Evolutionary history of Gymnocarpos (Caryophyllaceae) in the arid regions from North Africa to Central Asia

Gymnocarpos has only about ten species distributed in the arid regions of Asia and Africa, but it exhibits a geographical disjunction between eastern Central Asia and western North Africa and Minor Asia. We sampled eight species of the genus and sequenced two chloroplast regions (rps16 and psbB–psbH), and the nuclear rDNA (ITS) to study the phylogeny and biogeography. The results of the phylogenetic analyses corroborated that Gymnocarpos is monophyletic, in the phylogenetic tree two well supported clades are recognized: clade 1 includes Gymnocarpos sclerocephalus and G. decandrus, mainly the North African group, whereas clade 2 comprises the remaining species, mainly in the Southern Arabian Peninsula. Molecular dating analysis revealed that the divergence age of Gymnocarpos was c. 31.33 Mya near the Eocene and Oligocene transition boundary, the initial diversification within Gymnocarpos dated to c. 6.69 Mya in the late Miocene, and the intraspecific diversification mostly occurred during the Quaternary climate oscillations. Ancestral area reconstruction suggested that the Southern Arabian Peninsula was the ancestral area for Gymnocarpos. Our conclusions revealed that the aridification since mid‐late Miocene significantly affected the diversification of the genus in these areas.     

Species Diversity,Phylogeny and Large Scale Biogeographic Patterns of the Genus Padina (Phaeophyceae,Dictyotales)

The brown algal genus Padina (Dictyotales, Phaeophyceae) is distributed worldwide in tropical and temperate seas. Global species diversity and distribution ranges, however, remain largely unknown. Species‐level diversity was reassessed using DNA‐based, algorithmic species delineation techniques based on cox3 and rbcL sequence data from 221 specimens collected worldwide. This resulted in estimates ranging from 39 to 61 putative species (ESUs), depending on the technique as well as the locus. We discuss the merits, potential pitfalls, and evolutionary and biogeographic significance of algorithmic species delineation. We unveil patterns whereby ESUs are in all but one case restricted to either the Atlantic or Indo‐Pacific Ocean. Within ocean basins we find evidence for the vast majority of ESUs to be confined to a single marine realm. Exceptions, whereby ESUs span up to three realms, are located in the Indo‐Pacific Ocean. Patterns of range‐restricted species likely arise by repeated founder events and subsequent peripatric speciation, hypothesized to dominate speciation mechanisms for coastal marine organisms in the Indo‐Pacific. Using a three‐gene (cox3, psaA and rbcL), relaxed molecular clock phylogenetic analysis we estimated divergence times, providing a historical framework to interpret biogeographic patterns.     

Demystifying the marine‐terrestrial biodiversity gradient: response to Vermeij et al.

We respond to seven criticisms made by Vermeij et al. ( 2018 ) regarding Miller & Wiens ( 2017 ). Their criticisms generally reflect misunderstandings, unsupported speculations, and topics that were explicitly addressed in our paper.     

Climatic niche conservatism and ecological opportunity in the explosive radiation of arvicoline rodents (Arvicolinae,Cricetidae)

Climatic niche conservatism shapes patterns of diversity in many taxonomic groups, while ecological opportunity (EO) can trigger rapid speciation that is less constrained by the amount of time a lineage has occupied a given habitat. These two processes are well studied, but limited research has considered their joint and relative roles in shaping diversity patterns. We characterized climatic and biogeographic variables for 102 species of arvicoline rodents (Arvicolinae, Cricetidae), testing the effects of climatic niche conservatism and EO on arvicoline diversification as lineages transitioned between biogeographic regions. We found that the amount of time a lineage has occupied a precipitation niche is positively correlated with diversity along a precipitation gradient, suggesting climatic niche conservatism. In contrast, shift in diversification rate explained diversity patterns along a temperature gradient. Our results suggest that an indirect relationship exists between temperature and diversification that is associated with EO as arvicoline rodents colonized warm Palearctic environments. Climatic niche conservatism alone did not fully explain diversity patterns under density‐dependence, highlighting the additional importance of EO‐related processes in promoting the explosive radiation in arvicoline rodents and shaping diversity pattern among biogeographic regions and along climatic gradients.     

Distribution and microhabitats of native and non-native gammarids (Amphipoda, Crustacea) in Brittany, with particular reference to the endangered endemic sub-species Gammarus duebeni celticus

Aim  To assess temporal changes in gammarid distribution in Brittany and microhabitat-use overlap between the endangered endemic Gammarus duebeni celticus Stock & Pinkster, 1970 , the expanding natives G. pulex (Linnaeus, 1758) and Echinogammarus berilloni (Catta, 1878), and the introduced G. tigrinus Sexton, 1939.
Location  Brittany and adjacent regions in western France.
Methods  The spatial and temporal patterns in distribution of gammarids at the scale of Brittany were studied using 351 sites. Longitudinal distributions (from the source to the estuary of the river) and microhabitat-use (substratum type and water velocity) were also considered in selected rivers.
Results  At the regional scale, all species occurred together less often than expected statistically, with significant deviations from expected for G. pulex vs. both G. duebeni celticus and G. tigrinus , and for E. berilloni vs. both G. duebeni celticus and G. tigrinus . However, at the microhabitat scale, E. berilloni occurred significantly more often than expected with the endemic G. duebeni celticus , and this appears to be due to similar substratum and water velocity preferences, although at both the regional and microhabitat scales E. berilloni prefers wider streams than G. duebeni celticus . This study reveals a decline in the endangered G. duebeni celticus since 1970.
Main conclusions  The longitudinal and local distributions of G. duebeni celticus , and the higher-than-expected co-occurrence of the species with G. pulex , suggest that the decline of the endemic species may be due to changes in the environment and/or interference from native G. pulex , which is expanding its range in Brittany. The results are discussed as regards to the consequences for regional biodiversity.     

High tropical net diversification drives the New World latitudinal gradient in palm (Arecaceae) species richness

Aim Species richness exhibits striking geographical variation, but the processes that drive this variation are unresolved. We investigated the relative importance of two hypothesized evolutionary causes for the variation in palm species richness across the New World: time for diversification and evolutionary (net diversification) rate. Palms have a long history in the region, with the major clades diversifying during the Tertiary (65–2 Ma). Location Tropical and subtropical America (34° N–34° S; 33–120° W). Methods Using range maps, palm species richness was estimated in a 1° × 1° grid. Mean lineage net diversification was estimated by the mean phylogenetic root distance (MRD), the average number of nodes separating a species from the base of the palm phylogeny for the species in each grid cell. If evolutionary rate limits richness, then richness should increase with MRD. If time limits richness, then old, relict species (with low root distance) should predominantly occur in long‐inhabited and therefore species‐rich areas. Hence, richness should decrease with MRD. To determine the influence of net diversification across different time frames, MRD was computed for subtribe, genus and species levels within the phylogeny, and supplemented with the purely tip‐level measure, mean number of species per genus (MS/G). Correlations and regressions, in combination with eigenvector‐based spatial filtering, were used to assess the relationship between species richness, the net diversification measures, and potential environmental and geographical drivers. Results Species richness increased with all net diversification measures. The regression models showed that richness and the net diversification measures increased with decreasing (absolute) latitude and, less strongly, with increasing energy/temperature and water availability. These patterns therefore reflect net diversification at both deep and shallow levels in the phylogeny. Richness also increased with range in elevation, but this was only reflected in the MS/G pattern and therefore reflects recent diversification. Main conclusions The geographical patterns in palm species richness appear to be predominantly the result of elevated net diversification rates towards the equator and in warm, wet climates, sustained throughout most of the Tertiary. Late‐Tertiary orogeny has caused localized increases in net diversification rates that have also made a mark on the richness pattern.     

Probing Diversity in the Plankton: Using Patterns in Tintinnids (Planktonic Marine Ciliates) to Identify Mechanisms

In diversity research, the use of survey data appears to have declined in favour of experimental or modeling approaches because direct relationships are difficult to demonstrate. Here we show that use of field data can yield information concerning the mechanisms governing diversity. First, we establish that tintinnids display a global latitudinal pattern of diversity similar to other pelagic organisms; species numbers appear to peak between 20° and 30° north or south. This common large scale spatial trend has been attributed to the gradient in water column structure across the global ocean. We then examine the generality of a relationship between planktonic diversity and water column structure by considering data from the Mediterranean Sea, in which water column structure changes seasonally. Among populations of foraminifera, tintinnids, and the dinoflagellates of the genus Ceratium, we compare data from trans-Mediterranean sampling conducted at different times and monthly changes in species richness at single sites. We find that water column structure alone appears to be a poor predictor of temporal changes in diversity. Lastly, we present an example of temporal changes in tintinnid diversity based on data from an oceanographic sampling station in the N. W. Mediterranean where resources, as chlorophyll, appear distinctly unrelated to changes in water column structure. We show that short-tem temporal changes in diversity (week to week shifts) can be related to changes in chlorophyll concentration. We conclude that in tintinnids diversity can be directly linked to characteristics of food resources.     

West to east dispersal and subsequent rapid diversification of the mega‐diverse genus Begonia (Begoniaceae) in the Malesian archipelago

Aim The complex palaeogeography of the Malesian archipelago, characterized by the evolution of an ever‐changing mosaic of terrestrial and marine areas throughout the Cenozoic, provides the geographic backdrop for the remarkable diversification of Malesian Begonia (> 450 species). This study aimed to investigate the origin of Malesian Begonia, the directionality of dispersal events within the Malesian archipelago and the impact of ancient water gaps on colonization patterns, and to identify drivers of diversification. Location Asia, Southeast Asia, Malesia. Methods Plastid DNA sequence data of representatives of all families of the Cucurbitales and Fagales (matK, rbcL, trnL intron, trnL–F spacer, 4076 aligned positions, 92 taxa) and a sample of all major Asian Begonia sections (ndhA intron, ndhF–rpl32 spacer, rpl32–trnL spacer, 4059 aligned positions, 112 taxa) were analysed under an uncorrelated‐rates relaxed molecular clock model to estimate the age of the Begonia crown group divergence and divergence ages within Asian Begonia. Ancestral areas were reconstructed using a likelihood approach implementing a dispersal–extinction–cladogenesis model, and with a Bayesian approach to dispersal–vicariance analysis. Results The results indicated an initial diversification of Asian Begonia in continental Asia in the Miocene, and subsequent colonization of Malesia by multiple lineages. There was support for at least six independent dispersal events from continental Asia and western Malesia to Wallacea dating from the late Miocene to the Pleistocene. Begonia section Petermannia (> 270 species) originated in Western Malesia, and subsequently dispersed to Wallacea, New Guinea and the Philippines. Lineages within this section diversified rapidly since the Pliocene, coinciding with rapid orogenesis on Sulawesi and New Guinea. Main conclusions The predominant trend of Begonia dispersals between continental Asia and Malesia, and also within Malesia, has been from west to east. The water bodies separating the Sunda Shelf region from Wallacea have been porous barriers to dispersal in Begonia following the emergence of substantial land in eastern Malesia from the late Miocene onwards. We hypothesize two major drivers of the diversification of Malesian Begonia: (1) the formation of topographical heterogeneity and the promotion of microallopatry by orogenesis in the Pliocene and Pleistocene; and (2) cyclic vicariance by frequent habitat fragmentations and amalgamations due to climate and sea‐level fluctuations during the Pleistocene.     

Radiating in a river: systematics, molecular genetics and morphological differentiation of viviparous freshwater gastropods endemic to the Kaek River, central Thailand (Cerithioidea, Pachychilidae)

Speciation in the context of adaptive radiation is regarded as a key process in the creation of biodiversity. While several lacustrine species flocks provide ideal models for elucidating the underlying evolutionary mechanisms, riverine radiations are both rarely known and studied. The Kaek River, a third‐order tributary of the Nan River and Chao Praya drainage in central Thailand, harbours an exceptional endemic species assemblage of morphologically distinct, viviparous pachychilid gastropods. Our systematic revision, combining a morphological and molecular genetics approach, reveals the sympatric existence of at least seven species of the genus Brotia that is widespread in rivers of South‐east Asia where usually only two species at the most coexist. At eight locations along a 100‐kilometre stretch of the Kaek River, we found the syntopic occurrence of two to three species that are separated by specific habitat preferences and exhibit trophic specialization in their radula morphology. Phylogenetic analyses (using MP, NJ, ML and Bayesian inference statistics) of partial COI and 16S sequence data of 17 samples from six species occurring sympatrically and parapatrically, respectively, in the Kaek River drainage (plus the type species B. pagodula as outgroup) indicate monophyly of all these endemic species. Brotia solemiana, which also occurs in the Loei River, a tributary of the Mekong drainage system, was found to be sister to all other Kaek River pachychilids. The distinctive morphotypes, proposed here to represent biospecies, do not show high levels of genetic variation consistent with long periods of reproductive isolation. This suggests a relatively recent origin of this intrariverine radiation and rapid morphological divergence in the Kaek River Brotia. Recent diversification combined with ecological separation and trophic specialization parallels conditions found, albeit on a more specious level, in the lacustrine species flock of the closely related pachychilid genus Tylomelania, which is endemic to ancient lakes on the Indonesian island of Sulawesi. We discuss and compare the allopatric and ecological aspects of speciation in this unique riverine radiation and outline a putative historical biogeography of the Kaek River species, employing the most recent geological and palaeohydrological data for Thailand. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 82 , 275–311.     

Patterns of allozyme variation in relation to population size of the threatened plantMegaleranthis saniculifolia (Ranunculaceae) in Korea

Patterns of variation at 27 allozyme loci were investigated in the endangered endemic plantMegaleranthis saniculifolia. Levels of allozyme variation (A = 1.47,P = 40%,He = 0.088) were also compared with other endemic plant species. Genetic divergence between populations was very high (G ^st = 0.271 ), with moderate to high interpopulation differentiation, which probably arose through historical bottlenecks in a landscape of habitat fragmentation and/or human influence. The percentage of polymorphic loci, heterozygosity, and mean number of alleles per locus were positively related to population size, probably due to the stochastic loss of rare alleles in the smaller populations. Individuals in the small and marginal populations (TB, KD, and CJ) showed higher proportions of fixed loci. These ecologically marginal populations were typically more distant from the nearest neighboring population and were more genetically distinct from one another. The genetic structure of the current population ofM. saniculifolia is probably the result of local extinctions of intervening populations. This, in turn, is due to the Pleistocene climatic change and increased habitat destruction. A positive association appears to exist between genetic diversity and population size. Although these small population sizes are more sensitive to stochastic events, securing a certain number of individuals from the three larger populations (SB, JB, and TG) could be accomplished as part of a conservation strategy. In addition, it is important to prioritize populations in different regions in order to limit population declines caused by large-scale environmental catastrophes.