Biodiversity of marine sessile epifauna at an Aegean island subject to hydrothermal activity: Milos, eastern Mediterranean Sea

Sessile macroepifauna was sampled at six rocky sites between 2 and 90 m depth with a number of different methods involving both underwater photography and collection of specimens. A total of 212 species (or varieties) were identified, belonging to seven higher taxa: poriferans (24 species), cnidarians (32), molluscs (8), serpuloidean polychaetes (33), bryozoans (90), brachiopods (4) and ascidians (21). The combined use of a varied array of sampling methods was effective in obtaining a rich faunal inventory. Deep and offshore sites tended to be richer in species than shallow and inshore sites. In all cases species richness was higher at sites closest to hydrothermal vents on the sea floor. Although there are no comparable inventories of marine sessile epifauna in the Aegean, the high number of species found, with a relatively low sampling effort in a restricted area, indicates that the marine biodiversity of this sea is not as low as traditionally believed. Received: 20 January 1999 / Accepted: 25 August 1999     

The effects of feeding or addition of dissolved inorganic nutrients in maintaining the symbiosis between dinoflagellates and a tropical marine cnidarian

The availability of solid food (Artemia nauplii) and dissolved inorganic nutrients (ammonium, nitrate, phosphate) to the shallow-water marine hydroid Myrionema amboinense was manipulated for 1-8 days in order to investigate their role in the growth of intracellular symbiotic dinoflagellates (zooxanthellae) of the genus Symbiodinium. Symbionts from hydroids collected from the field or maintained under laboratory conditions (25°C, 12 h:12 h light:dark cycle, 80 µE m^-2 s^-1 fluorescent lighting) always exhibited a single peak in mitotic index (MI) at dawn. Symbionts in freshly collected field animals had an MI peak of about 15%. Symbiotic dinoflagellates in hydroids fed Artemia nauplii twice daily in the laboratory maintained this dawn peak of MI between 10% and 15%, but in the absence of feeding or added inorganic nutrients, this peak declined to less than 1% within 2-4 days. In contrast, when hydroids were placed in solutions containing ammonium (20 µM NH₄Cl), nitrate (10 µM NaNO₃), and a combination of ammonium and phosphate (2 µM Na₂HPO₄) immediately after collection, the algal MI remained between 5% and 15% for 4-7 days; the addition of 2 µM phosphate did not increase MI relative to unfed rates. When unfed animals were placed in dissolved nitrogen or fed Artemia, the symbiont MI increased from <1% to 10-17% within 2-3 days; P alone had no effect. However, the increase resulting from added inorganic nutrients was temporary, lasting only 5-7 days. These observations suggest that algal division in the host is maintained indefinitely in the field or by feeding particulate foods twice daily in the laboratory, but the addition of inorganic nutrients alone (ammonium, nitrate and ammonium/phosphate) appeared to support the completion of a maximum of one additional round of cell division. Nutrients required for continued growth and division of symbiotic dinoflagellates are linked to host feeding and host growth; without external food, neither host nor symbiont continue to grow. The same phenomenon is seen in zooxanthellate anemones, clams and corals, where total numbers of symbionts appear to be linked to changes in host-tissue biomass (protein), achieving relatively stable densities in M. amboinense, corals and other cnidarian symbioses, depending on their local environmental conditions. The results of the present study help explain the cellular responses of algal symbionts in reef-dwelling invertebrates to additions of dissolved inorganic nutrients to coral-reef ecosystems.     

To what extent does upright sessile epifauna affect benthic biodiversity and community composition?

Small-scale habitat complexity, including that caused by biological structures, is an important factor in structuring benthic communities and also sometimes in increasing biodiversity. The aim of this study was to determine if hydroid colonies have an effect on the composition of benthic communities in the Irish Sea, and if so, which components of the fauna are affected. Forty-six seabed core samples were taken by divers from two sites off Port Erin, Isle of Man, Irish Sea. Half of these were centred on hydroid colonies, half were not. All taxa retained by a 63-m sieve from the cores were identified and counted. Community composition and diversity were compared between hydroid and non-hydroid cores using multivariate and univariate methods. Benthic communities were significantly different between the two sample groups. This was almost entirely due to the presence of sessile and mobile epifaunal taxa in the hydroid cores. The tube-building amphipod, Ericthonius punctatus, was particularly abundant attached to the hydroid stems. Infauna was not significantly different between the two groups. Upright sessile epifauna may play a particularly important role in the Irish Sea as a settlement substrate for juvenile scallops (Pecten maximus and Aequipecten opercularis), which are an important fishery resource in this area. The 11-year closure of an area to dredging has not only enhanced scallop stocks but has had the added benefit of enhancing habitat complexity and biodiversity.Communicated by J.P. Thorpe, Port Erin     

Horizontal,vertical and seasonal distributions of epiphytic hydrozoa on the algaHalimeda tuna in the Northwestern Mediterranean Sea

The thalli ofHalimeda tuna (Ellis & Solander) grow horizontally from rocky surfaces and present an upper and lower surface which can be colonized by epiphytes. Thalli ofH. tuna were collected twice a month from March 1985 to March 1986 at Tossa de Mar (northeast Spain). Thirty five species of hydroid were recorded living onH. tuna thalli; of these, the 15 most abundant species were classified into three groups on the basis of their horizontal zonation on the thalli: those most abundant on the proximal, medial or distal articles. Those hydroids common on the proximal (oldest) articles were relatively large and were present throughout the year, those common on the medial articles were smaller and often occurred in dense monospecific patches, while those common on the distal (youngest) articles were present for only short periods and were the most opportunistic hydroids found onH. tuna. We hypothesize that this zonation reflects interspecific niche-selection, enabling successful competition for space with other hydroids, algae and bryozonas. Three hydroid species were more abundant on the upper surfaces of the algae than on the lower, probably because of gravitationally enhanced settlement of planula larvae coupled with (unusually) limited hydrorhizal growth on the underside. Three opportunistic species were more common on the lower surfaces of the thalli, possibly reflecting the greater availability of substratum in the shade, where epiphytic algae were less abundant. Many hydroids declined in abundance in summer because of the death of old thalli, the growth of new thalli and of apical articles on existing thalli, and possibly because of interspecific competition with epiphytic algae. The death of old thalli and the growth of new thalli reduced the abundance of epiphytic algae; this may have facilitated the increase in abundance of hydroids recorded in the autumn.     

Inbreeding and hermaphroditism in the sessile-brooding bryozoan Celleporella hyalina

This study examined the occurrence and effects of inbreeding in the sessile, colonial hermaphrodite Celleporella hyalina. The results are discussed with regard to theoretical explanations for the prevalence of hermaphroditism in sessile clonal organisms. C. hyalina exhibited inbreeding depression at all stages, including the pre-zygotic. Outcrossing, sib and half-sib matings produced offspring but selfing did not. There was inbreeding depression in embryo production and survival. Inbred colonies showed slower growth and later maturation, with fewer reproductive zooids. The relative numbers of male and female zooids were affected by inbreeding; increased production of males is a sign of stress in C. hyalina. When mated with an outbred non-relative, inbred colonies had lower success both as males and as females in embryo production and offspring survivorship. The low survivorship of embryos fathered by inbred colonies is a clear effect of inbreeding on the F2 generation. These results indicate that C. hyalina is unlikely to inbreed in the wild, supporting the "space-limited" model of hermaphroditism for this species. This study indicates that hermaphroditism, by avoiding much of the cost of sex, can confer optimal reproductive fitness for sessile brooding animals in a space-limited habitat even in the absence of inbreeding.     

Food division within two sympatric species-pairs of skates (Pisces: Rajidae)

Food habits of two sympatric species-pairs of skates (Raja erinacea-R. ocellata and R. radiata-R. senta), which occur off the east coast of North America were investigated. Stomachs from over 1600 specimens of the 4 species were collected during winter, summer, and autumn of 1969 and the winter of 1970. Diets of R. erinacea and R. ocellata consisted largely of amphipods, decapod crustaceans and polychaetes. However, R. ocellata consumed relatively more fishes and polychaetes and less decapods than R. erinacea. These two skates ate many of the same species but in different proportions; R. ocellata tended to feed on infauna and R. erinacea tended to feed on epifauna. Food preferences of the two species may be correlated to the difference in shape of the mouth and number of tooth rows. R. radiata and R. senta both fed heavily on decapod crustaceans and euphausids, but polychaetes were relatively more important to R. radiata and mysids were relatively more important to R. senta. R. radiata had a very diversified diet and fed on both epifauna and infauna. The diet of R. senta was very restricted and consisted almost entirely of epifauna. Diets of the two species-pairs were similar, but isopods and bivalves were more important to the R. erinacea-R. ocellata pair and euphausids and mysids were more important to the R. radiata-R. senta pair. These differences may reflect differences in the benthic communities with which the species-pairs are associated. Amount of overlap in resource utilization of the pairs of skates was compared with that of some other congeneric organisms.Virginia Institute of Marine Science Contribution No. 727.     

Long-term effects of ultraviolet radiation on growth and photosynthetic performance of polar and cold-temperate macroalgae

Long-term effects of artificial ultraviolet radiation (UV) and natural solar radiation on growth and photosynthetic activity, as measured by chlorophyll fluorescence, were investigated in 13 different polar and cold-temperate macroalgal species. Isolates of five different species from the Arctic and Antarctic were exposed to different light treatments of photosynthetically active radiation (PAR), PAR+UVA and PAR+UVA+UVB. Eight different species collected on the island of Helgoland, North Sea, Germany, were studied in the laboratory and under natural solar radiation conditions. Increase in fresh weight and changes in photosynthetic performance were monitored over a period of 3-4 weeks. The sublittoral polar species, particularly the Antarctic red algal species Gymnogongrus antarcticus and G. turquetii, the Arctic cold-temperate brown alga Alaria esculenta and, very drastically, the Arctic-endemic Laminaria solidungula, exhibited strong inhibiting effects of artificial UVB radiation on growth. In the cold-temperate sublittoral growth of the red algae Phycodrys rubens and, to a lesser extend, Membranoptera alata was substantially inhibited by UV radiation. In contrast, eulittoral species, e.g. Fucus serratus, did not show any differences in growth with respect to artificial irradiation conditions, with or without UV radiation. In the laboratory, some individuals of the green alga Codium fragile exhibited strong morphological changes of the whole thallus, particularly under UVB exposure. In the experimental outdoor set up, growth of most of the algal species was already inhibited by the full solar UV waveband, but, apart from Polyides rotundus, no additional UVB effect could be detected. Changes of in vivo fluorescence were not always consistent with the measurable changes in growth rate, indicating that physiological processes leading to an inhibition of growth may act independently of changes in photosynthetic activity. For the polar species, a general correlation between the natural vertical distribution in the field and the individual sensitivity towards UV radiation was indicated, while for eulittoral species from Helgoland no clear relationship was found. The obtained results show that measuring growth is a good ecological parameter to monitor long-term effects of UV radiation on single macroalgal species and the possible resulting changes of whole algal communities in coastal ecosystems.     

Molecular systematic and phylogenetic assessment of 34 calanoid copepod species of the Calanidae and Clausocalanidae

DNA sequences for a 639 bp region of mitochondrial cytochrome oxidase I (mtCOI) were determined for 34 species of ten genera in two families of calanoid copepods, including: Calanoides, Cosmocalanus, Meoscalanus, Nannocalanus, Neocalanus, and Undinula (family Calanidae); and Clausocalanus, Ctenocalanus, Drepanopus, and Pseudocalanus (family Clausocalanidae). MtCOI gene sequences proved to be diagnostic molecular systematic characters for accurate identification and discrimination of the species. Levels of mtCOI variation within species (range: 1-4%) were significantly less than those between species (9-25%). Higher levels of intraspecific variation (>2%) usually resulted from comparisons between ecologically distinct or geographically isolated populations. MtCOI sequence variation resolved evolutionary relationships among species of Clausocalanus, Neocalanus, and Pseudocalanus, although there was evidence of saturation at some variable sites. Phylogenetic relationships among 11 copepod genera (adding Calanus to the list above) were reconstructed using a 660 bp region of nuclear small-subunit 18S rRNA, a slowly evolving gene that showed no variability within a species and differed by <1-6% among the genera. The 18S rRNA molecular phylogeny was consistent with the accepted limits of the Calanidae and Clausocalanidae and clearly resolved relationships among genera within each family. This molecular systematic and phylogenetic study was part of the ZooGene project, an international partnership to create a DNA sequence database as a tool for uniform, molecularly based species identification of planktonic calanoid copepods and euphausiids.     

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.     

Ecology of neonate loggerhead turtles inhabiting lines of downwelling near a Gulf Stream front

Neonate sea turtles disperse from nesting beaches into the open ocean and develop during a multi-year growth period at sea, but data that characterize their behavior, feeding, and habitat during this developmental period have been few. Limited information has suggested that neonate sea turtles associate with lines of floating debris and biota at areas of surface downwelling. Data from the present study come from measurements of habitat, turtle behavior, and apparent foraging preferences in areas where neonate (post-hatchling) loggerhead turtles (Caretta caretta) were observed and captured. Turtles were observed (n=293) and captured (n=241 of the 293 observed) in downwelling lines that had formed in the slope water near the Gulf Stream front off east-central Florida, USA. Catch-per-unit-effort averaged 12.4 turtles/h from a vessel moving at 2.5 knots. Turtles were largely inactive and were closely associated with floating material, especially pelagic species of Sargassum. Turtles captured along with samples of floating material and given a gastric-esophageal lavage showed a preference for animal material (35.5% of volume in habitat, 70.9% in lavage) over plant material (60.3% of volume in habitat, 22.5% in lavage). Ingested anthropogenic debris included tar (20% of turtles) and plastics (15% of turtles). Ingested animals were principally small (most <1 mm) and were typically slow-moving or non-motile species or stages. Ingested plants were most commonly Sargassum fragments or seagrasses that bore epiphytic animals. Preferred or commonly ingested animals were hydroids, copepods, and pleuston such as Janthina, Creseis, Porpita, and Halobates. Data support a hypothesis describing post-hatchling loggerheads as facultatively active but principally low-energy float-and-wait foragers both within and outside of downwelling lines. Pelagic dispersal of turtles may be best predicted by a "smart" drifter analogy wherein turtle buoyancy, surface advection, and minimal oriented movement determine their distribution at sea. Conservation implications of plastic and tar ingestion are discussed.     

A re-evaluation of the diel feeding hypothesis for marine herbivorous fishes

We re-evaluated the "diel feeding hypothesis" by measuring diel variation in starch, protein, and floridoside in three algal "types" collected from a fringing coral reef at Lizard Island, Great Barrier Reef, Australia. Samples of two species of rhodophyte algae, Gracilaria arcuata and Acanthophora spicifera, and the turf assemblage from the territories of the herbivorous pomacentrid Stegastes nigricans were collected at four time periods through the day: 0630-0730, 1000-1100, 1330-1430, and 1630-1730 hours. We also measured the ability of several species of marine fish (the herbivores Acanthurus nigricans, A. lineatus, A. olivaceus, and Parma alboscapularis and the detritivore Ctenochaetus striatus) to hydrolyse floridoside by estimating !-galactosidase activity in tissue from the anterior intestine. We detected no diel pattern in protein content of the algae but found a significant steady increase in starch content throughout the day. Floridoside content increased in the morning and decreased in the afternoon, a pattern that may be driven by midday photoinhibition of the algae. All the fishes tested could utilise floridoside. Our results support the diel feeding hypothesis. Although floridoside content decreased in the afternoon, our results suggest floridoside was used during the day by the algae to synthesise starch. Thus the algae increased in nutritional value until photoinhibition occurred at midday then subsequently maintained their nutritional value during the afternoon. This pattern of algal nutrients increasing to a midday peak and remaining relatively constant throughout the afternoon correlates well with the diel feeding pattern in many species of marine herbivorous fish.     

Mechanisms of range expansion in the intertidal brown alga <Emphasis Type="Italic">Fucus</Emphasis><Emphasis Type="Italic">serratus</Emphasis> in northern Spain

The intertidal brown alga Fucus serratus is naturally expanding its distributional range along the north shore of Spain. Shores being colonised by F. serratus harbour saturated algal communities, with almost no free space. Processes responsible for the successful expansion were investigated in a locality where the alga was experimentally introduced. After a short period, the number of F. serratus plants increased exponentially at a monthly intrinsic rate of 0.111. At early and intermediate stages of invasion, colonisation occurred by the growth of patches of the invader around the transplant points. F. serratus is a bad disperser, with most of propagules settling in near proximity to parent plants. Colonisation of manipulated and control areas was influenced by the proximity of propagule sources. Experimental removals of canopies, irrespective of whether the turf was removed or left intact, accelerated the expansion, as this component of the community had a negative influence on recruitment of both F. serratus and F. vesiculosus. The unmanipulated algal beds were, however, invasible, as some control quadrats close (<10 m) to F. serratus patches had a 100% cover of F. serratus at the end of the study. Prior or concurrent to increases of F. serratus, a decrease in cover of F. vesiculosus and other resident macroalgae was evident. No formal competition experiment was done but results suggest that F. serratus outcompetes F. vesiculosus and Bifurcaria bifurcata at mid-tidal levels. In scraped quadrats, in which both Fucus species recruited at large densities, F. serratus grew at a higher rate than F. vesiculosus and eventually monopolised the experimental surfaces.     

Ecology of the bivalve-inhabiting hydroid Eugymnanthea inquilina in the coastal sounds of Taranto (Ionian Sea,SE Italy)

The life history of the bivalve-inhabiting hydroid Eugymnanthea inquilina was investigated in two different hosts, the mussel Mytilus galloprovincialis (collected from 1989–1991 from the Ionian Sea, Italy) and the clam Ruditapes decussatus (collected from 1991–1992, Ionian Sea, Italy). Hydroids living in mussels revealed a size selection for hosts longer than 40 mm, being almost completely absent in mussels below this size. This might be controlled by signals linked to the sexual maturity of the bivalve. The proportion of molluscs inhabited showed a seasonal trend in mussels only, with a sharp decline at the onset of medusoid liberation. Production of medusoids was high in mussels, whereas medusoids were rare and often abortive in clams. This indicates a higher degree of fitness of E. inquilina in mussels than in clams. The hydroid life cycle was not completed in clams, which therefore were presumably supplied by planulae from medusoids produced by mussel-inhabiting hydroids. Mussels played a key role in the stability and persistence of E. inquilina populations in the studied area. Selective ingestion of trematode sporocysts by E. inquilina hydroids indicated a protective role of the hydroid against mussel parasites, leading to reconsideration of this symbiotic association as a possible mutualism rather than a simple inquilinism.     

Effects of substrate type on growth and mortality of blue mussels (<Emphasis Type="Italic">Mytilus</Emphasis><Emphasis Type="Italic">edulis</Emphasis>) exposed to the predator <Emphasis Type="Italic">Carcinus</Emphasis><Emphasis Type="Italic">maenas</Emphasis>

Structure and complexity of the substrate are important habitat characteristics for benthic epifauna. The specific growth and mortality rates and inducible defence characters on medium-sized blue mussels (Mytilus edulis L.) exposed to shore crabs (Carcinus maenas L.) were examined on three different substrate types in combined field and laboratory experiments. The experiments showed that complexity of the substrate increased blue mussel survival significantly, through a decrease in predation pressure. However, increased intraspecific competition for food on the complex substrate resulted in significantly lower growth rates of the mussels. Inducible defence characters were also influenced by substrate type. Blue mussels were more affected by predators on the structurally simple substrate, where they developed thicker shells and a larger posterior adductor muscle.     

Cryptic species of Clavelina (Ascidiacea) in two different habitats: harbours and rocky littoral zones in the northwestern Mediterranean

Marinas and harbours provide ideal sites for the study of population genetics of marine invertebrates with restricted dispersal capabilities. They combine a confinement effect, particular ecological conditions (pollution, turbidity), and the possibility of high gene flow through ship-borne propagules, which greatly increases the natural dispersal capability of sexual and asexual propagules in many species with short-lived larvae. We studied the genetic structure of populations of the ascidian Clavelina lepadiformis living inside and outside harbours in the north-western Mediterranean. A 500-bp segment of the cytochrome c oxidase subunit I (COI) mitochondrial gene was sequenced in three populations from inside harbours (interior form) and in three populations from the rocky littoral (exterior form). Two congeneric Mediterranean species, Clavelina sp. and C. dellavallei, were used for comparison. We found that the interior and exterior forms of C. lepadiformis belong to two distinct clades, with a genetic divergence of 5%. Gene-flow values among these forms were insignificant. The lack of gene flow and the genetic divergence suggest that the interior and exterior forms of C. lepadiformis are in fact cryptic species rather than differentiated populations of the same species. Levels of gene flow were higher among interior habitats than among exterior habitats, a pattern likely maintained by genetic exchange through ships. We discuss the possible origins of the present-day distribution of these cryptic species. We contend that the study of species living both inside and outside these particular habitats will reveal more instances of genetic discontinuities allowing local adaptations.     

Animals on marine flowers: does the presence of flowering shoots affect mobile epifaunal assemblage in an eelgrass meadow?

Eelgrass, Zostera marina, produces two types of shoots: morphologically simple vegetative shoots and highly branched flowering (reproductive) shoots, the latter found only in summer months. We examined whether the abundance and diversity of mobile epifaunal assemblage are affected by the presence of flowering shoots in an eelgrass meadow of Otsuchi Bay, northeastern Japan. Comparisons of epifauna in natural vegetation revealed that density and species richness did not differ significantly between sites consisting of both flowering and vegetative shoots, and those only of vegetative shoots. A transplant experiment, conducted to examine the colonization rates of epifauna to defaunated eelgrass planted with different combination of vegetative and flowering shoots, showed no obvious variation in abundance and species richness. At species level, the density of some species such as a tanaid Zeuxo sp. and a polychaete Platynereis sp. was higher at sites and/or treatments with flowering shoots, whereas that of some gastropods, such as Lirularia iridescens and Siphonacmea oblongata was higher at sites without flowering shoots. The species-specific response led to dissimilarity of epifaunal assemblage between sites and among treatments with different densities of vegetative and flowering shoots. Similar patterns observed for natural vegetation and the transplant experiment suggest that the variation in assemblage structure is caused by habitat selection of each species, for example, the utilization of flowering shoots as feeding ground and nursery by Zeuxo sp.     

Deep-sea harpacticoid copepod diversity maintenance: The role of polychaetes

Several models to account for enhanced diversity in the deep sea have been proposed, but the available natural history information has been inadequate to distinguish among them. In particular, few data exist on patterns of co-occurrence among species. At 1220 m depth in the San Diego Trough (32°35.75N; 117°29.00W), harpacticoid copepod species covary significantly with polychaetes when the polychaetes are combined into functional groups on the basis of feeding type and mobility. In particular, harpacticoid species tend to avoid polychaetes which are sessile surface-deposit feeders. The results provide support for models in which disturbance/predation plays an important role in maintaining deep-sea diversity.Contribution No. 15 from Expedition Quagmire.     

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.     

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.