A preliminary analysis of distribution patterns in a large, pantropical genus, Barleria L. (Acanthaceae)

Barleria L. (Acanthaceae) is a large, polymorphic, widespread genus of herbs and shrubs comprising about 300 species, occurring mainly in Africa and Asia but with one species, Barleria oenotheroides Dum.Cours., extending to the New World tropics. Recent completion of a monographic infra-generic classification of the genus (in which seven sections are recognised, and the names of four of these validated in this paper—see Appendix 1), has facilitated a comprehensive analysis of distribution patterns on a global scale. The richest representation of Barleria is in Africa where there are two centres of diversity, one in tropical East Africa (about eighty species) and the other in southern Africa (about seventy species). The number of species tails off rapidly to both the Far East and the West. Barleria shows a marked trans-Atlantic disjunction between West Africa and the Neotropics, with B. oenotheroides shared by these two regions. This type of disjunction, which is known in other genera of the family, cannot be adequately explained in Barleria on the basis of long-distance dispersal or past continental movements. There is a high degree of regional endemism (e.g. 75% for the Indian subcontinent) at both the species and sectional levels within this genus. The degree of similarity between regions is correspondingly low. The endemics in each region tend to belong to only one or a few of the sections. There are few truly widespread taxa within the genus. East and West Africa are the only regions in which all sections are represented. Sections Barleria and Prionitis C.B. Cl. are the most widespread in the genus; Sections Somalia (Oliv.) Lindau, Fissimura M. Balkwill and Stellatohirtae M. Balkwill are mainly restricted to Africa and Sections Chrysothrix M. Balkwill and Cavirostrata M. Balkwill are the most restricted, occurring mainly in India and Sri Lanka. On a local scale, many of the species show highly restricted, clumped distributions; this is apparently related to particular soil types and possibly to the short-distance, ballistic mode of seed dispersal. This account of the biogeography of Barleria is to be regarded as preliminary, as much taxonomic work at the species level remains to be done before a full-scale cladistic biogeographic account can be undertaken. Particular areas worthy of future investigation include establishing the centre of origin of the genus and investigating the basis for the high degree of endemism shown by many of the species.     

Chloroplast phylogenomic data support Eocene amphi‐Pacific early radiation for the Asian Palmate core Araliaceae

Traditional phylogenies based on analysis of multiple genes have failed to obtain a well‐resolved evolutionary history for the backbone of the Asian Palmate group of Araliaceae, the largest clade of the family. In this study, we applied the genome skimming approach of next‐generation sequencing to address whether the lack of resolution at the base of the Asian Palmate tree is due to molecular sampling error or the footprint of ancient radiation. Twenty‐nine complete plastid genomes of Araliaceae (17 newly sequenced) were analyzed (RAxML, Beast, Lagrange, and BioGeoBears) to provide the first phylogenomic reconstruction of the group (95% of genera included). As a result, the early divergences of the Asian Palmate group have been clarified but the backbone of its core is not totally resolved, with short internal branches pointing to an ancient radiation scenario. East Asia is inferred as the most likely ancestral area for the Asian Palmate group (from late Paleocene to Eocene) from which early colonization of the Neotropics is inferred during the Eocene. The radiation of the core Palmate group took place during the late Eocene, most likely in the context of the Boreotropical hypothesis. Recurrent episodes of southward migration (to the tropics) coupled with northern latitude local extinctions (promoting geographic isolation of lineages) followed by northward expansion (promoting contact of lineages that erased the trace of preceding geographic isolation) are hypothesized to have linked to the alternation of the cold and warm periods of the Eocene.     

Polyphyletic Alyssum cuneifolium (Brassicaceae) revisited: Morphological and genome size differentiation of recently recognized allopatric taxa

Alyssum cuneifolium has been recognized as a perennial alpine species growing in five isolated European mountain ranges: the Pyrenees, Western Alps, Apennines, Pirin Mts and Mt Smolikas. Recent molecular systematic studies revealed that the disjunct populations from distant mountains are not closely related and belong to five independent species: A. cacuminum (Spain, Pyrenees), A. cuneifolium (Italy, Apennines), A. flexicaule (France, Western Alps), A. pirinicum (Bulgaria, Pirin Mts), and A. spruneri (Greece, Mt Smolikas). The present study brings the thorough morphometric analysis of the segregated taxa. We found minor morphological differences between them. Whereas A. pirinicum can be clearly distinguished, the other taxa are recognizable only at the level of population means of investigated characters. The morphological similarity of these distantly related species is obviously the result of adaptation to similar high‐alpine scree habitats. It is not clear, however, whether this adaptation is environmentally controlled or whether it is also genetically fixed and whether it reflects parallel evolution towards similar morphotypes. The observed morphological patterns and their assumed correlation with environmental factors are discussed using examples from other Alyssum taxa. Three different ploidy levels have been reported for the species under study. In the present article, we examine variation in relative nuclear genome size. The Alpine and Pyrenean species have larger relative monoploid genome sizes than the Apennine and Balkan ones, probably reflecting the evolutionary history of the group. A nomenclatural account of the study species is presented, and lectotypes of A. cuneifolium and of two other names are selected.     

Contrasting phylogeographies inferred for the two alpine sister species Cardamine resedifolia and C. alpina (Brassicaceae)

Aim We use Cardamine alpina and C. resedifolia as models to address the detailed history of disjunctions in the European alpine system. These species grow on siliceous bedrock: C. alpina in the Alps and Pyrenees, and C. resedifolia in several mountain ranges from the Sierra Nevada to the Balkans. We explore differentiation among their disjunct populations as well as within the contiguous Alpine and Pyrenean ranges, and compare the phylogeographical histories of these diploid sister species. We also include samples of the closely related, arctic diploid C. bellidifolia in order to explore its origin and post‐glacial establishment. Location European alpine system, Norway and Iceland. Methods We employed amplified fragment length polymorphisms (AFLPs). AFLP data were analysed using principal coordinates analysis, neighbour joining and Bayesian clustering, and measures of diversity and differentiation were computed. Results For the snow‐bed species C. alpina (27 populations, 203 plants) we resolved two strongly divergent lineages, corresponding to the Alps and the Pyrenees. Although multiple glacial refugia were invoked in the Pyrenees, we inferred only a single one in the Maritime Alps – from which rapid post‐glacial colonization of the entire Alps occurred, accompanied by a strong founder effect. For C. resedifolia (33 populations, 247 plants), which has a broader ecological amplitude and a wider distribution, the genetic structuring was rather weak and did not correspond to the main geographical disjunctions. This species consists of two widespread and largely sympatric main genetic groups (one of them subdivided into four geographically more restricted groups), and frequent secondary contacts exist between them. Main conclusions The conspicuously different histories of these two sister species are likely to be associated with their different ecologies. The more abundant habitats available for C. resedifolia may have increased the probability of its gradual migration during colder periods and also of successful establishment after long‐distance dispersal, whereas C. alpina has been restricted by its dependence on snow‐beds. Surprisingly, the arctic C. bellidifolia formed a very divergent lineage with little variation, contradicting a scenario of recent, post‐glacial migration from the Alps or Pyrenees.     

Evolution of the eastern Asian-North American biogeographic disjunctions in ferns and lycophytes

This paper reviews 31 groups in ferns and lycophytes hypothesized to show eastern Asian–North American disjunctions. Fourteen lineages have been supported by recent phylogenetic evidence: Lycopodium nikoense and Lycopodium sitchense; Isoëtes asiatica and the clade of the North American species complex closely allied to I. maritima; Osmundastrum cinnamomeum; Osmunda claytoniana; the Adiantum pedatum complex; the Cryptogramma acrostichoides complex; Diplaziopsidaceae; Cystopteris chinensis and the Cystopteris bulbifera clade; Asplenium rhizophyllum and Asplenium ruprechtii; diploid Phegopteris; Onoclea sensibilis; the Polypodium appalachianum clade; and the Polypodium glycyrrhiza clade. Phylogenetic and/or cytological evidence did not support the biogeographic disjunctions in six cases: (1) Isoëtes asiatica and I. truncata; (2) Botrychium ternatum; (3) Thelypteris beddomei and T. nipponica—Thelypteris noveboracensis and T. nevadensis; (4) Thelypteris glanduligera and Thelypteris. japonica—T. simulata; (5) Woodwardia japonica and W. virginica; and (6) Woodwardia orientalis and Woodwardia fimbriata. Both vicariance and dispersal have been suggested to be the mechanisms for the formation of the disjunct pattern; and the Beringian region has been an active pathway for the migration of ferns and lycophytes between Asia and North America. Disjunctions of ferns and lycophytes reviewed here have been dated in the Tertiary, and are similar to the ages of eastern Asian–North American disjunctions in seed plants, supporting the close biogeographic co‐diversification of ferns and seed plants. Future studies are needed to estimate divergence times and reconstruct biogeographic events in a broad phylogenetic framework, and to test the morphological stasis hypothesis in disjunct ferns and lycophytes.     

Ploidy race distributions since the Last Glacial Maximum in the North American desert shrub, Larrea tridentata

• 1 A classic biogeographic pattern is the alignment of diploid, tetraploid and hexaploid races of creosote bush (Larrea tridentata) across the Chihuahuan, Sonoran and Mohave Deserts of western North America. We used statistically robust differences in guard cell size of modern plants and fossil leaves from packrat middens to map current and past distributions of these ploidy races since the Last Glacial Maximum (LGM).
• 2 Glacial/early Holocene (26–10 ¹⁴C kyr bp or thousands of radiocarbon years before present) populations included diploids along the lower Rio Grande of west Texas, 650 km removed from sympatric diploids and tetraploids in the lower Colorado River Basin of south‐eastern California/south‐western Arizona. Diploids migrated slowly from lower Rio Grande refugia with expansion into the northern Chihuahuan Desert sites forestalled until after ~4.0 ¹⁴C kyr bp . Tetraploids expanded from the lower Colorado River Basin into the northern limits of the Sonoran Desert in central Arizona by 6.4 ¹⁴C kyr bp . Hexaploids appeared by 8.5 ¹⁴C kyr bp in the lower Colorado River Basin, reaching their northernmost limits (~37°N) in the Mohave Desert between 5.6 and 3.9 ¹⁴C kyr bp .
• 3 Modern diploid isolates may have resulted from both vicariant and dispersal events. In central Baja California and the lower Colorado River Basin, modern diploids probably originated from relict populations near glacial refugia. Founder events in the middle and late Holocene established diploid outposts on isolated limestone outcrops in areas of central and southern Arizona dominated by tetraploid populations.
• 4 Geographic alignment of the three ploidy races along the modern gradient of increasingly drier and hotter summers is clearly a postglacial phenomenon, but evolution of both higher ploidy races must have happened before the Holocene. The exact timing and mechanism of polyploidy evolution in creosote bush remains a matter of conjecture.

     

Impact of spatial disjunction within biophysical classes on plant species composition: implications for conservation planning

The use of surrogates for biodiversity is a practical tool to improve the cost effectiveness of regional conservation planning. However, there is still much uncertainty about the biological representativeness of surrogates. Using a biophysical classification system known as the Mitchell Landscapes, we compare plant species composition in contiguous versus disjunct units of nine Landscape types and hence the ability of this surrogate to capture patterns of plant species composition. We found that plant species homogeneity was higher within a contiguous Landscape than between non‐contiguous units of the same Landscape. Overall, the dissimilarity between non‐contiguous units and their contiguous counterparts was significant (P = 0.004). Biophysical classes with very high dissimilarities between non‐contiguous units of the same region may be of limited utility.     

Is Drosera meristocaulis a pygmy sundew? Evidence of a long-distance dispersal between Western Australia and northern South America

Background and aims

South America and Oceania possess numerous floristic similarities, often confirmed by morphological and molecular data. The carnivorous Drosera meristocaulis (Droseraceae), endemic to the Neblina highlands of northern South America, was known to share morphological characters with the pygmy sundews of Drosera sect. Bryastrum, which are endemic to Australia and New Zealand. The inclusion of D. meristocaulis in a molecular phylogenetic analysis may clarify its systematic position and offer an opportunity to investigate character evolution in Droseraceae and phylogeographic patterns between South America and Oceania.

Methods Drosera meristocaulis

was included in a molecular phylogenetic analysis of Droseraceae, using nuclear internal transcribed spacer (ITS) and plastid rbcL and rps16 sequence data. Pollen of D. meristocaulis was studied using light microscopy and scanning electron microscopy techniques, and the karyotype was inferred from root tip meristem.

Key Results

The phylogenetic inferences (maximum parsimony, maximum likelihood and Bayesian approaches) substantiate with high statistical support the inclusion of sect. Meristocaulis and its single species, D. meristocaulis, within the Australian Drosera clade, sister to a group comprising species of sect. Bryastrum. A chromosome number of 2n = approx. 32–36 supports the phylogenetic position within the Australian clade. The undivided styles, conspicuous large setuous stipules, a cryptocotylar (hypogaeous) germination pattern and pollen tetrads with aperture of intermediate type 7–8 are key morphological traits shared between D. meristocaulis and pygmy sundews of sect. Bryastrum from Australia and New Zealand.

Conclusions

The multidisciplinary approach adopted in this study (using morphological, palynological, cytotaxonomic and molecular phylogenetic data) enabled us to elucidate the relationships of the thus far unplaced taxon D. meristocaulis. Long-distance dispersal between southwestern Oceania and northern South America is the most likely scenario to explain the phylogeographic pattern revealed.