Morphological and genetic evidence for vicariance and refugium in Atlantic and Gulf of Mexico populations of the hermit crab Pagurus longicarpus

The number and wide variety of southeastern United States marine taxa with significant differentiation between Gulf of Mexico and Atlantic Ocean populations suggests that these taxa may have experienced major vicariance events, whereby populations were subdivided by geological or ecological barriers. The present study compared variation in morphology, allozymes, and mtDNA in Gulf of Mexico and western Atlantic populations of the longwrist hermit crab Pagurus longicarpus Say collected during 1997 and 1998. Combined Atlantic populations had significantly fewer denticles on the second segment of the third maxilliped than did Gulf of Mexico populations, and the mean ratio of dactyl length to propodus length was significantly greater in the Atlantic crabs than in the Gulf of Mexico crabs. Allozyme allele frequencies at three loci showed genetic differentiation between a Gulf of Mexico population and two Atlantic populations. Analysis of mtDNA sequence data revealed a clear reciprocal monophyly between Gulf and Atlantic populations, with an estimated divergence age of ~0.6 million years ago. This estimated age of divergence is significantly more recent than an age previously estimated for its congener Pagurus pollicaris (~4 million years ago), suggesting that species with a similar genetic break between Gulf and Atlantic populations may not necessarily share an identical history. Surprisingly, there is evidence of geographic subdivision within Atlantic populations of P. longicarpus along the east coast of North America. This differentiation is especially strong between Nova Scotia and southern populations, suggesting that the Nova Scotia population may represent survivors from a northern refugium during the last glacial maximum.     

Phylogeography of the marine interstitial nemertean <Emphasis Type="Italic">Ototyphlonemertes parmula</Emphasis> (Nemertea,Hoplonemertea) reveals cryptic diversity and high dispersal potential

We conducted a phylogeographic study of the meiofaunal nemertean Ototyphlonemertes parmula, an apparent species complex from the littoral zone of coarse-grained beaches, using a 494-bp fragment of the mitochondrial cytochrome oxidase 3 gene (cox3). Six populations from the Gulf and Atlantic coasts of Florida, two from New England, and one from the Caribbean were sampled in March and August 2005. Three major lineages were identified, separated by cox3 sequence divergence of 16–18%, with partially overlapping ranges. Tests for hybridization using ISSR markers detected nuclear gene exchange within but not between the major mitochondrial lineages, indicating the presence of cryptic species. One lineage dominating the Atlantic coast of Florida shows no evidence of geographic structuring. Another lineage shows a phylogenetic break between the Atlantic and Gulf coasts, suggesting that unsuitable habitat may act as a barrier to dispersal. Long-distance migration is evidenced by shared haplotypes between Florida and the eastern Caribbean. Overall, the widespread distribution of individual haplotypes and lack of structuring within geographic regions contrast with O. parmula’s strongly sediment-bound lifestyle. We speculate that dispersal of adults by storms and/or sediment transport may be more important than few and potentially short-lived planktonic larvae to explain geographic diversity in O. parmula and may be important for meiofauna in general.     

Diversity and community structure of symbiotic dinoflagellates from Caribbean coral reefs

A community ecology approach to the study of the most common group of zooxanthellae, dinoflagellates in the genus Symbiodinium, was applied to symbiotic invertebrate assemblages on coral reefs in the western Caribbean, off the Yucatan peninsula (Puerto Morelos, Mexico) and over 1000 km away in the northeastern Caribbean, at Lee Stocking Island, Bahamas. Sequence differences and intragenomic variation, as determined by denaturing gradient gel electrophoresis and sequencing of the internal transcribed spacer 2 (ITS 2) region, were used to classify these symbionts. Twenty-eight genetically distinct Symbiodinium types were identified, eleven of which were found in hosts from both Caribbean locations. A single symbiont population was detected in 72% of hosts from the Yucatan and 92% of hosts from the Bahamas. The reef-wide community distribution of these symbionts is dominated by a few types found in many different host taxa, while numerous rare types appear to have high specificity for a particular host species or genus. Clade or lineage A Symbiodinium spp. was restricted to compatible hosts located within 3-4 m of the surface, while Symbiodinium spp. types from other lineages displayed differences in vertical zonation correlated with ITS type but were independent of clade designation. A comparison of the symbiont types found in field-collected hosts with types previously cultured from these hosts indicates the existence of low density or "background"-symbiont populations and cryptic, potentially non-mutualistic types in some hosts.     

Are meiofaunal species cosmopolitan? Morphological and molecular analysis of Xenotrichula intermedia (Gastrotricha: Chaetonotida)

Many meiofaunal species are reported to be cosmopolitan, but due to uncertainties of identification, the affiliation of specimens from geographically distant areas to the same species-taxon is problematic. In this study, we examined morphological and molecular variation in samples of Xenotrichula intermedia Remane (Gastrotricha: Chaetonotida) from the Mediterranean Sea, the northwestern Atlantic and the northern Gulf of Mexico. Univariate analysis of 16 morphological traits was unable to detect differences among populations, except for the length of the pharynx, which was significantly shorter in the Gulf of Mexico specimens. Canonical discriminant analysis separated the Gulf of Mexico specimens from the other two populations, with pharynx length contributing about half of the total discrimination. Molecular analysis based on restriction-fragment length polymorphisms (RFLPs) in a 710-base pair polymerase chain-reaction (PCR) produet representing roughly half of the mitochondrial cytochrome oxidase I (COI) gene detected four haplotypes: one each from the Mediterranean and the Gulf of Mexico populations and two coexisting within the Atlantic population. The estimated nucleotide-sequence divergence calculated for each pairwise combination of haplotypes (based on the proportion of shared fragments) ranged from 5.3 to 11.5%. The high genetic divergence and the inability to clearly separate populations based on morphology suggest that individuals characterized by different haplotypes are genetically isolated sibling species.     

Cryptic isolation of Gulf of California shovelnose guitarfish evidenced by mitochondrial DNA

The shovelnose guitarfish Rhinobatos productus is an evolutionarily, ecologically, and economically important ray, with a continuous distribution from San Francisco, California (USA), to Mazatlan, Sinaloa, and in the Gulf of California (Mexico). Regional studies have revealed morphometric differences between shovelnose from the Gulf of California and the Pacific coast of Baja California, which may result from phenotypic plasticity in the presence of high levels of gene flow or from a degree of genetic differentiation in the presence of cryptic isolation within a continuous distribution. We used PCR-RFLP of the mitochondrial control region to assess the degree of genetic differentiation between Gulf of California and Pacific shovelnose guitarfish. We found very high levels of molecular diversity (averages: h=0.77, =1.19%), which may be associated with historically large and stable populations, as well as very significant levels of genetic differentiation between gulf and Pacific samples (²=64, P<0.0001; _ST=0.63, P<0.0001, mean nucleotide divergence d=2.47%). We found a deep phylogeographic break between haplotypes from the gulf and the Pacific, which may suggest the existence of cryptic species but clearly indicates more than one evolutionarily significant unit of R. productus. Our results show a pattern of genetic structure and levels of differentiation consistent with the geological history of the region. Furthermore, these findings have wide-ranging implications for the management and conservation of cartilaginous fish in Mexico, as they reveal the existence of biological diversity that will go unnoticed without the genetic scrutiny of intraspecific variation and that is highly relevant for much needed management and conservation efforts.Communicated by P.W. Sammarco, Chauvin     

Lack of phlorotannin induction in the kelp Laminaria hyperborea in response to grazing by two gastropod herbivores

Phlorotannins (polyphenolic compounds) are found exclusively in brown algae and are commonly regarded as an important component of algal defense against herbivores. The variation in phlorotannin concentration within and among individual seaweed specimens of the same species can be substantial, and seaweeds may respond to herbivore damage by an increased production of phlorotannins in order to prevent further herbivory. In the present study, we investigated the natural variation of phlorotannins in different parts of the kelp Laminaria hyperborea in an observational field survey. We also performed a manipulative induction experiment in which we studied the effect of natural grazing by the patellid limpet Ansates (=Helcion) pellucida and the littorinid snail Lacuna vincta, as well as artificial damage on the production of phlorotannins in L. hyperborea. There was a large variation in phlorotannin content both among and within L. hyperborea individuals and populations. The control plants in the induction experiment had a higher phlorotannin content compared to plants in natural populations, but grazing by A. pellucida or L. vincta, or artificial damage did not induce a higher phlorotannin concentration in the kelps. Instead, the phlorotannin content in wounded kelps was significantly lower than in the control plants, possibly due to removal of phlorotannin-rich outer layers of the kelp thallus, or to an induced compensatory growth response in the L. hyperborea plants. These results imply that L. hyperborea phlorotannins probably do not function as an inducible chemical defense against A. pellucida and L. vincta, instead grazing by these herbivores decreases the phlorotannin content of the kelps.     

Evidence of different reproductive traits in the transisthmian sister species, <Emphasis Type="Italic">Alpheus saxidomus</Emphasis> and <Emphasis Type="Italic">A. simus</Emphasis> (Decapoda,Caridea, Alpheidae): description of the first postembryonic stage

The emergence of the Isthmus of Panama, approximately 3.0-3.5 million years ago, established two very different marine systems (the Caribbean and the eastern Pacific) and separated previously continuous marine populations. The geographic isolation of transisthmian sister species provides an excellent basis for the study of divergence and speciation processes. Here we describe the morphology of the first postembryonic stage of Alpheus saxidomus and A. simus, two rock-boring alpheids; the estimated time since divergence for this transisthmian pair ranges from 4.4-6.1 to 3-9 million years. The presence of a first zoeal stage in A. saxidomus, e.g., without pleopods and a telson with 7+7 setae, indicates a prolonged developmental sequence. In contrast, hatchlings of A. simus are substantially more developed and resemble juveniles. Thus, the developmental modes of A. saxidomus and A. simus are strikingly different with abbreviated, most probably direct, development in the latter species. Reduced food availability in the Caribbean compared to the Pacific coast is likely to be a possible reason for the evolution of such important differences in life history traits of the two transisthmian sister species. It is suggested that a transition from prolonged to abbreviated development evolved gradually during the estimated time since divergence; however, such a shift may have taken place within a substantially smaller time span.     

Osmoregulation in fiddler crabs (<Emphasis Type="Italic">Uca</Emphasis>) from temperate Atlantic and Gulf of Mexico coasts of North America

Fiddler crabs (Crustacea: Decapoda; genus Uca) were collected from 16 locations along the temperate Atlantic and Gulf of Mexico coasts of eastern North America for osmoregulation studies. Three species, U. pugilator, U. pugnax and U. minax, were taken from habitats between Cape Cod, MA, and St. Mary's River, Nassau Co., GA Seven species, U. panacea, U. speciosa, U. spinicarpa, U. longisignalis, U. rapax, U. pugilator and U. minax, were collected in habitats between the Ochlochonee River, Liberty Co., and Thompson's Bayou, Escambia Co., along the Gulf coast of northwest Florida. To examine differences in osmoregulatory capabilities among the species, specimens were placed in various concentrations of artificial seawater (0-3450 mOsm) for 5 days. The oligohaline or "freshwater" species, U. minax, U. spinicarpa and U. longisignalis, possess the lowest average hemolymph osmolality. They are unable to control hemolymph osmolality above 2000 mOsm. On the other hand, the euryhaline species, U. speciosa, U. panacea, U. pugilator and U. pugnax, have much higher average hemolymph osmolality. They are able to withstand an osmotic challenge of 2200 mOsm or greater. Among the eight species, U. panacea and U. pugilator are able to osmoregulate across the broadest range of seawater concentrations (0-2800 mOsm). After an examination of the osmoregulation in several populations of U. minax, U. pugilator and U. pugnax, physiological adaptation is apparent in the two former species, but not the latter. Experimental evidence for capacity adaptation was examined in the freshwater species, U. minax. If slowly adapted to hypertonic seawater (1800 mOsm) for 14 days, this species is able to withstand much higher osmotic pressures than unadapted crabs. Although hemolymph isosmotic concentration remains the same, the limits for regulation are extended to higher osmolality. Consequently, as a result of studying osmoregulation in several species of Uca captured at various locations across their geographic range in the temperate zone, a clearer view of water-balance physiology is available for the genus. The Uca spp. are not uniform in osmoregulatory abilities. There is considerable inter- and intraspecific physiological variation associated with the ecological distribution of each species, respectively.     

Genetic evidence of cryptic speciation within hammerhead sharks (Genus Sphyrna)

Surveys of genetic variation within cosmopolitan marine species often uncover deep divergences, indicating historical separation and potentially cryptic speciation. Based on broad geographic (coastal eastern North America, Gulf of Mexico, western Africa, Australia, and Hawaii) and temporal sampling (1991–2003), mitochondrial (control region [CR] and cytochrome oxidase I [COI]) and nuclear gene (lactate dehydrogenase A intron 6 [LDHA6]) variation among 76 individuals was used to test for cryptic speciation in the scalloped hammerhead, Sphyrna lewini (Griffith and Smith). CR and COI gene trees confirmed previous evidence of divergence between Atlantic and Indo-Pacific scalloped hammerhead populations; populations were reciprocally monophyletic. However, the between-basin divergence recorded in the mtDNA genome was not reflected in nuclear gene phylogenies; alleles for LDHA6 were shared between ocean basins, and Atlantic and Indo-Pacific populations were not reciprocally monophyletic. Unexpectedly, CR, COI, and LDHA6 gene trees recovered a deep phylogenetic partition within the Atlantic samples. For mtDNA haplotypes, which segregated by basin, average genetic distances were higher among Atlantic haplotypes (CR: D _HKY=0.036, COI: D _GTR=0.016) than among Indo-Pacific haplotypes (CR: D _HKY=0.010, COI: D _GTR=0.006) and approximated divergences between basins for CR (D _HKY=0.036 within Atlantic; D _HKY=0.042 between basins). Vertebral counts for eight specimens representing divergent lineages from the western north Atlantic were consistent with the genetic data. Coexistence of discrete lineages in the Atlantic, complete disequilibrium between nuclear and mitochondrial alleles within lineages and concordant partitions in genetic and morphological characters indicates reproductive isolation and thus the occurrence of a cryptic species of scalloped hammerhead in the western north Atlantic. Effective management of large coastal shark species should incorporate this important discovery and the inference from sampling that the cryptic scalloped hammerhead is less abundant than S. lewini, making it potentially more susceptible to fishery pressure.     

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.     

Genetic structure of round scad mackerel <Emphasis Type="Italic">Decapterus macrosoma</Emphasis> (Carangidae) in the Indo-Malay archipelago

Round scad mackerel sampled in 1995-1998 were analysed for genetic variation using mitochondrial-DNA and nuclear-DNA markers. Sequence variation for a fragment of the cytochrome b gene (mitochondrial), amplified by polymerase chain reaction, was screened across individuals using single strand conformation polymorphism (SSCP). Sequence analysis of all SSCP haplotypes indicated two mitochondrial clades separated by, on average, 2.3% nucleotide divergence. The geographic distribution of haplotypes was homogeneous (Weir and Cockerham's [^(q)] \hat \theta =-0.002). Also, no geographic heterogeneity was detected for length polymorphism of Intron 1 of the gene encoding aldolase B ([^(q)] \hat \theta =0.005). Although homogeneity in allele frequencies throughout the Indo-Malay archipelago conformed to the expectations for a widely distributed pelagic fish in a highly connected habitat, this was at variance with the sharp geographic structure previously uncovered in Indian scad mackerel, Decapterus russelli, a fish with life-history characteristics similar to D. macrosoma. A remarkable similarity, however, was the occurrence of two similarly distinct clades within each species, suggesting a common history of geographic isolation. Low sea levels in the Pleistocene might have caused the separation and vicariance of populations within both D. macrosoma and D. russelli. Subsequent genetic exchange between populations would then have erased allele-frequency differences at the cytochrome b and aldolase B loci in D. macrosoma while some barrier to gene flow was maintained in D. russelli.     

Morphological, reproductive, and genetic variability among three populations of Crucibulum quiriquinae (Gastropoda: Calyptraeidae) in northern Chile

Crucibulum quiriquinae (Lesson, 1830) is the only species of Crucibulum currently recognized in northern Chile. Recent analysis of three Crucibulum populations obtained in northern Chile demonstrates the existence of morphological, genetic, and reproductive differences among populations. Two populations present in Bahía Tongoy (30°15'S), one inhabiting the shells of the snail Turritella cingulata and the other shells of the pectinid Argopecten purpuratus, showed morphological differences. However, both had planktonic larval development and show low genetic divergence (D=0.002). A third population from Bahía La Herradura (29°58'S), which also inhabits the shells of T. cingulata, did not show morphological differences compared with its counterpart from Bahía Tongoy. However the Bahía La Herradura population had intracapsular development and metamorphosis, and a larger genetic distance (D=0.06) from both Tongoy populations. The results of the reproductive and genetic analyses strongly suggested that the two Tongoy populations, although showing morphological differences, are biologically the same species, Crucibulum quiriquinae, whereas the La Herradura population is a new species.     

Extreme mitochondrial DNA divergence within populations of the deep-sea gastropod Frigidoalvania brychia

The deep sea supports a diverse and highly endemic invertebrate fauna, the origin of which remains obscure. Little is known about geographic variation in deep-sea organisms or the evolutionary processes that promote population-level differentiation and eventual speciation. Sequence variation at the 16 S rDNA locus was examined in formalin-preserved specimens of the common upper bathyal rissoid Frigidoalvania brychia (Verrill, 1884) to examine its population genetic structure. The specimens came from trawl samples taken over 30 years ago at depths of 457-1,102 m at stations in the Northwest Atlantic south of Woods Hole, Massachusetts, USA. Near the upper boundary of its bathymetric range (500 m), extremely divergent haplotypes comprising three phylogenetically distinct clades (average uncorrected sequence divergence among clades ~23%, ~3% within clades) were found at stations separated by a maximum distance of ~80 km, suggesting the presence of high levels of intraspecific divergence or the possibility of morphologically cryptic species. Only one of these clades was found at two stations in the mid- to lower part of F. brychia's depth distribution (800-1,100 m), suggesting lower clade diversity with increasing depth, although among-sample divergence, with a single exception, was minimal. One station was genetically divergent from all others sampled, containing a unique suite of haplotypes including two found only at this site. Steep vertical selective gradients, major oceanographic changes during the late Cenozoic, and habitat fragmentation by submarine canyons might have contributed to an upper bathyal region that is highly conducive to evolutionary change.     

Stoichiometry of mesozooplankton in N- and P-limited areas of the Baltic Sea

The Baltic Sea is a very suitable site for stoichiometric studies, since its subbasins differ in their concentration of elemental components, and primary production can therefore be either nitrogen or phosphorus limited. To reveal if the nutrient limitation of mesozooplankton mirrors that of the primary producers, carbon, nitrogen and phosphorus content of both seston and grazers (Acartia sp., Centropages hamatus, Daphnia cristata, Eurytemora affinis, Limnocalanus macrurus, Temora longicornis) were measured in midsummer in the Baltic proper, the Gulf of Finland and the Gulf of Bothnia. The mineral ratios of the different taxa were equal, apart from L. macrurus with notably higher C:P and N:P ratios. Molar C:N ratios were relatively stable (5.1-6.3), whereas C:P and N:P ratios fluctuated more (41-144 and 6.6-24). However, zooplankton elemental composition and limitation did not depend on the limiting nutrient of the phytoplankton, the seston mineral ratio or the sea area. Both the seston-zooplankton elemental imbalance and the food threshold ratio indicated phosphorus limitation of most of the grazers. While L. macrurus may be C or N limited, the possible P deficiency of the other studied taxa suggests that the Baltic Sea zooplankton may act as a potential phosphorus sink, as the freshwater secondary producers do.     

Genetic variability and level of differentiation in North Sea and Baltic Sea populations of the green alga<Emphasis Type="Italic"> Cladophora rupestris</Emphasis>

Cladophora rupestris is a perennial filamentous macroalga belonging to the Chlorophyta. It is widely distributed on both sides of the northern Atlantic Ocean and penetrates into the brackish Baltic Sea down to ca. 4 psu salinity. In this paper we present evidence for genetic differentiation of a Baltic form of this marine alga. We assessed genetic structure within and among 11 populations ranging along a salinity gradient from the Norwegian coast to the northern Baltic Sea proper. Samples of 328 individuals were studied using starch-gel protein electrophoresis to evaluate genetic variability and interpopulation differentiation based on allozymes. Of 11 loci examined, only one was polymorphic. For this locus, encoding superoxide dismutase (SOD-3), a total of seven alleles were distinguished. We found two genetically differentiated groups of populations of C. rupestris, one Baltic Sea group and one North Sea group, with a distinct border in the southern Kattegat near the entrance to the Baltic Sea. The genetic differentiation for SOD-3, expressed as pairwise F_ST values between the populations, was generally higher within the Baltic Sea group (0.10-0.43) than within the North Sea group (0.05-0.10); in the latter group also fewer pairs of populations differed significantly. Pairs of populations from different groups had the highest F_ST values (0.20-0.60). Hierarchical analysis of variance showed that 29.6% of the total variation in the SOD-3 locus was explained by variation between the two groups, while only 4.2% was explained by variation among the populations within the groups. The remaining variation (66.2%) was found within the populations.     

Identification of Illex coindetii, I. illecebrosus and I. argentinus (Cephalopoda: Ommastrephidae) throughout the Atlantic Ocean; by body and beak characters

Shortfinned squid species of the genus Illex support commercial fisheries throughout the Atlantic Ocean and Mediterranean Sea. Previous identification of interspecific and intraspecific populations by morphological and size-at-maturity studies have not provided conclusive results. We analysed morphometric body and beak variables (24 characters) in three species of the genus (I. coindetii, I. illecebrosus and I. argentinus), using a geographic and seasonal series of 33 populations for 1,500 specimens of I. coindetii, I. illecebrosus and I. argentinus. Residuals of the regression between each morphometric body and beak variable and mantle length were used as input in a stepwise discriminant analysis. Species discrimination by body and hectocotylus characters required at least eight variables and resulted in high correct-classification percentages for I. coindetii and I. argentinus (75% and 90%, respectively), whereas the best identification resulted from beak characters (83% correctly classified). Size of the suckerless basal arm, sucker-bearing length and beak lateral wall discriminated best among I. coindetii from northern Iberia, northwest Iberia (year-1996) and Ireland in the Atlantic and western Mediterranean versus middle and eastern Mediterranean samples. Canadian shelf and American samples were discriminated from Canadian slope I. illecebrosus. Winter/shelf and winter/slope samples of I. argentinus seemed to form a single biological group separated from Falkland Island, 46°S/autumn spawners and 46°S/1996 specimens along the Patagonian Shelf. No significant sexual or maturity polymorphism was obtained. Discriminant analysis optimised population diagnosis on a morphometric basis of interest in fisheries strategies. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s00227-002-0796-7.     

Seasonal patterns in size and abundance of <Emphasis Type="Italic">Phyllorhiza punctata</Emphasis>: an invasive scyphomedusa in coastal Georgia (USA)

Phyllorhiza punctata, commonly called the Australian white spotted jellyfish, invaded the Caribbean in the 1960s, becoming established there and subsequently in the United States in the northern Gulf of Mexico (by 2000) and eastern Florida (2001). With the prevailing Loop Current flowing clockwise around the Gulf of Mexico and joining the Gulf Stream along eastern Florida, potential transport of P. punctata along the eastern seaboard of the USA could be facilitated. P. punctata medusae were collected in small numbers along the entire Georgia coast during May–November in 2007 and 2008. Medusa bell diameters increased both years from ca. 10 cm in May to ca. 33 cm in autumn. Specimens lacked zooxanthellae, as reported for medusae in the northern Gulf of Mexico and Florida. It is possible that the P. punctata medusae observed were transported from established populations to the south; however, whether or not this species is established along the Georgia coast has yet to be determined.     

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.