Reef coral reproduction in the eastern Pacific: Costa Rica, Panamá and Galápagos Islands (Ecuador). IV. Agariciidae, recruitment and recovery of Pavona varians and Pavona sp.a

The reproductive ecology of two eastern Pacific zooxanthellate coral species was examined as part of a continuing series of studies relating bleaching/mortality events caused by the El Niño–Southern Oscillation disturbance, and is described for study sites in Costa Rica, Panamá, and the Galápagos Islands (Ecuador). This study deals with the sibling agariciid species Pavona varians and Pavona sp.a over a 13?yr period (1985 to 1997). Both Pavona species are broadcast-spawners with some gonochoric, but mostly sequential hermaphroditic colonies. Minimum colony sizes (and ages) at first reproduction were 5?cm (5?yr) and 3?cm (2 to 3?yr), respectively, in P. varians and Pavona sp.a. In the Panamá and Galápagos populations, gonochoric colonies spawn eggs or sperm at least monthly. Six fecundity attributes were not significantly different in the two species, but the eggs of P. varians are white to beige and positively buoyant, and those of Pavona sp.a are dark green and neutrally to negatively buoyant. Eggs of both species lack zooxanthellae. Both species are reproductively active year-round, with maximum activity in the dry season in the nonupwelling Gulf of Chiriquí, and in the wet season in the upwelling Gulf of Panamá. Spawning is predominantly during full moon, and possibly also at new moon at most study sites. Spawning in P. varians and Pavona sp.a is 12?h out of phase, with the former species spawning ～1?h before sunrise and the latter about 1?h after sunset. The fecundity of Pavona spp. at Caño and the Galápagos Islands was much greater (19?900 to 27?900 eggs cm^?2?yr^?1) than at all Panamá sites (14?800 to 19?800 eggs cm^?2?yr^?1). Intraspecific crosses in both species resulted in swimming planula larvae after 25 to 36?h. Recruitment of P. varians was highest in Panamá, moderate in Costa Rica, and nil in the Galápagos Islands, matching, respectively, the contributions of P. varians to the pre-1982/1983 El Niño coral-population abundances in these areas. Recruitment success of P. varians at Uva Island was significantly related to maximum monthly positive sea surface-temperature (SST) anomalies that occurred in the year preceding recruitment over the period 1982 to 1996; recruitment failed when SST anomalies exceeded 1.6 to 1.9?C° during the severe ENSO events of 1982/1983 and 1997/1998.     

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.     

Genetic evolution among selected members of the genus Naso (Nasinae), "unicornfishes" from Guam

A species-level phylogeny of selected surgeonfishes in the genus Naso (Acanthuridae: subfamily Nasinae) was constructed using allozyme data and found to be concordant with previous inferences drawn from morphological data. Naso spp. examined from waters surrounding Guam between 1992 and 1994 were: N. caesius (Randall and Bell, 1992), N. hexacanthus (Bleeker, 1855), N. lituratus (Schneider, 1801), N. thynnoides (Valenciennes, 1835), and N. unicornis (Forskål, 1775). Acanthurus lineatus (Linnaeus, 1758) (Acanthuridae: subfamily Acanthurinae) and Zanclus cornutus (Linnaeus, 1758) (Zanclidae) were included for the purpose of outgroup comparison. Analyses of the allozyme data set generated from 16 presumptive loci using both cladistic and phenetic tree-constructing techniques identified the sibling species N. caesius and N. hexacanthus as the most genetically similar taxa among the Naso species surveyed. The pair shared the most synapomorphic alleles and the lowest degree of genetic divergence with respect to other species studied. The unweighted pair-group method using the arithmetic mean (UPGMA) and the neighbor-joining (NJ) topologies, along with two synapomorphic alleles (Aat-2 and Gpi-A), characterize N. caesius, N. hexacanthus, N. lituratus, and N. unicornis as a monophyletic group. Many allelic autapomorphies were detected in N. thynnoides, a member of the subgenus Axinurus. This supports the morphological data, which suggest that one-plated unicornfishes form a monophyletic group. The proposed partial phylogeny inferred from the allozyme data is concordant with the morphological differences (i.e. dentition, osteology, and myology) that have been used to support the partition of the genus Naso into the subgenera Axinurus and Naso.     

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, 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.     

Competition,fluctuating asymmetry and sperm transfer in male gryllid crickets (<Emphasis Type="Italic">Gryllus bimaculatus</Emphasis> and <Emphasis Type="Italic">Gryllodes sigillatus</Emphasis>)

Sperm competition theory predicts that ejaculate expenditure by males should increase with the risk of competition from rivals. The tendency to modulate expenditure, however, may reflect various constraints acting on different individuals. We test this idea here by looking at the effects of competition and fluctuating asymmetry (FA) in two species of gryllid cricket. Male Gryllus bimaculatus and Gryllodes sigillatus modulated the sperm content of their spermatophores in relation to both apparent competition and FA in limb size. Males of both species increased the amount of sperm transferred when an apparent male competitor was present, sperm number increasing regardless of the species of apparent competitor in G. bimaculatus, but significantly more with a conspecific competitor in G. sigillatus. In addition, the relative size of the spermatophylax in G. sigillatus increased in the presence of apparent competition, while spermatophore transfer time decreased in G. bimaculatus. Sperm number showed opposite relationships with limb asymmetry in the two species, decreasing with increasing asymmetry in males in G. sigillatus, but increasing with asymmetry in both males and females in G. bimaculatus. G. bimaculatus, but not G. sigillatus, also transferred more sperm when paired with a larger female. Relationships between reproductive measures and limb asymmetry were significant only when competitors were present in G. bimaculatus, but were significant regardless of competition in G. sigillatus. The differences may reflect the additional burden of producing a spermatophylax in G. sigillatus.     

Characterization of urease activity in three marine phytoplankton species,<Emphasis Type="Italic"> Aureococcus anophagefferens</Emphasis>,<Emphasis Type="Italic"> Prorocentrum minimum</Emphasis>, and<Emphasis Type="Italic"> Thalassiosira weissflogii</Emphasis>

The availability of different forms of nitrogen in coastal and estuarine waters may be important in determining the abundance and productivity of different phytoplankton species. Although urea has been shown to contribute as much as 50% of the nitrogen for phytoplankton nutrition, relatively little is known of the activity and expression of urease in phytoplankton. Using an in vitro enzyme assay, urease activities were examined in laboratory cultures of three species: Aureococcus anophagefferens Hargraves et Sieburth, Prorocentrum minimum (Pavillard) Schiller, and Thalassiosira weissflogii (Grunow) Fryxell et Hasle. Cultures of P. minimum and T. weissflogii were grown on three nitrogen sources (NO₃^m, NH₄⁺, and urea), while A. anophagefferens was grown only on NO₃^m and urea. Urease was found to be constitutive in all cultures, but activity varied with growth rate and assay temperature for the different cultures. For A. anophagefferens, urease activity varied positively with growth rate regardless of the N source, while for P. minimum, urease activity varied positively with growth rate only for cultures grown on urea and NH₄⁺. In contrast, for T. weissflogii, activity did not vary with growth rate for any of the N sources. For all species, urease activity increased with assay temperature, but with different apparent temperature optima. For A. anophagefferens, in vitro activity increased from near 0-30°C, and remained stable to 50°C, while for P. minimum, increased in vitro activity was noted from near 0-20°C, but constant activity was observed between 20°C and 50°C. For T. weissfloggii, while activity also increased from 0°C to 20°C, subsequent decreases were noted when temperature was elevated above 20°C. Urease activity had a half-saturation constant of 120-165 wg atom N l^у in all three species. On both an hourly and daily basis, urease activity in A. anophagefferens exceeded nitrogen demand for growth. In P. minimum, urease activity on an hourly basis matched the nitrogen demand, but was less than the demand on a daily basis. For T. weissflogii, urease activity was always less than the nitrogen demand. These patterns in urease activity in three different species demonstrate that while apparently constitutive, the regulation of activity was substantially different in the diatom. These differences in the physiological regulation of urease activity, as well as other enzymes, may play a role in their ecological success in different environments.     

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.     

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.     

Interspecific infection of aposymbiotic juveniles of <Emphasis Type="Italic">Codakia</Emphasis><Emphasis Type="Italic">orbicularis</Emphasis> by various tropical lucinid gill-endosymbionts

Previous molecular phylogenetic analyses have shown that five tropical lucinid species living in or near Thalassia testudinum seagrass beds are colonized by the same bacterial symbiont species. In addition, a new lucinid species belonging to the genus Anodontia, which inhabits reducing sediment found near seagrass beds and in mangrove swamps, has been included in the present study. Endosymbiosis in Anodontia alba was examined according to symbiont phylogenetic and gill ultrastructural analysis. Phylogenetic analysis showed that partial 16S rDNA sequences of A. alba- and Codakia orbicularis-symbionts were 100% identical at all nucleotide positions determined, suggesting that A. alba also harbors the same symbiont species as C. orbicularis (and, consequently, as C. orbiculata, C. pectinella, Linga pensylvanica and Divaricella quadrisulcata). Based on light and electron microscopy, the cellular organization of the gill filament appeared similar to those already described in other lucinids. The most distinctive feature is the lack of "granule cells" in the lateral zone of A. alba gill filaments. In order to confirm the single-species hypothesis, purified fractions of gill bacterial symbionts obtained from the gills of each of the six tropical lucinids cited above were used to infect aposymbiotic juveniles of C. orbicularis. In each case, aposymbiotic juvenile batches were successfully infected by the gill-endosymbiont fractions, whereas, during the experiments, juveniles from the negative control were still uninfected. These experimental data confirm the phylogenetic data and also demonstrate that chemoautotrophic bacterial endosymbionts from their host cells can colonize aposymbiotic juveniles. The conclusion also follows that intracellular gill-endosymbionts still have the capacity to recognize and colonize new host generations. Lucinids provide a unique model for the study of sulfide-oxidizing symbiosis, even if symbionts remain unculturable.     

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.     

Identification of tenuis of four French Polynesian Carapini (Carapidae: Teleostei)

Four species of adult Carapini (Carapidae) occur on Polynesian coral reefs: Encheliophis gracilis, Carapus boraborensis, C. homei and C. mourlani. Samples collected in Rangiroa and Moorea allowed us to obtain different tenuis (larvae) during their settlement phases or directly inside their hosts. These were separated into four lots on the basis of a combination of pigmentation, meristic, morphological, dental and otolith (sagittae) features. Comparison of these characters with those of the adults allows, for the first time, taxonomic identification of these tenuis-stage larvae.     

Studies on pigments,proteins and photosynthetic rates in some mangroves and mangrove associates from Bhitarkanika,Orissa

Pigment contents, proteins and net photosynthesis were investigated in fully developed leaf of 1-year-old seedlings of six mangroves (Bruguiera gymnorrhiza, Rhizophora apiculata) and mangrove associates (Caesalpinia bonduc, Cerbera manghas, Derris heterophylla, Thespesia populnea), collected from Bhitarkanika, located on the east coast of India. Large variations in the photosynthetic rates (P_N) among the six species were observed, ranging from 10.16 µmol CO₂ m^-2 s^-1 in C. bonduc to 15.28 µmol CO₂ m^-2 s^-1 in R. apiculata. The total leaf protein content ranged from 12.09 mg g^-1 dry wt in T. populnea to 51.89 mg g^-1 dry wt in B. gymnorrhiza. The chlorophyll a/b ratio was typically about 3.0 in all the studied species, except C. bonduc (2.8). Photosynthetic rates and chl a/b ratio in the leaves were found to be correlated. Analysis of chlorophyll and xanthophyll spectra suggested: (1) variations in different forms and amounts of carotenes as well as xanthophylls and (2) the presence of high amounts of near-UV-absorbing substances in leaves, particularly in the two mangroves (B. gymnorrhiza, R. apiculata) and a mangrove associate (T. populnea), which appears to be an adaptive feature. Estimation of the chl a/b ratios in isolated thylakoids yielded a low value of 1.8 for R. apiculata and >2.6 for other species. The total protein/chlorophyll ratios in thylakoids varied considerably from 3.14 (D. heterophylla) to 10.88 (T. populnea) among the mangrove associates and from 16.09 to 18.88 between the members of the Rhizophoraceae. The chlorophyll/carotenoid ratios in thylakoids of the six species were more or less similar. The absorption spectra for washed thylakoids of C. manghas and D. heterophylla exhibited absorption characteristics typical for C₃-plant thylakoids. However, thylakoids isolated from R. apiculata, B. gymnorrhiza, C. bonduc and T. populnea exhibited an unusual increase in absorption in the blue region (380-410 nm) of the absorption spectrum. The presence of high-absorbing (in the short-wavelength, near-UV region) pigments appears to be closely associated with the thylakoids in R. apiculata and T. populnea. Our results, therefore, suggest a wide range of variation, not only in protein and pigment contents of photosynthetic tissues, but also in the spectral characteristics and composition of the pigments in mangrove species. An understanding of the nature of these pigments in mangroves and their associates, under their natural conditions and especially in relation to eco-physiological adaptations, is necessary, not only in relation to conservation, but also to allow propagation under different salinity conditions.     

Genetic structure of deep-water redfish, Sebastes mentella, populations across the North Atlantic

Patterns of genetic diversity of the marine deep-water redfish, Sebastes mentella Travin, from 21 sampling locations (n=46 per sample, from 1995 to 1998) were examined throughout its range in the North Atlantic. Eight microsatellite loci were used in order to: (1) define the genetic structure and consider the possible influence of dispersal, geographic distance, oceanic currents and historical factors; and (2) relate the structure to present management units and practices. Three divergent population units (mean Š=0.012) associated with distinct geographic areas were defined: Eastern (Norway and Barents Sea), Panoceanic and Western (Gulf of St. Lawrence and offshore Newfoundland). The most important observation was the lack of genetic differences (mean Š=-0.0004) and lack of genetic isolation by geographic distance (r=0.318, P=0.112) among samples within the Panoceanic zone, from the Faroe Islands to the Grand Banks (6,000 km). A combination of vicariance, historical introgression with a closely related species, S. fasciatus, and contemporary patterns of oceanic circulation are likely to have shaped the observed population structure. These results only partially support current management units, and call for more integrated practices for regulating the exploitation of S. mentella throughout its extensive range.     

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.     

Assessment of microhabitat preferences in juvenile dusky grouper (Epinephelus marginatus) by visual sampling

As a top-level predator, the brown grouper Epinephelus marginatus can play an important role in maintaining the ecological balance of hard-bottom ecosystems. However, to fulfil this role, the species must have a sufficient population density and a wide size range. The presence of such a "healthy" grouper population is one of the known benefits of the protection measures applied to marine protected areas. The availability, in marine reserves, of areas suitable for settlement and recruitment of early juveniles can contribute to preserving a well-structured population. Thus, knowledge of microhabitats preferred by juvenile groupers is an important step in locating, within protected or not yet protected areas, nursery sites that might need a specific protection regime. The aim of the present work was to evaluate habitat and microhabitat preferences of juvenile E. marginatus, in comparison with two other serranids of comparable size, Serranus cabrilla and S. scriba, in the marine reserve of Ustica Island (SW Mediterranean). At different sites, located along the shallow coastal area of the reserve, 329 individuals of the three serranid species were visually surveyed by means of skin- or SCUBA-diving in June 1999. The location of each encountered fish was characterised by both biotic and abiotic variables evaluated at two spatial scales. In order to assess interspecific differences in the use of the spatial environment, the results were analysed by correspondence analysis. According to observations on a small spatial scale, juvenile groupers showed a preference for cavities and recesses, in clear contrast with both Serranus species. When out of such sheltered places, juvenile groupers avoided visually exposed locations (convex substrates and very large visual fields), preferring flat or sub-horizontal rocky substrates. Conversely, S. scriba, and especially S. cabrilla, chose rather open microhabitats (flat to convex substrates, with large to very large visual fields). On a larger spatial scale, brown groupers and the two other serranids showed no marked differences in their habitat preferences.     

Seasonal variation in ecophysiological patterns in macroalgae from an Arctic fjord. I. Sensitivity of photosynthesis to ultraviolet radiation

Marine macroalgae inhabiting Arctic coastal ecosystems are exposed to pronounced seasonal variations in the radiation regime, including harmful UVB radiation. This study presents the first data on the seasonal changes in the sensitivity of macroalgal photosynthesis towards UV exposure by comparing under-ice, clear-water and turbid-water conditions characteristic for late winter, spring and summer. Various brown (Laminaria saccharina, L. digitata, L. solidungula, Saccorhiza dermatodea, Desmarestia aculeata), red (Palmaria palmata, Devaleraea ramentacea) and one green macroalgal species (Monostroma aff. arcticum) were collected at the same water depth throughout the seasons in the Kongsfjord (Spitsbergen, Svalbard, Norway). Maximum quantum yield (F_v/F_m) and maximum photosynthetic electron transport rates (ETR_max) were determined immediately after collection, after 2 h exposure to artificial UV radiation and after 18 h recovery in dim white light. Photosynthesis of the studied species showed different responses depending on their morpho-functional and physiological characteristics, their life strategies, phenology and depth distribution. Within the genus Laminaria, maximum quantum yield of adult specimens of the deep-water species L. solidungula was most strongly UV sensitive. Adult L. saccharina exhibited a lower UV sensitivity than a 6-month-old specimen. Inhibition of photosynthesis after UV exposure remained at the same level throughout the study period, both in adult L. saccharina and S. dermatodea. However, adult specimens of L. saccharina collected in May showed partial recovery only, whereas photosynthesis of specimens from both species collected later recovered fully. D. aculeata exhibited a remarkable decrease of UV sensitivity during the study period. Photosynthesis of specimens collected under the ice was strongly inhibited by UV, but the degree of inhibition decreased during spring and summer. Concomitantly ETR_max values were low after UV exposure in specimens collected in June, but increased later in the season. P. palmata exhibited a relatively flexible response. Photosynthesis in specimens collected under the ice in June or in turbid water in July/August was relatively strongly inhibited; specimens collected during sunny periods and in clear water in spring showed a much lower degree of photoinhibition after UV exposure. The seasonal pattern of low/high ETR_max values in spring/summer is probably a characteristic of the life strategy of this species. The UV sensitivity of D. ramentacea exhibited a similar seasonal pattern. In M. aff. arcticum, UV sensitivity increased and ETR_max values decreased during the study period, reflecting the life strategy of this annual late winter/spring species. The physiological basis for the seasonal changes in UV sensitivity of photosynthesis is presented in a companion paper (this issue).     

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.     

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.