Reconstructing ancestral ranges in historical biogeography: properties and prospects

Recent years have witnessed a proliferation of quantitative methods for biogeographic inference. In particular, novel parametric approaches represent exciting new opportunities for the study of range evolution. Here, we review a selection of current methods for biogeographic analysis and discuss their respective properties. These methods include generalized parsimony approaches, weighted ancestral area analysis, dispersal-vicariance analysis, the dispersal--extinction--cladogenesis model and other maximum likelihood approaches, and Bayesian stochastic mapping of ancestral ranges, including a novel approach to inferring range evolution in the context of island biogeography. Some of these methods were developed specifically for problems of ancestral range reconstruction, whereas others were designed for more general problems of character state reconstruction and subsequently applied to the study of ancestral ranges. Methods for reconstructing ancestral history on a phylogenetic tree differ not only in the types of ancestral range states that are allowed, but also in the various historical events that may change the ancestral ranges. We explore how the form of allowed ancestral ranges and allowed transitions can both affect the outcome of ancestral range estimation. Finally, we mention some promising avenues for future work in the development of model-based approaches to biogeographic analysis.     

Molecular phylogeny of Soims-laubachia （Brassicaceae） s.l., based on multiple nuclear and plastid DNA sequences,and its biogeographic implications

The Hengduan Mountains region of south-west China is a noted biodiversity,hotspot, but the geographic origins and historical assembly of its rich endemic flora, including the sky-island species ofSolms-laubachia Muschl. （Brassicaceae）, have been little studied. Previous molecular studies on the phylogeny of Solms-laubachia showed it to be paraphyletic, leading to considerable expansion not only of its taxonomic limits, but also its geographic range, with the inclusion of taxa from outside the Hengduan region. However, these studies provided little resolution of interspecific relationships, preventing inferences about historical biogeography within the clade. In the present study, new sequence data from two nuclear genes （LEAFY and G3pdh） and two chloroplast intergenic spacers （petN-psbM and psbM-trnD） were combined with existing markers to increase phylogenetic signals. Phaeonychium villosum （Maxim.） Al-Shehbaz was found to be nested within Solms-laubachia s.l. In general, phylogenetic relationships appear to be a good predictor of geography, with the Hengduan Mountain endemics embedded in a paraphyletic grade of species from the western Himalayas and central Asia, but they also imply morphological homoplasy, lncongruence was detected between the nuclear and chloroplast gene trees, perhaps resulting from incomplete lineage sorting of ancestral polymorphisms. The crown age ofSolms-laubachia s.l. was estimated to be approximately 1.42-3.68 mya, using Bayesian relaxed molecular clock analysis. Historical biogeographic analysis using a parametric dispersalextinction-cladogenesis model inferred central Asia and the western Himalayas as most probable ancestral range of Solms-laubachia s.l., and estimated higher rates of eastward expansion than westward during the diversification of descendant lineages. In summary, our results suggest that Solms-laubachia s.l. originated during the Pliocene in central Asia, and subsequently migrated eastward into the Hengduan Mountains, colonizing sky-island, alpine scre     

Proteotyping：A New Approach Studying Influenza Virus Evolution at the Protein Level

Phylogenetic methods have been widely used to detect the evolution of influenza viruses.However,previous phylogenetic studies of influenza viruses do not make full use of the genetic information at the protein level and therefore cannot distinguish the subtle differences among viral genes.Proteotyping is a new approach to study influenza virus evolution.It aimed at mining the potential genetic information of the viral gene at the protein level by visualizing unique amino acid signatures(proteotypes).Neuraminidase gene fragments of some H5N1 avian influenza viruses were used as an example to illustrate how the proteotyping method worked.Bayesian analysis confirmed that the NA gene tree was mainly divided into three lineages.The NA proteotype analysis further suggested there might be multiple proteotypes within these three lineages and even within single genotypes.At the same time,some proteotypes might even involve more than one genotype.In particular,it also discovered some amino acids of viruses of some genotypes might co-reassort.All these results proved this approach could provide additional information in contrast to results from standard phylogenetic tree analysis.     

Review of plant biogeographic studies in Brazil

Molecular phylogenetic studies have become a major area of interest in plant systematics, and their impacts on historical biogeographic hypotheses are not to be disregarded. In Brazil, most historical biogeographic studies have relied on animal phylogenies, whereas plant biogeographic studies have largely lacked a phylogenetic component, having a limited utility for historical biogeography. That country, however, is of great importance for most biogeographic studies of lowland tropical South America, and it includes areas from a number of biogeographic regions of the continent. Important biogeographic reports have been published as part of phylogenetic studies, taxonomic monographs, and regional accounts for small areas or phytogeographic domains, but the available information is subsequently scattered and sometimes hard to find. In this paper we review some relevant angiosperm biogeographic studies in Brazil. Initially we briefly discuss the importance of other continents as source areas for the South American flora. Then we present a subdivision of Brazil into phytogeographic domains, and we cite studies that have explored the detection of biogeographic units （areas of endemism） and how they are historically related among those domains. Examples of plant taxa that could be used to test some biogeographic hypotheses are provided throughout, as well as taxa that exemplify several patterns of endemism and disjunction in the Brazilian angiosperm flora.     

Habitat productivity is a poor predictor of body size in rodents

The"resource availability hypothesis"predicts occurrence of larger rodents in more productive habitats.This prediction was tested in a dataset of 1,301 rodent species.We used adult body mass as a measure of body size and normalized difference vegetation index(NDVI)as a measure of habitat productivity.We utilized a cross-species approach to investigate the association between these variables.This was done at both the order level(Rodentia)and at narrower taxonomic scales.We applied phylogenetic generalized least squares(PGLS)to correct for phylogenetic relationships.The relationship between body mas and NDVI was also investigated across rodent assemblages.We controlled for spatial autocorrelation using generalized least squares(GLS)analysis.The cross-species approach found extremely low support for the resource availability hypothesis.This was reflected by a weak positive association between body mass and NDVI at the order level.We find a positive association in only a minority of rodent subtaxa.The best fit GLS model detected no significant association between body mass and NDVI across assemblages.Thus,our results do not support the view that resource availability plays a major role in explaining geographic variation in rodent body size.     

Biogeographic diversification in Nolana （Solanaceae）, a ubiquitous member of the Atacama and Peruvian Deserts along the western coast of South America

The present paper reconstructs the biogeographic diversification for Nolana L.f. （Solanaceae）, a genus of 89 endemic species largely restricted to fog-dependent desert lomas formations of coastal Peru and Chile. Previous efforts have reconstructed a phylogenetic estimate for Nolana using a combination of molecular markers. Herein, we expand on those results to examine hypotheses of biogeographic origins and diversification patterns. Nolana occupies habitats within a continuous coastal desert and forms a terrestrial archipelago of discrete ＂islands＂ unique in size, topography, and species composition. Each locality contains at least one Nolana species and many contain multiple species in sympatry. The genus has a Chilean origin, with the basal clades confined to Chile with wide geographic and ecological distributions. Peru contains two strongly supported clades, suggesting two introductions with subsequent radiation. A Chilean clade of shrubby, small-flowered species appears to have had its origins from the same ancestors of the second line that radiated in Peru and northern Chile. Nolana galapagensis is endemic to the Islas Galapagos, with origins traced to Peruvian taxa with a divergence time of 0.35 mya. Rates of diversification over the past 4.02 mya in Nolana, in one of the driest habitats on Earth, suggest rapid adaptive radiation in several clades. Success in Nolana may be attributed to characters that confer a competitive advantage in unpredictable and water-dependent environments, such as succulent leaf anatomy and ecophysiology, and the reproductive mericarp unique to Nolana. The processes affecting or shaping the biota of western South America are discussed.     

Evolution of sexual size dimorphism and its relationship with sex ratio in carabid beetles of Genus Ceroglossus Solier

Although the degree of mate competition, given extreme differences in sex ratio, explains much of the pattern of male-biased size dimorphism among diverse taxa, it fails for some species which have potential for intense male competition for mates and yet exhibit little or no sexual size dimorphism （SSD）. This fact suggest that species with low SSD should be express the effect of evolutionary pressure in non-obvious geometrical shape promoted by sex ratio in an evolutionary time scale. To evaluate this hypothesis we used phylogenetic comparative method in a Bayesian framework to investigate the evolution of SSD and the role of sex ratio at inter-specific level in the species of Ceroglossus （Coleoptera： Carabidae）. In our results the proportion farthest from 1：1 is associated with more disparate body shape, even though the entire group has minimum variation in sex ratio, which is an intrinsic life history character of this group considering its phylogenetic conservatism or phylogenetic signal. We suggest that the sex ratio has determined the dimorphism degree during evolution of this group, since both traits have increased or decreased together during the species divergence （i.e. positive phylogenetic correlation： r2=0.85）. We suggest that morphological studies of SSD will benefit from using comparative method with Bayesian approaches to assess the effect of phylogenetic history and its uncertainty. Finally, this will be allow to researchers to quantify the uncertainty of specific evolutionary hypotheses accounting for observed sexual dimorphism patterns.     

A mini-review of advances in the study of the evolution of Perdix species

The Northern Eurasia origin hypothesis is one key point in the evolutionary model. This indicates that species riginated in Northern Eurasia. Perdix, belonging to Phasianidae of Galliformes, is a valuable type of wildfowl widely distributed Northern Eurasia and contains three species. This mini-review discusses the taxonomic status, phylogeny, and origin of Perdix. Morphological and molecular evidence suggest that the three species, P. hodgsoniae, P. dauuricae and P. perdix, form a monophyletic group and they should be classified as "pheasant" rather than "quail." The current review also shows that results were affected by the methods selected for data analysis. A taxonomic category should contain as many species as possible, have a clear system history, remain evolutionarily stable, and lack adaptive radiation. Appropriate type, number and length of genes are important as molecular markers and out-group affects the bootstrap support value. P. hodgsoniae is located at the root of the phylogenetic tree and has the phenotype of ancestral Perdix. The genus Perdix may originate in the Qinghai-Tibet Plateau and expanded into high latitude areas, towards the northern plateau. This phylogenetic and evolutionary statement differs from the Eurasian origin hypothesis.     

Global comparisons of beta diversity among mammals, birds, reptiles, and amphibians across spatial scales and taxonomic ranks

Beta diversity is the change in species composition among areas in a geographic region. The proportion of species shared between two areas often decreases when the distance separating them increases, leading to an increase in beta diversity. This study compares beta diversity among four classes of terrestrial vertebrates （mammals, birds, reptiles, and amphibians） at both regional （biogeographic realm） and global extents, using the same sets of faunal sample units for all four groups in each comparison. Beta diversity is lower for the two endothermic taxa （birds and mammals） than for the two ectothermic taxa （reptiles and amphibians） in all six biogeographic realms examined. When the four taxa in the six biogeographic realms are combined, beta diversity at the species rank is higher than that of the genus rank by a factor of 1.24, and is higher than that of the family rank by a factor of 1.85. The ratio of beta diversity at the genus rank to that at the family rank is 1.50. Beta diversity is slightly higher for ecoregions of 5000-99,999 km^2 than for ecoregions of 100,000-5,000,000 km^2.     

Phylogeny and Biogeography of Thuja L. （Cupressaceae）, an Eastern Asian and North American Disjunct Genus

In order to develop better insights into biogeographic patterns of eastern Asian and North American disjunct plant genera, sequences of nuclear ribosomal DNA internal transcribed spacer (nr DNAITS) region were used to estimate interspecific relationships of Thuja L. (Cupressaceae) and infer its biogeography based on the phylogeny. According to the phylogenetic analysis, two clades were recognized. The first clade included Thuja plicata D. Don (western North America) and T. koraiensis Nakai (northeastern Asia), and the second one contained T. occidentalis (Gord.) Cart. (Japan). The ancestral area of Thuja was inferred to be eastern Asia, and two dispersal events were responsible for the modern distribution of Thuja in North America. Both the North Atlantic land bridge and Bering land bridge were possible routes for the migration of ancestral populations to North America.     

Estimating ancestral distributions of lineages with uncertain sister groups: a statistical approach to Dispersal–Vicariance Analysis and a case using Aesculus L. (Sapindaceae) including fossils

Abstract We propose a simple statistical approach for using Dispersal–Vicariance Analysis (DIVA) software to infer biogeographic histories without fully bifurcating trees. In this approach, ancestral ranges are first optimized for a sample of Bayesian trees. The probability P of an ancestral range r at a node is then calculated as where Y is a node, and F(r_Y ) is the frequency of range r among all the optimal solutions resulting from DIVA optimization at node Y, t is one of n topologies optimized, and Pt is the probability of topology t. Node Y is a hypothesized ancestor shared by a specific crown lineage and the sister of that lineage “x”, where x may vary due to phylogenetic uncertainty (polytomies and nodes with posterior probability <100%). Using this method, the ancestral distribution at Y can be estimated to provide inference of the geographic origins of the specific crown group of interest. This approach takes into account phylogenetic uncertainty as well as uncertainty from DIVA optimization. It is an extension of the previously described method called Bayes‐DIVA, which pairs Bayesian phylogenetic analysis with biogeographic analysis using DIVA. Further, we show that the probability P of an ancestral range at Y calculated using this method does not equate to pp*F(r_Y ) on the Bayesian consensus tree when both variables are <100%, where pp is the posterior probability and F(r_Y ) is the frequency of range r for the node containing the specific crown group. We tested our DIVA‐Bayes approach using Aesculus L., which has major lineages unresolved as a polytomy. We inferred the most probable geographic origins of the five traditional sections of Aesculus and of Aesculus californica Nutt. and examined range subdivisions at parental nodes of these lineages. Additionally, we used the DIVA‐Bayes data from Aesculus to quantify the effects on biogeographic inference of including two wildcard fossil taxa in phylogenetic analysis. Our analysis resolved the geographic ranges of the parental nodes of the lineages of Aesculus with moderate to high probabilities. The probabilities were greater than those estimated using the simple calculation of pp*F(r_y) at a statistically significant level for two of the six lineages. We also found that adding fossil wildcard taxa in phylogenetic analysis generally increased P for ancestral ranges including the fossil's distribution area. The ΔP was more dramatic for ranges that include the area of a wildcard fossil with a distribution area underrepresented among extant taxa. This indicates the importance of including fossils in biogeographic analysis. Exmination of range subdivision at the parental nodes revealed potential range evolution (extinction and dispersal events) along the stems of A. californica and sect. Parryana.     

Fossil biogeography: a new model to infer dispersal,extinction and sampling from palaeontological data

Methods in historical biogeography have revolutionized our ability to infer the evolution of ancestral geographical ranges from phylogenies of extant taxa, the rates of dispersals, and biotic connectivity among areas. However, extant taxa are likely to provide limited and potentially biased information about past biogeographic processes, due to extinction, asymmetrical dispersals and variable connectivity among areas. Fossil data hold considerable information about past distribution of lineages, but suffer from largely incomplete sampling. Here we present a new dispersal–extinction–sampling (DES) model, which estimates biogeographic parameters using fossil occurrences instead of phylogenetic trees. The model estimates dispersal and extinction rates while explicitly accounting for the incompleteness of the fossil record. Rates can vary between areas and through time, thus providing the opportunity to assess complex scenarios of biogeographic evolution. We implement the DES model in a Bayesian framework and demonstrate through simulations that it can accurately infer all the relevant parameters. We demonstrate the use of our model by analysing the Cenozoic fossil record of land plants and inferring dispersal and extinction rates across Eurasia and North America. Our results show that biogeographic range evolution is not a time-homogeneous process, as assumed in most phylogenetic analyses, but varies through time and between areas. In our empirical assessment, this is shown by the striking predominance of plant dispersals from Eurasia into North America during the Eocene climatic cooling, followed by a shift in the opposite direction, and finally, a balance in biotic interchange since the middle Miocene. We conclude by discussing the potential of fossil-based analyses to test biogeographic hypotheses and improve phylogenetic methods in historical biogeography.     

A comparative study in ancestral range reconstruction methods: retracing the uncertain histories of insular lineages

Island systems have long been useful models for understanding lineage diversification in a geographic context, especially pertaining to the importance of dispersal in the origin of new clades. Here we use a well-resolved phylogeny of the flowering plant genus Cyrtandra (Gesneriaceae) from the Pacific Islands to compare four methods of inferring ancestral geographic ranges in islands: two developed for character-state reconstruction that allow only single-island ranges and do not explicitly associate speciation with range evolution (Fitch parsimony [FP; parsimony-based] and stochastic mapping [SM; likelihood-based]) and two methods developed specifically for ancestral range reconstruction, in which widespread ranges (spanning islands) are integral to inferences about speciation scenarios (dispersal-vicariance analysis [DIVA; parsimony-based] and dispersal-extinction-cladogenesis [DEC; likelihood-based]). The methods yield conflicting results, which we interpret in light of their respective assumptions. FP exhibits the least power to unequivocally reconstruct ranges, likely due to a combination of having flat (uninformative) transition costs and not using branch length information. SM reconstructions generally agree with a prior hypothesis about dispersal-driven speciation across the Pacific, despite the conceptual mismatch between its character-based model and this mode of range evolution. In contrast with narrow extant ranges for species of Cyrtandra, DIVA reconstructs broad ancestral ranges at many nodes. DIVA results also conflict with geological information on island ages; we attribute these conflicts to the parsimony criterion not considering branch lengths or time, as well as vicariance being the sole means of divergence for widespread ancestors. DEC analyses incorporated geological information on island ages and allowed prior hypotheses about range size and dispersal rates to be evaluated in a likelihood framework and gave more nuanced inferences about range evolution and the geography of speciation than other methods tested. However, ancestral ranges at several nodes could not be conclusively resolved, due possibly to uncertainty in the phylogeny or the relative complexity of the underlying model. Of the methods tested, SM and DEC both converge on plausible hypotheses for area range histories in Cyrtandra, due in part to the consideration of branch lengths and/or timing of events. We suggest that DEC model-based methods for ancestral range inference could be improved by adopting a Bayesian SM approach, in which stochastic sampling of complete geographic histories could be integrated over alternative phylogenetic topologies. Likelihood-based estimates of ancestral ranges for Cyrtandra suggest a major dispersal route into the Pacific through the islands of Fiji and Samoa, motivating future biogeographic investigation of this poorly known region.     

From Africa via Europe to South America: migrational route of a species‐rich genus of Neotropical lowland rain forest trees (Guatteria,Annonaceae)

Aim Several recent studies have suggested that a substantial portion of today’s plant diversity in the Neotropics has resulted from the dispersal of taxa into that region rather than by vicariance. In general, three routes have been documented for the dispersal of taxa onto the South American continent: (1) via the North Atlantic Land Bridge, (2) via the Bering Land Bridge, or (3) from Africa directly onto the continent. Here a species‐rich genus of Neotropical lowland rain forest trees (Guatteria, Annonaceae) is used as a model to investigate these three hypotheses. Location The Neotropics. Methods The phylogenetic relationships within the long‐branch clade of Annonaceae were reconstructed (using maximum parsimony, maximum likelihood and Bayesian inference) in order to gain insight in the phylogenetic position of Guatteria. Furthermore, Bayesian molecular dating and Bayesian dispersal–vicariance (Bayes‐DIVA) analyses were undertaken. Results Most of the relationships within the long‐branch clade of Annonaceae were reconstructed and had high support. However, the relationship between the Duguetia clade, the Xylopia–Artabotrys clade and Guatteria remained unclear. The stem node age estimate of Guatteria ranged between 49.2 and 51.3 Ma, whereas the crown node age estimate ranged between 11.4 and 17.8 Ma. For the ancestral area of Guatteria and its sister group, the area North America–Africa was reconstructed in 99% of 10,000 DIVA analyses, while South America–North America was found just 1% of the time. Main conclusions The estimated stem to crown node ages of Guatteria in combination with the Bayes‐DIVA analyses imply a scenario congruent with an African origin followed by dispersal across the North Atlantic Land Bridge in the early to middle Eocene and further dispersal into North and Central America (and ultimately South America) in the Miocene. The phylogenetically and morphologically isolated position of the genus is probably due to extinction of the North American and European stem lineages in the Tertiary.     

Accounting for phylogenetic uncertainty in biogeography: a Bayesian approach to dispersal-vicariance analysis of the thrushes (Aves: Turdus)

The phylogeny of the thrushes (Aves: Turdus) has been difficult to reconstruct due to short internal branches and lack of node support for certain parts of the tree. Reconstructing the biogeographic history of this group is further complicated by the fact that current implementations of biogeographic methods, such as dispersal-vicariance analysis (DIVA; Ronquist, 1997), require a fully resolved tree. Here, we apply a Bayesian approach to dispersal-vicariance analysis that accounts for phylogenetic uncertainty and allows a more accurate analysis of the biogeographic history of lineages. Specifically, ancestral area reconstructions can be presented as marginal distributions, thus displaying the underlying topological uncertainty. Moreover, if there are multiple optimal solutions for a single node on a certain tree, integrating over the posterior distribution of trees often reveals a preference for a narrower set of solutions. We find that despite the uncertainty in tree topology, ancestral area reconstructions indicate that the Turdus clade originated in the eastern Palearctic during the Late Miocene. This was followed by an early dispersal to Africa from where a worldwide radiation took place. The uncertainty in tree topology and short branch lengths seems to indicate that this radiation took place within a limited time span during the Late Pliocene. The results support the role of Africa as a probable source area for intercontinental dispersals as suggested for other passerine groups, including basal diversification within the songbird tree.     

An evaluation of new parsimony‐based versus parametric inference methods in biogeography: a case study using the globally distributed plant family Sapindaceae

Aim Recently developed parametric methods in historical biogeography allow researchers to integrate temporal and palaeogeographical information into the reconstruction of biogeographical scenarios, thus overcoming a known bias of parsimony‐based approaches. Here, we compare a parametric method, dispersal–extinction–cladogenesis (DEC), against a parsimony‐based method, dispersal–vicariance analysis (DIVA), which does not incorporate branch lengths but accounts for phylogenetic uncertainty through a Bayesian empirical approach (Bayes‐DIVA). We analyse the benefits and limitations of each method using the cosmopolitan plant family Sapindaceae as a case study. Location World‐wide. Methods Phylogenetic relationships were estimated by Bayesian inference on a large dataset representing generic diversity within Sapindaceae. Lineage divergence times were estimated by penalized likelihood over a sample of trees from the posterior distribution of the phylogeny to account for dating uncertainty in biogeographical reconstructions. We compared biogeographical scenarios between Bayes‐DIVA and two different DEC models: one with no geological constraints and another that employed a stratified palaeogeographical model in which dispersal rates were scaled according to area connectivity across four time slices, reflecting the changing continental configuration over the last 110 million years. Results Despite differences in the underlying biogeographical model, Bayes‐DIVA and DEC inferred similar biogeographical scenarios. The main differences were: (1) in the timing of dispersal events – which in Bayes‐DIVA sometimes conflicts with palaeogeographical information, and (2) in the lower frequency of terminal dispersal events inferred by DEC. Uncertainty in divergence time estimations influenced both the inference of ancestral ranges and the decisiveness with which an area can be assigned to a node. Main conclusions By considering lineage divergence times, the DEC method gives more accurate reconstructions that are in agreement with palaeogeographical evidence. In contrast, Bayes‐DIVA showed the highest decisiveness in unequivocally reconstructing ancestral ranges, probably reflecting its ability to integrate phylogenetic uncertainty. Care should be taken in defining the palaeogeographical model in DEC because of the possibility of overestimating the frequency of extinction events, or of inferring ancestral ranges that are outside the extant species ranges, owing to dispersal constraints enforced by the model. The wide‐spanning spatial and temporal model proposed here could prove useful for testing large‐scale biogeographical patterns in plants.     

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.     

Biogeography of ‘tropical Anagallis’ (Myrsinaceae) inferred from nuclear and plastid DNA sequence data

Aim ‘Tropical Anagallis’ corresponds to one of two evolutionary lineages within the genus Anagallis L. Generally, species within this lineage have a limited distribution in (sub‐)tropical regions in Africa or Madagascar. Two species, however, are endemic to South America, and exhibit a trans‐Atlantic disjunction with the rest of the species within the lineage. To investigate this disjunct distribution, as well as other dispersal events, the distribution of extant taxa was used to hypothesize the ancestral area(s) of distribution. Location Africa, Madagascar, Europe and South America. Methods Dispersal–vicariance analysis (DIVA) was used to optimize distribution areas onto parsimony and Bayesian phylogenies based on sequence data from four chloroplast loci and the nuclear internal transcribed spacers (ITS). Results Parsimony analysis gave one most parsimonious tree while Bayesian analysis resulted in a collapsed node due to alternative placements of Anagallis nummularifolia Baker, endemic to Madagascar. Optimization of the present distribution using DIVA, and the most parsimonious tree and six alternative topologies of the Bayesian analysis, show an origin of the lineage in Europe as most likely, although one topology indicates a broader ancestral distribution area. Dispersal to Africa appears to have been a single event, while two parallel dispersal events seem to have resulted in the American as well as Madagascan distributions. Main conclusions The lineage ‘tropical Anagallis’ evolved in Europe and may have been present in the Eocene boreotropical forests, although scarcity of fossils makes assessment of age difficult. Dispersal to South America is proposed to have been via the North Atlantic land bridge, or, more likely, through transport by the North Equatorial Current. Dispersal from Europe to Africa represents a single event, while dispersal to Madagascar from mainland Africa has occurred twice.     

Avian higher level biogeography: Southern Hemispheric origins or Southern Hemispheric relicts?

A recent analysis of a comprehensive phylogenetic data set suggested Southern Hemispheric origins for various higher level taxa of neornithine (crown group) birds. These results contrast with hypotheses derived from the fossil record, with the occurrence of successively branching stem group representatives of many “Southern Hemispheric” bird groups in the early Cenozoic of Europe and North America suggesting relict extant distributions. However, a selective consideration of the fossil record and the merging of extant ranges may have resulted in a skewed picture of the past biogeographical history of birds. For future studies, it is proposed that multiple taxa of successively branching Northern Hemispheric stem group representatives are considered to narrow down the ancestral areas of the crown group representatives.