Global mitochondrial DNA phylogeography and biogeographic history of the antitropically and longitudinally disjunct marine bryozoan Membranipora membranacea L. (Cheilostomata): another cryptic marine sibling species complex?

The origin of disjunct distributions in high dispersal marine taxa remains an important evolutionary question as it relates to the formation of new species in an environment where barriers to gene flow are not always obvious. To reconstruct the relationships and phylogeographic history of the antitropically and longitudinally disjunct bryozoan Membranipora membranacea populations were surveyed with mtDNA cytochrome oxidase 1 (COI) sequences across its cosmopolitan range. Maximum parsimony, maximum likelihood and Bayesian genealogies revealed three deep clades in the North Pacific and one monophyletic clade each in the southeast Pacific (Chile), southwest Pacific (Australia/New Zealand), North Atlantic and southeast Atlantic (South Africa). Human-mediated dispersal has not impacted M. membranacea’s large-scale genetic structure. M. membranacea did not participate in the trans-arctic interchange. Episodic long-distance dispersal, combined with climatic vicariance can explain the disjunct distribution. Dispersal led southward across the tropics perhaps 13 mya in the East Pacific and again northwards perhaps 6 mya in the Eastern Atlantic to colonize the North Atlantic from the south, and along the West Wind Drift to colonize Australia. The clades differentiated over evolutionary time in their respective ocean region, potentially forming a sibling species complex. The taxonomic status of the clades is discussed.     

Diversity and distribution of foliose Bangiales (Rhodophyta) in West Greenland: a link between the North Atlantic and North Pacific

Greenland is a continental island in the northern part of the North Atlantic where the foliose Bangiales flora is poorly known. It is an important area for the study of algal biogeography because of the region’s glacial history, in which Greenland has been alternately exposed to or isolated from the North Pacific via the Bering Strait. A molecular study using 3′ rbcL + 5′ rbcL–S sequences was undertaken to assess the diversity of foliose Bangiales on the west coast of Greenland and rbcL sequences were used to study the Greenland flora in a larger phylogenetic and floristic context. New and historic collections document seven species in four genera from the west coast of Greenland. All species had a close link to North Pacific species, being either conspecific with them or North Atlantic–North Pacific vicariant counterparts.     

Molecular lineage diversity and inter‐oceanic biogeographical history in Hiatella (Mollusca,Bivalvia)

Hiatella is one of the most widespread marine bivalve genera, occurring in diverse habitats from the temperate to polar latitudes in both hemispheres, and in fossil strata since almost 150 Myr ago. Despite variation in some biological and morphological traits, characters to resolve the current systematic structure consistently across the range of the genus are not known: all samples are often referred to a single species, Hiatella arctica (L.). Exploring the systematics of Hiatella using partial sequences of three genes (mitochondrial COI, and the nuclear ANT and 28S rRNA), we find high diversity of deep lineages (11–22% p‐distance in COI), and identify at least 13 distinct taxa both by heuristic criteria (congruence of the nuclear and mtDNA data) and by coalescence‐based analyses. At several localities, two or three of these cryptic species were found in sympatry. In the framework of previous fossil evidence and of hypotheses of paleoceanographical connections, scenarios of the phylogeny and biogeographical history of the identified species at a range of different time scales are outlined. The distinction between the main North Pacific and North Atlantic Hiatella clades and systematic diversification within each of them seem to have followed a Miocene trans‐Panamanian invasion. Apart from such earlier intra‐basin diversification, the data suggest that three successive counter‐invasions from the Pacific to the Atlantic via the Arctic Ocean route have later contributed to the current North Atlantic Hiatella diversity. These invasions probably took place in connection with (i) the Great Trans‐Arctic Biotic Interchange in the Pliocene, (ii) the last interglacial period c. 120 kya and (iii) the Holocene, postdating the last glaciation. This sequence of trans‐Arctic invasions is largely analogous to that hypothesized for some other boreal‐arctic bivalves.     

The Taxonomic and Evolutionary History of Fossil and Modern Balaenopteroid Mysticetes

Balaenopteroids (Balaenopteridae + Eschrichtiidae) are a diverse lineage of living mysticetes, with seven to ten species divided between three genera (Megaptera, Balaenoptera and Eschrichtius). Extant members of the Balaenopteridae (Balaenoptera and Megaptera) are characterized by their engulfment feeding behavior, which is associated with a number of unique cranial, mandibular, and soft anatomical characters. The Eschrichtiidae employ suction feeding, which is associated with arched rostra and short, coarse baleen. The recognition of these and other characters in fossil balaenopteroids, when viewed in a phylogenetic framework, provides a means for assessing the evolutionary history of this clade, including its origin and diversification. The earliest fossil balaenopterids include incomplete crania from the early late Miocene (7–10 Ma) of the North Pacific Ocean Basin. Our preliminary phylogenetic results indicate that the basal taxon, “Megaptera” miocaena should be reassigned to a new genus based on its possession of primitive and derived characters. The late late Miocene (5–7 Ma) balaenopterid record, except for Parabalaenoptera baulinensis and Balaenoptera siberi, is largely undescribed and consists of fossil specimens from the North and South Pacific and North Atlantic Ocean basins. The Pliocene record (2–5 Ma) is very diverse and consists of numerous named, but problematic, taxa from Italy and Belgium, as well as unnamed taxa from the North and South Pacific and eastern North Atlantic Ocean basins. For the most part Pliocene balaenopteroids represent extinct species and genera and reveal a greater degree of morphological diversity than at present. The Pleistocene record is very limited and, unfortunately, fails to document the evolutionary details leading to modern balaenopteroid species diversity. It is evident, however, that most extant species evolved during the Pleistocene. Morphological and molecular based phylogenies support two competing hypotheses concerning relationships within the Balaenopteroidea: (1) balaenopterids and eschrichtiids as sister taxa, and (2) eschrichtiids nested within a paraphyletic Balaenopteridae. The addition of fossil taxa (including a new Pliocene species preserving a mosaic of balaenopterid and eschrichtiid characters) in morphological and “total evidence” analyses, offers the potential to resolve the current controversy concerning the possible paraphyly of Balaenopteridae.     

The Bering Strait connection: dispersal and speciation in boreal macroalgae

A large number of boreal seaweeds have either sibling species or conspecific populations of a single species in the North Pacific and North Atlantic Oceans. This pattern is thought to have arisen from the dispersal between the two oceans through the Arctic Ocean after the opening of the Bering Strait in the mid-to-late Miocene or earliest Pliocene and from subsequent vicariant speciation as the Arctic Ocean froze and Bering Strait closed intermittently during glacial periods. Recent molecular studies of species in all three major seaweed phyla reveal patterns of vicariance. However, a number of lines of evidence point to differences in origins of these clades; some appear to be Pacific in origin whereas others appear to be derived from Atlantic stock. Different origins can be explained by recent stratigraphic finds that push the first Cenozoic opening of the Bering Strait back from 3.1–4.1 to 4.8–7.4 Ma (million years ago). Northern hemisphere ocean circulation models suggest that water flow would have been from the North Atlantic–Arctic south through the Bering Strait prior to the closure of the Panamanian Isthmus c. 3.5 Ma in contrast to the northward flow from the Pacific into the Arctic and North Atlantic, which developed after the closing of the Isthmus. Despite these differences in timing of the two invasions, there are no significant differences in levels of relationships among species with a North Atlantic origin compared with species with a North Pacific origin based on currently available data. More work is required to understand vicariance in seaweeds, especially in deciphering when a speciation event has occurred.     

Late Miocene Survival of a Hyper-Longirostrine Dolphin and the Neogene to Recent Evolution of Rostrum Proportions Among Odontocetes

Hyper-longirostry, the character of having extremely elongated rostra, emerged in the early and middle Miocene among several different clades of echolocating toothed whales (odontocetes) followed by a rapid decline near the end of the middle Miocene, and postdated by a much lower number of occurrences in the late Miocene and Pliocene and a complete absence among extant odontocetes. New finds of unreworked fossils of Xiphiacetus cristatus (Eurhinodelphinidae) in the middle Tortonian Diest Formation in Belgium (9.54–8.8 Ma) allow for the documentation of the survival of a hyper-longirostrine dolphin into the early late Miocene. An extensive dataset of the rostral index (calculated as the ratio between rostral length and condylobasal length) of Neogene and extant odontocetes is compiled and presented here, which facilitates discussion of evolutionary trends of rostrum proportions during a time period spanning 23 million years. Of interest, the iterative survival into the late Miocene of a single different species of hyper-longirostrine dolphins in a number of paleogeographic regions (North Sea Basin, Atlantic Coastal Plain, and probably the southeastern Pacific) is noted, whereas hyper-longirostrine morphologies only seem to re-appear by the late Messinian in the Northeastern Pacific. A correlation between this pattern and a decrease in habitat size for coastal to estuarine dolphins linked to a major sea level drop is tentatively proposed; such a process may also have played a role in the ecological shift in several dolphin families to freshwater habitats.

     

Phylogenetic relationships of assimineid gastropods of the Death Valley–lower Colorado River region: relicts of a late Neogene marine incursion?

Aim A small fauna of amphibious snails (genus Assiminea Fleming, 1828) living in association with highly mineralized springs in the Death Valley–lower Colorado River region (DVLCR) is thought to be a relict of the Bouse Embayment, a putative late Miocene–early Pliocene transgression of the ancestral Gulf of California along the lower Colorado River valley. We analysed the phylogenetic relationships of this fauna using mtDNA sequence data (1171 bp) to determine whether, as would be consistent with this hypothesis, it forms a substantially divergent unit sister to marine coastal congeners. Location South‐western Great Basin and lower Colorado River region, USA. Methods Two genes [mitochondrial cytochrome c oxidase subunit I (COI) and the mitochondrial 16S ribosomal RNA gene] were sequenced for 10 populations of DVLCR assimineas (Assiminea infima Berry, 1947 ; Assiminea sp.). We also sequenced an undescribed population from a spring in the Colorado River delta; western North American Pacific Coastal Assiminea californica (Tryon, 1865); the three other congeners that live on the continent; and three Old World assimineids (outgroups). Phylogenies based on the combined data set were obtained using Bayesian methods, and divergence times were estimated using a COI molecular clock for related gastropods. Results Composite haplotypes of the DVLCR assimineas, together with that observed in the Colorado River delta population, formed a weakly supported clade that was sister to a clade composed of populations of North American Pacific and Atlantic coastal species. The genetic distance between members of these two clades was 3.46 ± 0.47% for COI and 1.69 ± 0.38% for 16S. The former clade was composed of five subunits that differed from each other by 1.29–2.84% (COI) and 0.52–1.98% (16S) sequence divergence. Main conclusions Application of the COI clock suggests that progenitors of the DVLCR fauna diverged from coastal ancestors 2.13–1.89 Ma (late Pliocene), several million years after the Bouse Embayment would have been terminated by the establishment of the lower (freshwater) Colorado River. This finding, together with shallow genetic structuring of several DVLCR lineages that are widely distributed across the topographically complex regional landscape, suggests that the Assiminea fauna of this inland area was more likely to have been founded by coastal colonists transported on water birds than through a direct connection with the sea.     

Trans‐Pacific and trans‐Arctic pathways of the intertidal macroalga Fucus distichus L. reveal multiple glacial refugia and colonizations from the North Pacific to the North Atlantic

Aim We examined the phylogeography of the cold‐temperate macroalgal species Fucus distichus L., a key foundation species in rocky intertidal shores and the only Fucus species to occur naturally in both the North Pacific and the North Atlantic. Location North Pacific and North Atlantic oceans (42° to 77° N). Methods We genotyped individuals from 23 populations for a mitochondrial DNA (mtDNA) intergenic spacer (IGS) (n = 608) and the cytochrome c oxidase subunit I (COI) region (n = 276), as well as for six nuclear microsatellite loci (n = 592). Phylogeographic structure and connectivity were assessed using population genetic and phylogenetic network analyses. Results IGS mtDNA haplotype diversity was highest in the North Pacific, and divergence between Pacific haplotypes was much older than that of the single cluster of Atlantic haplotypes. Two ancestral Pacific IGS/COI clusters led to a widespread Atlantic cluster. High mtDNA and microsatellite diversities were observed in Prince William Sound, Alaska, 11 years after severe disturbance by the 1989 Exxon Valdez oil spill. Main conclusions At least two colonizations occurred from the older North Pacific populations to the North Atlantic between the opening of the Bering Strait and the onset of the Last Glacial Maximum. One colonization event was from the Japanese Archipelago/eastern Aleutians, and a second was from the Alaskan mainland around the Gulf of Alaska. Japanese populations probably arose from a single recolonization event from the eastern Aleutian Islands before the North Pacific–North Atlantic colonization. In the North Atlantic, the Last Glacial Maximum forced the species into at least two known glacial refugia: the Nova Scotia/Newfoundland (Canada) region and Andøya (northern Norway). The presence of two private haplotypes in the central Atlantic suggests the possibility of colonization from other refugia that are now too warm to support F. distichus. With the continuing decline in Arctic ice cover as a result of global climate change, renewed contact between North Pacific and North Atlantic populations of Fucus species is expected.     

Osteology and ontogeny of the wrymouths,genus Cryptacanthodes (Cottiformes: Zoarcoidei: Cryptacanthodidae)

The four species included in the family Cryptacanthodidae are eel‐like, burrowing fishes distributed in the cold‐temperate coastal waters of the North Pacific and the western North Atlantic. This study describes the osteology and aspects of the ontogenetic skeletal development of two species, Cryptacanthodes maculatus from the western North Atlantic and C. aleutensis from the eastern North Pacific. We discuss the relationships of Cryptacanthodidae among other zoarcoid families. The Cryptacanthodidae have been previously included in the Stichaeidae, but removed and classified as a separate family based on the skull, pectoral radial, and cephalic lateral‐line morphology. Our observations (similarities in gill arch and pectoral girdle morphology; specifically, a thin sheet‐like flange of bone from the posterior margin of the supracleithrum) suggest a close relationship to at least some of the members of the family Stichaeidae. J. Morphol. 276:185–208, 2015. © 2014 Wiley Periodicals, Inc.     

Out of the Pacific and back again: insights into the matrilineal history of Pacific killer whale ecotypes

Killer whales (Orcinus orca) are the most widely distributed marine mammals and have radiated to occupy a range of ecological niches. Disparate sympatric types are found in the North Atlantic, Antarctic and North Pacific oceans, however, little is known about the underlying mechanisms driving divergence. Previous phylogeographic analysis using complete mitogenomes yielded a bifurcating tree of clades corresponding to described ecotypes. However, there was low support at two nodes at which two Pacific and two Atlantic clades diverged. Here we apply further phylogenetic and coalescent analyses to partitioned mitochondrial genome sequences to better resolve the pattern of past radiations in this species. Our phylogenetic reconstructions indicate that in the North Pacific, sympatry between the maternal lineages that make up each ecotype arises from secondary contact. Both the phylogenetic reconstructions and a clinal decrease in diversity suggest a North Pacific to North Atlantic founding event, and the later return of killer whales to the North Pacific. Therefore, ecological divergence could have occurred during the allopatric phase through drift or selection and/or may have either commenced or have been consolidated upon secondary contact due to resource competition. The estimated timing of bidirectional migration between the North Pacific and North Atlantic coincided with the previous inter-glacial when the leakage of fauna from the Indo-Pacific into the Atlantic via the Agulhas current was particularly vigorous.     

Tandemly Repeated Sequences in the Mitochondrial DNA Control Region and Phylogeography of the Pike-PerchesStizostedion

DNA sequences from the mitochondrial DNA control region are used to test the phylogeographic relationships among the pike-perches,Stizostedion(Teleostei: Percidae) and to examine patterns of variation. Sequences reveal two types of variability: single nucleotide polymorphisms and 6 to 14 copies of 10- to 11-base-pair tandemly repeated sequences. Numbers of copies of the tandem repeats are found to evolve too rapidly to detect phylogenetic signal at any taxonomic level, even among populations. Sequence similarities of the tandem repeats amongStizostedionand other percids suggest concerted evolutionary processes. Predicted folding of the tandem repeats and their proximity to termination-associated sequences indicate that secondary structure mediates slipped-strand mispairing among the d-loop, heavy, and light strands. Neighbor-joining and maximum parsimony analyses of sequences indicate that the genus is divided into clades on the continents of North America and Eurasia. Calibrating genetic distances with divergence times supports the hypothesis thatStizostediondispersed from Eurasia to North America across a North Pacific Beringial land bridge approximately 4 million years before present, near the beginning of the Pliocene Epoch. The North AmericanS. vitreumandS. canadenseappear separated by about 2.75 million years, and the EurasianS. luciopercaandS. volgensisare diverged by about 1.8 million years, suggesting that speciation occurred during the late Pliocene Epoch.     

A tale of North and South America: time and mode of dispersal of the amphitropical genus Munroa (Poaceae,Chloridoideae)

Plant disjunctions have provided some of the most intriguing distribution patterns historically addressed by biogeographers. We evaluated the three hypotheses that have been postulated to explain these patterns [vicariance, stepping‐stone dispersal and long‐distance dispersal (LDD)] using Munroa, an American genus of grasses with six species and a disjunct distribution between the desert regions of North and South America. The ages of clades, cytology, ancestral characters and areas of distribution were investigated in order to establish relationships among species, to determine the time of divergence of the genus and its main lineages, and to understand further the biogeographical and evolutionary history of this genus. Bayesian inference recovered the North American M. pulchella as sister species to the rest. Molecular dating and ancestral area analyses suggest that Munroa originated in North America in the late Miocene–Pliocene (7.2 Mya; 8.2–6.5 Mya). Based on these results, we postulate that two dispersal events modelled the current distribution patterns of Munroa: the first from North to South America (7.2 Mya; 8.2–6.5 Mya) and the second (1.8 Mya; 2–0.8 Mya) from South to North America. Arid conditions of the late Miocene–Pliocene in the Neogene and Quaternary climatic oscillations in North America and South America were probably advantageous for the establishment of populations of Munroa. We did not find any relationship between ploidy and dispersal events, and our ancestral character analyses suggest that shifts associated with dispersal and seedling establishment, such as habit, reproductive system, disarticulation of rachilla, and shape and texture of the glume, have been important in these species reaching new areas. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 110–125.     

GENE SEQUENCE DIVERSITY AND THE PHYLOGENETIC POSITION OF ALGAE ASSIGNED TO THE GENERA PHAEOPHILA AND OCHLOCHAETE (ULVOPHYCEAE,CHLOROPHYTA)1

The phylogenetic position of microfilamentous marine green algae assigned to the species Phaeophila dendroides, Entocladia tenuis (Phaeophila tenuis, and Ochlochaete hystrix was examined through phylogenetic analyses of nuclear‐encoded small subunit rDNA and chloroplast‐encoded tufA gene sequences. These analyses placed the P. dendroides strains within the Ulvophyceae, at the base of a clade that contains representatives of the families Ulvaceae, Ulvellaceae, and the species Bolbocoleon piliferum, supporting an earlier hypothesis that P. dendroides constitutes a distinct lineage. Substantial divergence in both nuclear and plastid DNA sequences exists among strains of P. dendroides from different geographic localities, but these isolated strains are morphologically indistinguishable. The lineage may have an accelerated rate of gene sequence evolution relative to other microfilamentous marine green algae. Entocladia tenuis and O. hystrix are placed neither in the P. dendroides clade nor in the Ulvellaceae as previous taxonomic schemes predicted but instead form a new clade or clades at the base of the Ulvaceae. Ruthnielsenia gen. nov. is proposed to accommodate Kylin's species, which cannot be placed in Entocladia (=Acrochaete), Phaeophila, or Ochlochaete. Ruthnielsenia tenuis (Kylin) comb. nov., previously known only from Atlantic coasts, is reported for the first time from the Pacific coast of North America (San Juan Island, WA, USA). Isolates of R. tenuis from the Atlantic and Pacific coasts of North America have identical small subunit rDNA and tufA gene sequences.     

EVOLUTIONARY HISTORY OF NORTHERN HEMISPHERE NUCELLA (GASTROPODA,MURICIDAE): MOLECULAR,MORPHOLOGICAL, ECOLOGICAL,AND PALEONTOLOGICAL EVIDENCE

By combining data from a variety of sources we explore patterns of evolution and speciation in Nucella, a widely studied genus of shallow-water marine neogastropods. We present a hypothesis of phylogenetic relationships for all of the currently recognized species of northern hemisphere Nucella, based on an analysis of 718 base pairs of nucleotide sequence from the mitochondrial cytochrome b gene. The order of appearance of species in the fossil record is congruent with this hypothesis. The topology of the inferred phylogeny of Nucella, coupled with ecological, morphological, and fossil evidence, was used to address three main questions: (1) At what time and by which route was the North Atlantic invaded from the North Pacific compared to prior studies of the trans-Arctic interchange? (2) Do patterns of molecular variation within species corroborate the importance of climatic cycles in driving speciation in north temperate marine animals? (3) Was radiation in the direction of increased or decreased ecological specialization, body size, or vulnerability to predation? Molecular evidence confirmed that the sole North Atlantic species, N. lapillus, arose from a North Pacific ancestor. Biogeographic and paleontological evidence supported the dispersal of Nucella, and perhaps other interchange species, via the Eurasian Arctic. Rather intriguingly, the linkage of N. lapillus to a western as opposed to eastern Pacific clade, and the biogeographic origins of the eastern Pacific species, parallel closely similar patterns observed in another genus of rocky-shore gastropods, Littorina. This congruence, in conjunction with information on the climatic and geographic histories of the region, as well as the geographic arrangement of mtDNA haplotypes within Nucella species, supports a model of speciation in Nucella driven by cycles of climatic amelioration and deterioration that began during the Miocene. Calibrations from the fossil record of Nucella suggest that third position transitions and transversions accrue at a rate of 3–4% and 0.5% respectively per million yr. This supports an early participation by Nucella in the trans-Arctic interchange, as suggested by paleobiogeographic studies. Consistent with the unstable taxonomic history of species of Nucella, we found few nonmolecular traits to be phylogenetically informative. Among North Pacific species, more recently derived species (N. canaliculata and the N. emarginata clade) were more ecologically specialized (narrower diet and habitat range). Consistent with extensive intraspecific variation, shell traits were quite labile evolutionarily: neither overall size nor development of antipredatory traits exhibited consistent evolutionary trends over the history of the genus. Nurse eggs (unfertilized eggs consumed by developing embryos) were an ancestral trait that was lost evolutionarily in the two clades that also exhibited increased body size, suggesting that these two life-history traits may be coupled. The reduced number of chromosomes in N. lapillus is clearly a derived state and is consistent with White's (1978) observations on chromosome evolution in other clades.     

A mitogenomic approach to the taxonomy of pollocks: <Emphasis Type="Italic">Theragra chalcogramma</Emphasis> and <Emphasis Type="Italic">T. finnmarchica</Emphasis> represent one single species

Background  

The walleye pollock (Theragra chalcogramma) and Norwegian pollock (T. finnmarchica) are confined to the North Pacific and North Atlantic Oceans, respectively, and considered as distinct species within the family Gadidae. We have determined the complete mtDNA nucleotide sequence of two specimens of Norwegian pollock and compared the sequences to that of 10 specimens of walleye pollock representing stocks from the Sea of Japan and the Bering Sea, 2 specimens of Atlantic cod (Gadus morhua), and 2 specimens of haddock (Melanogrammus aeglefinus).     

Phylogenetic relationships of tropical western Atlantic snappers in subfamily Lutjaninae (Lutjanidae: Perciformes) inferred from mitochondrial DNA sequences

Phylogenetic relationships among 20 nominal species of tropical lutjanine snappers (Lutjanidae) (12 from the western Atlantic, one from the eastern Pacific, and seven from the Indo‐Pacific) were inferred based on 2206 bp (712 variable, 614 parsimony informative) from three protein‐coding mitochondrial genes. Also included in the analysis were DNA sequences from two individuals, identified initially as Lutjanus apodus, which were sampled off the coast of Bahia State in Brazil (western Atlantic), and from three individuals labelled as ‘red snapper’ in the fish market in Puerto Armuelles, Panama (eastern Pacific). Bayesian posterior probabilities and maximum‐likelihood bootstrap percentages strongly supported monophyly of all lutjanines sampled and the hypothesis that western Atlantic lutjanines are derived from an Indo‐Pacific lutjanine lineage. The phylogenetic hypothesis also indicated that oceans where lutjanines are distributed (western Atlantic, eastern Pacific, and Indo‐Pacific) are not reciprocally monophyletic for the species distributed within them. There were three strongly supported clades that included all western Atlantic lutjanines: one included six species of Lutjanus from the western Atlantic, two species of Lutjanus from the eastern Pacific, and the monotypic genera Rhomboplites and Ocyurus (western Atlantic); one that included three, probably four, species of Lutjanus in the western Atlantic; and one that included Lutjanus cyanopterus (western Atlantic), an unknown species of Lutjanus from the eastern Pacific, and three species of Lutjanus from the Indo‐Pacific. Molecular‐clock calibrations supported an early Miocene diversification of an Indo‐Pacific lutjanine lineage that dispersed into the western Atlantic via the Panamanian Gateway. Divergent evolution among these lutjanines appears to have occurred both by vicariant and ecological speciation: the former following significant geographic or geological events, including both shoaling and closure of the Panamanian Gateway and tectonic upheavals, whereas the latter occurred via phenotypic diversification inferred to indicate adaptation to life in different habitats. Taxonomic revision of western Atlantic lutjanines appears warranted in that monotypic Ocyurus and Rhomboplites should be subsumed within the genus Lutjanus. Finally, it appears that retail mislabelling of ‘red snapper’ in commercial markets extends beyond the USA. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 915–929.     

Origins, phytogeny and taxonomy of the gadfly petrels Pterodroma spp

Taxonomic characters of gadfly petrels (Procellariidae: Pterodroma spp.) are reviewed and the genus is redefined. The structure of the upper intestines, which have become helicoidally twisted to varying degrees in most species, is an important character not hitherto used. Results of a phylogenetic study of the genus based mainly on the development of helicoidal intestines agree substantially with findings from studies of the Mallophaga parasitizing these petrels. One species, the Kerguelen Petrel Lugensa brevirostris, previously classified in Pterodroma, is shown to have closer affinity with some fulmar genera. The Tahiti Petrel Pseudobulweria rostrata and its rare or extinct relatives, formerly placed in a superspecies of Pterodroma, are more closely related to the Procellaria and Bulweria petrels, and may be derived from the link between the subfamilies Procellariinae and Fulmarinae. From consideration of the fossil record, anatomical characters and Mallophaga, a phylogeny of the Procellariidae is proposed which supports its monophyletic origin. Gadfly petrels apparently descended from the fulmars through the ancestral lines of Snow Petrels Pagodroma, the Kerguelen Petrel and finally the Blue Petrel Halobaena, which may have given rise separately both to gadfly petrels and to the prions Pachyptila. A late Pliocene origin of Pterodroma in the vicinity of southern New Zealand is possible. The genus Pterodroma is divided into four subgenera, representing four connected radiations, and 29 species are recognised. In probable chronological order, Proaestrelata (new subgenus) contains five species and is restricted to the Pacific Ocean, Cookilaria comprises six species restricted as breeders to the South Pacific, Hallstroma has seven species mainly in the tropics of the Pacific, Indian and South Atlantic Oceans, whereas Pterodroma includes 11 species which extend as breeders from the southwest Pacific west to the South Atlantic and into the North Atlantic. Ecological, physical and physiological adaptations in the evolution of gadfly petrels are discussed.     

MITOCHONDRIAL DNA DIVERSITY IN THE SEA URCHINS STRONGYLOCENTROTUS PURPURATUS AND S. DROEBACHIENSIS

Restriction-fragment analysis was used to measure mitochondrial DNA (mtDNA) variability in 79 individuals of two species of temperate sea urchins. For the purple urchin Strongylocentrotus purpuratus, individuals were collected 1,500 km apart in 1985 and again from the same localities in 1988 (about one urchin generation). Twenty mtDNA genotypes belonging to four clades were found among 38 individuals. All four clades were found in both localities and in both years. Genetic structure was further tested by calculating the degree of interdeme genetic variation (G_ST) and comparing this value to the G_ST's from randomly shuffled data. No geographic structure was found. For S. droebachiensis, only six mtDNA genotypes were found among 41 individuals collected from the Pacific and Atlantic oceans. More than 80% of the individuals belonged to two genotypes. The genotype that dominated collections in the Pacific also occurred in the Atlantic; however, a common Atlantic genotype was never found in the Pacific. These two genotypes were identical at 64 of 65 restriction sites, and were only 0.2% divergent from each other. G_ST analysis confirmed that there were significant genetic differences between Atlantic and Pacific populations. The small divergence between genotypes suggests recent, but not continuous, migration. These marine species show smaller genotypic differences than terrestrial species over similar spatial and temporal scales. Both recruitment of adults from planktonic larval pools and the spread of sibling larvae over large distances from parents probably act as buffers to genetic differences in species with planktonic life-history phases.     

Globorotalia puncticulata: Population divergence, dispersal and extinction related to Pliocene–Quaternary water masses

The isolating effect of water mass partitioning of populations on the morphology, stratigraphic distribution and extinction of planktonic foraminifera is assessed from the Pliocene–Quaternary record of Globorotalia puncticulata. Southern Hemisphere, Mediterranean and North Atlantic data on these aspects of its history are examined and appear consistent with a limited dispersal biogeographic model wherein populations are largely confined by hydrographic barriers.Earliest populations appeared during the latest Miocene in Southern Hemisphere middle latitude water masses. However, morphometric analysis shows that significant differentiation in the axial shape of shells had developed by 4 Ma between Southwest Pacific populations from ODP Site 1123 (temperate water) and ODP Site 1119 (subantarctic water). These sites are in close proximity but separated by the Subtropical Front. At Site 1123 inflation of late-formed chambers and reduction in the number included in the outer whorl created shell profiles that anticipated the globose form of Globorotalia inflata. The latter's gradual evolution from G. puncticulata s.s. took place in this temperate water mass, with the earliest morphotypes with three chambers in the outer whorl present by 4.1 Ma. In contrast, subantarctic populations at Site 1119 retained four chambers but their axial shape was modified. The development of a large, highly arched aperture and increase in the number of chambers in the outer whorl in Mediterranean–North Atlantic Globorotalia puncticulata bononiensis is an example of population differentiation later in the Pliocene.Chronostratigraphy shows that the northward expansion of central temperate water populations commenced with their occupation of Southwest Pacific subtropical water about 4.8 Ma. The rather abrupt entry of substantial populations into Mediterranean and North Atlantic water at 4.5 Ma marked a major biogeographic expansion and established G. puncticulata as a bipolar species. It was widely distributed about 3 Ma, with major populations in several water masses during a period of middle Pliocene warmth.After this acme North Atlantic and Mediterranean G. puncticulata bononiensis populations collapsed as late Pliocene Northern Hemisphere glacials intensified. They were extinguished in MIS 96 (2.4 Ma). Concurrently, G. puncticulata s.s became extinct in the warm subtropical Southwest Pacific. Subantarctic populations persisted but in turn were decimated in severe glacials during the Middle Pleistocene Transition. Most had disappeared by MIS 16 (0.66 Ma). However, at Sites 594 and 1119 there was a small Lazarus-like revival in MIS 11 (0.41 Ma). The highest known occurrence is in MIS 9 (0.33 Ma) at Site 1119. Confinement of the species to subantarctic water in the Pleistocene may have raised its vulnerability to extinction.While stable isotope data indicate that the lineage's evolution is related to depth habitat selection about the thermocline, its biogeography suggests that hydrographic barriers significantly isolated populations and probably facilitated speciation. Eddies such as the North Brazil Current rings provide conduits for inter-water mass transfer of populations but the history of G. puncticulata suggests that such mechanisms seldom operated successfully.The morphology of the lectotype of G. puncticulata s.s., from beach sand at Rimini, Italy, is consistent with a lower Pliocene source. Reports of living occurrences are poorly documented and the species is considered to be extinct.