Environmental distribution of four Carex species (Cyperaceae) in an old-growth forest

We conducted an in-depth characterization of the range of micro-environments (1 m) in which four Carex species (C. backii, C. communis, C. plantaginea, and C. platyphylla) grow in the understory of an old-growth, deciduous forest in southern Québec, Canada. All four species occurred in significantly different micro-environments. Carex plantaginea was found at the wet end of a moisture gradient, in soils with high nitrate availability. Carex backii and C. platyphylla were found at the dry end of the moisture gradient, with C. backii occupying soils with higher phosphorus availability than C. platyphylla. Carex communis, the only ant-dispersed species studied, was found in the broadest range of environmental conditions. Our results suggest that environmental heterogeneity and interspecific microhabitat preferences are important for the maintenance of local species diversity in the forest understory, not only for common species as demonstrated in previous studies, but for infrequent species, and those within a functional group (upland Carex species). However, there was some evidence that the distributions of C. backii and C. communis were not in equilibrium with current environmental conditions, indicating that historical factors, such as dispersal and colonization events, may also have important effects on the distributions of these species and the maintenance of species diversity in old-growth forest.     

Diversification rates and chromosome evolution in the most diverse angiosperm genus of the temperate zone (Carex, Cyperaceae)

The sedge family (Cyperaceae: Poales; ca. 5600 spp.) is a hyperdiverse cosmopolitan group with centres of species diversity in Africa, Australia, eastern Asia, North America, and the Neotropics. Carex, with ca. 40% of the species in the family, is one of the most species-rich angiosperm genera and the most diverse in temperate regions of the Northern Hemisphere, making it atypical among plants in that it inverts the latitudinal gradient of species richness. Moreover, Carex exhibits high rates of chromosome rearrangement via fission, fusion, and translocation, which distinguishes it from the rest of the Cyperaceae. Here, we use a phylogenetic framework to examine how the onset of contemporary temperate climates and the processes of chromosome evolution have influenced the diversification dynamics of Carex. We provide estimates of diversification rates and map chromosome transitions across the evolutionary history of the main four clades of Carex. We demonstrate that Carex underwent a shift in diversification rates sometime between the Late Eocene and the Oligocene, during a global cooling period, which fits with a transition in diploid chromosome number. We suggest that adaptive radiation to novel temperate climates, aided by a shift in the mode of chromosome evolution, may explain the large-scale radiation of Carex and its latitudinal pattern of species richness.     

Isolation of 11 polymorphic tri- and tetranucleotide microsatellite loci in a North American sedge (Carex scoparia: Cyperaceae) and cross-species amplification in three additional Carex species

We report on the isolation and evaluation of 11 microsatellites from a widespread eastern North American wetland sedge, Carex scoparia. Loci exhibit 3-9 alleles over five populations and significant F(IS) (0.204-0.717) in most populations. All primers cross-amplify in at least two other species, and 10 cross-amplify in the more distantly related C. stipata. These markers will be used to examine population genetics and patterns of chromosomal diversification in this ecologically important sedge species and its relatives.     

Germination and establishment of forest sedges (Carex, Cyperaceae): tests for home-site advantage and effects of leaf litter

We investigated aspects of germination and seedling establishment that might influence the distribution and diversity of Carex species growing in the forest understory. We tested the ability of Carex communis and C. platyphylla to germinate and establish at plots where adult individuals of one of these two species, or one of two other Carex species (C. backii and C. plantaginea), occurred in an old-growth forest in southern Québec, Canada. We also tested for effects of leaf litter on germination and establishment of these sedges. From a series of experiments in the field and in a lath house, we found no evidence of home-site advantage with respect to germination or seedling establishment. Leaf litter had a negative effect on germination and establishment. The results emphasize the importance of dispersal and colonization events in determining local diversity and distribution of Carex species in upland forests. High frequency of occurrence of C. communis at our study site may result from relatively wide dispersal provided by ants, and a suite of traits associated with ant dispersal in some understory Carex species.     

Climatic niche evolution in Smilacaceae (Liliales) drives patterns of species diversification and richness between the Old and New World

Geographical variation in species richness in plant groups is determined by the interplay between historical, evolutionary, and ecological processes. However, the processes underlying the striking disparity in species richness between Asia and the Americas remain poorly understood. Here, we synthesize global phylogenetic and macroecological data on the diversification of Smilacaceae, deciphering potential drivers underlying the species diversity pattern biased toward Asia. We compiled global distributions of all Smilacaceae species, and reconstructed the biogeographic history and niche evolution using a new time-calibrated phylogeny (eight genes, 135 species). Integrating these data sets, we estimated evolutionary histories and diversification rates for each region, and tested correlations among species diversification, niche evolution, and niche divergence. Smilacaceae probably originated during the Late Cretaceous/Early Palaeocene and began to diversify in middle to low latitudes in Central America and Eurasia during the Late Eocene. Both the Old and New World clades exhibited a steady, albeit slight, increase of species diversification from the Late Eocene to Early Miocene. However, the Old World clade experienced an abrupt increase in net diversification during the Late Miocene. Our findings also revealed that species diversification rates were positively correlated with ecological niche evolution and niche divergence. Niche shifts and climatic niche evolution since the Middle Miocene played crucial roles in species diversification dynamics within Smilacaceae. The high plant richness in Asia may be explained by greater diversification in this region, potentially promoted by heterogeneous environments.     

Complex patterns of environmental niche evolution in Iberian columbines (genus Aquilegia,Ranunculaceae)

Aims This study explores the patterns of niche differentiation in a group of seven closely related columbines (genus Aquilegia, Ranunculaceae) from the Iberian Peninsula. Populations of these columbines are subject to complex patterns of divergent selection across environments, which partly explain the taxonomic structure of the group. This suggests the hypothesis that niche divergence must have occurred along the process of diversification of the group.Methods We used MaxEnt to build environmental niche models of seven subspecies belonging to the three species of Aquilegia present in the Iberian Peninsula. From these models, we compared the environmental niches through two different approaches: ENMtools and multivariate methods.Important findings MaxEnt distributions conformed closely to the actual distribution of the study taxa. ENMtools methods failed to uncover any clear patterns of niche differentiation or conservatism in Iberian columbines. Multivariate analyses indicate the existence of differentiation along altitudinal gradients and along a gradient of climatic conditions determined by the summer precipitation and temperatures. However, climatic conditions related to winter temperature and precipitation, as well as soil properties, were equally likely to show conservatism or divergence. The complex patterns of niche evolution we found suggest that Iberian Columbines have not been significantly constrained by forces of niche conservatism, so they could respond adaptively to the fast and profound climate changes in the Iberian Peninsula through the glacial cycles of the Pleistocene.     

中国莎草科新记录种对马薹草及其与近缘种的比较研究

报道了中国莎草科薹草属一新记录种：对马薹草（Carex tsushimensis（Ohwi）Ohwi）。该种现知分布于日本九州的对马岛和中国浙江省遂昌县的九龙山国家级自然保护区,生长于林下水沟边。根据小坚果先端具僧帽状膨大的环盘特征,对马薹草属于灰帽薹草组（sect.Mitratae）,形态接近中华薹草（C.chinensis）、伴生薹草（C.sociata）和龙奇薹草（C.longkiensis）。对马薹草与伴生薹草最为近缘,区别主要在于前者叶片宽2~4 mm,小穗较少,4或5个,顶生雄小穗与其下的侧生小穗稍疏远,雌花鳞片先端延伸成长约3 mm的粗糙长芒。     

Congruent spatial patterns of species richness and phylogenetic diversity in karst flora: Case study of Primulina (Gesneriaceae)

The karst landform in southern China is renowned for its high levels of species diversity and endemism. Globally, karst ecosystems are under threat from unsustainable anthropogenic disturbance and climate changes and are among the most threatened ecosystems worldwide. In this study, we used the typical karst endemic genus in southern China, Primulina Hance, as a model to identify areas within the karst landform with high diversity and to investigate congruence between phylogenetic and species‐based measures of diversity. Using phylogenetic information and species distribution data, we measured geographical patterns of diversity with four metrics: species richness (SR), corrected weighted endemism (CWE), phylogenetic diversity (PD), and phylogenetic endemism (PE). Our results revealed a high spatial congruence among SR, PD, and PE, with hotspot areas identified in the Nanling Mountains (i.e., north Guangdong and northeast Guangxi) and southeast Yungui Plateau (i.e., north and southwest Guangxi), whereas the hotspots of CWE are comparatively uniform throughout the geographic extent. The categorical analysis of neo‐ and paleoendemism identified a pattern of mixed neo‐ and paleoendemism in numerous grid cells, suggesting that karst areas in southern China have acted as both “museums” and “cradles” of plant evolution. Conservation gap analysis of hotspots revealed that the majority of prioritized hotspots (>90%) of the genus are outside of protected areas, therefore indicating the limited effectiveness of national nature reserves for the karst flora. Overall, our results suggest that the karst flora merits more conservation attention and SR can be an effective surrogate to capture PD in conservation planning.     

Model‐based approach to test hard polytomies in the Eulaemus clade of the most diverse South American lizard genus Liolaemus (Liolaemini,Squamata)

Lack of resolution in a phylogenetic tree is usually represented as a polytomy, and often adding more data (loci and taxa) resolves the species tree. These are the ‘soft’ polytomies, but in other cases additional data fail to resolve relationships; these are the ‘hard’ polytomies. This latter case is often interpreted as a simultaneous radiation of lineages in the history of a clade. Although hard polytomies are difficult to address, model‐based approaches provide new tools to test these hypotheses. Here, we used a clade of 144 species of the South American lizard clade Eulaemus to estimate phylogenies using a traditional concatenated matrix and three species tree methods: *BEAST, BEST, and minimizing deep coalescences (MDC). The different species tree methods recovered largely discordant results, but all resolved the same polytomy (e.g. very short internodes amongst lineages and low nodal support in Bayesian methods). We simulated data sets under eight explicit evolutionary models (including hard polytomies), tested these against empirical data (a total of 14 loci), and found support for two polytomies as the most plausible hypothesis for diversification of this clade. We discuss the performance of these methods and their limitations under the challenging scenario of hard polytomies. © 2015 The Linnean Society of London     

Species coherence in the face of karyotype diversification in holocentric organisms: the case of a cytogenetically variable sedge (Carex scoparia,Cyperaceae)

Background and Aims

The sedge genus Carex, the most diversified angiosperm genus of the northern temperate zone, is renowned for its holocentric chromosomes and karyotype variability. The genus exhibits high variation in chromosome numbers both among and within species. Despite the possibility that this chromosome evolution may play a role in the high species diversity of Carex, population-level patterns of molecular and cytogenetic differentiation in the genus have not been extensively studied.

Methods

Microsatellite variation (11 loci, 461 individuals) and chromosomal diversity (82 individuals) were investigated in 22 Midwestern populations of the North American sedge Carex scoparia and two Northeastern populations.

Key Results

Among Midwestern populations, geographic distance is the most important predictor of genetic differentiation. Within populations, inbreeding is high and chromosome variation explains a significant component of genetic differentiation. Infrequent dispersal among populations separated by >100 km explains an important component of molecular genetic and cytogenetic diversity within populations. However, karyotype variation and correlation between genetic and chromosomal variation persist within populations even when putative migrants based on genetic data are excluded.

Conclusions

These findings demonstrate dispersal and genetic connectivity among widespread populations that differ in chromosome numbers, explaining the phenomenon of genetic coherence in this karyotypically diverse sedge species. More generally, the study suggests that traditional sedge taxonomic boundaries demarcate good species even when those species encompass a high range of chromosomal diversity. This finding is important evidence as we work to document the limits and drivers of biodiversity in one of the world''s largest angiosperm genera.     

Abundance and species richness patterns of diving beetles (Coleoptera, Dytiscidae) from exposed and protected sites in 98 northern Swedish lakes

Forty species of predaceous diving beetles (Coleoptera, Dytiscidae) were collected in sweepnet samples from 98 boreal lakes in northern Sweden. Samples from protected sites with vegetation had significantly more specimens and species than those from exposed sites without vegetation in the same lakes. No geographically based differences, e.g. latitudinal or in distance from the coast, were found. These gradients were 200 and 150 km long, respectively. Species' distribution in terms of occurence at number of sites was positively correlated with the mean abundance of the species. The relationship between species' abundance and body length was characterized by the lack of large, abundant species. Partial least square regressions on dytiscid abundance and species richness showed low predictive power of lake area, altitude and water chemistry. The seven environmental variables used explained at best 15.3 and 24.4% of the total variance in abundance and species richness, respectively. The results are compared to activity trap catches of dystiscids in lakes in an adjacent region. A majority of the species occured in both materials. However, species occuring in high proportion in one of the materials, were rare in the other material. The mean body length of the species caught more efficiently with traps was not larger than that of those overpresented in net sampling. Based on this study and the available literature data, the regional species pool of boreal lake dytiscids is estimated to 30–40 species. It is still an open question if lake assemblages are markedly poorer than those found in the development of vegetation, whereas the impact of water chemistry is small.     

Linking ecology,morphology, and metabolism: Niche differentiation in sympatric populations of closely related species of the genus Littorina (Neritrema)

Divergence of ecological niches in phylogenetically closely related species indicates the importance of ecology in speciation, especially for sympatric species are considered. Such ecological diversification provides an advantage of alleviating interspecies competition and promotes more efficient exploitation of environmental resources, thus being a basis for ecological speciation. We analyzed a group of closely related species from the subgenus Neritrema (genus Littorina, Caenogastropoda) from the gravel‐bouldery shores. In two distant sites at the Barents and Norwegian Sea, we examined the patterns of snail distribution during low tide (quantitative sampling stratified by intertidal level, presence of macrophytes, macrophyte species, and position on them), shell shape and its variability (geometric morphometrics), and metabolic characteristics (metabolomic profiling). The studied species diversified microbiotopes, which imply an important role of ecological specification in the recent evolution of this group. The only exception to this trend was the species pair L. arcana / L. saxatilis, which is specifically discussed. The ecological divergence was accompanied by differences in shell shape and metabolomic characteristics. Significant differences were found between L. obtusata versus L. fabalis and L. saxatilis / L. arcana versus L. compressa both in shell morphology and in metabolomes. L. saxatilis demonstrated a clear variability depending on intertidal level which corresponds to a shift in conditions within the occupied microhabitat. Interestingly, the differences between L. arcana (inhabiting the upper intertidal level) and L. compressa (inhabiting the lower one) were analogous to those between the upper and lower fractions of L. saxatilis. No significant level‐dependent changes were found between the upper and lower fractions of L. obtusata, most probably due to habitat amelioration by fucoid macroalgae. All these results are discussed in the contexts of the role of ecology in speciation, ecological niche dynamics and conservatism, and evolutionary history of the Neritrema species.     

Phylogenomics,biogeography, and evolution of morphology and ecological niche of the eastern Asian–eastern North American Nyssa (Nyssaceae)

Nyssa (Nyssaceae, Cornales) represents a classical example of the well‐known eastern Asian–eastern North American floristic disjunction. The genus consists of three species in eastern Asia, four species in eastern North America, and one species in Central America. Species of the genus are ecologically important trees in eastern North American and eastern Asian forests. The distribution of living species and a rich fossil record of the genus make it an excellent model for understanding the origin and evolution of the eastern Asian–eastern North American floristic disjunction. However, despite the small number of species, relationships within the genus have remained unclear and have not been elucidated using a molecular approach. Here, we integrate data from 48 nuclear genes, fossils, morphology, and ecological niche to resolve species relationships, elucidate its biogeographical history, and investigate the evolution of morphology and ecological niches, aiming at a better understanding of the well‐known EA–ENA floristic disjunction. Results showed that the Central American (CAM) Nyssa talamancana was sister to the remaining species, which were divided among three, rapidly diversified subclades. Estimated divergence times and biogeographical history suggested that Nyssa had an ancestral range in Eurasia and western North America in the late Paleocene. The rapid diversification occurred in the early Eocene, followed by multiple dispersals between and within the Erasian and North American continents. The genus experienced two major episodes of extinction in the early Oligocene and end of Neogene, respectively. The Central American N. talamancana represents a relic lineage of the boreotropical flora in the Paleocene/Eocene boundary that once diversified in western North America. The results supported the importance of both the North Atlantic land bridge and the Bering land bridge (BLB) for the Paleogene dispersals of Nyssa and the Neogene dispersals, respectively, as well as the role of Central America as refugia of the Paleogene flora. The total‐evidence‐based dated phylogeny suggested that the pattern of macroevolution of Nyssa coincided with paleoclimatic changes. We found a number of evolutionary changes in morphology (including wood anatomy and leaf traits) and ecological niches (precipitation and temperature) between the EA–ENA disjunct, supporting the ecological selection driving trait evolutions after geographic isolation. We also demonstrated challenges in phylogenomic studies of lineages with rapid diversification histories. The concatenation of gene data can lead to inference of strongly supported relationships incongruent with the species tree. However, conflicts in gene genealogies did not seem to impose a strong effect on divergence time dating in our case. Furthermore, we demonstrated that rapid diversification events may not be recovered in the divergence time dating analysis using BEAST if critical fossil constraints of the relevant nodes are not available. Our study provides an example of complex bidirectional exchanges of plants between Eurasia and North America in the Paleogene, but “out of Asia” migrations in the Neogene, to explain the present disjunct distribution of Nyssa in EA and ENA.     

Species diversity,ecology and evolution in a primitive Angiosperm genus:Hibbertia (Dilleniaceae)

Among the approximately 130 species ofHibbertia found in Australia, there are tall shrubs, low or trailing shrubs and vines bearing a diversity of leaves as to shape and venation pattern. Flowers are solitary, in leafy cymes or in false spikes, and display various gradual and abrupt transitions from vegetative to reproductive appendages. In the androecium, stamen number is highly variable both between and within species. Some sections have radial symmetry, others bilateral symmetry of the androecium and gynoecium. Follicle number varies from 10 to 1. Basic chromosome numbers of n = 4, 5, 8, 9, 10, 12 and 13 have been found in various sections, and occasional higher numbers, up to n = 64, indicate the presence of polyploidy. Habitats vary from tropical savanna through rain forest margins, wet and dry sclerophyll forests, heaths, sphagnum swamps, and mallee scrub to desert margins. The principal center of diversity is southwestern Australia, less diverse centers are in southeastern and northern Australia. With respect to leaf size, structure and venation; floral symmetry; and chromosome numbers; the diversity found among the species ofHibbertia exceeds that found in all but a few genera of Angiosperms, and is greater than that in any other exclusively woody genus. Nevertheless, individual species are relatively constant with respect to both morphology and ecological preferences.Presented at the symposium Speciation and the Species Concept during the XIIth International Botanical Congress, Leningrad, July 8, 1975.     

Male differentiation patterns in two polyphenic sister species of the genus Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae): a geometric morphometric approach

This paper focuses on morphological (both shape and size ) differences that quite similar polyphenic sister species evolve during divergence processes. Traits were analysed using a geometrical morphometric approach, which has the ability to evidence also very subtle differences in shape. As a case study, we considered males of the dung beetle sister species pair Onthophagus taurus and Onthophagus illyricus (Coleoptera, Scarabaeidae); these species represent a typical example of polyphenic trait expression concerning the facultative development of horns and considerable body size differences. External shape morphology failed to discriminate O. taurus from O. illyricus , whereas the reproductive system shape showed significant interspecific discrimination power. However, the head of O. taurus was significantly larger than that of O. illyricus and the reverse was true for the elytra. The two species also showed different allometric values of the head with respect to body size. This complex pattern of interspecific morphological divergence is discussed in the light of the differential trait divergence rate hypothesis. In both species, differences between major and minor forms concern the overall shape of head and pronotum: we suggest that such different forms, which likely reflect morphological readjustment to accommodate horns of considerable bulk and disproportionate length, may be nevertheless advantageously used by the two male morphs in their alternative reproductive tactics. Male genitalia sizes were virtually constant with respect to body size; however, the ratio between phallotheca and body size was significantly higher in minor males, in keeping with the hypothesis of a higher investment in genitalia borne by this morph.