Cryptic speciation in a high gene flow scenario in the oviparous marine sponge Chondrosia reniformis

Sponge systematics has been traditionally based on the study of the skeleton (spicules and spongin fibres). However, sponges of the genus Chondrosia are devoid of those skeletal features, making it difficult to distinguish between different species in the genus. Chondrosia reniformis Nardo, 1847, the type species of the genus, was described from the Mediterranean Sea. The lack of distinguishing morphological features may have been responsible for the widespread assignment of specimens of the genus to this species; as a result C. reniformis is considered to be a cosmopolitan species. In this work, populations of C. reniformis from the western Mediterranean (France) and the West Atlantic (Bermuda and Brazil) were analysed using allozyme electrophoresis for 13 enzyme loci. Levels of mean heterozygosity were high (Bermuda and Brazil H=0.27 and W Mediterranean H=0.12), as is often observed in sponge species. Gene identities observed between West Atlantic and Mediterranean populations were low (I=0.40-0.52, typical values for congeneric species), including the presence of four diagnostic loci. This level of divergence clearly shows that they are not conspecific. Hence, a worldwide or cosmopolitan distribution of C. reniformis would seem improbable. However, the West Atlantic samples (Bermuda and Brazil) were genetically similar (gene identity, I=0.88-0.95) over a distance of 8,000 km. This is the first report of genetic homogeneity in a sponge species over such a large geographical distance.     

Short-term temporal patterns of early recruitment of coral reef fishes in the tropical eastern Pacific

Short-term temporal patterns of recruitment have been described in a variety of coral reef fishes and have often been related with lunar and tidal cycles. While the relative importance of lunar and tidal factors in determining recruitment patterns has been difficult to assess, most studies have been done in the Caribbean and Indo-Pacific, where tidal amplitudes are small. We studied the short-term temporal dynamics of fish recruitment at Gorgona Island (tropical eastern Pacific), where there is a large tidal amplitude (~4.4 m). Every other day during three consecutive months in 1998, we directly measured the magnitude of reef fish recruitment to standardized coral units (SCUs) isolated from natural reefs. A total of 40 species from 21 families settled on the SCUs. Of 11 species with sufficient numbers for meaningful statistical analyses, two (Lutjanus guttatus and Pomacanthus zonipectus) had lunar recruitment with peaks near the new moon; three combined species of antennariids showed semilunar recruitment with peaks near moon quarters; and eight other species showed sporadic and aperiodic recruitment pulses. The contribution of lunar (moonlight intensity) and tidal factors (tidal amplitude and net tidal flow) to recruitment dynamics varied among species, although it was generally low (<18%) even among species with periodic patterns, except perhaps in L. guttatus. In this species, recruitment magnitude correlated negatively with moonlight intensity, accounting for 34.5% of the variance. Post-settlement predation by roving predators may be one cause of this relationship. In the remaining species, particularly those with sporadic and aperiodic recruitment pulses, stochastically varying weather and oceanographic events may be more important in determining temporal variation in recruitment.     

Mitochondrial cytochrome b variation in sleeper sharks (Squaliformes: Somniosidae)

Sleeper sharks are a poorly studied group of deep-sea sharks. The subgenus, Somniosus, contains three morphologically similar species: S. microcephalus found in the Arctic and North Atlantic; S. pacificus in the North Pacific; and S. antarcticus in the Southern Ocean. These sharks have been reported mainly in temperate to polar waters and occasionally in subtropical locations. They have not been recorded in tropical waters. This study investigates the relationships among the accepted species of Somniosus through analysis of mitochondrial cytochrome b sequence variation. Seventy-five samples were examined from four sampling locations in the North Pacific, Southern Ocean and North Atlantic. Twenty-one haplotypes were found. A minimum spanning parsimony network separated these haplotypes into two divergent clades, an S. microcephalus and an S. pacificus/antarcticus clade, strongly supporting the distinction of S. microcephalus as a separate species from the Pacific sleeper shark species. Analysis of genetic structure within the S. pacificus/antarcticus clade (analysis of molecular variance, allele frequency comparisons, and a nested clade analysis) showed limited or no differences amongst three populations. Further examination of genetic variation at more variable mtDNA and nuclear markers is needed to examine the species status of S. pacificus and S. antarcticus.     

Oculina patagonica: a non-lessepsian scleractinian coral invading the Mediterranean Sea

The scleractinian coral Oculina patagonica De Angelis is a new immigrant from the Southwest Atlantic to the Mediterranean Sea, having established itself only recently along the Israeli coast. This species is the only scleractinian coral reported to have invaded a new region. In order to understand the swift establishment of this species along the Israeli coast, from 1994 to 1999 we studied its distribution, abundance, reproduction, recruitment, survival, and the effect of bleaching events on its population abundance. In addition, population studies of O. patagonica were performed at several localities along the eastern and western Mediterranean coasts. Highest abundance was recorded along the Spanish coast, reaching 30lj colonies per 10 m line transect in shallow water. Second in abundance was the Israeli coast, with a maximum of 10DŽ colonies per transect. O. patagonica was rare along the coast of Italy, and absent along the Mediterranean coast of France. During the study, both mortality and recruitment along the Israeli coast were very low. In contrast, recruitment along the Spanish coast was very high. Reproduction of the species was studied using gonadal histology. O. patagonica is gonochoric. Female gonads were first observed in May and male gonads in July, both reaching maturity in late August and early September. Naturally occurring azooxanthellate colonies of O. patagonica inhabiting small dark caves developed gonads and spawned in parallel to zooxanthellate colonies exposed to light. No gonads were found in zooxanthellate colonies that underwent bleaching during the reproduction season. The high incidence of bleaching events along the Israeli coast observed throughout the years of this study may explain the low recruitment of new colonies during the same period. In view of its current recruitment patterns, we expect further expansion of O. patagonica in range and abundance in the western Mediterranean, but very small expansion of the population in the eastern Mediterranean, due to repetitive annual bleaching events.     

Large-scale oceanic distribution and population structure of Calanus finmarchicus, in relation to physical environment, food and predators

The investigation was carried out from 62°N to 73°N and from 14°E to 11°W in the Norwegian Sea during 19 June-12 July 1997. Regional differences in the phase of the seasonal development of the plankton community were evident, most pronounced across the Arctic front. In the Coastal and eastern Atlantic domains, post-bloom conditions prevailed, characterised by low chlorophyll a (chl a) levels and a phytoplankton assemblage dominated by coccolithophorids and small flagellates. During the study period, egg production rates of Calanus finmarchicus were low (<10 eggs female^-1 day^-1), older copepodite stages dominated, and the seasonal descent to deeper waters had started. In the Arctic domain, bloom conditions were evident by high chl a levels and a high abundance of large diatoms. Egg production rates were higher (a maximum of 29 eggs female^-1 day^-1), but the dominance of stages CI-CIII indicated that considerable spawning had already occurred prior to the spring bloom. The seasonal descent had barely started. Both invertebrate and fish predators were most abundant in the Coastal and eastern Atlantic domains, with abundance strongly decreasing north-westwards. No tight relationship between total abundance of invertebrate or fish predators and that of C. finmarchicus was apparent. However, a weak, but significant, relationship between abundance of young stages of chaetognaths and Euchaeta spp. versus young stages of C. finmarchicus was found, indicating that these invertebrate predators develop parallel to the development of the new cohort of C. finmarchicus. In early summer, C. finmarchicus had reached overwintering stages, and had started to accumulate in deeper waters in areas with the highest abundance of horizontally migratory planktivorous fish.     

Population genetic structure of escolar (<Emphasis Type="Italic">Lepidocybium flavobrunneum</Emphasis>)

Escolar (Lepidocybium flavobrunneum) is a large, mesopelagic fish that inhabits tropical and temperate seas throughout the world, and is a common bycatch in pelagic longline fisheries that target tuna and swordfish. Few studies have explored the biology and natural history of escolar, and little is known regarding its population structure. To evaluate the genetic basis of population structure of escolar throughout their range, we surveyed genetic variation over an 806 base pair fragment of the mitochondrial control region. In total, 225 individuals from six geographically distant locations throughout the Atlantic (Gulf of Mexico, Brazil, South Africa) and Pacific (Ecuador, Hawaii, Australia) were analyzed. A neighbor-joining tree of haplotypes based on maximum likelihood distances revealed two highly divergent clades (δ = 4.85%) that were predominantly restricted to the Atlantic and Indo-Pacific ocean basins. All Atlantic clade individuals occurred in the Atlantic Ocean and all but four Pacific clade individuals were found in the Pacific Ocean. The four Atlantic escolar with Pacific clade haplotypes were found in the South Africa collection. The nuclear ITS-1 gene region of these four individuals was subsequently analyzed and compared to the ITS-1 gene region of four individuals from the South Africa collection with Atlantic clade haplotypes as well as four representative individuals each from the Atlantic and Pacific collections. The four South Africa escolar with Pacific mitochondrial control region haplotypes all had ITS-1 gene region sequences that clustered with the Pacific escolar, suggesting that they were recent migrants from the Indo-Pacific. Due to the high divergence and geographic separation of the Atlantic and Pacific clades, as well as reported morphological differences between Atlantic and Indo-Pacific specimens, consideration of the Atlantic and Indo-Pacific populations as separate species or subspecies may be warranted, though further study is necessary.     

Is the Great Barracuda (<Emphasis Type="Italic">Sphyraena barracuda</Emphasis>) a reef fish or a pelagic fish? The phylogeographic perspective

Current taxonomy indicates a single global species of the Great Barracuda (Sphyraena barracuda) despite differences in color and behavior between Atlantic and Pacific forms. To investigate these differences and qualify the dispersal characteristics of this unique coastal–pelagic teleost (bony fish), we conducted a global phylogeographic survey of 246 specimens from thirteen sampling locations using a 629-base pair fragment of mtDNA cytochrome b. Data indicate high overall gene flow in the Indo-Pacific over large distances (>16,500 km) bridging several biogeographic barriers. The West Atlantic population contains an mtDNA lineage that is divergent from the Indo-Pacific (d = 1.9%), while the East Atlantic (N = 23) has two mutations (d = 0.6%) apart from the Indo-Pacific. While we cannot rule out distinct evolutionary partitions among ocean basins based on behavior, coloration, and near-monophyly between Atlantic and Indo-Pacific subpopulations, more investigation is required before taxonomic status is revised. Overall, the pattern of high global dispersal and connectivity in S. barracuda more closely resembles those reported for large oceanic predators than reef-associated teleosts.     

The relationship between dissolved oxygen concentration and maximum size in deep-sea turrid gastropods: an application of quantile regression

Bathymetric gradients in body size are the most well-known patterns of geographic variation in deep-sea organisms. The causes of size-depth relationships remain uncertain, but most have been attributed to rates of nutrient input. Chapelle and Peck (1999, Nature 399:114-115) recently hypothesized that body size in marine invertebrates is a function of dissolved oxygen concentration. We tested this hypothesis by using quantile regression techniques to assess the relationship of dissolved oxygen levels to maximum size in deep-sea turrid gastropods collected from the North Atlantic. Relationships were examined for a group of nine turrid species and within the abundant lower bathyal species Benthomangelia antonia (Dall, 1881). We controlled the analysis for depth because size in deep-sea gastropods varies bathymetrically. When the effects of depth are accounted for statistically, maximum size in B. antonia increases with increasing levels of dissolved oxygen. In turrids as a group, both depth and oxygen appear to explain significant proportions of the variance in maximum size. These findings suggest that a suite of factors, including dissolved oxygen concentration, may influence maximum size in deep-sea organisms.     

Molecular taxonomy of vestimentiferans of the western Pacific and their phylogenetic relationship to species of the eastern Pacific. I. Family Lamellibrachiidae

Phylogenetic relationships among vestimentiferans in the family Lamellibrachiidae collected from ten sites in the western Pacific were analyzed on the basis of the nucleotide sequence of part of a mitochondrial gene for cytochrome oxidase I. The 103 individuals analyzed were tentatively classified into five species: namely, Lamellibrachia satsuma, three tentative species inhabiting Japanese waters and one tentative species from the Manus Basin. Phylogenetic analysis of the four tentative species, L. satsuma, and two lamellibrachiids, whose sequences had been reported previously, namely L. columna from the Lau Basin and L. barhami from the Oregon Margin and the Middle Valley, revealed that the lamellibrachiids from Japanese waters were not monophyletic. While two tentative species from Japanese waters and L. columna formed a monophyletic group, the other tentative species from Japanese waters was closely related to the tentative species from the Manus Basin. L. satsuma was shown to be phylogenetically distinct from other lamellibrachiids of the western Pacific. A lamellibrachiid that had been collected from the Nikko Seamount was also demonstrated to be L. sastuma.     

Molecular taxonomy of vestimentiferans of the western Pacific and their phylogenetic relationship to species of the eastern Pacific

The nucleotide sequence of part (624 bp) of a mitochondrial gene for cytochrome oxidase I was determined for 46 escarpiid vestimentiferans collected from seven sites in the western Pacific and 49 individual specimens of Arcovestia ivanovi from two sites in the Manus Basin. Phylogenetic analysis, based on the newly obtained and previously reported sequences, indicated that escarpiids in the western Pacific can be divided into two tentative species, as we proposed in a previous report. While members of the first tentative species have been collected exclusively from a seep area at a depth of 300 m off the coast of central Japan, the members of the second species inhabit some sites at depths greater than 1,100 m, namely, seep areas in Japanese and Papua-New Guinean waters as well as hydrothermal vent fields in the Okinawa Trough and the Manus Basin. We detected no genetic structure among populations of the second tentative species. The first tentative species was more closely related to a species in the eastern Pacific, Escarpia spicata, and to a species in the Gulf of Mexico, Escarpia laminata, than to the second tentative species in the western Pacific. Sequences obtained from all arcovestiids were identical with the exception of those from three individuals, each of which included a single synonymous nucleotide substitution relative to the dominant haplotype, and no genetic differences were detected between specimens from the two sites in the Manus Basin.     

Molecular evidence for multiple lineages in the gorgonian family Plexauridae (Anthozoa: Octocorallia)

Octocorals are diverse and abundant on many marine hard substrates, and, within this group, members of the family Plexauridae are an important component of tropical reef assemblages, especially in the Caribbean. To understand historical relationships within this large and diverse assemblage, and to test the monophyly of the family and some of its genera, DNA sequences of two mitochondrial loci (msh1 and ND2, ~1,185 bp) were analyzed from 46 species in 21 genera from deep and shallow waters in the tropical western Atlantic and in the tropical western and eastern Pacific (plus 9 taxa in the closely related Gorgoniidae and 1 species of the more distantly related Alcyoniidae). Five strongly supported clades were recovered. Three large clades correspond roughly to the Plexauridae, Paramuriceidae, and Gorgoniidae, and two smaller clades were comprised of taxa previously assigned to several families. Astrogorgia sp. did not group with any of the clades. The mutual relationships among the five clades remain unclear. Several genera previously regarded as unrelated appear to be grouped among the three families; e.g. Hypnogorgia sp. (Paramuriceidae) falls within a clade consisting of both Pacific and Atlantic Muricea spp. (Plexauridae), while Swiftia sp., Scleracis sp., and an Atlantic Thesea sp. (all Paramuriceidae) group with the gorgoniids. In several instances, genera containing Atlantic and Pacific species were recovered as monophyletic (Muricea spp., Bebryce spp.). However, in at least three cases (Echinomuricea spp., Thesea spp., Villogorgia spp.), placement of Atlantic and Pacific species in the same genus may reflect convergence of sclerite morphology. The results indicate a strong need for reexamination of octocoral taxonomy using a combination of molecular, morphological, and chemical evidence.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00227-005-1592-y.Communicated by J.P. Grassle, New Brunswick     

Experimental effect of cold,La Niña temperatures on the survival of reef fishes from Gorgona Island (eastern Pacific Ocean)

The eastern tropical Pacific (ETP) reefs are affected at irregular times by extremely cold temperatures that occur principally during La Niña events. The effects of these low temperatures on the survival of reef fishes were experimentally assessed by determining the critical thermal minimum (CTM) of 15 reef fish species from Gorgona Island (ETP), and comparing these CTMs with the records of temperature during past La Niña events. Among species, mean CTMs ranged from 10.8°C to 16.3°C, which were lower than the coldest temperature recorded during the last La Niña event (18°C during La Niña 1998-1999). However, the observed ranges of CTM for two species (Thalassoma lucassanum and Eucinostomus gracilis) extended above 18°C. These results suggest that most of the reef fishes we studied are physiologically tolerant to the cold temperatures encountered during La Niña, though decreases in at least two populations may be expected as a result of the mortality of less tolerant individuals. Although tolerant to cold temperatures, reef fish populations may still experience negative changes during La Niña, because other determinants in population maintenance (e.g. reproduction and recruitment) are more temperature sensitive. The effects of other cold phenomena on reef fish survival are also discussed herein.     

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.     

Evolutionary relationships of deep-sea mussels inferred by mitochondrial DNA sequences

In order to elucidate the evolutionary process of deep-sea Bathymodiolus mussels, we investigated the phylogenetic relationships of 16 species worldwide by analyzing nucleotide sequences of the mitochondrial COI and ND4 genes. Deep-sea mussels were clustered into three groups by basal trichotomous divergence. The first was composed of four species found in Japanese waters and one species from the Gulf of Mexico, which contain methanotrophic endosymbiotic bacteria. The second included nine species distributed in the West and East Pacific, Indian, and Atlantic Oceans. Members of the second group were trichotomously divided into the Indo-West Pacific, Atlantic, and East Pacific subclusters. The Indo-West Pacific subcluster was composed of three very closely related species with mutual genetic distances at the intraspecific level (av. 0.019 in COI and 0.009 in ND4 relative to av. 0.156 in COI and 0.265 in ND4 among Bathymodiolus species other than Cluster A species), suggesting some gene flow among these species. The third consisted of two West Pacific species. Species in the second and third groups contain mainly thioautotrophic endosymbionts, including some species harboring both methanotrophs and thioautotrophs.     

Energy budgets and feeding rates of Coryphaenoides acrolepis and C. armatus

This study develops energy budgets and estimates feeding rates for two macrourid fishes, Coryphaenoides acrolepis, dominant in the bathyal eastern North Pacific, and the abyssal cosmopolitan species, Coryphaenoides armatus. Daily energy expenditure by C. acrolepis was nearly twice that of C. armatus. C. acrolepis allocated nearly equal amounts of energy to metabolism and growth. Once sexual maturity was reached reproduction became the dominant energetic cost. Either these costs are necessary to retain adequate numbers of eggs and larvae on the continental slopes, or this fish does not reproduce on an annual basis and the calculated costs are an overestimate. C. armatus allocated relatively more energy to metabolism than growth. It may be semelparous, and this strategy would be of great energetic savings in its food-poor but stable environment. Individual daily ration for C. acrolepis decreased from 0.31% to 0.07% of body weight (BW) and for C. armatus from 0.12% to 0.02% BW with increasing fish length. These rates are substantially lower than those for fishes living in cold waters on the continental shelves. The population feeding rates for C. acrolepis ranged from 0.8 to 15 kg km^-2 day^-1 and for C. armatus from 5 to 2,800 g km^-2 day^-1. The scavenging behaviour of C. acrolepis was used to investigate the role of carrion as a food supply to the deep-sea benthos. It was estimated that the carrion eaten by C. acrolepis is equivalent to 0.04 mg C m^-2 day^-1 or only 0.2-0.4% of the average small particulate flux. Carrion consumption is important for scavengers like C. acrolepis, but it is not an important component of the carbon flux into the deep-sea benthic environment.     

Population genetics of the nurse shark (Ginglymostoma cirratum) in the western Atlantic

The nurse shark, Ginglymostoma cirratum, inhabits shallow, tropical, and subtropical waters in the Atlantic and the eastern Pacific. Unlike many other species of sharks, nurse sharks are remarkably sedentary. We assayed the mitochondrial control region and eight microsatellite loci from individuals collected primarily in the western Atlantic to estimate the degree of population subdivision. Two individuals from the eastern Atlantic and one from the Pacific coast of Panama also were genotyped. Overall, the mtDNA haplotype (h = 48 ± 5%) and nucleotide (π = 0.08 ± 0.06%) diversities were low. The microsatellite data mirror the mitochondrial results with the average number of alleles ([`(N)]<sub>A</sub> \bar{N}_{A}  = 9) and observed heterozygosity ([`(H)]_O \bar{H}_{O}  = 0.58) both low. The low levels of diversity seen in both the mtDNA and the microsatellite may be due to historical sea level fluctuations and concomitant loss of shallow water habitat. Eight of the 10 pair-wise western Atlantic F _ST estimates for mtDNA indicated significant genetic subdivision. Pair-wise F _ST values for the microsatellite loci indicated a similar pattern as the mtDNA. The western Atlantic population of nurse sharks is genetically subdivided with the strongest separation seen between the offshore islands and mainland Brazil, likely due to deep water acting as a barrier to dispersal. The eastern and western Atlantic populations were closely related. The eastern Pacific individual is quite different from Atlantic individuals and may be a cryptic, sister species.     

New insight on population genetic connectivity of widespread amphidromous prawn Macrobrachium lar (Fabricius, 1798) (Crustacea: Decapoda: Palaemonidae)

Due to the sparse and unstable nature of insular freshwater habitats, marine larval dispersal of amphidromous species is considered a critical element of population persistence. We assessed population genetic structure of freshwater prawn Macrobrachium lar across its range that encompasses two biogeographic barriers: the vast open ocean separating Western and Central Pacific regions and the Indo-Malay archipelago separating Indian and Pacific oceans. A total of 173 samples collected from 21 islands throughout the Indo-Pacific were sequenced at 16S and 28S rDNA. We observed distinct genetic isolation of populations located at the eastern and southwestern edge of the species range but no evidence of an effect of the Indo-Pacific barrier. Differentiation patterns are consistent with a stepping-stone model of dispersal. Genetic differences of Central Pacific populations may reflect founder events associated with colonization of isolated islands, or be a signature of a past bottleneck after population depletion caused by drastic climatic events.     

Analysis of ribosomal ITS1 sequences indicates a deep divergence between Rhodactis (Cnidaria: Anthozoa: Corallimorpharia) species from the Caribbean and the Indo-Pacific/Red Sea

This paper explores the implications of ribosomal ITS1-sequence analysis for evolutionary relationships between four species of corallimorphs which are morphologically very similar but have distinct distributions. The analyses highlight the unreliability of morphological criteria in establishing systematic relationships amongst tropical corallimorphs. Both parsimony and distance methods of phylogenetic analysis strongly support a deep divergence between the Caribbean species Rhodactics sanctithoma and the three morphologically-related species R. howesii and R. indosinensis from the Indo-Pacific and R. rhodostoma from the Red Sea. The analyses also strongly support a closer relationship between the Red Sea species, R. rhodostoma, and one of the Indo-Pacific species, R. howesii, than between these two and the second Indo-Pacific species, R. indosinensis. The first of these results is presumably a consequence of the closure of the Central American Seaway, the event which led to the separate development of Atlantic and Indo-Pacific coral faunas. However, the lack of detailed distribution data represents a major barrier to reconstructing the evolutionary history of the genus.     

Phylogeography of two squid (Loligo pealei and L. plei) in the Gulf of Mexico and northwestern Atlantic Ocean

The loliginid squids Loligo pealei LeSueur and L. plei Blaineville (both recently proposed for reclassification as Doryteuthis) are commercially important, similar in appearance, and sympatric throughout much of the northwestern Atlantic Ocean, the Gulf of Mexico, and the Caribbean Sea. To investigate possible cryptic speciation and population structure, we examined samples (collected from 1995 to 1997) of both species for restriction fragment length polymorphisms (RFLPs) in PCR products of the mitochondrial gene cytochrome c oxidase (subunit I). RFLP haplotypes were further characterized by direct sequencing. In North American waters, cryptic speciation was rejected by the far greater nucleotide sequence divergence between species (~14%) versus within species (<1%). Each species displayed about a dozen RFLP haplotypes, but only three of their respective haplotypes were found among 90% of L. pealei specimens (n=356) and 97% of L. plei specimens (n=431). For L. pealei, a genetic break existed between the northern Gulf of Mexico and the Atlantic Ocean; among sample units within each population, gene flow was consistent with panmixia. The phylogeography of L. pealei is likely a consequence of the eastward currents of the Florida Straits, the elevated temperatures of those surface waters, and the restriction of this species to the continental shelf. For L. plei, a genetic break existed between longitudes 88°W and 89°W, with the northwestern Gulf of Mexico and the northeastern Gulf-Atlantic Ocean comprising separate populations; among sample units within each population, gene flow fit an isolation-by-distance model. If the genetic break found for L. plei represents resident populations separated by nearshore physical parameters (e.g. effects of the Mississippi River and the sediment boundary at longitude 88°W), the lack of structure within the Gulf for L. pealei might be due to its distribution farther from shore. However, the two populations of L. plei probably represent annual recolonization from the southwestern Gulf of Mexico and from the eastern Caribbean Sea, whereas the populations of L. pealei probably are permanent residents within their respective regions.     

Global phylogeography of the fissiparous sea-star genus <Emphasis Type="Italic">Coscinasterias</Emphasis>

The fissiparous starfish genus Coscinasterias (Verrill) is represented in shallow waters around many of the world's continents. This wide distribution could be explained by dispersal, vicariance, or translocation associated with shipping, and represents an excellent system for marine biogeographic research. We conducted a global phylogeographic analysis of 42 Coscinasterias mtDNA cytochrome oxidase 1 sequences (15 haplotypes) from 18 sites, including representatives of all four recognised species. Phylogenetic analysis yielded a robust phylogeny, with strong support for the monophyly of the genus (90% bootstrap support) and of each separate species (99-100%). Haplotypes exhibited strong phylogeographic structure, with robust mtDNA clades often associated with distinct land masses. A general lack of genetic differentiation within sites may reflect fissiparity. However, shared haplotypes over larger distances (e.g. across Japan), and the presence of related haplotypes on adjacent land masses (e.g. Tasmania, New Zealand; 1.6-1.8%) suggest that long-distance dispersal is an important biogeographical process for Coscinasterias. The 4.0-4.4% divergence between Japanese and South African sister groups may relate to transequatorial dispersal around the onset of the Pleistocene. Divergent Atlantic and Mediterranean populations of C. tenuispina (maximum 1.5%) may deserve subspecific recognition, and high divergences within Australian C. muricata (maximum 8.0%) suggest a species complex.