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Structure, biodiversity, and historical biogeography of nematode faunas in holarctic ruminants: morphological and molecular diagnoses for Teladorsagia boreoarcticus n. sp. (Nematoda: Ostertagiinae), a dimorphic cryptic species in muskoxen (Ovibos moschatus).

Discovery of the ostertagiine nematode Teladorsagia boreoarcticus n. sp. in muskoxen, Ovibos moschatus, from the central Canadian Arctic highlights the paucity of knowledge about the genealogical and numerical diversity of nematode faunas characteristic of artiodactyls at high latitudes across the Holarctic. Teladorsagia boreoarcticus is a dimorphic cryptic species distinguished from Teladorsagia circumcincta/Teladorsagia trifurcata in domestic sheep by a 13% divergence in the ND4 region of mitochondrial DNA, constant differences in the synlophe, and significantly longer esophageal valve, spicules, gubernaculum, and bursa. Teladorsagia boreoarcticus represents an archaic component of the North American fauna and may have a Holarctic distribution in muskoxen and caribou. Recognition of T. boreoarcticus in muskoxen, in part, corroborates hypotheses for the existence of a cryptic species complex of Teladorsagia spp. among Caprinae and Cervidae at high latitudes and indicates the importance of climatological determinants during the late Tertiary and Pleistocene on diversification of the fauna. Also reinforced is the concept of the North American fauna as a mosaic of endemic and introduced species. Discovery of a previously unrecognized species of Teladorsagia has additional implications and clearly indicates that (1) our knowledge is incomplete relative to potentially pathogenic nematodes that could be exchanged among domestic and wild caprines; (2) we do not have sufficient knowledge of the fauna to understand the ecological control mechanisms (limitations) on dissemination and host range; and (3) an understanding of historical and geographical influences on the genealogical diversity and distribution of nematode faunas in domestic and wild ruminants is requisite to define the interface between agricultural and natural ecosystems across the Holarctic.

Abomasum↗

Phylogeny and biogeography of Eupatorium (Asteraceae: Eupatorieae) based on nuclear ITS sequence data.

The classification of the predominantly Neotropical Eupatorieae depends upon the circumscription of the core genus Eupatorium. The recently proposed narrowing of Eupatorium to ∼42 species in eastern temperate North America, Europe, and eastern Asia was tested with phylogenetic analysis of nucleotide sequence variation in the internal transcribed spacer (ITS) region of nuclear ribosomal DNA. A total of 40 samples (36 species) of Eupatorieae was analyzed. Several species from North America, South America, and Eurasia that were formerly recognized within a large Eupatorium s.l. (sensu lato) were included in the study. Other taxa included were representative of the majority of the subtribes native to eastern temperate North America. Parsimony analysis supported the contention that Eupatorium be defined narrowly and suggested that Eupatoriadelphus is distinct. The tree topology suggested that Eupatorium and Eupatoriadelphus share a common North American ancestor with Liatris relative to other Eupatorieae. It was apparent that the presumed sister taxa in Eupatoriinae from South America belong to a different clade. These results suggest that, following initial divergence in North America, Eupatorium reached Europe via dispersal during the late Pliocene with subsequent radiation in Asia.

Journal Article↗

Phylogeny and biogeography of Juglans (Juglandaceae) based on matK and ITS sequence data.

We investigated phylogenetic and biogeographic relationships within Juglans (walnuts), a Tertiary disjunct genus, using 15 species of Juglans and related (Juglandaceae) outgroups. The relationships were analyzed using nucleotide sequences of the chloroplast gene matK and its flanking spacers and of the internal transcribed spacers (ITS) and 5.8S gene of the nuclear ribosomal DNA. The DNA sequences provided 246 informative characters for parsimony analysis. ITS data supported as monophyletic groups the four generic sections, Cardiocaryon, Dioscaryon, Rhysocaryon, and Trachycaryon. Within Rhysocaryon, the temperate black walnuts and the tropical black walnuts were supported as monophyletic groups. When the two data sets were combined, J. cinerea was nested within Cardiocaryon. Combined analysis with published nuclear DNA restriction site data placed J. cinerea in a monophyletic group with Cardiocaryon. These analyses consistently supported Juglans as a monophyletic group and as the sister group to the genus Pterocarya. The results of this work are consistent with the known geological history of Juglans. The fossil record suggests that the butternuts had evolved by the early Oligocene in North America. The presence of butternuts in Eurasia could be the result of migration from North America to Eurasia during the warming trend of the mid Oligocene.

Journal Article↗

Molecular phylogeny and biogeography of Linanthus (Polemoniaceae).

To better understand the evolutionary history of Linanthus (Polemoniaceae) and its relatives, molecular phylogenies based on DNA sequence data from the internal transcribed spacer (ITS) region of nrDNA and the chloroplast gene matK were estimated using several methods. Our data suggest two separate and well-supported lineages of Linanthus in close association with two other genera-Leptodactylon and Phlox. These results agree with previous molecular systematic work on the Polemoniaceae, but do not support the traditional classification of the genus as a natural group, nor do they support the sectional classification within the genus. With a distribution centered primarily in western North America and a high degree of endemism in the California Floristic Province, it has been suggested by Raven and Axelrod that the origin and diversification of Linanthus and its relatives were tied to the development of a summer-dry climate in western North America, which began around 13-15 million years ago (mya). Increased drying during the Pliocene (1.2-5 mya) has also been hypothesized by Axelrod to have led to an increase in plant speciation in California and adjacent areas. Divergence times within the Linanthus lineages were estimated from the ITS and matK gene trees. A log-likelihood ratio test could not reject clock-like evolution for the matK data; however, the clock was strongly rejected for the ITS data set. Although ITS molecular evolution was not clock-like, the estimated times of divergence were similar to those of the matK data set. Within both lineages of Linanthus there seems to have been considerable diversification that has occurred since the Pliocene.

Journal Article↗

[Fishes of the Cabo Pulmo reef, Gulf of California, Mexico: systematic list and aspects of abundance and biogeography].

The Cabo Pulmo reef is the most important coral formation of the Gulf of California; however, its ichthyological fauna has been poorly studied. To produce a systematic list with data on relative abundance and frequency, and biogeographical affinities, we relied on visual census, field observations, analysis of commercial and sport fisheries (from 1986 to 1998), and the literature. A total of 236 species have been recorded at Cabo Pulmo (155 genera and 60 families). This number doubles previous compilations and represents 65.1% of all reef fishes known from the Gulf of California, and about 35% of its entire shallow-water fishes. Of the total species number, 68.3% are from the Panamic Province, 11.0% Indo Pacific colonizers and the same percentage gulf endemics, 7.6% are circumtropical, 1.7% Atlantic and 0.5% cosmopolitan; none are endemic to the reef. The most abundant taxa are the labrids Thalassoma lucasanum, T. grammaticum and the pomacentrid Chromis atrilobata. Only eleven species (4.7% of total) appeared in 75% to 100% of census, and 36 (15.3% of total) had high levels of both abundance and frequency, evidencing that the community is dominated by few taxa. Local species richness exceeds the number reported for most rocky or coral reefs of the Pacific coast of México, and indicates that Cabo Pulmo is a key area in the gulf and the entire Mexican Pacific, from the ichthyological point of view.

Animals↗

Biogeography and origin of Lilium longiflorum and L. formosanum (Liliaceae) endemic to the Ryukyu Archipelago and Taiwan as determined by allozyme diversity.

Allozyme diversity on 13 isozyme loci was investigated for two bulbous species, Lilium longiflorum and L. formosanum, endemic to the subtropical archipelago of continental origin located in East Asia. Degrees of allozyme variability and divergence for L. longiflorum were very high for insular endemic species, indicating relatively longtime persistence of the present widespread distribution across many islands in this phenotypically little-changed species. Lilium formosanum exhibited rather lower variability and divergence than did L. longiflorum and was genetically close to the southern peripheral populations of L. longiflorum with 0.978 as its highest genetic identity value. Combined with other biological and insular geohistorical information, our results suggest that L. longiflorum was established around the end of the Pliocene when the current distribution area was still a continuous part of the ancient Asian continent, and L. formosanum was derived from southern populations of L. longiflorum around the late Pleistocene when the mainland of Taiwan was completely separated from the adjacent islands and the main continent. Depauperization of allozyme variability in some L. longiflorum populations was found on islands with lower altitudes. This reflects bottleneck effects after the complete or almost complete submergence of such low islands during the archipelago's development.

Journal Article↗

Historical biogeography of Melastomataceae: the roles of Tertiary migration and long-distance dispersal.

Melastomataceae and Memecylaceae are pantropically distributed sister groups for which an ndhF gene phylogeny for 91 species in 59 genera is here linked with Eurasian and North American fossils in a molecular clock approach to biogeographical reconstruction. Nine species from the eight next-closest families are used to root phylogenetic trees obtained under maximum likelihood criteria. Melastomataceae comprise ∼3000 species in the neotropics, ∼1000 in tropical Asia, 240 in Africa, and 225 in Madagascar in 150-166 genera, and the taxa sampled come from throughout this geographic range. Based on fossils, ranges of closest relatives, tree topology, and calibrated molecular divergences, Melastomataceae initially diversified in Paloecene/Eocene times in tropical forest north of the Tethys. Their earliest (Eocene) fossils are from northeastern North America, and during the Oligocene and Miocene melastomes occurred in North America as well as throughout Eurasia. They also entered South America, with earliest (Oligocene) South American fossils representing Merianieae. One clade (Melastomeae) reached Africa from the neotropics 14-12 million years ago and from there spread to Madagascar, India, and Indochina. Basalmost Melastomataceae (Kibessieae, Astronieae) are species-poor lineages restricted to Southeast Asia. However, a more derived Asian clade (Sonerileae/Dissochaeteae) repeatedly reached Madagascar and Africa during the Miocene and Pliocene. Contradicting earlier hypotheses, the current distribution of Melastomataceae is thus best explained by Neogene long-distance dispersal, not Gondwana fragmentation.

Journal Article↗

The historic biogeography of India: isolation or contact?

Geophysical maps depicting continental movement have consistently shown India, as it moved northward, to be located far out in the Tethys Sea. India split off from the African east coast about 148 m.y.a. From that time onward, according to almost all geophysical accounts, India was isolated from all of other continents until the early Miocene when it made contact with Eurasia. But the biological data, both fossil and Recent, indicate that this concept cannot be correct. If India had really existed as an isolated, oceanic continent for about 100 m.y., it should have developed a peculiar biota with many endemic genera and families in its terrestrial and shallow marine habitats. But there are virtually no remains of organisms indicating that India was isolated for any substantial time (millions of years). Instead, we find that almost all Indian taxa were possessed in common with other continents. As time went on, the northern relationships became stronger and the southern ones weaker. Most of the recent geophysical accounts show India not making contact with Eurasia until the early Miocene, but fossil materials show that this event must have taken place by the early Eocene. It has been postulated that, as India moved northward, it created a biogeographic barrier that separated marine fish populations and resulted in the east-west provinces that are now apparent in the Indian Ocean. At the same time, the barrier effect was supposed to have resulted in the formation of sister species that are now located far apart. Information currently available indicates that most living, tropical marine species are probably not over 3 m.y. old. Consequently, the northward movement of India, which took place primarily between 148 and 50 m.y.a., could have no bearing on the relationships of modern species.

Animals↗

Biogeography, a dirty word?

The newly developing fields of phylogeography and macroecology are ignoring historical patterns at their peril.

Animals↗

Introduction to Physidae (Gastropoda: Hygrophila); biogeography, classification, morphology.

Physidae, a world-wide family of freshwater snails with about 80 species, are reclassified by progressive characters of the penial complex (the terminal male reproductive system): form and composition of penial sheath and preputium, proportions and structure of penis, presence or absence of penial stylet, site of pore of penial canal, and number and insertions of penial retractor muscles. Observation of these characters, many not recognized previously, has been possible only by the technique used in anesthetizing, fixing, and preserving. These progressive characters are the principal basis of 23 genera, four grades and four clades within the family. The two established subfamilies are divided into seven new tribes including 11 new genera, with diagnoses and lists of species referred to each. Proposed as new are: in Aplexinae, Austrinautini, with Austrinauta g.n. and Caribnautu harryi g.n., nom.nov.; Aplexini; Amecanautini with Amecanauta jaliscoensis g.n., sp.n., Mexinauta g.n., and Mayabina g.n., with M. petenensis, polita, sanctijohannis, tempisquensis spp.nn., Tropinauta sinusdulcensis g.n., sp.n.; and Stenophysini, with Stenophysa spathidophallus sp.n.; in Physinae, Haitiini, with Haitia moreleti sp.n.; Physini, with Laurentiphysa chippevarum g.n., sp.n., Physa mirollii nom.nou.; and Physellini, with Chiapaphysa g.n., and C. grijalvae, C. pacifica spp.nn., Utahphysa g.n., Archiphysa g.n., with A. ashmuni, A. sonomae spp.nn., Physella hemphilli sp.n., and Ultraphysella sinaloae g.n., sp.n. The simplest reproductive system is found in Austrinauta of the Aplexinae; its penial complex approaches that in the related family Lymnaeidae. Within Physinae a close approximation is found in Haitia. By these two genera the two subfamilies are drawn close together. Four grades of progressive complexity are recognized: (I) penial sheath entirely muscular; (II) penial sheath with both glandular and muscular tissue; (III) penis with penial stylet or other specialization of the tip of the penis; and (IV) pore of penial canal lateral rather than terminal as in the lower grades. In both subfamilies there are clades with glandular tissue in the penial sheath, a penial sheath subdivided into two parts, and tip of penis specialized in various ways. These clades are formalized as new tribes. Of 23 genera of Physidae, 17 occur in Pacific drainages of North and Central America, eight of these restricted to the region. Concentration of primitive genera along the Pacific coast from Mexico to Costa Rica conforms to previous observations that primitive pulmonate families are concentrated within, or along the continental margins of, the Pacific Ocean. An ancestral origin of Physidae along an ancient eastern Pacific coast is probable. From this region the several lineages have spread north, south and east in the Americas, and through Siberia to Europe. Although Physinae have fewer genera than Aplexinae (11 v. 12), they have more species (47 v. 34). Greater land area in the temperate zone has provided more opportunity for speciation of Physinae, in contrast to the generally tropical and warm-temperate range of Aplexinae. Furthermore, 10 species of Physinae are localized in individual lakes, whereas Aplexinae are not lake-dwellers. Both well-developed egg strings and capsular strings are found in the spawn of Sibirenauta elongatus. These structures have been known in Lymnaeidae, but not hitherto in Physidae; they are a link with some marine groups, such as Siphonariidae. Spiral color bands and white streaks in the shell of Mexinauta recall those in Lancidae (Lymnaeacea), whereas the radula of Physidae is like that of Chilinidae. Physidae thus show affinities to various basal stocks of aquatic pulmonates; no clear-cut sister-group can be recognized. Most species have a restricted range; out of 55 with sufficiently detailed information for analysis, 25 are limited to a single 1 degrees x 1 degrees quadrangle. Only a few species are widespread, on one or even two continents. Accordingly, more species of Physidae are threatened by habitat destruction than in other families of Hygrophila with generally wider distributions. Other features are a key to genera; catalog of more than 430 names applied to living Physidae, with original reference, type locality, and location of type specimens; summary of museums with types; and glossary.

Animals↗

Psychrophilic prokaryote structural-functional relationships, biogeography and evolution within marine sediment.

Prokaryote diversity has been found to be surprisingly high in cold marine sediments with numerous clades detected spread throughout many phyla. Marine benthic sediment clades are largely ecotypically distinct and autochthonous. Since almost all marine sediment prokaryotic taxa have yet to be cultivated, functionality is currently overwhelmingly cryptic for most benthic prokaryotic taxa except those falling into specific lineages for which there is cultivation or detailed biogeochemical data. Multivariate statistical comparisons of 16S rRNA gene sequence and denaturing gradient gel electrophoresis (DGGE) data show distinct distribution patterns of prokaryotic communities in sediment layers. By comparison geographical differences and differences related to the physical texture and organic content seem to result in generally smaller differences.

Archaea↗

Phylogeny, biogeography, and molecular dating of cornelian cherries (Cornus, Cornaceae): tracking Tertiary plant migration.

Data from four DNA regions (rbcL, matK, 26S rDNA, and ITS) as well as extant and fossil morphology were used to reconstruct the phylogeny and biogeographic history of an intercontinentally disjunct plant group, the cornelian cherries of Cornus (dogwoods). The study tests previous hypotheses on the relative roles of two Tertiary land bridges, the North Atlantic land bridge (NALB) and the Bering land bridge (BLB), in plant migration across continents. Three approaches, the Bayesian, nonparametric rate smoothing (NPRS), and penalized likelihood (PL) methods, were employed to estimate the times of geographic isolations of species. Dispersal and vicariance analysis (DIVA) was performed to infer the sequence and directionality of biogeographic pathways. Results of phylogenetic analyses suggest that among the six living species, C. sessilis from western North America represents the oldest lineage, followed by C. volkensii from Africa. The four Eurasian species form a clade consisting of two sister pairs, C. mas-C. officinalis and C. chinensis-C. eydeana. Results of DIVA and data from fossils and molecular dating indicate that the cornelian cherry subgroup arose in Europe as early as the Paleocene. Fossils confirm that the group was present in North America by the late Paleocene, consistent with the DIVA predictions that, by the end of the Eocene, it had diversified into several species and expanded its distribution to North America via the NALB and to Africa via the last direct connection between Eurasia and Africa prior to the Miocene, or via long-distance dispersal. The cornelian cherries in eastern Asia appear to be derived from two independent dispersal events from Europe. These events are inferred to have occurred during the Oligocene and Miocene. This study supports the hypothesis that the NALB served as an important land bridge connecting the North American and European floras, as well as connecting American and African floras via Europe during the early Tertiary.

Base Sequence↗

Global population genetic structure and biogeography of the oceanic copepods Eucalanus hyalinus and E. spinifer.

Although theory dictates that limited gene flow between populations is a necessary precursor to speciation under allopatric and parapatric models, it is currently unclear how genetic differentiation between conspecific populations can arise in open-ocean plankton species. I examined two recently distinguished sympatric, circumglobal sister species, Eucalanus hyalinus and Eucalanus spinifer, for population genetic structure throughout their global biogeographic ranges. Here I show that oceanic zooplankton species can be highly genetically structured on macrogeographic spatial scales, despite experiencing extensive gene flow within features of the large-scale ocean circulation. Mitochondrial DNA analyses of 450 and 383 individuals of E. hyalinus and E. spinifer, respectively, revealed that habitat discontinuities at the boundaries of subtropical gyres in the North and South Pacific, as well as continental land masses, acted as effective barriers to gene flow for both species. However, the impact of specific barriers on population genetic structure varied between the sister species, despite their close phylogenetic relationship and similar circumglobal biogeogeographic distributions. The sister species differed in their oceanographic distributions, with E. spinifer dominating oligotrophic waters of the subtropical gyres and E. hyalinus more abundant along central water mass boundaries and in frontal zones and upwelling systems. This species-specific difference in the oceanographic habitat is an important factor determining the historical and contemporary patterns of dispersal of the two species. I suggest that species-specific ecological differences are likely to be a primary determinant of population genetic structure of open-ocean plankton.

Animals↗