A tale of two continents: Baker's rule and the maintenance of self-incompatibility in Lycium (Solanaceae)

Over 50 years ago, Baker (1955, 1967) suggested that self-compatible species were more likely than self-incompatible species to establish new populations on oceanic islands. His logic was straightforward and rested on the assumption that colonization was infrequent; thus, mate limitation favored the establishment of self-fertilizing individuals. In support of Baker's rule, many authors have documented high frequencies of self-compatibility on islands, and recent work has solidified the generality of Baker's ideas. The genus Lycium (Solanaceae) has ca. 80 species distributed worldwide, and phylogenetic studies suggest that Lycium originated in South America and dispersed to the Old World a single time. Previous analyses of the S-RNase gene, which controls the stylar component of self-incompatibility, have shown that gametophytically controlled self-incompatibility is ancestral within the genus, making Lycium a good model for investigating Baker's assertions concerning reproductive assurance following oceanic dispersal. Lycium is also useful for investigations of reproductive evolution, given that species vary both in sexual expression and the presence of self-incompatibility. A model for the evolution of gender dimorphism suggests that polyploidy breaks down self-incompatibility, leading to the evolution of gender dimorphism, which arises as an alternative outcrossing mechanism. There is a perfect association of dimorphic gender expression, polyploidy, and self-compatibility (vs. cosexuality, diploidy, and self-incompatibility) among North American Lycium. Although the association between ploidy level and gender expression also holds for African Lycium, to date no studies of mating systems have been initiated in Old World species. Here, using controlled pollinations, we document strong self-incompatibility in two cosexual, diploid species of African Lycium. Further, we sequence the S-RNase gene in 15 individuals from five cosexual, diploid species of African Lycium and recover 24 putative alleles. Genealogical analyses indicate reduced trans-generic diversity of S-RNases in the Old World compared to the New World. We suggest that genetic diversity at this locus was reduced as a result of a founder event, but, despite the bottleneck, self-incompatibility was maintained in the Old World. Maximum-likelihood analyses of codon substitution patterns indicate that positive Darwinian selection has been relatively strong in the Old World, suggesting the rediversification of S-RNases following a bottleneck. The present data thus provide a dramatic exception to Baker's rule, in addition to supporting a key assumption of the Miller and Venable (2000) model, namely that self-incompatibility is associated with diploidy and cosexuality.     

Association of ploidy and sexual system in Lycium californicum (Solanaceae)

In North American Lycium (Solanaceae), the evolution of gender dimorphism has been proposed as a means of restoring outcrossing after polyploidization causes the loss of self-incompatibility. Previous studies of this process in Lycium focused on comparisons between species that differ in ploidy. We examined intraspecific variation in floral morphology and DNA content in populations of L. californicum to determine correlations between sexual system and cytotype. We also used nuclear ITS and GBSSI sequence data to determine whether diploid and polyploid forms represent the same phylogenetic species, and the phylogeographic relationships among populations and ploidy levels. Within populations, no variation in ploidy was found, although among populations there was a perfect correspondence between sexual system and cytotype. Diploid populations were all hermaphroditic, whereas tetraploid populations were all gender dimorphic. There was no clear geographic pattern to the occurrence of diploid and tetraploid forms. Phylogenetic analysis confirms that L. californicum, regardless of ploidy, forms a monophyletic group within the genus Lycium. Sequences from diploid and polyploid individuals did not form reciprocally monophyletic clades, indicating either multiple gains of polyploidy, ongoing gene flow between cytotypes, or lack of lineage sorting since the evolution of polyploidy. The correspondence between ploidy and sex expression is consistent with the hypothesis that polyploidization triggers the evolution of gender dimorphism in this and other Lycium species.     

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.     

Drift in the tropics: Phylogenetics and biogeographical patterns in Combretaceae

Aim

The aim of this study was to further advance our understanding of the species-rich, and ecologically important angiosperm family Combretaceae to provide new insights into their evolutionary history. We assessed phylogenetic relationships in the family using target capture data and produced a dated phylogenetic tree to assess fruit dispersal modes and patterns of distribution.

Location

Tropical and subtropical regions.

Time Period

Cretaceous to present.

Major Taxa Studied

Family Combretaceae is a member of the rosid clade and comprises 10 genera and more than 500 species, predominantly assigned to genera Combretum and Terminalia, and occurring on all continents and in a wide range of ecosystems.

Methods

We use a target capture approach and the Angiosperms353 universal probes to reconstruct a robust dated phylogenetic tree for the family. This phylogenetic framework, combined with seed dispersal traits, biome data and biogeographic ranges, allows the reconstruction of the biogeographical history of the group.

Results

Ancestral range reconstructions suggest a Gondwanan origin (Africa/South America), with several intercontinental dispersals within the family and few transitions between biomes. Relative abundance of fruit dispersal types differed by both continent and biome. However, intercontinental colonizations were only significantly enhanced by water dispersal (drift fruit), and there was no evidence that seed dispersal modes influenced biome shifts.

Main Conclusions

Our analysis reveals a paradox as drift fruit greatly enhanced dispersal distances at intercontinental scale but did not affect the strong biome conservatism observed.     

Evolution of fruit types and seed dispersal:A phylogenetic and ecological snapshot

Success of flowering plants is greatly dependent on effective seed dispersal. Specific fruit types aid different mechanisms of seed dispersal. However, little is known about what evolutionary forces have driven the diversification of fruit types and whether there were phylogenetic constraints on fruit evolution among angio-sperm lineages. To address these questions, we first surveyed the orders and families of angiosperms for fruit types and found no clear association between fruit types and major angiosperm lineages, suggesting there was little phylogenetic constraint on fruit evolution at this level. We then surveyed fruit types found in two contrasting habitats: an open habitat including the Indian desert and North American plains and prairies, and a closed forest habitat of Australian tropical forest. The majority of genera in the survey of tropical forests in Australia were fleshy fruit trees, whereas the majority of genera in the survey of prairies and plains in central North America were herbs with capsules and achenes. Both capsules and achenes are frequently dispersed by wind in the open, arid habitat, whereas fleshy fruits are generally dispersed by animals. Since desert and plains tend to provide continuous wind to aid dispersal and there are more abundant mammal and bird dispersers in the closed forest, this survey suggests that fruit evolution was driven at least in part by dispersal agents abundant in particular habitats.     

果实类型和种子传播的进化:系统发育和生态学简论

Success of flowering plants is greatly dependent on effective seed dispersal. Specific fruit types aid different mechanisms of seed dispersal. However, little is known about what evolutionary forces have driven the diversification of fruit types and whether there were phylogenetic constraints on fruit evolution among angiosperm lineages. To address these questions, we first surveyed the orders and families of angiosperms for fruit types and found no clear association between fruit types and major angiosperm lineages, suggesting there was little phylogenetic constraint on fruit evolution at this level. We then surveyed fruit types found in two contrasting habitats an open habitat including the Indian desert and North American plains and prairies, and a closed forest habitat of Australian tropical forest. The majority of genera in the survey of tropical forests in Australia were fleshy fruit trees, whereas the majority of genera in the survey of prairies and plains in central North America were herbs with capsules and achenes. Both capsules and achenes are frequently dispersed by wind in the open, arid habitat, whereas fleshy fruits are generally dispersed by animals. Since desert and plains tend to provide continuous wind to aid dispersal and there are more abundant mammal and bird dispersers in the closed forest, this survey suggests that fruit evolution was driven at least in part by dispersal agents abundant in particular habitats.     

Historical biogeography of amphibian parasites, genus Polystoma (Monogenea: Polystomatidae)

Aim  The present-day geographical distribution of parasites with a direct biological life cycle is guided mostly by the past dispersal and vicariance events that have affected their hosts. The Amphibia– Polystoma association (which satisfies these criteria) also exhibits original traits, such as host specificity and world-wide distribution. This biological model was thus chosen to investigate the common historical biogeography of its widespread representatives.
Location  North and South America, Eurasia and Africa.
Methods  We investigated the phylogeny of 12 species of neobatrachian parasites sampled from North and South America, Eurasia and Africa. Hosts belonged mostly to hyloids and ranoids of families Bufonidae, Hylidae, Leptodactylidae, Ranidae and Hyperoliidae. Phylogenetic reconstructions were inferred from maximum likelihood and maximum parsimony analyses from complete ITS1 sequences.
Results  The group of American species appeared paraphyletic with one species at the base of a Eurafrican clade, within which two lineages were seen: one composed of only Eurasian species, and the other of European and African species, with the two European species basal to an African clade.
Main conclusions  The route of Polystoma evolution is deduced from the phylogenetic tree and discussed in the light of host evolution. We conclude that Polystoma originated in South America on hyloids, after the separation of South America from Africa. The genus must have colonized North America in Palaeocene times and Eurasia by the mid-Cainozoic, taking advantage of the dispersal of either ancestral bufonids or hylids. Africa, however, appears to have been colonized more recently, during the Messinian period.     

Phylogeography of the Microcoleus vaginatus (Cyanobacteria) from three continents--a spatial and temporal characterization

It has long been assumed that cyanobacteria have, as with other free-living microorganisms, a ubiquitous occurrence. Neither the geographical dispersal barriers nor allopatric speciation has been taken into account. We endeavoured to examine the spatial and temporal patterns of global distribution within populations of the cyanobacterium Microcoleus vaginatus, originated from three continents, and to evaluate the role of dispersal barriers in the evolution of free-living cyanobacteria. Complex phylogeographical approach was applied to assess the dispersal and evolutionary patterns in the cyanobacterium Microcoleus vaginatus (Oscillatoriales). We compared the 16S rRNA and 16S-23S ITS sequences of strains which had originated from three continents (North America, Europe, and Asia). The spatial distribution was investigated using a phylogenetic tree, network, as well as principal coordinate analysis (PCoA). A temporal characterization was inferred using molecular clocks, calibrated from fossil DNA. Data analysis revealed broad genetic diversity within M. vaginatus. Based on the phylogenetic tree, network, and PCoA analysis, the strains isolated in Europe were spatially separated from those which originated from Asia and North America. A chronogram showed a temporal limitation of dispersal barriers on the continental scale. Dispersal barriers and allopatric speciation had an important role in the evolution of M. vaginatus. However, these dispersal barriers did not have a permanent character; therefore, the genetic flow among populations on a continental scale was only temporarily present. Furthermore, M. vaginatus is a recently evolved species, which has been going through substantial evolutionary changes.     

Evolution of the Madrean–Tethyan disjunctions and the North and South American amphitropical disjunctions in plants

Abstract The present paper reviews advances in the study of two major intercontinental disjunct biogeographic patterns: (i) between Eurasian and western North American deserts with the Mediterranean climate (the Madrean–Tethyan disjunctions); and (ii) between the temperate regions of North and South America (the amphitropical disjunctions). Both disjunct patterns have multiple times of origin. The amphitropical disjunctions have largely resulted from long‐distance dispersal, primarily from the Miocene to the Holocene, with available data indicating that most lineages dispersed from North to South America. Results of recent studies on the Mediterranean disjuncts between the deserts of Eurasia and western North America support the multiple modes of origin and are mostly consistent with hypotheses of long‐distance dispersal and the North Atlantic migration. Axelrod's Madrean–Tethyan hypothesis, which implies vicariance between the two regions in the early Tertiary, has been favored by a few studies. The Beringian migration corridor for semiarid taxa is also supported in some cases.     

Evolution of the Madrean-Tethyan disjunctions and the North and South American amphitropical disjunctions in plants

The present paper reviews advances in the study of two major intercontinental disjunct biogeographic patterns： （i） between Eurasian and western North American deserts with the Mediterranean climate （the Madrean- Tethyan disjunctions）; and （ii） between the temperate regions of North and South America （the amphitropical disjunctions）. Both disjunct patterns have multiple times of origin. The amphitropical disjunctions have largely resulted from long-distance dispersal, primarily from the Miocene to the Holocene, with available data indicating that most lineages dispersed from North to South America. Results of recent studies on the Mediterranean disjuncts between the deserts of Eurasia and western North America support the multiple modes of origin and are mostly consistent with hypotheses of long-distance dispersal and the North Atlantic migration. Axelrod＇s Madrean-Tethyan hypothesis, which implies vicariance between the two regions in the early Tertiary, has been favored by a few studies. The Beringian migration corridor for semiarid taxa is also supported in some cases.     

A tale of North and South America: time and mode of dispersal of the amphitropical genus Munroa (Poaceae,Chloridoideae)

Plant disjunctions have provided some of the most intriguing distribution patterns historically addressed by biogeographers. We evaluated the three hypotheses that have been postulated to explain these patterns [vicariance, stepping‐stone dispersal and long‐distance dispersal (LDD)] using Munroa, an American genus of grasses with six species and a disjunct distribution between the desert regions of North and South America. The ages of clades, cytology, ancestral characters and areas of distribution were investigated in order to establish relationships among species, to determine the time of divergence of the genus and its main lineages, and to understand further the biogeographical and evolutionary history of this genus. Bayesian inference recovered the North American M. pulchella as sister species to the rest. Molecular dating and ancestral area analyses suggest that Munroa originated in North America in the late Miocene–Pliocene (7.2 Mya; 8.2–6.5 Mya). Based on these results, we postulate that two dispersal events modelled the current distribution patterns of Munroa: the first from North to South America (7.2 Mya; 8.2–6.5 Mya) and the second (1.8 Mya; 2–0.8 Mya) from South to North America. Arid conditions of the late Miocene–Pliocene in the Neogene and Quaternary climatic oscillations in North America and South America were probably advantageous for the establishment of populations of Munroa. We did not find any relationship between ploidy and dispersal events, and our ancestral character analyses suggest that shifts associated with dispersal and seedling establishment, such as habit, reproductive system, disarticulation of rachilla, and shape and texture of the glume, have been important in these species reaching new areas. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 110–125.     

Phylogenetic relationships and evolution in Chrysosplenium (Saxifragaceae) based on matK sequence data

Chrysosplenium (Saxifragaceae) consists of 57 species widely distributed in temperate and arctic regions of the Northern Hemisphere, with two species restricted to the southern part of South America. Species relationships within the genus are highly problematic. The genus has traditionally been divided into two groups, sometimes recognized as sections (Oppositifolia and Alternifolia), based on leaf arrangement, or, alternatively, into 17 series. Based on morphological features, Hara suggested that the genus originated in South America and then subsequently migrated to the Northern Hemisphere. We conducted phylogenetic analyses of DNA sequences of the chloroplast gene matK for species of Chrysosplenium to elucidate relationships, test Hara's biogeographic hypothesis for the genus, and examine chromosomal and gynoecial diversification. These analyses revealed that both sections Oppositifolia and Alternifolia are monophyletic and form two large sister clades. Hence, leaf arrangement is a good indicator of relationships within this genus. Hara's series Pilosa and Macrostemon are each also monophyletic; however, series Oppositifolia, Alternifolia, and Nepalensia are clearly not monophyletic. MacClade reconstructions suggest that the genus arose in Eastern Asia, rather than in South America, with several independent migration events from Asia to the New World. In one well-defined subclade, species from eastern and western North America form a discrete clade, with Old World species as their sister group, suggesting that the eastern and western North American taxa diverged following migration to that continent. The South American species forms a clade with species from eastern Asia; this disjunction may be the result of ancient long-distance dispersal. Character mapping demonstrated that gynoecial diversification is dynamic, with reversals from inferior to half-inferior ovaries, as well as to ovaries that appear superior. Chromosomal evolution also appears to be labile with several independent origins of n = 12 (from an original number of n = 11) and multiple episodes of aneuploidy.     

West Wind Drift revisited: testing for directional dispersal in the Southern Hemisphere using event-based tree fitting

Aim Recent studies suggest that if constrained by prevailing wind or ocean currents dispersal may produce predictable, repeated distribution patterns. Dispersal mediated by the West Wind Drift (WWD) and Antarctic Circumpolar Current (AAC) has often been invoked to explain the floristic similarities of Australia, South America and New Zealand. If these systems have been important dispersal vectors then eastward dispersal – from Australia to New Zealand and the western Pacific to South America – is expected to predominate. We investigate whether phylogenies for Southern Hemisphere plant groups provide evidence of historical dispersal asymmetry and more specifically whether inferred asymmetries are consistent with the direction of the WWD/AAC. Location Southern Hemisphere. Methods We assembled a data set of 23 published phylogenies for plant groups that occur in New Zealand, Australia and/or South America. We used parsimony‐based tree fitting to infer the number and direction of dispersals within each group. Observed dispersal asymmetries were tested for significance against a distribution of expected values. Results Our analyses suggest that dispersal has played a major role in establishing present distributions and that there are significant patterns of asymmetry in Southern Hemisphere dispersal. Consistent with the eastward direction of the WWD/ACC, dispersal from Australia to New Zealand was inferred significantly more often than in the reverse direction. No significant patterns of dispersal asymmetry were found between the western Pacific landmasses and South America. However, eastward dispersal was more frequently inferred between Australia and South America, while for New Zealand–South American events westward dispersal was more common. Main conclusions Our results suggest that eastward circumpolar currents have constrained the dispersal of plants between Australia and New Zealand. However, the WWD/ACC appear to have had less of an influence on dispersal between the western Pacific landmasses and South America. This observation may suggest that differences in dispersal mechanism are important – direct wind or water dispersal vs. stepping‐stone dispersal along the Antarctic coast. While our analyses provide useful preliminary insights into dispersal asymmetry in the Southern Hemisphere we will need larger data sets and additional methodological advances in order to test fully these dispersal patterns and infer processes from phylogenetic data.     

Pollen grain morphology of Nolana L. (Solanaceae: Nolanoideae: Nolaneae) and related genera of southern South American Solanaceae

Nolana comprises 89 species almost entirely restricted to the Atacama and Peruvian deserts. These species are adapted to the arid lomas environment of northern Chile and southern Peru, where they depend on marine fog locally known as garúa or camanchaca for their development. Recent molecular phylogenetic studies provide support for the placement of Nolana in the Solanaceae rather than Nolanaceae. Additionally, a sister relationship with the tribe Lycieae has been suggested. The pollen morphology of 111 species from 18 genera of southern South American Solanaceae was investigated under light and scanning electron microscopes in order to establish the palynological affinities of Nolana within the Solanaceae and to investigate if pollen morphology is congruent with the phylogenetic relationships and former taxonomic classification of the species of Nolana. The following genera were studied: Nolana, Cestrum, Combera, Dunalia, Fabiana, Jaborosa, Latua, Lycium (incl. Grabowskia, Phrodus), Nicandra, Nicotiana, Nierenbergia, Petunia, Physalis, Reyesia, Salpiglossis, Schizanthus, Solanum and Vestia. Two pollen types that differ in exine structure were distinguished within Nolana; these pollen types are congruent with recent phylogenetic studies, thus, we propose to reinstate Nolana sect. Sorema.     

The biogeography of the austral, subalpine genus Ourisia (Plantaginaceae) based on molecular phylogenetic evidence: South American origin and dispersal to New Zealand and Tasmania

Molecular phylogenetic analyses of 26 of the 28 species of Ourisia , including eight of ten subspecies and two purported natural hybrids, are presented and used to examine the biogeography of the genus, which is distributed in subalpine to alpine habitats of South America, New Zealand and Tasmania. Gondwanan vicariance, often cited as the cause of this classic austral biogeographical pattern, was rejected by parametric bootstrapping of our combined dataset. Alternatively, various lines of evidence are presented in favour of a South American origin of Ourisia and subsequent dispersal to Australasia. Specifically, the genus likely arose in the Andes of central Chile and spread to southern Chile and Argentina, to the north-central Andes, and finally to Tasmania and New Zealand. The ancestor of the New Zealand species probably first arrived on the South Island, where the New Zealand species of Ourisia are most diverse, and migrated to the North and Stewart Islands. Because the Tasmanian and New Zealand species are sister to one another, the direction of dispersal between these two areas is equivocal. These results agree with other molecular phylogenetic studies that show that past dispersal between southern hemisphere continents has played an important role in the evolutionary history of many high-elevation austral plants. Our data also show that within South America, many of the geographical barriers (with the exception of the Atacama Desert) that have played a role in the evolution of other plant groups have not affected Ourisia species. Within New Zealand, the phylogeny and biogeography of species of Ourisia coincide with the geological history of the country and patterns of other alpine plants. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 479–513.     

Convergence, divergence, and homogenization in the ecological structure of emydid turtle communities: the effects of phylogeny and dispersal

Studies that have explored the origins of patterns of community structure from a phylogenetic perspective have generally found either convergence (similarity) in community structure between regions through adaptive evolution or lack of convergence (dissimilarity) due to phylogenetic conservatism in the divergent ecological characteristics of lineages inhabiting different regions. We used a phylogenetic approach to document a third pattern in the structure of emydid turtle communities. Emydid communities in southeastern North America tend to have a higher proportion of aquatic species than those in the northeast. This pattern reflects phylogenetic conservatism in the ecology and biogeography of two basal emydid clades, limiting convergence in community structure between these regions. However, differences in community structure between northeastern and southeastern North America have also been homogenized considerably by the dispersal of species with phylogenetically conserved ecological characteristics between regions. This pattern of ecologically conservative dispersal may be important in many continental and oceanic systems.     

The evolutionary origin of a second radiation of annual Castilleja (Orobanchaceae) species in South America: The role of long distance dispersal and allopolyploidy

Considerable attention has been directed toward understanding the wide gaps in range that are common among many groups of closely related organisms. By placing their biology and geography in a phylogenetic context, we may gain a broader knowledge of the series of historical events that have led to present species distributions. In addition to the North American annuals, a second radiation of annual Castilleja species is in Andean Peru and central Chile. Phylogenetic analyses of chloroplast and nuclear DNA regions revealed a complex history for the origin and diversification of annual Castilleja species in South America. In addition to at least three independent long-distance dispersal events from North America, allopolyploidy has played a significant role in this disjunct radiation. Only C. attenuata occurs in both California and South America, and these results support its recent arrival to central Chile. Two Peruvian species are inferred to be allopolyploids; hybridization between annual lineages derived from independent long-distance dispersal events from North America gave rise to C. profunda, and hybridization between South American annual and perennial species gave rise to C. cerroana. The relative importance these events are discussed with reference to the observed morphological, ecological, and distributional patterns.     

The evolution of body size in extant groups of North American freshwater fishes: speciation,size distributions,and Cope's rule

Change in body size within an evolutionary lineage over time has been under investigation since the synthesis of Cope's rule, which suggested that there is a tendency for mammals to evolve larger body size. Data from the fossil record have subsequently been examined for several other taxonomic groups to determine whether they also displayed an evolutionary increase in body size. However, we are not aware of any species-level study that has investigated the evolution of body size within an extant continental group. Data acquired from the fossil record and data derived from the evolutionary relationships of extant species are not similar, with each set exhibiting both strengths and weaknesses related to inferring evolutionary patterns. Consequently, expectation that general trends exhibited in the fossil record will correspond to patterns in extant groups is not necessarily warranted. Using phylogenetic relationships of extant species, we show that five of nine families of North American freshwater fishes exhibit an evolutionary trend of decreasing body size. These trends result from the basal position of large species and the more derived position of small species within families. Such trends may be caused by the invasion of small streams and subsequent isolation and speciation. This pattern, potentially influenced by size-biased dispersal rates and the high percentage of small streams in North America, suggests a scenario that could result in the generation of the size-frequency distribution of North American freshwater fishes.     

BIOGEOGRAPHY OF MARINE RED ALGAE: MYTHS AND REALITIES

Hommersand, M. H. Department of Biology, Coker Hall, University of North Carolina, Chapel Hill, NC 27599-3280 USA Theories about the geographical distribution of marine algae fall roughly into two categories: (1) a concept of biogeographical regions in which algal distribution is determined primarily by growth, reproductive and lethal temperature boundaries (Setchell, van den Hoek, Breeman, Lüning) and (2) an historical perspective in which distribution is determined primarily by patterns of dispersal and the establishment of barriers to dispersal (vicariance biogeography) (Svedelius, Garbary, Lindstrom, Hommersand). Setchell proposed the 5° isotherm rule in 1920, and in 1924 Svedelius advocated a worldwide distribution for tropical and subtropical groups followed by discontinuous distribution upon closure of the connection between the Indian Ocean and Mediterranean Sea and, later, between North and South America (Wegener's theory). Transarctic dispersal routes have received special attention in recent years (Lindstrom, Lüning, van Oppen, Olsen, Stam), as have special relationships between Australasia, South Africa and South America (Hommersand). Less well understood are the climatic changes that have taken place in the Cenozoic which are strategic to an understanding vicariant biogeography. The advent of molecular methods combined with the tools of phylogenetic systematics now make it possible to identify ancestral taxa, test the consistency of tree topologies, and calculate mean branch lengths between sister lineages diverging from an interior node of a tree. With such methods it may be possible to infer ancestral areas, identify dispersal pathways, determine the chronology of isolating events, assess the impact of multiple invasions, and generally relate dispersal and vicariance models to phylogenetic hypotheses for red, brown and green algal taxa.     

Amphisbaenian paleobiogeography: Evidence of vicariance and geodispersal patterns

Paleobiogeographic patterns within the Amphisbaenia were evaluated using the modified Brooks Parsimony Analysis (BPA) and recently published morphological and molecular phylogenies. Extant amphisbaenians are present in Africa, South America, North America, Europe, and the Middle East. The modified BPA was used to determine the relative effects of Pangean breakup, sea-level change, and climate change on evolutionary and distributional patterns within the Amphisbaenia. The modified BPA also tested the biogeographic effect of the Rhineuridae's phylogenetic position as either most basal in the morphologic phylogeny or most derived in the molecular phylogeny. The morphological and molecular analyses resulted in two different biogeographic hypotheses. The morphological analysis indicated three major biogeographic regions for the Amphisbaenia: 1) Africa, South America, and the Caribbean, 2) western Asia, and 3) North America. The molecular analysis indicated two major biogeographic regions: 1) Africa, western Asia, and North America, and 2) South America. The morphological biogeographic pattern corresponds with the known timing of the breakup of Pangea and the resulting paleogeographic reconstructions of the Mesozoic and Early Cenozoic. While the molecular pattern is similar to patterns recovered from dinosaurian biogeographic studies, the closer connection of Africa with North America rather than South America does not match well-constrained geologic evidence for the sequence of Pangean breakup. Both paleobiogeographic analyses, however, resulted in congruent patterns of speciation through vicariance and geodispersal. This suggests that in addition to the breakup of Pangea, such cyclical Earth history processes as sea-level and climate changes played an important role in the biogeographic patterns of the Amphisbaenia.