Mitochondrial DNA analysis of the genetic relationships among populations of scad mackerel (Decapterus macarellus, D. macrosoma, and D. russelli) in South-East Asia

The genetic relationships among South-East Asian populations of the scad mackerels Decapterus macarellus, D. macrosoma and D. russelli (Pisces: Carangidae) were investigated. In 1995 and 1996, 216 fish were sampled in seven localities spanning the seas of Indonesia and were examined for restriction-site polymorphisms using ten restriction enzymes for the mitochondrial (mt) DNA control region, amplified by the polymerase chain-reaction. The inferred phylogeny of haplotypes led to the recognition of three distinct mitochondrial lineages or phylads consistent with the distinctions of current taxonomy. All 15 mtDNA haplotypes found in D. macarellus and all 9 haplotypes found in D. macrosoma were arranged as star-like clusters, suggesting recent evolutionary history. In contrast, the phylad formed by 6 haplotypes in D. russelli from the Sulawesi Sea exhibited diffuse topology, suggesting that ancestral lineages of this species have been retained to the present. Average nucleotide-divergence estimates between haplotypes of different phylads were between 0.042 and 0.135, suggesting ancient separation, in consistency with published allozyme data. High levels of haplotype diversity, but no geographical heterogeneity, was detected within D. macarellus from the Molucca Sea and the Banda Sea. Populations of D. macrosoma exhibited both significant differences between adjacent regions (Sunda Strait and Java Sea), and broadscale genetic homogeneity from the South China Sea to the Sulawesi Sea via the Java Sea and Makassar Strait. The geographic isolation of the D. macrosoma population sampled in the Sunda Strait suggests that this region constitute a sharp transition zone between the Indian Ocean and the Sunda Shelf. Near-monomorphism of haplotypes and low nucleotide diversity (d _X) were observed in the samples of D. macrosoma from the continental shelf (haplotype-diversity estimates, h, = 0.00 to 0.25 ± 0.08 and d _X = 0.000 to 0.002). This was in contrast to the comparatively high haplotype and nucleotide diversities observed in other pelagic fish species including D. macarellus (h = 0.82 ± 0.05, d _X = 0.012 to 0.015) and D. russelli (h = 0.63 ± 0.12, d _X = 0.016), and in the oceanic D. macrosoma population sampled in the Sunda Strait (h = 0.67 ± 0.31, d _X = 0.005). We hypothesise that this may be the consequence of recent and perhaps repeated bottleneck events that have affected the D. macrosoma population sampled on the continental shelf. Received: 29 September 1997 / Accepted: 3 September 1999     

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.     

Sympatry of distinct mitochondrial DNA lineages in a copepod inhabiting estuarine creeks in the southeastern USA

The population genetic structure of the meiobenthic harpacticoid copepod Microarthridion littorale (Poppe) was examined with a geographic survey of a 348 bp fragment of the mitochondrial cytochrome b gene. Copepods were collected from ten locations on the coast from North Carolina to Georgia, USA, from January 1997 to November 1998. Sequence divergence among 198 individuals was as much as 4.3%, and three divergent mitochondrial clades were uncovered that differed by six to nine nucleotide changes. A rapid assay was developed to distinguish among mitochondrial clades, and an additional 333 specimens were surveyed. The three lineages co-occurred in seven of ten sampling locations. Data analyses were carried out separately for individuals assayed by DNA sequencing as well as for a combined data set that included individuals typed by restriction endonuclease digestion. An analysis of molecular variance indicated that a significant proportion of the total genetic variance could be partitioned among populations, although no significant correlation between geographical and genetic distance was detected.     

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.     

Phylogeography of the pipefish, <Emphasis Type="Italic">Urocampus carinirostris</Emphasis>, suggests secondary intergradation of ancient lineages

We assayed the pattern of mitochondrial DNA evolution in the live bearing, seagrass specialist pipefish, Urocampus carinirostris, in eastern Australia. These life history attributes were predicted to result in strong phylogeographic structure in U. carinirostris. Phylogenetic analysis of cytochrome b sequences detected two monophyletic mtDNA clades that differed by 8.69% sequence divergence - a large level of intraspecific divergence for a marine fish. The geographical distribution of clades was non-random and resembled clinal secondary intergradation over a 130-km stretch of coastline. Contrary to phylogeographic predictions, this large phylogeographic break does not occur across a traditionally recognised biogeographic boundary. Analyses of historical demography suggested that individuals belonging to the most widespread clade underwent a population expansion from a small refuge population during the Pleistocene.     

Mitochondrial DNA variation in Eritrean hamadryas baboons (Papio hamadryas hamadryas): life history influences population genetic structure

The hamadryas baboon, Papio hamadryas hamadryas, represents a rare exception from the pattern of female philopatry and male-biased dispersal predominant in mammals including primates. To elucidate the possible consequences of the dispersal pattern on the population genetic structure of hamadryas baboons, we sequenced the maternally transmitted mitochondrial hypervariable region I of 74 individuals from ten sampling locations in different ecogeographic zones of Eritrea. To this end, individual fecal samples were collected at sleeping cliffs. Upon comparing the individual sequences by means of phylogenetic tree reconstructions and AMOVA, we could not detect a population genetic structure corresponding to a geographic pattern. Tree reconstructions revealed the existence of two profoundly different lineages both present at most of the sampling locations. These findings and Mantel correlations of genetic distances and the frequency of shared haplotypes to geographic distances point to the presence of female dispersal. Female-mediated gene flow is detectable over geographic distances exceeding those between neighboring subpopulations. Our study therefore corroborates local behavioral observations on a broad geographic scale. After inclusion of geographically closely situated olive baboons, P. h. anubis, in the analyses, all anubis sequences fell within one hamadryas clade. Possible scenarios leading to this situation including long-term hybridization processes are discussed.     

Geographical genetic structure and phylogeography of the <Emphasis Type="Italic">Sargassum horneri</Emphasis>/<Emphasis Type="Italic">filicinum</Emphasis> complex in Japan,based on the mitochondrial <Emphasis Type="Italic">cox3</Emphasis> haplotype

The genetic structure and phylogeography of the brown seaweed Sargassum horneri/filicinum complex in Japan were studied based on the mitochondrial cox3 haplotype. The cox3 haplotypes found were divided into three clades in a statistical parsimony network, among which there were large numbers of steps. Contrary to the reported large amount of drifting S. horneri along the Japanese coast, the three clades were dividedly distributed on the Japanese coast: the northern Pacific, the central Pacific, and western Japan. The western Japan S. horneri had haplotypes that were phylogenetically closer to those of S. filicinum than to the northern and central Pacific S. horneri populations. The S. filicinum populations were included within the western Japan clade and grouped together with the S. horneri samples from western Japan. Taken together with the unstable morphological diagnosis, this result suggests that S. filicinum should be reduced into a synonymy of S. horneri. The TMRCA analysis suggested that the divergence time of each clade may go back to the last interglacial period and a skyline plot suggested that the last glacial maximum had only a small effect on the population size of S. horneri. The geographic subdivision of the three groups, in spite of a large amount of drifting mats, suggests a limited contribution of drifting mats to gene flow on a large geographic scale. On a small geographic scale, a small number of haplotypes were shared between S. horneri-type and S. filicinum-type populations. This result suggests that populations of these two types are partially, though not completely, isolated from each other, possibly by selfing in S. filicinum-type populations or by a difference in peak reproduction.     

Maturation and diel reproductive periodicity of round scad (Carangidae: Decapterus punctatus)

The diel reproductive periodicity of Carangidae is poorly known but appears to be highly variable between species. Some species spawn during the day, others are believed to spawn at night, and it is demonstrated here that round scad, Decapterus punctatus, spawn at dusk. We collected D. punctatus in the eastern Gulf of Mexico during three April cruises (1995, 1996, and 1997). Based on histological criteria, size at 50% maturity was 113 mm fork length (FL) for males and 128 mm FL for females. The gonad-somatic index (GSI) of mature males was significantly different between hours and appeared to show diel periodicity. Diel periodicity was also observed in changes in female GSIs, whole oocyte diameters, and ovarian histology. The average GSI of mature females fluctuated two-fold between day and night, and the size distribution of whole oocytes in some fish was bimodal (at 0.3-0.4 and 0.7-0.8 mm diameter) at dusk rather than unimodal during most of the diel cycle. Histological preparations revealed that these rapid changes in ovarian GSIs and oocyte size distributions were the result of final oocyte maturation. Germinal vesicle migration was observed from 0900 to 1400 hours eastern standard time (EST), germinal vesicle breakdown was evident as early as 1100 hours EST, and ovulation occurred as early as 1800 hours EST. Spawning frequency (approximately every 5 days) was similar whether calculated from the proportion of females with hydrated oocytes during the afternoon or from the proportion of females with postovulatory follicles during the morning. Batch fecundity correlated with fish size and ranged from 5,500 to 34,700 hydrated eggs per individual. These findings do not support published hypotheses that young-of-the-year D. punctatus reproduce before their first winter or that D. punctatus reproductive output is bimodal within a year.     

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.     

Influence of environmental factors on burrow irrigation and oxygen consumption in the mudflat invertebrate Urechis caupo

We examined burrow irrigation activity by the mudflat worm Urechis caupo in response to suspended food, ambient hypoxia (down to 3.3 kPa PO₂), hydrogen sulfide exposure (up to 100 µmol l^-1), and short-term temperature change (range 10-22°C). In normoxic, nutrient-free water at 14°C, O₂ consumption ( [(M)\dot]O₂ ) \left( {\dot M{\rm O}_2 } \right) was 45 nmol min^-1 g^-1, water flow rate ( [(V)\dot]_W ) \left( {\dot V_{\rm W} } \right) was 27 ml min^-1 (0.66 ml min^-1 g^-1), frequency of peristaltic waves (F_P) was 2.6 contractions min^-1, stroke volume (SV) was 11 ml, and O₂ extraction coefficient (EO₂) was 0.27. Adding suspended food to the burrow water occasionally elicited stereotypical feeding behavior but had no effect on any measured variables during nonfeeding periods. Hypoxia greatly decreased [(M)\dot]O₂ \dot M{\rm O}_2 (75% reduction at 3.3 kPa PO₂) but did not affect [(V)\dot]_W \dot V_{\rm W} , F_P, SV, or EO₂. Sulfide at 50 µmol l^-1 or less had no effect on burrow irrigation activity, whereas 100 µmol l^-1 sulfide decreased [(V)\dot]_W \dot V_{\rm W} by 58% and F_P by 50% but had no effect on SV. Temperature strongly affected [(V)\dot]_W \dot V_{\rm W} (Q₁₀ of 1.9 from 10°C to 22°C). We propose that U. caupo's ability to live in the hypoxic, sulfidic mud of productive mudflat environments, combined with its very efficient mucous net, allows it to process much less water for feeding than other suspension-feeding invertebrates. This, in turn, necessitates an efficient O₂ extraction mechanism, which is provided by the water lung activity of U. caupo's unique hindgut.     

Phylogenetic relationship among several genera of Dictyotaceae (Dictyotales, Phaeophyceae) based on 18S rRNA and partial rbcL gene sequences

The partial sequences of the 18S rRNA gene and the rbcL gene, RuBisCo spacer region of representatives of the Dictyotaceae were determined and compared to resolve their phylogenetic relationships. A total of 1,861 base pairs of 18S rDNA sequences were aligned and examined. There was a high similarity of nucleotide sequences within the tribe Dictyoteae (99.4-99.6%) and the tribe Zonarieae (92.8-99.8%). The largest intergeneric divergence within the Dictyotaceae was 7.31%, between Dilophus and Zonaria. The aligned 422 bases of partial rbcL gene sequences gave a similar phylogenetic relationship among taxa to those of the 18S rDNA sequences, although our alignment covered less than 30% of the whole gene. All genera of Dictyotales consisted of several clades forming a polychotomy in MP trees based on partial rbcL gene sequences and on 18S rDNA sequences. The genera within the tribe Dictyoteae formed a clade, a Dictyota-Phacydictyon-Dilophus clade, while genera of the tribe Zonarieae formed four clades: Padina; Distromium-Lobophora-Zonaria; Spatoglossum-Lobospira; and Dictyopteris-Spatoglossum. All genera of the Dictyotaceae included in these sequence analyses are classified into five groups, which have little phylogenetic resolution among them. The genera classified in the tribe Zonarieae according to Womersley's criteria do not appear to be monophyletic, based on our analyses. Although not all genera of Dictyotaceae were analysed, our data suggest that the status of the meristem cells should not be a criterion for separating tribes within the family Dictyotaceae and that other hierarchical systems should be adopted to properly reflect phylogenetic relationships.     

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.     

Behavioral adaptations of the cubozoan medusa <Emphasis Type="Italic">Tripedalia cystophora</Emphasis> for feeding on copepod (<Emphasis Type="Italic">Dioithona oculata</Emphasis>) swarms

The cubozoan medusa Tripedalia cystophora preys on dense swarms of the copepod Dioithona oculata in the mangrove prop-root habitat of Puerto Rico. The copepod swarms form in shafts of sunlight that penetrate the mangrove canopy during the day. T. cystophora are found primarily within the same illuminated areas, feeding heavily on the dense swarms of copepods. Laboratory studies were conducted to examine the behavioral adaptations of T. cystophora that allow them to remain within these dense copepod swarms. In the presence of vertical light shafts, T. cystophora showed both a significant change in swimming speed and a significant increase in turning rate. T. cystophora turn abruptly when they leave a vertical shaft of light and quickly re-enter. The presence or absence of copepod prey, either in swarms or dispersed, had no significant effect on T. cystophora swimming speed or turning behavior. Feeding rates of T. cystophora on the copepod D. oculata increase rapidly with copepod density, and maximum rates are not achieved until the high densities associated with swarms are reached. It seems doubtful that these small medusae could survive without their ability to encounter and exploit these dense swarms of copepods.     

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.     

Nuclear and mitochondrial DNA markers indicate unidirectional gene flow of Indo-Pacific to Atlantic bigeye tuna (<Emphasis Type="Italic">Thunnus obesus</Emphasis>) populations,and their admixture off southern Africa

A sharp genetic break separates Atlantic from Indo-Pacific bigeye tuna (Thunnus obesus) populations, as the frequencies of two major mitochondrial (mt) DNA types ( and ) found in this species are different across the tip of southern Africa. The level of nucleotide divergence between mtDNA types and is of the same order as that between reproductively isolated taxa. To further investigate the genetic structure of bigeye tuna over its distribution range and in the contact zone off southern Africa, bigeye tuna samples collected between 1992 and 2001 (including samples from a previous mtDNA survey) were characterized for four nuclear DNA loci and for mtDNA. Nuclear markers did not support the hypothesis that and mitochondria characterize sibling species. Significant allele-frequency differences at one intronic locus (GH2) and one microsatellite locus (µ208) were found between Atlantic and Indo-Pacific samples, although the level of nuclear genetic differentiation (Weir and Cockerhams =0.025 to 0.042) was much lower than in mtDNA ( =0.664 to 0.807). Probabilistic Bayesian assignment of individuals to a population confirmed that southern African bigeye tuna samples represent a simple mixture of individuals from Atlantic and Indian stocks that do not interbreed, with a higher contribution from Indian Ocean individuals (about 2/3 vs. 1/3).Communicated by M.S. Johnson, Crawley     

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.     

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

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

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.     

Dynamics of pigment degradation by the copepodite stage of <Emphasis Type="Italic">Pseudodiaptomus</Emphasis><Emphasis Type="Italic">euryhalinus</Emphasis> feeding on <Emphasis Type="Italic">Tetraselmis</Emphasis><Emphasis Type="Italic">suecica</Emphasis>

Short-term (3 h) changes in concentration of chlorophylls and their derivatives in stage V Pseudodiaptomus euryhalinus and their fecal material were followed by HPLC during a 24 h experiment. Copepodites were fed with the prasinophyte Tetraselmis suecica. Intact chlorophyll a and b were found in animals and fecal material and had similar dynamics of accumulation over time. The extent of transformation of chlorophyll a and b to colorless compounds was different with chlorophyll a being more extensively degraded. Additionally, several chlorophyll derivatives (pheophytin and pyropheophytin-like pigments) were found. Pyropheophytin a was the most abundant followed by pheophytin b, pheophytin a, and pheophorbide a. Relative amounts of pheopigments were different in copepodites and fecal material, and pheophytin a, pheophorbide a, and pheophytin b concentrations were low and variable. The amount of ingested chlorophyll recovered as chlorophyll a and its derivatives in fecal and copepodite pools was generally low (<5%), with one exception occurring after 9 h, when it accounted for >70%. These data suggest individual pheopigments are produced at different rates and that chemical or enzymatic mechanisms in the gut of copepodites act on the two chlorophylls in different ways.