaraveo, such as a low population size and the impacts of fishing on population structure. Analysis of population structure using both D-loop and microsatellite analysis indicates low to moderate, but significant, genetic differentiation between populations at a regional level. This study supports studies on other deep-sea fish species that indicate that hydrographic or topographic barriers prevent dispersal of adults and/or larvae between populations at regional and oceanographic scales. The implications for the management and conservation of deep-sea fish populations are discussed.Communicated by J.P. Thorpe, Port Erin     

Association of Syscenus infelix (Crustacea: Isopoda: Aegidae) with benthopelagic rattail fishes, Nezumia spp. (Macrouridae), along the western North Atlantic continental slope

During submersible surveys along the continental slope (summers of 1991 and 1992, 184–847 m) between False Cape, Virginia, and Cape Hatteras, North Carolina, USA, we observed the aegid isopod, Syscenus infelix Harger, attached to the macrourid Nezumia bairdii (Goode and Bean). This is the first report of S. infelix attached to fishes in the western North Atlantic. The association of this blind isopod with its host appears species specific. The large, conspicuous isopod always attached to a fish in the same location, the dorsal midline, immediately behind the first dorsal fin. Attachment appears to be long term, with the isopod forming a characteristic scar consisting of a distinct discolored oval depression with seven small, dark impressions that coalesce as the fish grows. Only one S. infelix was found on each host fish. The isopod occurred on 23.7% of N. bairdii observed from submersible on the middle continental slope off Virginia and North Carolina, compared with 16.6% of 1236 museum specimens of the same species (based on inspection for scars) collected at latitudes 26°–64°N. Prevalence of the fish–isopod association was not correlated with depth or latitude. We also found identical scars on preserved specimens of N. aequalis (2.6% of 660 specimens), N. sclerorhynchus (1.2% of 86 specimens), and N. suilla (14.3% of 7 specimens), mostly from areas outside the range of N. bairdii. No scars were found on museum specimens of N. atlantica (n=27), N. cyrano (n=57), or N. longebarbata (n=7). The low incidence of isopod attachment on these species suggests that N. bairdii is the preferred host. Infestation by the isopod appears to result in erosion of host fish scales and tissue. We propose that S. infelix is an obligate associate of its host fish and should be considered parasitic. Received: 9 June 2000 / Accepted: 21 October 2000     

Growth of the deep-sea irregular sea urchins Echinosigra phiale and Hemiaster expergitus in the Rockall Trough (N.E. Atlantic Ocean)

Growth in the deep-sea irregular sea urchins Echinosigra phiale (family Pourtalesiidae) and Hemiaster expergitus (family Hemiasteridae) was studied from deep-sea samples taken during the years 1973 to 1985 from two stations at 2900 and 2200 m depth in the Rockall Trough (N.E. Atlantic Ocean). Growth zones, similar to those described from sea urchins in shallow water, are present as a series of wide white bands separated by narrow, dark rings in the calcite stereom of the test plates after heating to 350°C. In shallow water, such growth zones seem to result from seasonally varying growth rates. In the supposedly constant conditions in the deep sea, a seasonal growth pattern is unexpected but may occur in response to recently discovered annual pulses in downward flux of detritus from the euphotic zone, providing a seasonally varying food supply for such deposit-feeding species living in the bottom sediment. On this assumption, growth curves were fitted to counts of growth zones (as representing age in years), in the larger lateral and ventral test plates of E. phiale and H. expergitus. The opportunity was also taken to fit growth curves derived from counts of growth zones in samples of the inshore spatangoids Spatangus purpureus and Echinocardium pennatifidum. Plots of counts against test length of Echinosigra phiale and H. expergitus, although scattered and not clearly asymptotic, indicate, growth to be slower than in the two inshore spatangoids, and than in the coastal species Echinocardium cordatum, for which there are good recent growth data, available.     

Short- and long-term importance of small sharks in the diet of the rare deep-sea shark Dalatias licha

Knowing the trophic ecology of marine predators is essential to develop an understanding of their ecological role in ecosystems. Research conducted on deep-sea and threatened shark species is limited. Here, by combining analyses of individual stomach contents and stable isotope values, we examined the trophic ecology (dietary composition and trophic position) of the kitefin shark Dalatias licha, a deep-sea shark considered as near threatened globally and as data deficient in the Mediterranean Sea. Results revealed the importance of small sharks in the diet of the kitefin shark at short- and long-term scales, although fin-fish, crustaceans and cephalopods were also found. Predation on sharks reveals the high trophic position of the kitefin shark within the food web of the western Mediterranean Sea. Stable isotope values from liver and muscle tissues confirmed our results from stomach content analysis and the high trophic position.     

Importance of macroalgal fields as coral reef fish nursery habitat in north-west Australia

Macroalgal fields are a feature of the shallow tropical benthos, yet their importance for coral reef fish population dynamics remains poorly understood. The abundance of fish recruits was recorded using underwater visual census at six macroalgal and 11 coral reef sites in the Montebello and Barrow Islands. Surveys identified 6,935 individual recruit fish from 105 species, 54 genera and 20 families. Of these, 1,401 recruits from 48 species, 31 genera and 14 families were observed in macroalgal sites. Sixteen of the 105 recruit species (15.2 %) were observed exclusively at macroalgal sites. Forty-two (87.5 %) of these species have been observed as adults on adjacent coral reefs. Species composition of fish recruits differed significantly between the two habitats. Corallivore, small omnivore and zooplanktivore recruits had significantly higher numbers in the coral sites, while the results clearly demonstrate that juveniles, within the genera Lethrinus and Choerodon, as well as large algal croppers, are predominantly found at macroalgal (74–100 %) rather than coral-dominated sites. High-canopy macroalgae cover was positively correlated with abundance of these taxa, particularly Lethrinids (r ² = 0.40). This study is the first to highlight the important attributes of tropical macroalgal fields and suggests that they have a similar role to seagrass meadows as essential juvenile habitat, thus warranting greater attention in conservation planning and ecological studies.     

Occurrence of a cold-water coral along natural pH gradients (Patagonia, Chile)

Increasing dissolution of anthropogenic-released carbon dioxide into the world’s oceans is causing ocean acidification (OA). OA is thought to negatively affect most marine-calcifying organisms, notably cold-water corals (CWC), which may be especially sensitive due to the deep and cold waters they normally thrive in. However, the impact of OA on CWC is difficult to predict. Recorded distributions of CWC are rarely linked to in situ water chemistry, and the boundaries of their distributions are not clearly defined. The fjord Comau in Chilean Patagonia features pronounced pH gradients, and up to 0.5 pH units have been recorded both vertically (at some sites within 50 m depth) and less distinct horizontally (from head to mouth). The cosmopolite coral Desmophyllum dianthus grows along the course of the fjord and of the entire pH range. It occurs in shallow depths (below 12 m, pH 8.1) as part of a deep-water emergence community, but also in 225 m depth at a pH of 7.4. Based on pH and total alkalinity, data calculations of the associated carbonate chemistry revealed that this CWC thrives commonly close the aragonite (the orthogonal crystal form of calcium carbonate, the mineral structure of coral skeletons) saturation horizon and even below. This suggests a high adaptation potential of D. dianthus to adjust its calcification performance to conditions thermodynamically unfavourable for the precipitation of aragonite.     

Morphological divergence of Eastern Pacific and Caribbean isopods: effects of a land barrier and the Panama Canal

The Isthmus of Panama rose approximately 3 million years before the present (mybp) and isolated biotas in the tropical eastern Pacific from those in the Caribbean Sea. Populations that were split by the Isthmus and have evolved in allopatry since that time are known as geminates. The surf zone/beach isopod Excirolana braziliensis Richardson was examined between 1984 and 1989 to test the hypothesis that divergence in geminate isopod morphology has occurred, and that geminate divergence is greater than divergence between local populations from the same coastline. Three morphs of Excirolana braziliensis, one in the Caribbean and two in the eastern Pacific, were discovered using numerical taxonomic methods that adjust for body size. The two Pacific morphs have overlapping large-scale distributions, but those morphs are segregated on a smaller scale by beach. We inferred that one Pacific morph and one Caribbean morph were geminates, based on their relative similarity in shape, their geographical ranges, and natural history information about the organism's dispersal capabilities. The origin of the third morph probably predates the Isthmus of Panama, given its relative dissimilarity from the geminate morphs. The presumed geminates differ primarily with respect to the rostrum, antennae and one male reproductive structure. Divergence between geminates is greater than divergence between local populations of any morph along a coastline. Because only one morph occurs in the Caribbean, that region contains less morphological variation than the eastern Pacific, which contains two morphs. There was weak evidence that some introductions may have taken place in the last century from the Caribbean to the Pacific; however, introductions have not masked the pattern of divergence that has developed over millions of years.