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Molecular phylogeography of Dolichopoda cave crickets (Orthoptera, Rhaphidophoridae): a scenario suggested by mitochondrial DNA.

This study focuses on the phylogenetic relationships among a number of West-Mediterranean cave crickets species belonging to Dolichopoda; primarily a Mediterranean genus, distributed from eastern Pyrenees to Caucasus. In this paper, 11 Dolichopoda species from the French Pyrenees (D. linderi), the island of Corsica (D. bormansi and D. cyrnensis), and northern, central, and southern Italy (D. ligustica, D. schiavazzii, D. aegilion, D. baccettii, D. laetitiae, D. geniculata, D. capreensis, and D. palpata) were studied. Two more species, one from the Caucasus, D. euxina, and one from Greece, D. remyi, were also included in the analyses, together with more distant species within the same family to be used as outgroups. Fifteen hundred base pairs of mitochondrial DNA, corresponding to the small subunit of the ribosomal RNA (16S rRNA) and to the subunit I of the cytochrome oxidase I (COI), were sequenced in order to clarify the phylogenetic relationships and biogeography of this group of Mediterranean cave crickets. The molecular data are congruent with a phylogeographic pattern; with the geographically close species also the most related ones. Based on mtDNA divergence, the present-day distribution of genetic diversity seems to have been impacted by climatic events due to glacial and interglacial cycles that have characterized the Pleistocene era.

Animals↗

The phylogenetic relationships among non-diplomystid catfishes as inferred from mitochondrial cytochrome b sequences; the search for the ictalurid sister taxon (Otophysi: Siluriformes).

The relationships among families of catfishes are poorly understood and have yet to be the subject of a comprehensive investigation with molecular data. Existing phylogenetic hypotheses are based on morphological data and incompletely resolved. This study analyzed complete sequences of mitochondrial gene cytochrome b for 170 species from 29 of 33 extant families, and focused on the relationships of Ictaluridae to other catfishes. In addition to previous phylogenetic studies, the fossil record, paleogeography, biogeography, and distribution of extant catfish families collectively suggest the location (if extant) of the ictalurid sister taxon to be Northern or Eastern Asia. Of the extant catfishes currently native to this area and included in this analysis, parsimony and Bayesian likelihood analyses recovered Cranoglanis bouderius as the most proximal sister taxon of Ictaluridae. Seemingly, ictalurids and cranoglanidids represent another biogeographic component linking freshwater fishes of North America and eastern Asia, e.g., catostomids and paddlefishes. The results coupled with present-day catfish distributions and inferences from the fossil record collectively suggest the ancestor of Ictaluridae to have invaded freshwaters of North America at the close of the Cretaceous through northeastern Asia and northwestern North America. Other superfamilial nodes supported the results of previous phylogenetic studies of narrower taxonomic scope. Several novel relationships were recovered (including a clade composed of Pimelodidae, Pseudopimelodidae, and Heptapteridae) and these along with sources of systematic error are discussed. A broad sampling of Bagridae permitted an examination of intergeneric relationships within this family and in light of recent morphological and molecular studies.

Animals↗

Molecular phylogenetic relationship of toads distributed in the Far East and Europe inferred from the nucleotide sequences of mitochondrial DNA genes.

The toads of the Bufo bufo species group are widely distributed in the Eurasian continent and Japanese Archipelago. In this study, we analyzed the mtDNA gene sequences of this species group and estimated the divergence time to clarify the evolutionary relationships and biogeography of toads distributed in the Far East and Europe. The phylogenetic tree indicated that this group produced Bufo bufo in Europe, whereas it produced B. japonicus in the Far East. B. japonicus was divided into three major clades corresponding to a group consisting of B. j. gargarizans in China, B. j. bankorensis in Taiwan, and B. j. miyakonis on Miyako Isl. and eastern and western groups of Japanese B. j. japonicus subspecies group. The eastern and western groups were divided into several subclades which tended to reflect the region-specific geographic distribution of all localities except B. j. japonicus from Hakodate. The estimated branching times of these clades suggest that geological events may have influenced the divergence of the toads distributed in the Far East and Europe.

Animals↗

Analysis of family-level relationships in bees (Hymenoptera: Apiformes) using 28S and two previously unexplored nuclear genes: CAD and RNA polymerase II.

We analyzed a combined data set of two protein-coding nuclear genes (CAD and RNA polymerase II) and a nuclear ribosomal gene (28S D2-D4 region) for 68 bee species and 11 wasp outgroups. Our taxon sampling included all seven extant bee families, 17 of 20 subfamilies, and diverse tribes. Wasp outgroups included the two families most closely related to bees: Crabronidae and Sphecidae. We analyzed the combined and single gene data sets using parsimony and Bayesian methods, which yielded largely congruent results. Our results provide reasonably strong support for family and subfamily-level relationships among bees. Our data set strongly supports the sister-group relationship of the Colletidae and Stenotritidae, and places Halictidae as sister to this clade combined. Our analyses place the Melittidae and the long-tongued (LT) bee clade (Apidae+Megachilidae) near the base of the tree with Colletidae (and Stenotritidae) in a fairly highly derived position. This topology ("Melittidae-LT basal") was obtained in previous morphological studies under certain methods of character coding. A more widely accepted tree topology that places Colletidae (and/or Stenotritidae) as sister to all other bees ("Colletidae basal") is not supported by our data. The "Melittidae-LT basal" hypothesis may better explain patterns in the bee fossil record as well as historical biogeography of certain bee groups. Our results provide new insights into higher-level bee phylogeny and indicate that CAD, RNA polymerase II, and 28S are useful data sets for resolving Cretaceous-age divergences in bees and other Hymenoptera.

Animals↗

Phylogenetic relationships of horned lizards (Phrynosoma) based on nuclear and mitochondrial data: evidence for a misleading mitochondrial gene tree.

It has proven remarkably difficult to obtain a well-resolved and strongly supported phylogeny for horned lizards (Phrynosoma) because of incongruence between morphological and mitochondrial DNA sequence data. We infer the phylogenetic relationships among all 17 extant Phrynosoma species using >5.1 kb of mtDNA (12S rRNA, 16S rRNA, ND1, ND2, ND4, Cyt b, and associated tRNA genes), and >2.2kb from three nuclear genes (RAG-1, BDNF, and GAPD) for most taxa. We conduct separate and combined phylogenetic analyses of these data using maximum parsimony, maximum likelihood, and Bayesian methods. The phylogenetic relationships inferred from the mtDNA data are congruent with previous mtDNA analyses based on fewer characters and provide strong support for most branches. However, we detected strong incongruence between the mtDNA and nuclear data using comparisons of branch support and Shimodaira-Hasegawa tests, with the (P. platyrhinos+P. goodei) clade identified as the primary source of this conflict. Our analysis of a P. mcalliixP. goodei hybrid suggests that this incongruence is caused by reticulation via introgressive hybridization. Our preferred phylogeny based on an analysis of the combined data (excluding the introgressed mtDNA data) provides a new framework for interpreting character evolution and biogeography within Phrynosoma. In the context of this improved phylogeny we propose a phylogenetic taxonomy highlighting four clades: (1) Tapaja, containing the viviparous short-horned lizards P. ditmarsi, P. hernandesi, P. douglasii, and P. orbiculare; (2) Anota, containing species with prominent cranial horns (P. solare, P. mcallii, and the P. coronatum group); (3) Doliosaurus, containing three species lacking antipredator blood-squirting (P. modestum, P. platyrhinos, and P. goodei); and (4) Brevicauda, containing two viviparous species with extremely short tails that lack blood-squirting (P. braconnieri and P. taurus).

Animals↗

A multidisciplinary approach reveals cryptic diversity in Western Palearctic Tetramorium ants (Hymenoptera: Formicidae).

Diversity of ants of the Tetramorium caespitum/impurum complex was investigated in a multidisciplinary study. Focusing on morphologically hardly distinguishable Western Palearctic samples, we demonstrate the genetic and phenotypic diversity, demarcate phylogenetic entities, and discuss the clades in terms of biogeography. Sequences of 1113bp of the mitochondrial COI gene revealed 13 lineages. COII data, worker morphometry and male genitalia morphology corroborated the COI results for seven lineages; the remaining six were disregarded because of small sample size. A comparison with published data on cuticular hydrocarbons showed correspondence. The seven entities show different distribution patterns, though some ranges overlap in Central Europe. Since no major discrepancy between the results of the different disciplines became apparent, we conclude that the seven entities within the T. caespitum/impurum complex represent seven species. Geographical evidence allows the identification of T. caespitum and T. impurum, and we therefore designate neotypes and redescribe the two species in terms of morphology and mtDNA. As the revision of about 50 taxon names would go beyond the scope of this study, we refer to the remaining five species under code names. We discuss our findings in terms of plesiomorphy and convergent evolution by visualizing the mtDNA phylogeny in morphological space.

Animals↗

Molecular phylogeny of Banza (Orthoptera: Tettigoniidae), the endemic katydids of the Hawaiian Archipelago.

The extant endemic katydids (Orthoptera: Tettigoniidae) of the Hawaiian Archipelago include one to three species per high island and a single species on Nihoa, all currently placed in the genus Banza. These acoustic insects provide an excellent opportunity for investigating the evolution of reproductive isolation and speciation, but such studies require an understanding of phylogenetic relationships within the group. We use maximum parsimony, likelihood-based Bayesian inference, and maximum likelihood to infer phylogenetic relationships among these taxa, based on approximately 2kb of mitochondrial cytochrome oxidase I and cytochrome b. Our results strongly support two distinct high island clades: one clade ("Clade I") composed of species from Kauai, Oahu, Molokai, and Lanai and another clade ("Clade II") composed of species from Maui and Hawaii (Banza unica, from Oahu, may be basal to both these clades, but its placement is not well resolved). Within these clades, some inferred relationships are strongly supported, such as the sister status of B. kauaiensis (Kauai) and B. parvula (Oahu) within Clade I, but other relationships remain more ambiguous, such as the relative position of B. brunnea (Maui) within Clade II. Although a detailed reconstruction of the historical biogeography of the Hawaiian katydids is difficult, we use our genetic data combined with the known geological history of the Hawaiian Islands to set limits on plausible historical scenarios for diversification of this group. Beyond these historical biogeographic inferences, our results indicate possible cryptic speciation on both Oahu and Hawaii, as well as what may be unusually high average rates of nucleotide substitution. The present work sets the stage for future genetic and experimental investigations of this group.

Animals↗

An analysis of species boundaries and biogeographic patterns in a cryptic species complex: the rotifer--Brachionus plicatilis.

Since the advent of molecular phylogenetics, there is increasing evidence that many small aquatic and marine invertebrates--once believed to be single, cosmopolitan species--are in fact cryptic species complexes. Although the application of the biological species concept is central to the identification of species boundaries in these cryptic complexes, tests of reproductive isolation do not frequently accompany phylogenetic studies. Because different species concepts generally identify different boundaries in cryptic complexes, studies that apply multiple species concepts are needed to gain a more detailed understanding of patterns of diversification in these taxa. Here we explore different methods of empirically delimiting species boundaries in the salt water rotifer Brachionus plicatilis by comparing reproductive data (i.e., the traditional biological species concept) to phylogenetic data (the genealogical species concept). Based on a high degree of molecular sequence divergence and largely concordant genetic patterns in COI and ITS1, the genealogical species hypothesis indicates the existence of at least 14 species--the highest estimate for the group thus far. A test of the genealogical species concept with biological crosses shows a fairly high level of concordance, depending on the degree of reproductive success used to draw boundaries. The convergence of species concepts in this group suggests that many of the species within the group may be old. Although the diversity of the group is higher than previously understood, geographic distributions remain broad. Efficient passive dispersal has resulted in global distributions for many species with some evidence of isolation by distance over large geographic scales. These patterns concur with expectations that micro-meiofauna (0.1-1mm) have biogeographies intermediate to microbial organisms and large vertebrates. Sympatry of genetically distant strains is common.

Animals↗

Molecular phylogeny of New World Myotis (Chiroptera, Vespertilionidae) inferred from mitochondrial and nuclear DNA genes.

Recent studies have shown that species in the genus Myotis have evolved a number of convergent morphological traits, many of which are more related to their mode of food procurement than to their phylogeny. Surprisingly, the biogeographic origins of these species are a much better predictor of phylogenetic relationships, than their morphology. In particular, a monophyletic clade that includes all New World species was apparent, but only a third of the 38 species have been analysed. In order to better understand the evolution of this clade, we present phylogenetic reconstructions of 17 Nearctic and 13 Neotropical species of Myotis compared to a number of Old World congeners. These reconstructions are based on mitochondrial cytochrome b (1140 bp), and nuclear Rag 2 genes (1148 bp). Monophyly of the New World clade is strongly supported in all analyses. Two Palaearctic sister species, one from the west (M. brandtii) and one from the east (M. gracilis), are embedded within the New World clade, suggesting that they either moved across the Bering Strait, or that they descended from the same ancestor that reached the New World. An emerging feature of these phylogenetic reconstructions is that limited faunal exchanges have occurred, including between the North and South American continents, further emphasizing the importance of biogeography in the radiation of Myotis. A fossil-calibrated, relaxed molecular-clock model was used to estimate the divergence time of New World lineages to 12.2+/-2.0 MYA. Early diversification of New World Myotis coincides with the sharp global cooling of the Middle Miocene. Radiation of the temperate-adapted Myotis may have been triggered by these climatic changes. The relative paucity of species currently found in South America might result from a combination of factors including the early presence of competitors better adapted to tropical habitats.

Animals↗

A multilocus phylogeny of New World jay genera.

We studied phylogenetic relationships of the New World Jays (NWJs) based on DNA sequences from three mitochondrial and two nuclear loci. Sampling included at least two individuals from each of the seven NWJ genera and four outgroups of closely related corvids, as well as six of the 16 Cyanocorax species (including two representatives of the previously recognized "Cissilopha"). Phylogenetic analyses were conducted using maximum parsimony, maximum likelihood, and Bayesian analyses for individual genes and a combined dataset. The combined phylogenetic analysis supports the basal position of Cyanolyca to all other NWJs, a (Cyanocorax (Calocitta, Psilorhinus)) clade, and a ((Cyanocitta, Aphelocoma) Gymnorhinus) clade that agrees with a novel morphological synapomorphy uniting Cyanocitta and Aphelocoma. Within Cyanocorax, C. yncas (former "Xanthoura") is basal to a split among former "Cyssilopha" species and the rest of the Cyanocorax species. To explore implications for the historical biogeography of the NJWs, we used Dispersal-Vicariance Analysis, which indicated that NWJs originated either in Mesoamerica or North America+Mesoamerica, with South American NWJs dispersing three times independently from Mesoamerica.

Adenylate Kinase↗

Molecular biology techniques in parasite ecology.

Molecular techniques are increasingly being used to study the ecology of a variety of organisms. These techniques represent important tools for the study of the systematics, population genetics, biogeography and ecology of parasites. Here, we review the techniques that have been employed to study the ecology and systematics of parasites (including bacteria and viruses). Particular emphasis is placed on the techniques of isoenzyme electrophoresis, in situ hybridisation and nucleic acid amplification to characterise parasite/microbial communities. The application of these techniques will be exemplified using ticks, bacterial endosymbionts and parasitic protozoa.

Animals↗

Genetic structure of populations of the red sea urchin, Strongylocentrotus franciscanus.

Population subdivision was evaluated in the red sea urchin, Strongylocentrotus franciscanus, using DNA sequence data from 134 adult individuals collected in 1995 and 1996. On average 22 individuals were sequenced from six geographic locations between Alaska and Baja California (N=134), nearly the full extent of the species range. DNA sequence data was obtained from direct sequencing of a 273 base pair region of the bindin gene, which encodes a sperm fertilization protein. Results indicate that bindin is sufficiently polymorphic to serve as a genetic marker. We identified 14 unique alleles present in the entire range sampled with a maximum of eight alleles at a specific site. Mean pairwise comparison of the 14 unique alleles indicates moderate sequence diversity (p-distance=1.06). Although there is conflicting evidence to suggest that Alaska populations may deviate from the Hardy-Weinberg expectations, analysis of bindin genotype frequencies indicate that it is not possible to reject the null hypothesis of random mating throughout the species range. The results of a chi-square test with pooling conform to Hardy-Weinberg expectations for all populations (P>0.05) except for the Alaska population (P=0.037). Inbreeding coefficients are consistent with this result and suggest that for the bindin locus, there is high gene flow. These results are compared with previously published results of genetic substructuring in sea urchins to examine relationships among population structure, dispersal potential and biogeography.

Journal Article↗

Persistence and transport of fauna on drifting kelp (Macrocystis pyrifera (L.) C. Agardh) rafts in the Southern California Bight.

Drifting rafts of Macrocystis pyrifera may connect isolated kelp forests in the Southern California Bight. To determine which species might utilize this dispersal mechanism, faunal samples from natural drifting rafts and attached M. pyrifera plants were collected during five cruises between March 1995 and December 1997. These rafts, which can be considered as floating islands, were aged and the macroinvertebrate assemblage enumerated. There was no significant relationship between raft age and species richness, or between species richness and distance offshore, which contrasts with predictions based on island biogeography. Species richness, however, was related to raft weight. Patterns of species presence and density were investigated relative to raft age for the species most frequently associated with rafts. Only one species, the isopod Idotea resecata, was found on all sampled rafts. Some species increased in frequency with raft age and others decreased, but only one relationship, a decline in the frequency of the anemone Epiactis prolifera with raft age was significant. When species density was examined over all cruises, only I. resecata had a significant change in density (an increase) with raft age, but additional significant relationships were found when species density patterns were considered by cruise. The results of all the tests were combined to provide a measure of "raft success". Nine of the most frequent 19 species had a positive score, indicating a favorable response to rafting, seven were unaffected, and two species had negative responses to rafting. Extinction times were calculated using species abundance and raft age relationships. Two species (E. prolifera and Paracerceis cordata), were predicted to persist on rafts for only about 100 days, which is the maximum estimated raft lifetime. All other species were predicted to persist for longer periods if the rafts floated longer. Kelp fauna that begin rafting appear to be largely unaffected by rafting, and hence dispersal on kelp rafts is possible for many members of the kelp forest community.

Journal Article↗

Isolation, characterization and quantification of microcystins (heptapeptides hepatotoxins) in Microcystis aeruginosa dominated bloom of Lalla Takerkoust lake-reservoir (Morocco).

This paper presents the first data on the identification, characterization and quantification of microcystins isolated from both an extract of a cyanobacteria natural bloom, collected from a eutrophic Moroccan reservoir (Lalla Takerkoust, Marrakesh) and an isolated strain cultivated under laboratory conditions. The isolation and purification of toxins was performed by reverse phase HPLC and then characterized by amino acid analysis and fast atom bombardment mass spectrometry (FAB-MS). Chemical characterization of the toxins from the bloom revealed variants of microcystins such as Mcyst-LR, Mcyst-RR, Mcyst-YR and [D-Asp3]Mcyst-LR. However, the Microcystis aeruginosa strain produced only Mcyst-RR. Using an ELISA assay the total microcystin contents of eight bloom samples collected from 1994 to 1997 ranged from 0.7 to 8.8 microg/mg of lyophilized material. The two isolated Microcystis strains contained higher amounts of microcystins (0.65 microg/ mg of dry weight) than the Pseudanabaena strains (0.021 microg/mg of dry weight). Our results show that the presence of cyanobacteria toxins in water used for drinking in a North African country may be regarded as an health hazard. These results are a contribution to the knowledge of the biogeography of toxic cyanobacteria and their toxins, namely in north African countries.

Bacterial Toxins↗

Fecundity and life-history strategies in marine invertebrates.

The reproductive strategies of an organism play a major role in the dynamics of the population and the biogeography and continuity of the species. Numerous processes are involved in reproduction leading to the production of offspring. Although diverse processes are involved in oogenesis (the production of eggs) and spermatogenesis (the production of sperm), the basic patterns of gametogenesis are similar amongst invertebrates, with the proliferation and differentiation of germ cells leading to the final production of mature gametes. The production of gametes, especially eggs, is energetically expensive, and therefore strongly sensitive to selective pressures. An organism can ingest and assimilate a limited amount of energy from the environment. The different ways by which energy is allocated to growth and reproduction in order to maximize fitness forms the basis of the differing life-history strategies that have developed in marine invertebrates. Fecundity is defined as the number of offspring produced by a female in a determined time period. The term fecundity needs to be explicitly defined in each study in order to obtain the maximum information from the data analysed. Because of the variety of egg production patterns found among marine invertebrates, a wide range of methodologies has been developed to quantify fecundity. These include direct egg counts in brooding species, spawning induction in live individuals and histological studies of preserved material. Specific considerations need to be taken into account for colonial organisms, because of their modular organization. The production of eggs requires an optimal allocation of energy into growth and reproduction for the maximization of parental fitness. Fecundity is central in studies of life-history theory and in the development of life-history models because it is directly related to energy allocation and partitioning. There are important relationships and trade-offs between fecundity and other life-history traits, such as egg size, female size and age, age at first reproduction, reproductive effort and residual reproductive value. These trade-offs, together with morpho-functional constraints and genetic variation determine the evolution of life histories through natural selection. Fecundity is a highly plastic character within the limits defined by the bioenergetics and life-history strategy of the organism. Egg production is affected mainly by environmental factors such as food quantity and quality, temperature or presence of toxic elements in the habitat. The differences in fecundity found among closely related species from different biogeographical locations reflect, at least in part, the differing environmental conditions of their habitat.

Adaptation, Physiological↗

Biology of early life stages in cephalopod molluscs.

Recent literature on embryonic and post-embryonic development, biology and behavioural ecology of juvenile cephalopods is reviewed. Emphasis is placed on biological processes. Life-history patterns and phylogenetic systematics, which are important for a proper understanding of the evolutionary history of the cephalopods, are only briefly touched upon. Egg sizes in cephalopods range from less than 1 mm to about 30 mm in diameter, so the hatchlings emerging from the largest eggs are bigger than the adults of pygmy squid, the smallest known cephalopods. Developmental durations from spawning to hatching range from a few days (for very small eggs developing at high temperatures) to one or possibly several years (for very large eggs developing at low temperatures). Such important differences notwithstanding, the morphogenetic processes are very similar in all cephalopod embryos, the major variant being the size of the so-called outer yolk sac, which may be rudimentary in extremely small embryos. Several questions concerning the timing of hatching in relation to the developmental stage attained, especially in terms of yok absorption, need clarification. These questions concern the elimination of the transient closure of the mouth, the final differentiation of digestive gland cells, and the removal of the tranquilliser effect of the perivitelline fluid necessary for the onset of the hatching behaviour. Cephalopod hatchlings are active predators. They refine their behavioural repertoires by learning from individual experience in dealing with prey and would-be predators. There is no truly larval phase, and the ecologically defined term paralarva should be used with caution. Given the considerable resource potential of cephalopods, investigations into dispersal and recruitment are of particular interest to fishery biology, but they are also important for ecological biogeography. The related studies of feeding and growth involve field sampling and tentative age determination of caught specimens, in combination with laboratory studies to test food quality, measure feeding rates, and validation of periodicities in accretional growth structures (e.g. "daily rings" in statoliths).

Animals↗

Paleoecology and coalescence: phylogeographic analysis of hypotheses from the fossil record.

The application of principles from coalescence theory to genealogical relationships within species can provide insights into the process of diversification and the influence of biogeography on distributional patterns. There are several features that make some organisms more suitable for detailed studies of historical processes; in particular, limited dispersal, which serves to conserve the patterns of genetic variation that developed during colonization. We describe the potential benefits of studies that integrate analyses of genetic variation with information from the fossil pollen record and present recent examples of the application of quantitative methods of phylogeographic analysis.

Journal Article↗

Life at the front: history, ecology and change on southern ocean islands.

Terrestrial ecosystems of southern ocean islands have enjoyed renewed attention recently owing to the discovery that their climates are changing dramatically. This has led to an enhanced understanding of the biogeography of this region, and an increased awareness that these ecosystems provide unrivalled opportunities for investigating the impacts of environmental change on interactions between invasive and indigenous species. Recent studies have revealed increases in the abundance of established alien species and in the strength of their negative impacts on local biota, especially through indirect interactions. Also, increases in island temperature and human visitor frequency are likely to result in increasing numbers of successful alien colonization events.

Journal Article↗