The diversity and origin of exotic ants arriving in New Zealand via human-mediated dispersal

The number of exotic ant species being dispersed to new regions by human transportation and the trade pathways responsible for this are poorly understood. In this study, the taxonomic diversity, trade pathways, and origin of exotic ants intercepted at the New Zealand border were examined for the period 1955–2005. Overall, there were a total 4355 interception records, with 115 species from 52 genera. The 10 most frequently intercepted genera, and the 20 most frequently intercepted species contributed > 90% of all records. Many of the species frequently intercepted are regarded as invasive species, and several are established in New Zealand. The most intercepted species was Pheidole megacephala . Despite a relatively low trade relationship, a high proportion (> 64%) of the exotic ants which were intercepted originated from the Pacific region. However, the majority of species intercepted from the Pacific was exotic to the region (71%), or to a lesser extent, wide-ranging Pacific native species. No endemic species from the Pacific were intercepted. The effectiveness of detecting exotic ant species at the New Zealand border ranged from 48–78% for different trade pathways, indicating a number of species remain undetected. Trade routes associated with specific geographical regions represent a major filter for the arrival of exotic ant species. Despite the importance of the Pacific as a frequent pathway, we suggest that the future establishment of exotic ant species in New Zealand is likely to be mitigated by a renewed focus on trade routes with cool temperate regions, particularly Australia.     

Marine biodiversity of Costa Rica: Crustacea: Infraorder Anomura

The anomuran crabs are among the best known crustacean groups from the Pacific coast. However, this group is poorly known from the Caribbean coast of Costa Rica. In this compilation based on the literature and the collection at the Zoology Museum, Biology School, University of Costa Rica, we report the presence of 114 species of the Infraorder Anomura for Costa Rica, 20 species from the Caribbean, 96 species from the Pacific (two are present on both coasts). Twenty-nine species are new reports for Costa Rica, 15 from the Caribbean coast (74% of the total of species from that coast) and 14 from the Pacific (15% of the total from the Pacific). The range often species is extended to Costa Rica, siete from the Caribbean and three from the Pacific. Six species are reported for the first time from Cocos Island, where there are also four endemic species.     

Indo-Pacific population structure and evolutionary history of the coconut crab Birgus latro

Mitochondrial DNA variation was used to examine population structure in a widespread, marine-dispersed species, Birgus latro . Crabs were collected from eight locations throughout the species' Indo-Pacific distribution. Purified mtDNA from 160 individuals was cut with five restriction enzymes, revealing high haplotype diversity (0.96) and moderate nucleotide diversity (0.75%). Island populations from the Indian Ocean (Christmas I.) and Pacific Ocean were significantly different ( G _ST= 0.37) and had distinct mtDNA lineages with a net sequence divergence of 1.4%. Pacific island populations had diverged in a manner consistent with isolation by distance, with only the most peripheral populations being significantly different. The results for mtDNA are largely concordant with those from allozymes, although estimates of gene flow between the Indian and Pacific Oceans were much lower when based on mtDNA. The mtDNA phylogeny also permitted a deeper examination of the evolutionary and demographic history of Birgus latro . Long-term separation of populations is evident in the complete phylogenetic subdivision of mtDNA lineages between the Indian and Pacific Ocean populations sampled. The starlike phylogeny of alleles from the Pacific suggests a rapid population expansion in the Pacific during the Pleistocene. Including information about allele phylogeny, as well as distribution and frequency, obscured contemporary population structure, but provided unique insights into the evolutionary history of the species.     

Oceanic barriers as indicated by scombrid fishes and their parasites

Four genera of scombrid fishes (26 species) and their copepod (32 species) and monogenean ectoparasites (25 species) were used to test the hypothesis that the East Pacific Barrier is responsible for the most pronounced break in the circum-tropical warm water fauna of the continental shelves, and not the New World Barrier. Analysis at the species level showed that there is a primary centre of diversity in the West Pacific, and a secondary centre in the West Atlantic. The former, almost entirely, shares its species of the largely coastal Scomberomorus and Grammatorcynus and their parasites with seas located to the west. Only four parasites (all copepods) are shared by the East and West Pacific, and they are circum-tropical. In contrast, the West Pacific shares species of the more pelagic Scomber and their parasites with seas both to the east and west, although at the genus level, only two circum-tropical monogenean genera are shared by the E and W Pacific. We conclude that the East Pacific Barrier has been a 100% or almost 100% effective barrier to dispersal of species of Scomberomorus, Grammatorcynus and their parasites, whereas for species of Scomber and their parasites, the East Pacific has been a less effective barrier.     

Autolytinae, Eusyllinae an Exogoninae (Syllidae: Polychaeta) from Coiba National Park, Panama)

This is the second paper on the Syllidae (Polychaeta) from hard substrata of the Coiba National Park (Pacific coast of Panama) (the first paper treated Syllinae). On this second paper nineteen species belonging to eleven genera of the subfamilies Autolytinae, Eusyllinae and Exogoninae (Syllidae: Polychaeta) are reported. The samples were collected by SCUBA diving, either by removing 4 kg blocks of dead coral (Pocillopora spp.) or scraping off 25 x 25 cm quadrats of Telesto multiflora or algae (Dyctiota cf. flavellata, Padina cf. durvillaei and another currently unidentified species). They were collected during four expeditions carried out between 1996 and 1998. Four species are newly reported for the Eastern Pacific: Eusyllis lamelligera Marion & Bobretzky, 1875, Exogone exmouthensis Hartmann-Schr?der, 1980, Proceraea cf. cornuta (Agassiz, 1863) and Sphaerosyllis (S.) magnidentata Perkins, 1981. Ten species are reported for the first time from the Central America Pacific coast: Amblyosyllis cf. granosa Ehlers, 1897 (sensu Westheide 1974), Amblyosyllis speciosa Izuka, 1912, Autolytus multidenticulatus Westheide, 1974, Exogone naidinoides Westheide, 1974, Grubeosyllis concinna (Westheide, 1974), Odontosyllis fulgurans (Audouin & Milne Edward, 1833), Opisthodonta mitchelli Kudenov & Harris, 1995, Pionosyllis articulata Kudenov & Harris, 1995, Sphaerosyllis (S.) californiensis Hartman, 1966 and Syllides cf. reishi Dorsey, 1978. The family Syllidae is represented in the Coiba National Park by 14% of cosmopolitan species, 14% of circumtropical species, 22% of Atlantic and Pacific species and 25% of Pacific species restricted to warm waters. In samples of dead coral the dominant species are S. armillaris (Muller, 1771) and S. gracilis (Campoy, 1982); in samples of telestaceans Grubeosyllis concinna dominates.     

Ichthyofauna of the Navidad Bay, Jalisco, México]

A check-list and the relative abundance of the ichthyofauna from Bahia de Navidad, Jalisco, Mexico is presented. This poorly known area of the eastern Tropical Pacific ocean is characterized by its great fish diversity. A total of 245 fish species (163 genera, 72 families) were collected during the 1994-1995 annual cycle. Vouchers were deposited in Centro de Ecologia Costera, Universidad de Guadalajara. The most important families in number of species were Carangidae (20 species), Haemulidae (20), Serranidae (17), Sciaenidae (13) and Lutjanidae (8); 20 species represented 85% and 80% of relative abundance and biomass respectively. Microlepidotus brevipinnis, Caranx caninus, Caranx caballus, Kiphosus analogus and Caranx sexfasciatus, were the dominant species. Most fish species collected in Bahia de Navidad have zoogeographical affinity with the Panamic Province (65%), and in lesser proportion with the Eastern Pacific Ocean (19%), Circumtropical (7%), Mexican Province (5%), Indo-Pacific Ocean (3%) and Gulf of California (0.4%). The great ichthyological affinity between the study area and the southern Gulf of California suggests an extension of the of Gulf southern limit.     

Genetic isolation between the Western and Eastern Pacific populations of pronghorn spiny lobster Panulirus penicillatus

The pronghorn spiny lobster, Panulirus penicillatus, is a circumtropical species which has the widest global distribution among all the species of spiny lobster, ranging throughout the entire Indo-Pacific region. Partial nucleotide sequences of mitochondrial DNA COI (1,142-1,207 bp) and 16S rDNA (535-546 bp) regions were determined for adult and phyllosoma larval samples collected from the Eastern Pacific (EP)(Galápagos Islands and its adjacent water), Central Pacific (CP)(Hawaii and Tuamotu) and the Western Pacific (WP)(Japan, Indonesia, Fiji, New Caledonia and Australia). Phylogenetic analyses revealed two distinct large clades corresponding to the geographic origin of samples (EP and CP+WP). No haplotype was shared between the two regional samples, and average nucleotide sequence divergence (Kimura's two parameter distance) between EP and CP+WP samples was 3.8±0.5% for COI and 1.0±0.4% for 16S rDNA, both of which were much larger than those within samples. The present results indicate that the Pacific population of the pronghorn spiny lobster is subdivided into two distinct populations (Eastern Pacific and Central to Western Pacific), with no gene flow between them. Although the pronghorn spiny lobster have long-lived teleplanic larvae, the vast expanse of Pacific Ocean with no islands and no shallow substrate which is known as the East Pacific Barrier appears to have isolated these two populations for a long time (c.a. 1MY).     

Molecular phylogenetics of Floridosentis ward, 1953 (Acanthocephala: Neoechinorhynchidae) parasites of mullets (Osteichthyes) from Mexico, using 28S rDNA sequences

Species of Floridosentis (Acanthocephala) are common parasites of mullets (Mugil spp., Mugilidae) found in tropical marine and brackish water in the Americas. Floridosentis includes 2 species distributed in Mexico, i.e., Floridosentis pacifica, restricted to the Pacific Ocean near Salina Cruz, Oaxaca, and Floridosentis mugilis, distributed along the coast of the Pacific Ocean and the Gulf of Mexico. We sampled 18 populations of F. mugilis and F. pacifica (12 from the Pacific and 6 from the Gulf of Mexico) and sequenced a fragment of the rDNA large subunit to evaluate phylogenetic relationships of populations of Floridosentis spp. from Mexico. Species identification of museum specimens of F. mugilis from the Pacific Ocean was confirmed by examination of morphology traits. Phylogenetic trees inferred with maximum parsimony, maximum likelihood, and Bayesian inference indicate that Floridosentis is monophyletic comprising of 2 major well-supported clades, the first clade corresponding to F. mugilis from the Gulf of Mexico, and the second to F. pacifica from the Pacific Ocean. Genetic divergence between species ranged from 7.68 to 8.60%. Intraspecific divergence ranged from 0.14 to 0.86% for F. mugilis and from 1.72 to 4.49% for F. pacifica. Data obtained from diagnostic characters indicate that specimens from the Pacific Ocean in Mexico have differences in some traits among locations. These results are consistent with the phylogenetic hypothesis, indicating that F. pacifica is distributed in the Pacific Ocean in Mexico with 3 major lineages.     

Endemic and Indo-Pacific plankton in the Mediterranean Sea: a study based on dinoflagellate records

Aim To investigate biogeographical patterns based on published dinoflagellate records from the Mediterranean and Black Seas, and to provide a tentative list of endemic and Indo‐Pacific dinoflagellates in the Mediterranean Sea. Location Mediterranean Sea, Black Sea. Methods Checklists of dinoflagellates of the Mediterranean and Black Seas were compared with worldwide literature records. Only species reported in the Indo‐Pacific Ocean or exclusively known in the Mediterranean Sea were selected for biogeographical analysis. Results Dinoflagellates in the Mediterranean Sea comprised c. 43% of the world marine species and c. 88% of the dinoflagellate genera. Species richness among the Mediterranean sub‐basins showed marked differences due to the less reliable records of unarmoured (athecate) and rare dinoflagellates. These differences disappeared when only the more easily identifiable taxa were considered. Of the 673 dinoflagellates cited in the Mediterranean, 87% were also reported in the Atlantic Ocean. Only 40 taxa (6% of the total) were considered to be potential Indo‐Pacific species. Most were reported from the Ligurian Sea (21), and only two species from the Levantine basin. The other 48 taxa (7% of total) were known exclusively from the Mediterranean Sea, mainly from the Ligurian Sea. Half of these taxa were reported by a single author. Main conclusions Substantial dinoflagellates species richness can be attributed, in part, to the historical tradition of taxonomic studies in the Mediterranean Sea. The list of species of both Indo‐Pacific and exclusively Mediterranean species included taxa of dubious taxonomic validity or that were insufficiently known. The exclusion of these questionable taxa revealed the near absence of endemic dinoflagellates in the Mediterranean Sea compared with macroscopic organisms. This could be related to: (1) continuous replenishment of the plankton populations by the inflow of Atlantic water through the Strait of Gibraltar, (2) the possibility that species introduced during the Pliocenic flooding after the Messinian salinity crisis have not had enough time to diverge from their Atlantic ancestors, and/or (3) the reliance on traditional taxonomy based on morphological characters, which precludes the detection of cryptic speciation.     

Deep mitochondrial divergence in Baja California populations of an aquilopelagic elasmobranch: the golden cownose ray

Assessing the realized effect of dispersal in the genetic makeup of a species has significant evolutionary, ecological, and economical consequences. Here, we investigate the genetic diversity and population differentiation in the aquilopelagic golden cownose ray Rhinoptera steindachneri from the Gulf of California (GC) and the Pacific coast of Baja California (PCBC) using the mitochondrial NADH2 gene. Low levels of genetic diversity were found with only 4 polymerase chain reaction-restriction fragment length polymorphism haplotypes among 76 specimens. Pacific coast organisms were fixed for a unique haplotype not shared with rays from the gulf; 92% of GC rays possessed a single NADH2 haplotype not found in the Pacific. This produced significant differentiation between the GC and the PCBC (Φ(CT) = 0.972, P < 0.001). A pronounced phylogeographic pattern was found in which GC haplotypes were reciprocally monophyletic relative to a very divergent Pacific lineage (d = 10%). Our results indicate that despite high dispersal potential, GC and PCBC golden cownose ray populations are characterized by highly divergent mitochondrial lineages. Although more evidence is needed to corroborate the genetic isolation and systematic status of PCBC and GC golden cownose rays, our results suggest a possible cryptic species in the region.     

Mu opioid receptor-like sequences are present throughout vertebrate evolution

The sequence of the mu opioid receptor is highly conserved among human, rat, and mouse. In order to gain insights into the evolution of the mu opioid receptor, polymerase chain reaction (PCR) was used to screen genomic DNA from a number of different species using degenerate oligonucleotides which recognize a highly conserved region. DNA was assayed from representative species of both the protostome and deuterostome branches of the metazoan phylogenetic tree. Mu opioid receptor-like sequences were found in all vertebrate species that were analyzed. These species included bovine, chicken, bullfrog, striped bass, thresher shark, and Pacific hagfish. However, no mu opioid receptor-like sequences were detected from protostomes or from any invertebrates. The PCR results demonstrate that the region of the mu opioid receptor gene between the first intracellular loop and the third transmembrane domain (TM3) has been highly conserved during evolution and that mu opioid receptor-like sequences are present in the earliest stages of vertebrate evolution. Additional opioid receptor-like sequence was obtained from mRNA isolated from Pacific hagfish brain using rapid amplification of cDNA ends (RACE). The sequence of the Pacific hagfish was most homologous with the human mu opioid receptor (72% at the amino acid level between intracellular loop 1 and transmembrane domain 6) although over the same region high homology was also observed with the delta opioid receptor (69%), the kappa receptor (63%), and opioid receptor-like (ORL1) (59%). The hagfish sequence showed low conservation with the mammalian opioid receptors in the first and second extracellular loops but high conservation in the transmembrane and intracellular domains. Received: 5 January 1996 / Accepted: 7 March 1996     

A biogeographic analysis and review of the far eastern Pacific coral reef region

New information on the presence and relative abundances of 41 reef-building (zooxanthellate) coral species at 11 eastern Pacific and 3 central Pacific localities is examined in a biogeographic analysis and review of the eastern Pacific coral reef region. The composition and origin of the coral fauna and other reef-associated taxa are assessed in the context of dispersal and vicariance hypotheses. A minimum variance cluster analysis using coral species presence–absence classification data at the 14 localities revealed three eastern Pacific reef-coral provinces: (1) equatorial– mainland Ecuador to Costa Rica, including the Galápagos and Cocos Islands; (2) northern– mainland México and the Revillagigedo Islands; (3) island group– eastern Pacific Malpelo Island and Clipperton Atoll, and central Pacific Hawaiian, Johnston and Fanning Islands. Coral species richness is relatively high in the equatorial (17–26 species per locality) and northern (18–24 species) provinces, and low at two small offshore island localities (7–10 species). A high proportion (36.6%, 15 species) of eastern Pacific coral species occurs at only one or two localities; of these, three disappeared following the 1982–83 ENSO event, three occur as death assemblages at several localities, and five are endangered with known populations of ten or fewer colonies. Principal component analysis using ordinal relative density data for the 41 species at the 14 localities indicated three main species groupings, i.e., those with high, mid, and narrow spatial distributions. These groupings correlated with species population-dynamic characteristics. These results were compared with data for riverine discharges, ocean circulation patterns, shoreline habitat characteristics, and regional sea surface temperature data to help clarify the analyses as these measures of environmental variability affect coral community composition. Local richness was highest at localities with the highest environmental variability. Recent information regarding the strong affinity between eastern and central Pacific coral faunas, abundance of teleplanic larvae in oceanic currents, high genetic similarity of numerous reef-associated species, and appearances of numerous Indo-west Pacific species in the east Pacific following ENSO activity, suggest the bridging of the east Pacific filter bridge (formerly east Pacific barrier). Accepted: 20 September 1999     

Mitochondrial genomics of gadine fishes: implications for taxonomy and biogeographic origins from whole-genome data sets.

Phylogenetic analysis of 13 substantially complete mitochondrial DNA genome sequences (14,036 bp) from 10 taxa of gadine codfishes and pollock provides highly corroborated resolution of outstanding questions on their biogeographic evolution. Of 6 resolvable nodes among species, 4 were supported by >95% of bootstrap replications in parsimony, distance, likelihood, and similarly high posterior probabilities in bayesian analyses, one by 85%-95% according to the method of analysis, and one by 99% by one method and a majority of the other two. The endemic Pacific species, walleye pollock (Theragra chalcogramma), is more closely related to the endemic Atlantic species, Atlantic cod (Gadus macrocephalus), than either is to a second Pacific endemic, Pacific cod (Gadus macrocephalus). The walleye pollock should thus be referred to the genus Gadus as originally described (Gadus chalcogrammus Pallas 1811). Arcto-Atlantic Greenland cod, previously regarded as a distinct species (G. ogac), are a genomically distinguishable subspecies within pan-Pacific G. macrocephalus. Of the 2 endemic Arctic Ocean genera, Polar cod (Boreogadus) as the outgroup to Arctic cod (Arctogadus) and Gadus sensu lato is more strongly supported than a pairing of Boreogadus and Arctogadus as sister taxa. Taking into consideration historical patterns of hydrogeography, we outline a hypothesis of the origin of the 2 endemic Pacific species as independent but simultaneous invasions through the Bering Strait from an Arcto-Atlantic ancestral lineage. In contrast to the genome data, the complete proteome sequence (3830 amino acids) resolved only 3 nodes with >95% confidence, and placed Alaska pollock outside the Gadus clade owing to reversal mutations in the ND5 locus that restore ancestral, non-Gadus, amino acid residues in that species.     

The ‘Expansion–Contraction’ model of Pleistocene biogeography: rocky shores suffer a sea change?

Approximately 20 000 years ago the last glacial maximum (LGM) radically altered the distributions of many Northern Hemisphere terrestrial organisms. Fewer studies describing the biogeographic responses of marine species to the LGM have been conducted, but existing genetic data from coastal marine species indicate that fewer taxa show clear signatures of post-LGM recolonization. We have assembled a mitochondrial DNA (mtDNA) data set for 14 co-distributed northeastern Pacific rocky-shore species from four phyla by combining new sequences from ten species with previously published sequences from eight species. Nuclear sequences from four species were retrieved from GenBank, plus we gathered new elongation factor 1-α sequences from the barnacle Balanus glandula . Results from demographic analyses of mtDNA for five (36%) species ( Evasterias troschelii, Pisaster ochraceus, Littorina sitkana, L. scutulata, Xiphister mucosus ) were consistent with large population expansions occurring near the LGM, a pattern expected if these species recently recolonized the region. However, seven (50%) species ( Mytilus trossulus, M. californianus, B. glandula, S. cariosus, Patiria miniata, Katharina tunicata , X. atropurpureus ) exhibited histories consistent with long-term stability in effective population size, a pattern indicative of regional persistence during the LGM. Two species of Nucella with significant mtDNA genetic structure showed spatially variable demographic histories. Multilocus analyses for five species were largely consistent with mtDNA: the majority of multilocus interpopulation divergence times significantly exceeded the LGM. Our results indicate that the LGM did not extirpate the majority of species in the northeastern Pacific; instead, regional persistence during the LGM appears a common biogeographic history for rocky-shore organisms in this region.     

Records of anglerfishes (Lophiiformes: Lophiidae) from the western South Pacific Ocean, with descriptions of two new species

Species of Lophiidae collected from the western South Pacific Ocean are examined. Nine nominal species are recognized, all but one species with their distributions extended eastwards and southwards from the western Pacific Ocean. Two new species are described from Polynesia. Lophiodes iwamotoi n. sp. is described from five specimens collected from Savannah Seamount. It is characterized by a relatively long third dorsal-fin spine (56.9-70.8% standard length, L(S)) that extends (when retracted) to between the end of the soft dorsal fin and caudal-fin base; a relatively short fifth dorsal-fin spine (10.5-13.1% L(S)); a relatively short and narrow head; 19-20 pectoral-fin rays. Lophiodes maculatus n. sp. is described from 20 specimens collected from Marquesas Islands. It is characterized by an extremely long third dorsal-fin spine (71.9-87.2% L(S)), extending well beyond the caudal-fin base; a relatively short fifth dorsal-fin spine (11.4-16.5% L(S)); 17-18 pectoral-fin rays. A key to the lophiids in the western South Pacific Ocean is provided.     

Molecular phylogeny and phylogeography of free-living Bryozoa (Cupuladriidae) from both sides of the Isthmus of Panama

Genetic data were used to identify Recent species of free-living bryozoans (Cupuladriidae) from both sides of the Isthmus of Panama, and to examine their phylogenetic relationships, species richness, and population structures. An approximately 480bp fragment of the 16S mitochondrial rRNA gene was sequenced from 182 individuals from Panama, the Gulf of Mexico, and El Salvador. Ten haplotype groups (Cupuladria 4, 5, and 6; Discoporella 1, 2, 3A, 3B, 3C, 7, and 8) were identified. Genetic distances between haplotype groups (3.2-26.5%; K2P+Gamma) were 1-2 orders of magnitude greater than within groups (0.1-1.4%). Seven of the haplotype groups represent morphologically distinct species; Discoporellas 3A-C appear to be cryptic species. Phylogenetic analyses identified two pairs of transisthmian sister clades. An average divergence rate derived from other taxa suggests that Cupuladrias 4 and 5 diverged approximately 7Ma, a Discoporella 7 clade diverged from a 3A-C clade approximately 11Ma, and the 3A-C clade radiated approximately 6-4Ma; these events all predated final closure of the isthmus? 3Ma. The Caribbean side of the isthmus, with 5 species, is only marginally richer in cupuladriids than the Pacific side, with 4, but has greater phylogenetic depth. The Caribbean retains lineages stemming from a New World Miocene radiation that are not represented in the eastern Pacific; extant eastern Pacific cupuladriids share most recent common ancestry with only two of the Caribbean lineages. Species in the eastern Pacific tend to show shallow population structures, with high levels of gene flow between geographically separate populations, whereas Caribbean species tend to show deeper populations structures, with indications of restricted gene flow between Bocas del Toro/Gulf of Mosquitos and Costa Arriba/San Blas. The population structures derive from Pleistocene histories and may be of limited value in interpreting the macroevolutionary pattern, as our results provide no evidence of speciation on either side of the isthmus following closure in the late Pliocene.     

Diversity of mesophilic clostridia in Costa Rican soils

Costa Rica is located in the Tropic, one of the most biologically diverse regions of the world; its soil is an important epicenter of biodiversity and Clostridium spp. are among the most frequent bacteria. The diversity of clostridia in Costa Rican soils and its possible association with geographic zone, pH or type of soil was studied in 117 soil samples: 18 from the Atlantic Zone, 30 from the Central Plateau, 30 from the Dry Pacific, 13 from the North Zone, and 26 from the South Pacific. The pH and the mesophilic clostridia species were determined for each sample. For bacterial isolation, a selective methodology for spores and pre-reduced anaerobically sterilized media were used. A total of 1945 strains of clostridia were isolated, 98% were identified and corresponded to 54 species; the most frequent species were C. subterminale (56%), C. oceanicum (51%), C. bifermentans and C. glycolicum (50%, each), C. sporogenes (49%), and C. sordellii (42%). An average of 7.1 species per sample was obtained; the Atlantic Zone showed the greatest diversity: 8.6 species per sample and a total of 45 species. Except for C. chauvoei, all described toxigenic clostridia species were isolated; C. sordellii (42%) and C. perfringens (38%) were the most frequent. No statistical relation could be established between geographic zone or type of soil and any species, showing that clostridia had a high adaptation capability to grow in different soil conditions; only some clostridia were isolated from very acidic samples while others from soils with a wide range of pH. In general, a uniform distribution of most species and a high variety of clostridia in Costa Rican soils were observed, in agreement with the high biodiversity described for other living beings in this country.     

THE BIOGEOGRAPHIC ORIGIN OF ARCTIC ENDEMIC SEAWEEDS: A THERMOGEOGRAPHIC VIEW1

The Arctic is geologically and biogeographically young, and the origin of its seaweed flora has been widely debated. The Arctic littoral biogeographic region dates from the latest Tertiary and Pleistocene. Following the opening of Bering Strait, about 3.5 mya, the “Great Trans‐Arctic Biotic Interchange” populated the Arctic with a fauna strongly dominated by species of North Pacific origin. The Thermogeographic Model (TM) demonstrates why climate and geography continued to support this pattern in the Pleistocene. Thus, Arctic and Atlantic subarctic species of seaweeds are likely to be evolutionarily “based” in the North Pacific, subarctic species are likely to be widespread in the warmer Arctic, and species of Atlantic Boreal or warmer origin are unlikely in the Arctic and Subarctic. Although Arctic seaweeds have been thought to have a greater affinity with the North Atlantic, we have reanalyzed the Arctic endemic algal flora, using the Thermogeographic Model and evolutionary trees based on molecular data, to demonstrate otherwise. There are 35 congeneric species of the six, abundant Arctic Rhodophyta that we treat in this paper; 32 of these species (91%) occur in the North Pacific, two species (6%) occur in the Boreal or warmer Atlantic Ocean, and a single species is panoceanic, but restricted to the Subarctic. Laminaria solidungula J. Agardh, a kelp Arctic “endemic” species, has 18 sister species. While only eleven (61%) occur in the North Pacific, this rapidly dispersing and evolving genus is a terminal member of a diverse family and order (Laminariales) widely accepted to have evolved in the North Pacific. Thus, both the physical/time‐based TM and the dominant biogeographic pattern of relatives of Arctic macrophytes suggest strong compliance with the evidence of zoology, geology, and paleoclimatology that the Arctic marine flora is largely of Pacific origin.     

Characterization of diatom-cyanobacteria symbioses on the basis of nifH, hetR and 16S rRNA sequences

Richelia intracellularis is a symbiotic heterocystous cyanobacterium that is capable of forming associations with several genera of diatoms. nifH, 16S rRNA and hetR sequences were amplified and cloned from field populations of Richelia associated with Hemiaulus hauckii (N. Atlantic), with Rhizosolenia clevei (N. Pacific), and from a cultivated isolate of Calothrix associated with Chaetoceros from station ALOHA (N. Pacific). Sequence identity was highest (98.2%) among the 16S rRNA sequences, and more divergent for the hetR (83.8%) and nifH (91.1%) sequences. The hetR and nifH DNA and amino acid sequences obtained from the symbionts associated with the three different diatom genera diverged into three separate lineages supported by high bootstrap values. The data indicate that symbionts in the different hosts are distinct species or strains. Furthermore, three previously unidentified heterocystous-like nifH sequence groups recently reported from station ALOHA in the subtropical Pacific, het-1, het-2 and het-3, were linked to Richelia associated with R. clevei, H. hauckii and the Calothrix symbiont of Chaetoceros sp. respectively.     

A mitochondrial phylogeography of Brachidontes variabilis (Bivalvia: Mytilidae) reveals three cryptic species

This study examined genetic variation across the range of Brachidontes variabilis to produce a molecular phylogeography. Neighbour joining (NJ), minimum evolution (ME) and maximum parsimony (MP) trees based on partial mitochondrial DNA sequences of 16S-rDNA and cytochrome oxidase (COI) genes revealed three monophyletic clades: (1) Brachidontes pharaonis s.l. from the Mediterranean Sea and the Red Sea; (2) B. variabilis from the Indian Ocean; (3) B. variabilis from the western Pacific Ocean. Although the three clades have never been differentiated by malacologists employing conventional morphological keys, they should be ascribed to the taxonomic rank of species. The nucleotide divergences between Brachidontes lineages (between 10.3% and 23.2%) were substantially higher than the divergence between congeneric Mytilus species (2.3–6.7%) and corresponded to interspecific divergences found in other bivalvia, indicating that they should be considered three different species. Analysis of the 16S-rDNA sequences revealed heteroplasmy, indicating dual uniparental inheritance (DUI) of mtDNA in the species of Brachidontes collected in the Indian Ocean, but not in the species in the Pacific nor in the species in the Red Sea and the Mediterranean Sea. When we employed the conventional estimate of the rate of mitochondrial sequence divergence (2% per million years), the divergence times for the three monophyletic lineages were 6–11 Myr for the Indian Ocean and Pacific Ocean Brachidontes sp. and 6.5–9 Myr for the Red Sea and Indian Ocean Brachidontes sp . Thus, these species diverged from one another during the Miocene (23.8–5.3 Myr). We infer that a common ancestor of the three Brachidontes species probably had an Indo-Pacific distribution and that vicariance events, linked to Pleistocene glaciations first and then to the opening of the Red Sea, produced three monophyletic lineages.