Nuclear ribosomal DNA and karyotypes indicate a NW African origin of South American Hypochaeris (Asteraceae, Cichorieae)

Hypochaeris has a disjunct distribution, with more than 15 species in the Mediterranean region, the Canary Islands, Europe, and Asia, and more than 40 species in South America. Previous studies have suggested that the New World taxa have evolved from ancestors similar to the central European H. maculata. Based on internal transcribed spacer (ITS) sequences and fluorescence in situ hybridization (FISH) with 5S and 18S-25S rDNA of the previously overlooked Hypochaeris angustifolia from Moyen Atlas, Morocco, we show that it is sister to the entire South American group. A biogeographic analysis supports the hypothesis of long-distance dispersal from NW Africa across the Atlantic Ocean for the origin of the South American taxa rather than migration from North America, through the Panamian land bridge, followed by subsequent extinction in North America. With the assumption of a molecular clock, the trans-Atlantic dispersal from NW Africa to South America is roughly estimated to have taken place during Pliocene or Pleistocene.     

Kerguelen Plateau and the Late Cretaceous southern-continent bioconnection hypothesis: tales from a topographical ocean

Aim  To evaluate rigorously an influential palaeobiogeographical hypothesis which states that in the Late Cretaceous (until c. 80 Ma) the Kerguelen Plateau provided a terrestrial causeway between East Antarctica and India that, in turn, formed part of a longer overland route between South America and Madagascar.
Location  Southern Ocean, Indian Ocean, East Antarctica, India and Madagascar.
Methods  Palaeogeographical modelling drawing on geological and geophysical data, bathymetric charts and plate tectonic reconstructions.
Results  During the Late Cretaceous, only small portions of the present-day Kerguelen Plateau were sub-aerial. Additionally, the plateau's north-north-west and south-south-east ends did not directly abut India and Antarctica, but instead were separated by large gaps. Thus, the notion that the two continents were then linked by a land route running the entire length of the edifice is almost certainly incorrect.
Main conclusions  The currently available physical evidence indicates that the Late Cretaceous southern-continent connection hypothesis, which is based exclusively on biological data, is untenable. Assuming the fossil and/or extant biological records of Madagascar–India are closely related to those of South America, alternative palaeogeographical scenarios need to be explored to explain this conundrum. Overwater dispersal and/or an alternative passage involving a more direct route via Africa (with crossings of the Mozambique Channel and a then appreciably narrower Central Atlantic) should be considered.     

Anchialine podocopid Ostracoda of the Galapagos Islands

Water-filled fissures and lava tubes on Isla Santa Cruz and Isla Isabela provide anchialine habitats for 11 species of podocopid and platycopid Ostracoda, most with Atlantic and West Indian affinities. Their ancestors may have been carried to earlier islands in the Galapagos system by trans-Isthmian currents, on vegetation or larger animals, or by migrating waterfowl. None are derived from present-day Pacific coastal faunas of North, Central or South America. Two species of Cytheracea, a new species of Anchistrocheles (Bairdiacea), and one of Cytherella (Platycopida) are described.     

The evolution of the Brachycytheride Ostracoda in the context of the breakup of Pangea

The ostracode Family Brachycytheridae is one of the most common taxa of Late Cretaceous shallow marine microfossils, and occurs in the marginal basins of Africa, Madagascar, the Middle East, South and North America and the Caribbean region. Their migration pathways were controlled largely by the plate tectonic events associated with the breakup of Pangea and the formation of the North and South Atlantic Oceans, the Caribbean Sea and the Gulf of Mexico. This study is an attempt to combine specific information on the plate tectonic events as evidenced by the timings of continental-scale structural movements, sea level dynamics (largely the product of mantle processes), the creation and destruction of migration pathways, and taxonomy of the group, and thus relate deep-seated Earth processes, surficial expression of these movements, and biological evolution. Although the Late Cretaceous Brachycytheridae are very widely distributed, their use for correlation has been hampered due to their conservative external morphology, with all of them having a posteriorly-sloped dorsal margin and a swollen ventrolateral carapace, and all of them were long assigned to a single genus, Brachycythere. In 2002, Puckett defined a new genus, Acuminobrachycythere, based on a distinctive clade of North American brachycytherids. It was also recognized at that time that the members of the family from Gondwana (mainly Africa and South America) were distinct in having a single (unsplit) second adductor muscle scar, whereas all of those from North America had a split second adductor. This tiny difference holds significant paleogeographic—and presumably evolutionary—information. Two new genera are therefore described herein, Sapucariella, which is restricted to Gondwana, and Tricostabrachycythere, which is an early, short-lived genus that occurred around the margins of Gondwana. Three new species are described and one is re-described. In all, there are six genera in the Family Brachycytheridae, which include Acuminobrachycythere, Brachycythere, Kaesleria (which is restricted to the Middle East), Opimocythere (which evolved in the Paleogene of North America and is the last of the Brachycytheridae), Sapucariella and Tricostabrachycythere. Taxonomic information is presented for 85 species, including new images of many of the type specimens. A summary of the breakup of Pangea is also presented, with precise paleogeographic reconstructions based on global magnetic anomalies.     

Inferring the biogeographic origins of inter‐continental disjunct endemics using a Bayes‐DIVA approach

The arcto‐Tertiary relictual flora is comprised of many genera that occur non‐contiguously in the temperate zones of eastern Asia, Europe, eastern North America, and western North America. Within each distributional area, species are typically endemic and may thus be widely separated from closely related species within the other areas. It is widely accepted that this common pattern of distribution resulted from of the fragmentation of a once more‐continuous arcto‐Tertiary forest. The historical biogeographic events leading to the present‐day disjunction have often been investigated using a phylogenetic approach. Limitations to these previous studies have included phylogenetic uncertainty and uncertainty in ancestral range reconstructions. However, the recently described Bayes‐DIVA method handles both types of uncertainty. Thus, we used Bayes‐DIVA analysis to reconstruct the stem lineage distributions for 185 endemic lineages from 23 disjunct genera representing 17 vascular plant families. In particular, we asked whether endemic lineages within each of the four distributional areas more often evolved from (1) widespread ancestors, (2) ancestors dispersed from other areas, or (3) endemic ancestors. We also considered which of these three biogeographic mechanisms may best explain the origins of arcto‐Tertiary disjunct endemics in the neotropics. Our results show that eastern Asian endemics more often evolved from endemic ancestors compared to endemics in Europe and eastern and western North America. Present‐day endemic lineages in the latter areas more often arose from widespread ancestors. Our results also provide anecdotal evidence for the importance of dispersal in the biogeographic origins of arcto‐Tertiary species endemic in the neotropics.     

The uplift of Qinghai-Xizang (Tibet) Plateau and the vicariance speciation of glyptosternoid fishes (Siluriformes: Sisoridae)

Based on the phylogenetic and biogeographical studies of the glyptosternoid fishes in Qinghai-Tibet area, the following hypothesis is proposed: the speciation of this group has a direct relationship with the three uplift intervals of the Qinghai-Tibet Plateau. This process was explained by the theory of vicariance of biogeography. The ancestor of this group was similar to Bagarus and/or Glyptothorax, which still have a wide distribution. At the moment when the Tethys sea closed, the Indian tectonic plate collided with the Eurasian tectonic plate, so the Glyptothorax-like and Bagarus-like ancestors entered Eurasia and gradually became widely distributed. After the Pleistocene, with the enforced colliding, the gradual uplift of the Qinghai-Tibet Plateau brought about the current water environment, and the Glyptosternoids were generated from Glyptotho-rax-like fish under this environment. The present Glyptosternum, distributed across the Himalayas is the ancestor of Glyptosternoids. In the three uplift inte     

Pakaraimaea dipterocarpacea is ectomycorrhizal, indicating an ancient Gondwanaland origin for the ectomycorrhizal habit in Dipterocarpaceae

The consistent association of Paleotropical Dipterocarpaceae with ectomycorrhizal (ECM) fungi suggests that ECM status is an ancestral character in the family. Despite its distinctive morphology, Pakaraimaea dipterocarpacea, a Neotropical Dipterocarpaceae endemic to the Guayana Region, is phylogenetically related to the Paleotropical Dipterocarpaceae. The confirmation of P. dipterocarpacea ECM status would indicate that Paleotropical Dipterocarpaceae and P. dipterocarpacea probably had a common ECM ancestor. Mycorrhizal colonization of P. dipterocarpacea was assessed, and ECMs were recorded using histological and molecular methods. P. dipterocarpacea was highly colonized by typical ECMs, and several ECM fungal taxa belonging to Clavulinaceae, Sebacinaceae, Cortinariaceae and Amanitaceae were identified. This paper provides the first documented evidence of ECM in a neotropical genus of Dipterocarpaceae and indicates that ECMs possibly evolved in Gondwana in ancestors of Dipterocarpaceae before the separation of South America from Africa by the Atlantic, c. 135 million years ago. The observation of Sebacinaceae and Clavulinaceae suggests that broad host range fungi are important components of P. dipterocarpacea ECM communities.     

How isolated is Antarctica?

The traditional view of Antarctica and the surrounding Southern Ocean as an isolated system is now being challenged by the recent discovery at the Antarctic Peninsula of adult spider crabs Hyas areneus from the North Atlantic and of larvae of subpolar marine invertebrates. These observations question whether the well described biogeographical similarities between the benthic fauna of the Antarctic Peninsula and the Magellan region of South America result from history (the two regions were once contiguous), or from a previously unrecognized low level of faunal exchange. Such exchange might be influenced by regional climate change, and also exacerbated by changes in human impact.     

Biogeographical and geological evidence for a smaller, completely-enclosed Pacific Basin in the Late Cretaceous

Aim To use biogeographical, palaeomagnetic, palaeosedimentary, and plate circuit data from Late Cretaceous regions in and around the Pacific to test the plate tectonic hypothesis of a pre‐Pacific superocean. Location East Asia, Australia, Antarctica, the western Americas, and the Pacific. Methods Literature surveys of the distributions of Cretaceous, circum‐Pacific taxa were compared with palaeomagnetic and palaeosedimentary data. Uncontroversial plate motions based on seafloor spreading data were also used to test the results of the biogeographical and palaeomagnetic analyses. Results The distributions of Cretaceous terrestrial taxa, mostly dinosaurs, imply direct, continental connections between Australia and East Asia, East Asia and North America, North America and South America, South America and Antarctica, and Antarctica and Australia. Palaeomagnetic, palaeosedimentary, and basic plate circuit analyses require little to no latitudinal motion of the Pacific plate with respect to the surrounding continents. Specifically, the data implies that western North America, East Asia, and the Pacific plate all increased in latitude by roughly the same amount (c. 11 ± 5°) since the Campanian – and that the Pacific Ocean Basin has increased in length north‐to‐south. Main conclusions Each of the analyses provides independent corroboration for the same conclusion: the Late Cretaceous Pacific plate was completely enclosed by the surrounding continents and has not experienced significant latitudinal motion with respect to North America, East Asia, or the Bering land bridge. This contrasts significantly with the plate tectonic history of the Pacific, implying instead that the Pacific plate formed in situ, pushing the continents apart as the plate and basin expanded. These results also substantiate recent biogeographical analyses that have concluded that a narrower Pacific Ocean Basin in the Mesozoic and early Tertiary provides the most reasonable explanation for the great number of trans‐Pacific disjunctions of poor dispersing taxa.     

Trans-oceanic dispersal and evolution of early composites (Asteraceae)

How did Asteraceae (the daisy family) expand from its area of origin and become so widespread? This question has challenged generations of evolutionary botanists. Molecular phylogenetic and biogeographic analyses indicate a South American origin of Asteraceae, a view supported by the recent discovery of the earliest fossils of the family in Middle Eocene (ca. 50 Ma) deposits in southern South America. The early-branching lineages in the phylogenetic tree of Asteraceae are South American and African, suggesting that the earliest successful colonization of areas outside South America may have involved long-distance dispersal to Africa. However, one particularly challenging unanswered question is how early members of Asteraceae reached Africa at a time when the Atlantic Ocean constituted a barrier between the two continents. Morphological, phylogenetic, geographic, paleogeographic, and paleontologic data have been combined to propose scenarios on possible geographical and dispersal routes and vectors of dispersion of early-branching lineages of Asteraceae from South America to Africa. Of the different scenarios proposed here, two concern alternative geographical routes: (1) via the Rio Grande Rise-Walvis Ridge axis in the South Atlantic; or (2) via Antarctica, possibly including the Subantarctic islands. Three scenarios consider different dispersal routes: (1) stepping-stones; (2) single-step; and (3) sweepstakes. Finally, three vectors of dispersion are considered: (1) birds; (2) wind; and (3) floating islands. Evaluation of these scenarios suggests that early-branching lineages of Asteraceae probably dispersed from South America to Africa along an island chain formed by the Rio Grande Rise and the Walvis Ridge, transported by birds, possibly combined with rafting and/or sweepstakes. Morphological changes typically associated with evolution on islands characterize many African carduoid descendants, providing indirect evidence for step-wise dispersal along the island chain.     

Panbiogeographical analysis of distribution patterns in hagfishes (Craniata: Myxinidae)

Aim To analyse the worldwide distribution patterns of hagfishes using panbiogeographical track analysis, and to attempt to correlate these patterns with the tectonic history of the ocean basins. Location Atlantic and Pacific oceans. Method The distributions of 47 out of 70 species of hagfish (in the genera Eptatretus, Myxine, Nemamyxine, Neomyxine, and Paramyxine) were studied by the panbiogeographical method of track analysis. The analysis was performed using distributional data obtained from the collections included in the Ocean Biogeographic Information System (OBIS, http://www.iobis.org ) and FishBase ( http://www.fishbase.org ), with additional records from the literature. Individual tracks were obtained for each species by plotting localities and connecting them by minimum‐spanning trees. Generalized tracks were determined from the spatial overlap between individual tracks. Results Six generalized tracks were found: in the Gulf of Mexico, Caribbean Sea, South‐eastern Atlantic, Western Pacific, North‐eastern Pacific and South‐eastern Pacific. Main conclusions The distribution patterns of myxinids are marked by a high degree of endemism and vicariance, and are correlated with the tectonic features involved in many of the events that led to the development of oceanic basins. The main massing of the group is around the Pacific Basin. In the Atlantic Ocean, the distribution of Myxine glutinosa seems to correspond to a classic trans‐oceanic track and vicariance resulting from the opening of the Atlantic Ocean during the Cretaceous. In the Pacific Ocean, the distribution of the Eptatretus and Paramyxine species is clearly associated with the margins of the Pacific tectonic plate. The generalized tracks of hagfishes are shared by several other groups of marine organisms, including many from shallow tropical waters, implying a common history for this marine biota. Overall, vicariance is a major feature of hagfish distribution, suggesting vicariant differentiation of widespread ancestors as a result of sea‐floor spreading between continents in connection with ocean formation.     

The evolution of South American endemic canids: a history of rapid diversification and morphological parallelism

The origin of endemic South American canid fauna has been traditionally linked with the rise of the Isthmus of Panama, suggesting that diversification of the dog fauna on this continent occurred very rapidly. Nevertheless, despite its obvious biogeographic appeal, the tempo of Canid evolution in South America has never been studied thoroughly. This issue can be suitably tackled with the inference of a molecular timescale. In this study, using a relaxed molecular clock method, we estimated that the most recent common ancestor of South American canids lived around 4 Ma, whereas all other splits within the clade occurred after the rise of the Panamanian land bridge. We suggest that the early diversification of the ancestors of the two main lineages of South American canids may have occurred in North America, before the Great American Interchange. Moreover, a concatenated morphological and molecular analysis put some extinct canid species well within the South American radiation, and shows that the dental adaptations to hypercarnivory evolved only once in the South American clade.     

Tracing the diversification history of a Neogene rodent invasion into South America

We investigated spatial patterns of evolutionary relatedness and diversification rates to test hypotheses about the historical biogeographic processes underlying the radiation of Neotropical rats and mice (Sigmodontinae, ~400 species). A negative correlation between mean phylogenetic distance and diversification rates of rodent assemblages reveals a pattern of species co‐occurrence in which assemblages of closely related species are also the fastest diversifying ones. Subregions of the Neotropics occupied by distantly related species that are on average more slowly diversifying include Central America, northern South America, and the Atlantic forest. In southern South America, recent species turnover appears to have been higher. Ancestral locations for the main tribes of sigmodontines were also estimated, suggesting eastern South America and the Amazonian lowlands were colonized before some central Andean regions, even though the latter are now centers of species richness for these rodents. Moreover, a past connection between the tropical Andes and the Atlantic Forest is suggested by our results, highlighting a role for a hypothetical arc connecting the two biomes, which would have impacted many other groups of organisms. Whether rapid, recent speciation in some regions is related to Quaternary climatic fluctuations and the young age of sigmodontines (~12.7 Ma crown age) or instead to intrinsic traits of these rodents remains an open question. If the former is true, we hypothesize that contrasting trends will characterize older Neotropical clades.     

Evolution of locomotion in Anthropoidea: the semicircular canal evidence

Our understanding of locomotor evolution in anthropoid primates has been limited to those taxa for which good postcranial fossil material and appropriate modern analogues are available. We report the results of an analysis of semicircular canal size variation in 16 fossil anthropoid species dating from the Late Eocene to the Late Miocene, and use these data to reconstruct evolutionary changes in locomotor adaptations in anthropoid primates over the last 35 Ma. Phylogenetically informed regression analyses of semicircular canal size reveal three important aspects of anthropoid locomotor evolution: (i) the earliest anthropoid primates engaged in relatively slow locomotor behaviours, suggesting that this was the basal anthropoid pattern; (ii) platyrrhines from the Miocene of South America were relatively agile compared with earlier anthropoids; and (iii) while the last common ancestor of cercopithecoids and hominoids likely was relatively slow like earlier stem catarrhines, the results suggest that the basal crown catarrhine may have been a relatively agile animal. The latter scenario would indicate that hominoids of the later Miocene secondarily derived their relatively slow locomotor repertoires.     

Hubrecht,Macfarlane, Jensen and Willmer: on the nature and testability of four versions of the nemertean theory of vertebrate origins

Four authors (A. A. W. Hubrecht, J. M. Macfarlane, D. D. Jensen and E. N. Willmer) have argued that nemerteans are the proximal invertebrate ancestors of vertebrates. While all these authors accept certain homologies (i.e. vertebrate pituitary and notochord with nemertean proboscis invagination and proboscis sheath or mesoderm), there is considerable variation among them in the derivation of the vertebrate nervous system, sense organs and foregut apertures. For example, Hubrecht suggests that the vertebrate nervous system developed from a solid dorsal nerve found in some nemerteans; Macfarlane suggests it developed by fusion of the left and right solid brains of nemerteans; Jensen suggests that the vertebrate tubular nervous system developed by the backward extension of the nemertean frontal organ and fusion with the nemertean dorsal and ventral brain ganglia. Modern EM, histochemical and biochemical methods can test empirically the four theories. Research testing these theories will also test the more popular acraniate theory, since that theory implies that vertebrate head structures evolved de novo in chordates and that no homologies should exist between vertebrate and nemertean head structures.     

Phylogenetic relationships within the aplocheiloid fish genus Rivulus (Cyprinodontiformes, Rivulidae): implications for Caribbean and Central American biogeography

We examined the phylogenetic relationships of 16 northern species of the aplocheiloid genus Rivulus inhabiting the Caribbean, Central America, and South America. A total of 714 base pairs per taxon were sequenced from two segments of the mitochondrial genome, 12S rRNA and cytochrome b. Both parsimony and neighbor-joining analyses suggest an ancient vicariant origin of the Greater Antillean taxa, in addition to a quite recent dispersal of species into the Lesser Antilles from the South American mainland. Combined analyses support the monophyly of the northern South American assemblage as the sister group of a Central American/Columbian biota. However, the monophyly of the Central American biota remains uncertain. Divergence estimates for the Central American taxa are calibrated from the Last Cretaceous separation of the proto-Antilles from the Americas. These data suggest that the extant Central American taxa represent the descendants of at least two separate invasions during the Cenozoic, prior to the closing of the Panamanian isthmus. Times are consistent with the extensive evidence for reptilian and mammalian exchange throughout the Cenozoic.      

Molecular phylogeny of the spiny lobster genus Palinurus (Decapoda: Palinuridae) with hypotheses on speciation in the NE Atlantic/Mediterranean and SW Indian Ocean

Sequence data derived from the mitochondrial DNA 16S rRNA and COI genes were used to determine the phylogenetic relationships among six Palinurus spiny lobster species. Three species (P. charlestoni, P. elephas, and P. mauritanicus) occur in the northeastern Atlantic/Mediterranean, and the others (P. barbarae, P. delagoae and P. gilchristi) inhabit the southwestern Indian Ocean. Parsimony and model based phylogenetics strongly supported the monophyly of the genus. A combined parsimony analysis based on 1001bp and 274 parsimony informative characters recovered the most resolved phylogeny with >70% bootstrap support for associations among species. The Atlantic P. charlestoni consistently clusters nested within the Indian Ocean clade, and the mtDNA sequence divergence between the two most distant species is 8.24%. If the northward collision of Africa with Eurasia in the Miocene caused the final physical separation between the Atlantic and Indian Ocean taxa, then the Palinurus mtDNA (COI and 16S combined) evolved no faster than 0.18% (lower bound) to 0.36% (upper bound) per lineage per million years. The six extant species occur in the pathways of the North Atlantic and South Indian Ocean gyres, and hypotheses on their radiation are developed relative to the strengthening of boundary currents in the Miocene and life history traits congruent with survival in strong ocean currents.     

GEOGRAPHIC GENETIC STRUCTURE IN THE FRANCISCANA DOLPHIN (PONTOPORIA BLAINVILLEI)

The franciscana, Pontoporia blainvillei , is endemic to the South Atlantic coast of South America. The species is of conservation concern because it suffers elevated mortality rates due to incidental captures in fishing nets, and perhaps it is one of the most threatened small cetaceans in this region. Previous morphological and genetic studies have suggested the existence of at least two distinct stocks to the north and south of Santa Catarina Island in Brazil. Fixed differences were found between a sample from Rio de Janeiro and one from Rio Grande do Sul, in southern Brazil. Using 94 sequences of the control region of mitochondrial DNA, we examined the genetic structure of the species. We found no shared haplotypes between Rio de Janeiro and samples from the southern range of the species distribution. However, a phylogenetic analysis suggests that the former population is diphyletic with respect to the southern samples. This suggests that the populations have not been isolated long enough to reach reciprocal monophyly. Furthermore, genetic differentiation is broadly consistent with a simple model of isolation by distance, that therefore appears as an alternative to a model of strict isolation of two stocks. The estimated levels of gene flow are higher among neighboring populations, and decrease as more distant localities are compared. Finally, the molecular data suggest that franciscanas have expanded in Rio de Janeiro.     

Origin and radiation of Southern Hemisphere coastal dolphins (genus Cephalorhynchus)

The genus Cephalorhynchus (Gray 1846) consists of four species of small coastal dolphins distributed in cool temperate waters around the Southern Hemisphere. Each species is sympatric with other members of the subfamily Lissodelphininae but widely separated from other congeners. To describe the origin and radiation of these species, we examined 442 bp of mitochondrial DNA control region sequences of 307 individuals from the genus Cephalorhynchus and compared these to sequences from other members of the subfamily Lissodelphininae. We investigate the hypotheses that Cephalorhynchus is a monophyletic genus or, alternatively, that the four species have arisen separately from pelagic Lissodelphine species and have converged morphologically. Our results support the monophyly of Cephalorhynchus within the Lissodelphininae and a pattern of radiation by colonization. We confirm a pattern of shallow but diagnosable species clades with Heaviside's dolphin as the basal branch. We further examine the monophyly of maternal haplotypes represented by our large population sample for each species. Based on this phylogeographic pattern, we propose that Cephalorhynchus originated in the waters of South Africa and, following the West Wind Drift, colonized New Zealand and then South America. The Chilean and Commerson's dolphins then speciated along the two coasts of South America, during the glaciation of Tierra del Fuego. Secondary radiations resulted in genetically isolated populations for both the Kerguelen Island Commerson's dolphin and the North Island Hector's dolphin. Our results suggest that coastal, depth-limited odontocetes are prone to population fragmentation, isolation and occasionally long-distance movements, perhaps following periods of climatic change.