Evolutionary biology: genetics and bisexuality.
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Alleles↗
Biomedical subjects
Vincent Savolainen
Publications and source records attributed to Vincent Savolainen.
The mahogany family "out-of-Africa": divergence time estimation, global biogeographic patterns inferred from plastid rbcL DNA sequences, extant, and fossil distribution of diversity.
With information on fossils and extant distribution of diversity/endemism in the mahogany family, we perform a global biogeographic study of Meliaceae using plastid rbcL data for all subfamilies, tribes and nearly all genera. Our study indicates that: (1) Meliaceae are of western Gondwanan origin; (2) dispersal played an important role for the current distribution of mahogany biota; and (3) the direction of dispersal was most likely an "out-of-Africa" scenario with important dispersal routes across Eurasia and between Eurasia and North America provided by Beringia and the North Atlantic land bridge and North America and South America via island chains and/or direct land connections. Populations in North America, Europe, and East Asia were presumably eliminated as tropical climates disappeared from these areas during the Miocene. Extensive Meliaceae fossil findings confirm that the entry of megathermal (frost-intolerant) angiosperms into southern continents from Oligocene to Pliocene must be considered as an important means of establishing pantropical distribution patterns.
Africa↗
Sympatric speciation in palms on an oceanic island.
The origin of species diversity has challenged biologists for over two centuries. Allopatric speciation, the divergence of species resulting from geographical isolation, is well documented. However, sympatric speciation, divergence without geographical isolation, is highly controversial. Claims of sympatric speciation must demonstrate species sympatry, sister relationships, reproductive isolation, and that an earlier allopatric phase is highly unlikely. Here we provide clear support for sympatric speciation in a case study of two species of palm (Arecaceae) on an oceanic island. A large dated phylogenetic tree shows that the two species of Howea, endemic to the remote Lord Howe Island, are sister taxa and diverged from each other well after the island was formed 6.9 million years ago. During fieldwork, we found a substantial disjunction in flowering time that is correlated with soil preference. In addition, a genome scan indicates that few genetic loci are more divergent between the two species than expected under neutrality, a finding consistent with models of sympatric speciation involving disruptive/divergent selection. This case study of sympatric speciation in plants provides an opportunity for refining theoretical models on the origin of species, and new impetus for exploring putative plant and animal examples on oceanic islands.
Altitude↗
Neutral theory, phylogenies, and the relationship between phenotypic change and evolutionary rates.
The neutral theory of molecular evolution predicts that rates of phenotypic change are largely independent from genotypic change. A recent study by Bromham et al. (2002) confirmed this expectation, finding no evidence for correlated phenotypic and molecular evolutionary rates in animals. We reevaluate this hypothesis, sampling at different taxonomic levels in plants and animals, using Bayesian inference to reconstruct phylogenetic trees and estimate rates of molecular evolution. We use independent contrasts in branch lengths to maximize the information extracted from each of the trees and nodal posterior probabilities to assess the influence of phylogenetic error. Our results indicate that in vascular plants between 2% and 11% of the variation in phenotypic rates of change can be explained by the rate of genotypic change. These results may be explained by the idea that processes that affect general evolutionary rates, such as body size, may also be expected to influence rates of morphological change.
Animals↗
60 million years of co-divergence in the fig-wasp symbiosis.
Figs (Ficus; ca 750 species) and fig wasps (Agaoninae) are obligate mutualists: all figs are pollinated by agaonines that feed exclusively on figs. This extraordinary symbiosis is the most extreme example of specialization in a plant-pollinator interaction and has fuelled much speculation about co-divergence. The hypothesis that pollinator specialization led to the parallel diversification of fig and pollinator lineages (co-divergence) has so far not been tested due to the lack of robust and comprehensive phylogenetic hypotheses for both partners. We produced and combined the most comprehensive molecular phylogenetic trees to date with fossil data to generate independent age estimates for fig and pollinator lineages, using both non-parametric rate smoothing and penalized likelihood dating methods. Molecular dating of ten pairs of interacting lineages provides an unparalleled example of plant-insect co-divergence over a geological time frame spanning at least 60 million years.
Animals↗
Land plants and DNA barcodes: short-term and long-term goals.
Land plants have had the reputation of being problematic for DNA barcoding for two general reasons: (i) the standard DNA regions used in algae, animals and fungi have exceedingly low levels of variability and (ii) the typically used land plant plastid phylogenetic markers (e.g. rbcL, trnL-F, etc.) appear to have too little variation. However, no one has assessed how well current phylogenetic resources might work in the context of identification (versus phylogeny reconstruction). In this paper, we make such an assessment, particularly with two of the markers commonly sequenced in land plant phylogenetic studies, plastid rbcL and internal transcribed spacers of the large subunits of nuclear ribosomal DNA (ITS), and find that both of these DNA regions perform well even though the data currently available in GenBank/EBI were not produced to be used as barcodes and BLAST searches are not an ideal tool for this purpose. These results bode well for the use of even more variable regions of plastid DNA (such as, for example, psbA-trnH) as barcodes, once they have been widely sequenced. In the short term, efforts to bring land plant barcoding up to the standards being used now in other organisms should make swift progress. There are two categories of DNA barcode users, scientists in fields other than taxonomy and taxonomists. For the former, the use of mitochondrial and plastid DNA, the two most easily assessed genomes, is at least in the short term a useful tool that permits them to get on with their studies, which depend on knowing roughly which species or species groups they are dealing with, but these same DNA regions have important drawbacks for use in taxonomic studies (i.e. studies designed to elucidate species limits). For these purposes, DNA markers from uniparentally (usually maternally) inherited genomes can only provide half of the story required to improve taxonomic standards being used in DNA barcoding. In the long term, we will need to develop more sophisticated barcoding tools, which would be multiple, low-copy nuclear markers with sufficient genetic variability and PCR-reliability; these would permit the detection of hybrids and permit researchers to identify the 'genetic gaps' that are useful in assessing species limits.
Biodiversity↗
Towards writing the encyclopedia of life: an introduction to DNA barcoding.
An international consortium of major natural history museums, herbaria and other organizations has launched an ambitious project, the 'Barcode of Life Initiative', to promote a process enabling the rapid and inexpensive identification of the estimated 10 million species on Earth. DNA barcoding is a diagnostic technique in which short DNA sequence(s) can be used for species identification. The first international scientific conference on Barcoding of Life was held at the Natural History Museum in London in February 2005, and here we review the scientific challenges discussed during this conference and in previous publications. Although still controversial, the scientific benefits of DNA barcoding include: (i) enabling species identification, including any life stage or fragment, (ii) facilitating species discoveries based on cluster analyses of gene sequences (e.g. cox1 = CO1, in animals), (iii) promoting development of handheld DNA sequencing technology that can be applied in the field for biodiversity inventories and (iv) providing insight into the diversity of life.
Conservation of Natural Resources↗
Environment, area, and diversification in the species-rich flowering plant family Iridaceae.
Phylogenetic analyses provide a means to explore evolutionary explanations for regional variation in species richness. The environment might also explain much of the previously unexplained imbalance of phylogenetic trees. We use data on geographic distribution and phylogenetic affinity to examine correlates of species richness among genera of irises (family: Iridaceae). Irises display strong phylogenetic imbalance, with a few clades containing a disproportionate number of species, most notably those found in the dry Mediterranean climate of the Cape of South Africa. The abiotic environment and area are strong predictors of iris species richness, but environment alone is insufficient to explain the high diversity of Cape clades. One possible explanation is that the interaction between biological traits and environment resulted in the unusually high diversification rates in the region.
Biodiversity↗
Environmental causes for plant biodiversity gradients.
One of the most pervasive patterns observed in biodiversity studies is the tendency for species richness to decline towards the poles. One possible explanation is that high levels of environmental energy promote higher species richness nearer the equator. Energy input may set a limit to the number of species that can coexist in an area or alternatively may influence evolutionary rates. Within flowering plants (angiosperms), families exposed to a high energy load tend to be both more species rich and possess faster evolutionary rates, although there is no evidence that one drives the other. Specific environmental effects are likely to vary among lineages, reflecting the interaction between biological traits and environmental conditions in which they are found. One example of this is demonstrated by the high species richness of the iris family (Iridaceae) in the Cape of South Africa, a likely product of biological traits associated with reproductive isolation and the steep ecological and climatic gradients of the region. Within any set of conditions some lineages will tend to be favoured over others; however, the identity of these lineages will fluctuate with a changing environment, explaining the highly labile nature of diversification rates observed among major lineages of flowering plants.
Biodiversity↗
Environmental energy and evolutionary rates in flowering plants.
The latitudinal gradient in species richness is a pervasive feature of the living world, but its underlying causes remain unclear. We evaluated the hypothesis that environmental energy drives evolutionary rates and thereby diversification in flowering plants. We estimated energy levels across angiosperm family distributions in terms of evapotranspiration, temperature and UV radiation taken from satellite and climate databases. Using the most comprehensive DNA-based phylogenetic tree for angiosperms to date, analysis of 86 sister-family comparisons shows that molecular evolutionary rates have indeed been faster in high-energy regions, but that this is not an intermediate step between energy and diversity. Energy has strong, but independent effects on both species richness and molecular evolutionary rates.
Biodiversity↗
Darwin's abominable mystery: Insights from a supertree of the angiosperms.
Angiosperms are among the major terrestrial radiations of life and a model group for studying patterns and processes of diversification. As a tool for future comparative studies, we compiled a supertree of angiosperm families from published phylogenetic studies. Sequence data from the plastid rbcL gene were used to estimate relative timing of branching events, calibrated by using robust fossil dates. The frequency of shifts in diversification rate is largely constant among time windows but with an apparent increase in diversification rates within the more recent time frames. Analyses of species numbers among families revealed that diversification rate is a labile attribute of lineages at all levels of the tree. An examination of the top 10 major shifts in diversification rates indicates they cannot easily be attributed to the action of a few key innovations but instead are consistent with a more complex process of diversification, reflecting the interactive effects of biological traits and the environment.
Biodiversity↗
Genome-scale data, angiosperm relationships, and "ending incongruence": a cautionary tale in phylogenetics.
As systematists grapple with assembling the Tree of Life, recent studies have encouraged a genomic-scale approach, obtaining DNA sequence data for entire nuclear, plastid or mitochondrial genomes for a few exemplar taxa. Some have proclaimed that this comparative genomic strategy heralds the end of incongruence in phylogeny reconstruction. Although we applaud the use of many genes to resolve phylogenetic patterns, there is a significant caveat. In spite of, or even because of, the abundant data per taxon, whole-genome sequencing for a few exemplars can provide completely resolved and strongly supported, but incorrect, evolutionary reconstructions. We provide a conspicuous example that includes Amborella, the putative sister of all other extant angiosperms, highlighting the limits of phylogenetics when whole genomes are used but taxon sampling is poor.
Codon↗
Phylogeny and evolution of basils and allies (Ocimeae, Labiatae) based on three plastid DNA regions.
A phylogeny of basils and allies (Lamiaceae, tribe Ocimeae) based on sequences of the trnL intron, trnL-trnF intergene spacer and rps 16 intron of the plastid genome is presented. Several methods were used to reconstruct phylogenies and to assess statistical support for clades: maximum parsimony with equally and successively weighted characters, bootstrap resampling, and Bayesian inference. The phylogeny is used to investigate the distribution of morphological, pericarp anatomy, chemical, and pollen characters as well as the geographical distribution of the clades. Tribe Ocimeae is monophyletic and easily diagnosable with morphological synapomorphies. There are monophyletic clades within Ocimeae that broadly correspond to currently recognised subtribes: Lavandulinae, Hyptidinae, Ociminae, and Plectranthinae. Only Lavandulinae has clear non-molecular synapomorphies. Several currently recognised genera are not monophyletic. Floral morphology consistent with sternotribic pollination is most common in Ocimeae, but there are independent departures from this model. Buzz pollination is likely in some species, the only postulated occurrence of this within Lamiaceae. Quinone diterpenoids and flavones in the leaf exudates differ in their distributions across the phylogeny and this could contribute to differences in the recorded medicinal as well as pesticidal uses of the species in the different clades. Mapping geographic distribution on to an ultrametric phylogenetic tree produced using non-parametric rate smoothing supports an Asiatic origin for Ocimeae. There are several secondary occurrences in Asia arising from the African Ociminae and Plectranthinae clades. Colonisation of Madagascar occurred at least five times, and New World colonisation occurred at least three times.
Bayes Theorem↗
Europe's fight for the tree of life.
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Animals↗
A decade of progress in plant molecular phylogenetics.
Over the past decade, botanists have produced several thousand phylogenetic analyses based on molecular data, with particular emphasis on sequencing rbcL, the plastid gene encoding the large subunit of Rubisco (ribulose bisphosphate carboxylase). Because phylogenetic trees retrieved from the three plant genomes (plastid, nuclear and mitochondrial) have been highly congruent, the "Angiosperm Phylogeny Group" has used these DNA-based phylogenetic trees to reclassify all families of flowering plants. However, in addition to taxonomy, these major phylogenetic efforts have also helped to define strategies to reconstruct the "tree of life", and have revealed the size of the ancestral plant genome, uncovered potential candidates for the ancestral flower, identified molecular living fossils, and linked the rate of neutral substitutions with species diversity. With an increased interest in DNA sequencing programmes in non-model organisms, the next decade will hopefully see these phylogenetic findings integrated into new genetic syntheses, from genomes to taxa.
DNA, Plant↗
Assessing internal support with large phylogenetic DNA matrices.
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Data Interpretation, Statistical↗
Molecular systematics, GISH and the origin of hybrid taxa in Nicotiana (Solanaceae).
Phylogenetic relationships in the genus Nicotiana were investigated using parsimony analyses of the internal transcribed spacer (ITS) regions of nuclear ribosomal DNA (nrDNA). In addition, origins of some amphidiploid taxa in Nicotiana were investigated using the techniques of genomic in situ hybridization (GISH), and the results of both sets of analyses were used to evaluate previous hypotheses about the origins of these taxa. Phylogenetic analyses of the ITS nrDNA data were performed on the entire genus (66 of 77 naturally occurring species, plus three artificial hybrids), comprising both diploid and polyploid taxa, and on the diploid taxa only (35 species) to examine the effects of amphidiploids on estimates of relationships. All taxa, regardless of ploidy, produced clean, single copies of the ITS region, even though some taxa are hybrids. Results are compared with a published plastid (matK) phylogeny using fewer, but many of the same, taxa. The patterns of relationships in Nicotiana, as seen in both analyses, are largely congruent with each other and previous evolutionary ideas based on morphology and cytology, but some important differences are apparent. None of the currently recognized subgenera of Nicotiana is monophyletic and, although most of the currently recognized sections are coherent, others are clearly polyphyletic. Relying solely upon ITS nrDNA analysis to reveal phylogenetic patterns in a complex genus such as Nicotiana is insufficient, and it is clear that conventional analysis of single data sets, such as ITS, is likely to be misleading in at least some respects about evolutionary history. ITS sequences of natural and well-documented amphidiploids are similar or identical to one of their two parents-usually, but not always, the maternal parent-and are not in any sense themselves 'hybrid'. Knowing how ITS evolves in artificial amphidiploids gives insight into what ITS analysis might reveal about naturally occurring amphidiploids of unknown origin, and it is in this perspective that analysis of ITS sequences is highly informative.
DNA, Plant↗