Holistic Pattern Analysis as an Alternative to Pattern Cladistics in Hypothesizing Crustacean Phylogenetic Sequences

There has been a growing realization that ‘true’ phylogenetic relationships cannot be determined using pattern cladistical methods alone. The applications of such techniques to the problem of crustacean evolution is first examined using the results presented by Wilson (in this issue). It is found that the degree of correlation of characters in the head vs trunk regions explains very well the grouping of taxa in the cladograms, suggesting that the groups are nothing more than mathematical constructs. The argument is presented that functions of different parts of the crustacean body need to be determined, along with an understanding of how the body morphology is designed in accordance with the dominant functions. A method for producing hypothesized phylogenetic sequences from an analysis of body morphology is proposed and examined using the amphipod family Iphimediidae. It is suggested that cladograms are merely hypotheses and can be tested using functional morphological information of the type outlined.     

Patterns of gammaridean amphipod abundance and species composition associated with dominant subtidal macroalgae from the western Antarctic Peninsula

The communities of gammaridean amphipods associated with eight dominant macroalgal species were examined near Palmer Station, Western Antarctic Peninsula. A total of 78,415 individuals belonging to 32 amphipod taxa were identified with mean densities ranging up to 20 individuals/g algal wet wt. The most abundant amphipod taxon, Metaleptamphopus pectinatus, was found to associate predominately with the brown alga Desmarestia menziesii, while the second most common taxon, Jassa spp. occurred primarily on the red alga Gigartina skottsbergii. Non-metric multidimensional scaling analysis demonstrated that the population densities of each amphipod species and amphipod species composition were similar on the same algal species but dissimilar on different species of algae. Comparisons of amphipod communities associated with a given algal species but from different sampling sites indicated that although the structure of species-specific macroalgal-associated amphipod communities can vary across spatial scales of 3 km, 50% of the macroalgal species examined showed no significant inter-site differences in associated amphipod community structure. Spearman rank correlation analyses showed that higher abundances of amphipods occurred on the macroalgae with the highest number of branches. As many Antarctic amphipods are known consumers of macroalgae, their remarkable abundances are likely to play a significant role in mediating energy and nutrient transfer in nearshore Antarctic Peninsular macroalgal communities.     

Potential impact of the main benthic amphipods on the eastern Weddell Sea shelf ecosystem (Antarctica)

As they represent one of the most diversified taxonomic groups on Antarctic bottoms, amphipods are likely to play a complex role in biogeochemical fluxes that occur within benthic ecosystems. The aim of this paper is to present, using both digestive tract analyses and relative species abundance data, the impact of amphipod feeding on different potential preys of the Weddell Sea. The study is based on data obtained for 29 representative amphipod species collected at 130 stations distributed along the eastern shelf of the Weddell Sea (depth range: 60-2,000 m) during three summer cruises, from 1989 to 1998. Sedimenting plankton particles (10-27%), crustaceans (22-32%) and fish carrion (5-18%) are the main food resources. Other abundant potential preys, such as molluscs or tunicates, do not seem to be consumed. Variations in the proportions of the different preyed food items are observed, mainly related to differences in relative amphipod species composition in samples. Presented results will help in refining ecological models of the prospected area, but also underline the need for accurate and reliable measurements of the feeding rates of Antarctic benthic organisms.     

Trophic diversity within the eastern Weddell Sea amphipod community

Amphipod crustaceans form one of the most diversified animal groups within the Antarctic macrozoobenthos, both from the taxonomic point of view (more than 800 species have been recorded in the Southern Ocean) as by niche occupation and at the community level. Thus, amphipods are likely to play an important role in the organic matter fluxes that occur on the Antarctic sea floor. The dietary behaviour of these peracarids is still poorly known, and only few species have been analysed. This paper describes the trophic preferences of some dominant amphipod species of the Eastern Weddell Sea benthos, deduced from stomach content analyses and behavioural observations in aquaria. More than 1000 specimens, belonging to 40 species (representing 27 genera and 15 families) were dissected; and several thousands of individuals were kept in aquaria for 6–9 weeks and presented with various potential foods. These two approaches revealed at least eight different feeding types: suspension-feeding, deposit-feeding, deposit-feeding coupled with predation, opportunistic predation, micropredatory browsing, macropredation coupled with scavenging, opportunistic necrophagy and true necrophagy. These different behaviours cover almost all the possible feeding types with the exception of macroherbivorous browsing. Among the eight described feeding types, no particular one is dominant. In the same way, types involving microphagy and macrophagy are equally represented. Predatory types (opportunistic or exclusive) account for 64% of the species analysed, while scavenging types (facultative or obligate) account for 60%. The overlap suggests that many amphipod species have a wide dietary spectrum and are able to take advantage of different food resources.     

The effects of temperature on walking and righting in temperate and Antarctic crustaceans

Antarctic marine invertebrates live in a cold, thermally stable environment and cannot tolerate large changes in body temperature (i.e. they are stenothermal). Their temperate relatives, by contrast, are eurythermal, living in warmer and thermally more variable environments. Have these different environments influenced how specific behaviours are affected by changes of temperature? This question was addressed in two temperate crustaceans, the decapod Carcinus maenas and isopod Ligia oceanica, and two Antarctic crustaceans, the isopod Glyptonotus antarcticus and amphipod Paraceradocus gibber. The thermal dependence of walking speed was analysed by contrasting the slopes of the linear part of each species’ behavioural curve. Over the temperature ranges analysed, the temperature sensitivity of walking speed in the stenotherms was 13–23% that of the eurytherms when measured in body lengths s^?1. There was a linear relationship between walking speed and temperature up to +4.5°C in the Antarctic species G. antarcticus and P. gibber. Elevating temperature by up to 3.5°C above the maximum temperature experienced in the Antarctic (+1°C), does not lead to an acute breakdown of motor coordination. We describe for the first time the righting behaviour of G. antarcticus. The mean time-to-right tended to a minimum on warming from ?2 to+5°C, but this trend was not statistically significant. Our results suggest that the physiological adaptations which permit continued activity at low Antarctic temperatures have resulted in a lower thermal dependence of activity in Antarctic species, compared to related temperate species.     

Molecular and morphological evolution of the amphipod radiation of Lake Baikal

Lake Baikal, in Siberia, Russia, contains the highest biodiversity of any extant lake, including an impressive radiation of gammaroidean amphipods that are often cited as a classic case of adaptive radiation. However, relationships among Baikal's amphipods remain poorly understood. The phylogenetic history of 32 Lake Baikal amphipod species, representing most major lineages of the endemic fauna, was examined using three genes (COI, 16S rRNA, and 18S rRNA), and 152 morphological characters. Results support monophyly of the largest and most diverse of the Baikalian families, the Acanthogammaridae. Analyses suggest that a second Baikalian family, the fossorial Micruropodidae, is paraphyletic and composed of two divergent clades, one of which includes Macrohectopus branickii, a morphologically specialized pelagic planktivore traditionally assigned its own family. The extreme morphological and ecological divergence of Macrohectopus from its close genetic relatives, and conversely, the large genetic distances among other morphologically similar micruropodids, suggest that morphological and molecular evolution have often been uncoupled during the radiation of Baikal's amphipods. This study suggests that the amphipod fauna of Lake Baikal is polyphyletic; originating from two independent invasions of the lake.     

Chemical defence against predators in a sub-Arctic fjord

The development of chemical defences in marine organisms is supposed to be driven by intense pressure of predation and competition. While benthic communities in tropical and also Antarctic regions are thought to be mainly structured by intra- and interspecific interactions, these factors are proposed to be less important in northern high latitudes. Consequently, selective pressure for chemical defence should be low in these regions. To investigate the incidence of chemical defence against predation in northern high latitudes, crude extracts of 18 abundant sessile or slow-moving invertebrate species (4 sponges, 3 actinians, 1 soft coral, 4 bryozoans, 3 ascidians and the egg mass of a gastropod) from Kongsfjord (Spitsbergen) were tested for feeding deterrent activity. Laboratory assays were performed by offering artificial food with extracts to two different predators, the amphipod Anonyx nugax which is a common species in Kongsfjord, and the starfish Asterias rubens from the North Sea. Of the 18 extracts tested, 2 (Haliclona viscosa, Hormathia nodosa) exhibited significant feeding deterrent effects in the amphipod assay. Furthermore, 6 extracts had a significantly stimulating effect on the amphipod feeding, and 10 extracts did not affect consumption. In the starfish assay, only the crude extract of H. viscosa was significantly rejected. For H. viscosa, feeding deterrence could be established for two pure compounds, and for H. nodosa for one fraction. The present data show that feeding deterrent compounds are present in sub-Arctic marine invertebrates from Kongsfjord but are less abundant than in temperate, tropical and Antarctic species.     

DNA analyses reveal abundant homoplasy in taxonomically important morphological characters of Eusiroidea (Crustacea,Amphipoda)

Eusiroidea is one of the 20 amphipod superfamilies that were erected to subdivide the very large and controversial suborder Gammaridea. Yet, the definition of the superfamily is not based on synapomorphies, but on a combination of diagnostic phenetic similarities that hold more or less consistently across families. Moreover, many of the characters used to define eusiroid families are suspected to show convergent evolution. The current classification of the Eusiroidea may therefore not reflect evolutionary relationships accurately. Here, we present a molecular phylogenetic re‐analysis of the Eusiroidea based on a comparison of 18S and 28S rDNA sequences of 73 species, representing 47 genera and 16 families that potentially belong to the superfamily. The results suggest that at least species belonging to 14 of these traditional families would be part of a eusiroid clade, increasing by more than twofold the species and generic richness of the group. However, most of the eusiroid families surveyed do not appear monophyletic. Finally, the analyses show that several important morphological characteristics, traditionally used in eusiroid taxonomy, are homoplastic.     

Complete mitochondrial genome of the Antarctic amphipod Gondogeneia antarctica (Crustacea, amphipod)

The complete sequence of the mitochondrial genome of the Antarctic amphipod Gondogeneia antarctica was determined to be 18,424 bp in length, and to contain 13 protein-coding genes (PCGs), 22 tRNA genes, and large (rrnL) and small (rrnS) rRNA genes. Its total A+T content is 70.1%. The G. antarctica mitogenome is the largest known among those of crustaceans, due to the existence of two relatively large intergenic non-coding sequences. The PCG arrangement of G. antarctica is identical to that of the ancestral pancrustacean ground pattern, although the tRNA arrangement differs somewhat. The complete mitogenome sequences of 68 species of pancrustacea have been added to the NCBI database, only 4 of which represent complete mitogenome sequences from amphipods. This is the first report of a mitogenome sequence of an Antarctic amphipod, and provides insights into the evolution of crustacean mitochondrial genomes, particularly in amphipods.     

Phylogeny of Antarctic dragonfishes (Bathydraconidae,Notothenioidei, Teleostei) and related families based on their anatomy and two mitochondrial genes

Although Antarctic teleosts of the suborder Notothenioidei are well studied, the status of some families remains unclear because of limited taxonomic sampling and sometimes poor statistical support from molecular phylogenies. It is true for the Bathydraconidae, the sister-family of the famous haemoglobin-less icefishes, the Channichthyidae. The present study is aimed at clarifying bathydraconid phylogeny and the interrelationships of higher notothenioid families, taking nototheniids as the outgroup. For this purpose, about 300 positions in the mitochondrial control region, 750 positions in the cytochrome b, and a matrix of morphological characters were employed for separate and simultaneous phylogenetic analyses. We conclude that (1) molecular data strongly support the split of bathydraconids into three clades, here called the Bathydraconinae (Bathydraco, Prionodraco, Racovitzia), the Gymnodraconinae (Gymnodraco, Psilodraco, Acanthodraco), and the Cygnodraconinae (Cygnodraco, Gerlachea, Parachaenichthys). Interrelationships between these three and the Channichthyidae remain unclear. Molecular data support neither paraphyly nor monophyly of the bathydraconids, while morphology leads to the monophyly of the family based on the synapomorphic loss of the spinous dorsal fin; (2) The Channichthyidae, the Harpagiferidae, and the Artedidraconidae are monophyletic families; (3) the phylogeny of the haemoglobin-less channichthyids is completely resolved and congruent with the conclusions of based on anatomical characters; (4) The present molecular results as well as other molecular studies favour the hypothesis that harpagiferids are the sister-group of artedidraconids, though our morphological matrix puts harpagiferids as the sister-group of all other families on the basis of a single character. With regard to harpagiferid relationships, it is interesting to notice that, when analysed simultaneously, morphological characters are not automatically "swamped" within molecular ones: in the tree based on the simultaneous analysis of all available data, morphological characters impose their topology on molecules.     

Chemo–ecological interactions in Antarctic bryozoans

The role of bioactive metabolites in ecological interactions involving Antarctic bryozoans has been scarcely studied. Bryozoans are one of the most abundant and diverse members of the Antarctic benthos and are preyed upon by diverse kinds of predators. They seem to be casual food items of the common Antarctic sea urchin Sterechinus neumayeri and the ubiquitous omnivorous amphipod Cheirimedon femoratus. In this study, the cytotoxic activity against embryos and sperm of the sea urchin S. neumayeri and the substrate preferences of the amphipod C. femoratus were assessed using organic extracts from Antarctic bryozoans, in order to determine the presence of chemical defenses. New adapted protocols were designed using a solidifying gel for simulating bryozoans’ surface. We analyzed 32 organic extracts from 16 samples that belonged to 13 different bryozoan species. No cytotoxicity was detected against embryos of S. neumayeri, while 12 of the 13 bryozoan species were cytotoxic to sperm at natural concentrations. In the substrate preference assays, the amphipod C. femoratus was repelled by ten species. The variable bioactivities found in both types of organic partitions of extracts indicated the presence of both lipophilic and hydrophilic defenses. Inter- and intraspecific variability of chemical defenses was detected also, suggesting environmental-induced responses, symbiotic production, and/or genetic variability. Possible alternative defensive mechanisms are also discussed for species with low or no chemical defense. Our results clearly support the fact that chemically mediated bioactivity in Antarctic bryozoans is common, and there is a likely ecological role of cytotoxic and repellent compounds for their protection.     

Looking beneath the tip of the iceberg: diversification of the genus <Emphasis Type="Italic">Epimeria</Emphasis> on the Antarctic shelf (Crustacea,Amphipoda)

The amphipod genus Epimeria is very speciose in Antarctic waters. Although their brooding biology, massive and heavily calcified body predict low dispersal capabilities, many Epimeria species are documented to have circum-Antarctic distributions. However, these distribution records are inevitably dependent on the morphological species definition. Yet, recent DNA evidence suggests that some of these Epimeria species may be complexes of species with restricted distributions. Mitochondrial COI and nuclear 28S rDNA sequence data were used to infer evolutionary relationships among 16 nominal Epimeria species from the Antarctic Peninsula, the eastern Weddell Sea and the Adélie Coast. Based on this phylogenetic framework, we used morphology and the DNA-based methods GMYC, bPTP and BPP to investigate species boundaries, in order to revise the diversity and distribution patterns within the genus. Most of the studied species appeared to be complexes of pseudocryptic species, presenting small and previously overlooked morphological differences. Altogether, 25 lineages were identified as putative new species, increasing twofold the actual number of Antarctic Epimeria species. Whereas most of the species may be geographically restricted to one of the three studied regions, some still have very wide distribution ranges, hence suggesting a potential for large-scale dispersal.     

Evidence of hidden biodiversity, ongoing speciation and diverse patterns of genetic structure in giant Antarctic amphipods

Recent molecular research on Antarctic benthic organisms has challenged traditional taxonomic classifications, suggesting that our current perceptions of Antarctic biodiversity and species distributions must be thoroughly revised. Furthermore, genetic differentiation at the intraspecific level remains poorly understood, particularly in eastern Antarctica. We addressed these issues using DNA sequence data for two sibling amphipod species that could be collected on a circum-Antarctic scale: Eusirus perdentatus and Eusirus giganteus. Haplotype networks and Bayesian phylogenies based on mitochondrial (COI, CytB) and nuclear (ITS2) DNA provided strong evidence of multiple cryptic species of Eusirus, with several occurring in sympatry and at least one likely to have a true circum-Antarctic distribution. Within species, gene flow was often highly restricted, consistent with a brooding life history and in some cases suggestive of current or future allopatric speciation. Patterns of genetic structure were not always predictable: one cryptic species showed preliminary evidence of high genetic differentiation across ～150 km in eastern Antarctica (F(ST) > 0.47, P < 0.01), yet another was remarkably homogenous across ～5000 km (F(ST) = 0.00, P = 1.00). Genetic diversity also varied among cryptic species, independent of sample size (π = 0.00-0.99). These results indicate several hidden levels of genetic complexity in these Antarctic amphipods that are neither apparent from previous taxonomic or ecological studies nor predictable from their life history. Such genetic diversity and structure may reflect different modes of survival for Antarctic benthic organisms during historic glacial cycles, and/or subsequent re-establishment of populations on the shelf, and highlight our misunderstanding of Antarctic marine species diversity.     

Phylogenetic investigations of Antarctic notothenioid fishes (Perciformes: Notothenioidei) using complete gene sequences of the mitochondrial encoded 16S rRNA

The Notothenioidei dominates the fish fauna of the Antarctic in both biomass and diversity. This clade exhibits adaptations related to metabolic function and freezing avoidance in the subzero Antarctic waters, and is characterized by a high degree of morphological and ecological diversity. Investigating the macroevolutionary processes that may have contributed to the radiation of notothenioid fishes requires a well-resolved phylogenetic hypothesis. To date published molecular and morphological hypotheses of notothenioids are largely congruent, however, there are some areas of significant disagreement regarding higher-level relationships. Also, there are critical areas of the notothenioid phylogeny that are unresolved in both molecular and morphological phylogenetic analyses. Previous molecular phylogenetic analyses of notothenioids using partial mtDNA 12S and 16S rRNA sequence data have resulted in limited phylogenetic resolution and relatively low node support. One particularly controversial result from these analyses is the paraphyly of the Nototheniidae, the most diverse family in the Notothenioidei. It is unclear if the phylogenetic results from the 12S and 16S partial gene sequence dataset are due to limited character sampling, or if they reflect patterns of evolutionary diversification in notothenioids. We sequenced the complete mtDNA 16S rRNA gene for 43 notothenioid species, the largest sampling to-date from all eight taxonomically recognized families. Phylogenetic analyses using both maximum parsimony and maximum likelihood resulted in well-resolved trees with most nodes supported with high bootstrap pseudoreplicate scores and significant Bayesian posterior probabilities. In all analyses the Nototheniidae was monophyletic. Shimodaira–Hasegawa tests were able to reject two hypotheses that resulted from prior morphological analyses. However, despite substantial resolution and node support in the 16S rRNA trees, several phylogenetic hypotheses among closely related species and clades were not rejected. The inability to reject particular hypotheses among species in apical clades is likely due to the lower rate of nucleotide substitution in mtDNA rRNA genes relative to protein coding regions. Nevertheless, with the most extensive notothenioid taxon sampling to date, and the much greater phylogenetic resolution offered by the complete 16S rRNA sequences over the commonly used partial 12S and 16S gene dataset, it would be advantageous for future molecular investigations of notothenioid phylogenetics to utilize at the minimum the complete gene 16S rRNA dataset.     

Two new Amphipod families recorded in South America shed light on an old biogeographical enigma

The known diversity of freshwater amphipods in South America is substantially lower than on other continents. This has puzzled biologists for decades. Two hypotheses have been proposed in attempts to explain this pattern. According to the first one, the majority of amphipod lineages never dispersed across South America. The alternative hypothesis is that the recently diversified hyalellids have outcompeted and depleted the ancestral amphipod fauna. The recently discovered freshwater amphipod species Seborgia potiguar sp. nov. (Seborgidae) and Potiberaba porakuara gen. nov., sp. nov. (Mesogammaridae) from Brazil reveals the existence of two additional families of amphipods in South America. In the light of these discoveries we have analysed the amphipod faunistic structure of South America to test the above two biogeographic hypotheses. First, the number of amphipod families in South America is not as low as was thought. Falklandellididae are limited to the Falkland Islands and Chile. All other families (Ingolfiellidae, Bogidiellidae, Phreatogammaridae, Paraleptamphopidae, Pseudoingolfiellidae, Paracorpohiinae, Mesogammaridae and Seborgidae) but one (Dogielinotidae) share two properties: (1) they have a transoceanic distribution and (2) they are from subterranean waters. Since the dispersal ability of amphipods is limited, trans-oceanic disjunctions are best explained by plate tectonics, which implies their early origin, negating the first biogeographical hypothesis. These ancient families, for unknown reasons, survived only in a stable subterranean environment which can be regarded as a refuge. The only recent colonizer of the continent might be Dogielinotidae with the species-rich genus Hyalella. Although it cannot be determined whether hyallelids truly out-competed ancient amphipods, we suggest that the second hypothesis fits better to the data. Further findings of amphipods are expected in South America, especially from subterranean waters. This habitat is highly endangered in Brazil, and should be more rigorously protected. http://www.zoobank.org/urn:lsid:zoobank.org:pub:CBE21C1E-4748-4237-9EAD-FBD240A7D501     

What is a Suiforme (Artiodactyla)?: Contribution of Cranioskeletal and Mitochondrial DNA Data

Suiformes (Artiodactyla) traditionally includes three families: Suidae, Tayassuidae, and Hippopotamidae but the monophyly of this suborder has recently been questioned from molecular data. A maximum parsimony analysis of molecular, morphological, and combined data was performed on the same set of taxa including representatives of the three Artiodactyla suborders (Suiformes, Ruminantia, and Tylopoda) and Perissodactyla as outgroup. Mitochondrial (cytochromeband 12S rRNA) sequence comparisons support the monophyly of Suina (Suidae and Tayassuidae) and Ancodonta (Hippopotamidae) but not the monophyly of Suiformes. Inversely, our preliminary morphological analysis supports the monophyly of Suiformes whereas relationships among the three families are not resolved. The combined data set does not resolve the relationships between Suina, Ancodonta, and Ruminantia. These results are discussed in relation to morphological characters and paleontological data. Some improvements are suggested to clarify the morphological definition of Suiformes and relationships among them.     

Amphipoda (crustacea) from palau, micronesia: families ampeliscidae, ampithoidae, aoridae, colomastigidae and cyproideidae

12 species of amphipod in 5 families, collected from shallow reefs in Palau by S. DeGrave during 2002, are reported here. Of these, five species are new to science and Microdeutopus tridens Schellenberg (1938) is redescribed and transferred to the genus Bemlos Shoemaker (1925). The collection included several additional species in the genera Amphilochus Bate, 1862, Ampithoe Leach (1814), Bemlos, Byblis Boeck (1871), Colomastix Grube (1861) and Notopoma Lowry & Berents (1996), that were either incomplete or juvenile and could therefore not adequately be described. In addition, two new species of Plumithoe Barnard & Karaman (1991) are erected from the literature. Other families collected in Palau will be considered in later contributions.     

Island endemism, morphological stasis, and possible cryptic speciation in two coral reef, commensal Leucothoid amphipod species throughout Florida and the Caribbean

Coral reefs are believed to be one of the most diverse ecosystems, but the true magnitude of their biodiversity and patterns of endemism is uncertain. This uncertainty stems partly from the relative paucity of investigations on the small, difficult to collect taxa (cryptofauna) that may make up the majority of reef biodiversity and require specialized expertise for morphological identification. To assess the extent of diversity in some of the reef micro-cryptofauna, we analyzed 414 bp of the mitochondrial cytochrome oxidase subunit I gene from 556 individuals representing two brooding amphipod species (Leucothoe ashleyae and Leucothoe kensleyi). These amphipods are commensal inside the branching vase sponge Callyspongia vaginalis, and were sampled throughout Florida and the Caribbean. Phylogenetic analyses revealed 11 deeply divergent, strongly supported lineages (seven L. ashleyae and four L. kensleyi) each with very narrow geographic ranges. The level of intraspecific lineage divergence for both morphospecies was among the highest reported for any marine crustacean (12.4–26.0% uncorrected), and exceeded that of congeners from nine diverse amphipod families, as well as the patristic genetic distance suggested as a threshold for crustacean species delineation. These findings suggest a history of cryptic speciation within each morphospecies, concomitant with a pronounced period of morphological stasis involving each of the morphotypes. The observation of multiple, highly divergent, evolutionary significant units, each endemic to Florida and Caribbean island locations, supports the emerging view that coral reef biodiversity, especially in the cryptofaunal component, is likely vastly underestimated.     

Chromosomes of the Antarctic amphipod Waldeckia obesa Chevreux

Mitotic and meiotic chromosomes of the Antarctic lysianassoid amphipod Waldeckia obesa are described. The modal chromosome number is n = 25 and 2n = 50. The potential applications of cytogenetical studies in this group are discussed.