Phylogeography of North Atlantic intertidal tardigrades: refugia,cryptic speciation and the history of the Mid‐Atlantic Islands

Aim To analyse the phylogeographical history of intertidal tardigrades in the North Atlantic in order to improve our understanding of geographical differentiation in microscopic organisms, and to understand the potential importance of the Mid‐Atlantic Islands as stepping stones between the American and European coasts of the Atlantic Ocean. Location Twenty‐four localities from the Mid‐Atlantic Islands (Greenland, Iceland and the Faroe Islands) and both sides of the North Atlantic Ocean. Methods A mitochondrial marker (cytochrome c oxidase subunit I) was sequenced from individual tardigrades belonging to the genus Echiniscoides. The existence of cryptic species was detected using generalized mixed Yule coalescence analysis; lineage ages were estimated with relaxed clock methods; and the degree of geographical differentiation was analysed with samova analyses, haplotype networks and Mantel tests. Results Echiniscoides hoepneri, previously known only from Greenland, was recovered throughout the Mid‐Atlantic Islands. The Faroe Islands population was isolated from Greenland and Iceland, but overall genetic variation was low. The morphospecies Echiniscoides sigismundi had high genetic variation and consisted of at least two cryptic species. A northern and a southern species were both recovered on both sides of the Atlantic, but only the northern species was found on the Mid‐Atlantic Islands. The northern species showed signs of long‐term isolation between the Western and Eastern Atlantic, despite the potential of the Mid‐Atlantic islands to act as stepping‐stones. There was no sign of long‐term isolation in the southern species. The Mid‐Atlantic individuals of the northern species were of Eastern Atlantic origin, but Greenland and Iceland showed signs of long‐term isolation. The genetic pattern found in the southern species is not clearly geographical, and can probably be best explained by secondary contact between former isolated populations. Main conclusions North Atlantic intertidal tardigrades from the genus Echiniscoides showed strong geographical differentiation, and the Mid‐Atlantic Islands seemed unimportant as stepping stones across the Atlantic. The geographical variation of the northern species of E. sigismundi suggests post‐glacial recolonization from several refugia.     

Genetic divergence between the sympatric queen morphs of the ant Myrmica rubra

Pairs of obligate social parasites and their hosts, where some of the parasites have recently diverged from their host through intraspecific social parasitism, provide intriguing systems for studying the modes and processes of speciation. Such speciation, probably in sympatry, has also been propounded in the ant Myrmica rubra and its intraspecific social parasite. In this species, parasitism is associated with queen size dimorphism, and the small microgyne has become a social parasite of the large macrogyne. Here, we investigated the genetic divergence of the host and the parasite queen morphs in 11 localities in southern Finland, using nuclear and mitochondrial markers of queens and workers. We formulated and tested four speciation‐related hypotheses that differed in the degree of genetic divergence between the morphs. The queen morphs were genetically distinct from each other with little hybridization. In the nuclear data, when localities were nested within queen morphs in the hierarchical amova , 39% of the genetic variation was explained by the queen morph (standardized F'_CT = 0.63, uncorrected F_CT = 0.39), whereas 18% was explained by the locality (F'_SC = 0.39, F_SC = 0.29). This result corroborated the hypothesis of advanced sympatric speciation. In contrast, the mitochondrial DNA could not settle between the hierarchical levels of locality and queen morph, thus substantiating equally the hypotheses of incipient and advanced sympatric speciation. Together, our results support the view that the microgynous parasite has genetically diverged from its macrogynous host to the level of a nascent species.     

Computational prediction of drug response in short QT syndrome type 1 based on measurements of compound effect in stem cell-derived cardiomyocytes

Short QT (SQT) syndrome is a genetic cardiac disorder characterized by an abbreviated QT interval of the patient’s electrocardiogram. The syndrome is associated with increased risk of arrhythmia and sudden cardiac death and can arise from a number of ion channel mutations. Cardiomyocytes derived from induced pluripotent stem cells generated from SQT patients (SQT hiPSC-CMs) provide promising platforms for testing pharmacological treatments directly in human cardiac cells exhibiting mutations specific for the syndrome. However, a difficulty is posed by the relative immaturity of hiPSC-CMs, with the possibility that drug effects observed in SQT hiPSC-CMs could be very different from the corresponding drug effect in vivo. In this paper, we apply a multistep computational procedure for translating measured drug effects from these cells to human QT response. This process first detects drug effects on individual ion channels based on measurements of SQT hiPSC-CMs and then uses these results to estimate the drug effects on ventricular action potentials and QT intervals of adult SQT patients. We find that the procedure is able to identify IC₅₀ values in line with measured values for the four drugs quinidine, ivabradine, ajmaline and mexiletine. In addition, the predicted effect of quinidine on the adult QT interval is in good agreement with measured effects of quinidine for adult patients. Consequently, the computational procedure appears to be a useful tool for helping predicting adult drug responses from pure in vitro measurements of patient derived cell lines.     

Extinction risk and diversification are linked in a plant biodiversity hotspot

It is widely recognized that we are entering an extinction event on a scale approaching the mass extinctions seen in the fossil record. Present-day rates of extinction are estimated to be several orders of magnitude greater than background rates and are projected to increase further if current trends continue. In vertebrates, species traits, such as body size, fecundity, and geographic range, are important predictors of vulnerability. Although plants are the basis for life on Earth, our knowledge of plant extinctions and vulnerabilities is lagging. Here, we disentangle the underlying drivers of extinction risk in plants, focusing on the Cape of South Africa, a global biodiversity hotspot. By comparing Red List data for the British and South African floras, we demonstrate that the taxonomic distribution of extinction risk differs significantly between regions, inconsistent with a simple, trait-based model of extinction. Using a comprehensive phylogenetic tree for the Cape, we reveal a phylogenetic signal in the distribution of plant extinction risks but show that the most threatened species cluster within short branches at the tips of the phylogeny--opposite to trends in mammals. From analyzing the distribution of threatened species across 11 exemplar clades, we suggest that mode of speciation best explains the unusual phylogenetic structure of extinction risks in plants of the Cape. Our results demonstrate that explanations for elevated extinction risk in plants of the Cape flora differ dramatically from those recognized for vertebrates. In the Cape, extinction risk is higher for young and fast-evolving plant lineages and cannot be explained by correlations with simple biological traits. Critically, we find that the most vulnerable plant species are nonetheless marching towards extinction at a more rapid pace but, surprisingly, independently from anthropogenic effects. Our results have important implications for conservation priorities and cast doubts on the utility of current Red List criteria for plants in regions such as the Cape, where speciation has been rapid, if our aim is to maximize the preservation of the tree-of-life.     

Nuclear-cytoplasmic interactions reduce male fertility in hybrids of Arabidopsis lyrata subspecies

We examined the level of postzygotic reproductive isolation in F(1) and F(2) hybrids of reciprocal crosses between the Arabidopsis lyrata subspecies lyrata (North American) and petraea (European). Our main results are: first, the percentage of fertile pollen was significantly reduced in the F(1) and F(2) compared to the parental populations. Second, mean pollen fertility differed markedly between reciprocal crosses: 84% in the F(2) with ssp. lyrata cytoplasm and 61% in the F(2) with ssp. petraea cytoplasm. Third, 17% of the F(2) with ssp. petraea cytoplasm showed male sterility (produced less than 30 pollen grains in our subsample). The hybrids were female fertile. We used QTL mapping to find the genomic regions that determine pollen fertility and that restore cytoplasmic male sterility (CMS). In the F(2) with ssp. lyrata cytoplasm, an epistatic pair of QTLs was detected. In the reciprocal F(2) progeny, four QTLs demonstrated within-population polymorphism for hybrid male sterility. In addition, in the F(2) with ssp. petraea cytoplasm, there was a strong male fertility restorer locus on chromosome 2 where a cluster of CMS restorer gene-related PPR genes have been found in A. lyrata. Our results underline the importance of cytonuclear interactions in understanding genetics of the early stages of speciation.     

Phylogenetic relationships among arecoid palms (Arecaceae: Arecoideae)

Background and Aims

The Arecoideae is the largest and most diverse of the five subfamilies of palms (Arecaceae/Palmae), containing >50 % of the species in the family. Despite its importance, phylogenetic relationships among Arecoideae are poorly understood. Here the most densely sampled phylogenetic analysis of Arecoideae available to date is presented. The results are used to test the current classification of the subfamily and to identify priority areas for future research.

Methods

DNA sequence data for the low-copy nuclear genes PRK and RPB2 were collected from 190 palm species, covering 103 (96 %) genera of Arecoideae. The data were analysed using the parsimony ratchet, maximum likelihood, and both likelihood and parsimony bootstrapping.

Key Results and Conclusions

Despite the recovery of paralogues and pseudogenes in a small number of taxa, PRK and RPB2 were both highly informative, producing well-resolved phylogenetic trees with many nodes well supported by bootstrap analyses. Simultaneous analyses of the combined data sets provided additional resolution and support. Two areas of incongruence between PRK and RPB2 were strongly supported by the bootstrap relating to the placement of tribes Chamaedoreeae, Iriarteeae and Reinhardtieae; the causes of this incongruence remain uncertain. The current classification within Arecoideae was strongly supported by the present data. Of the 14 tribes and 14 sub-tribes in the classification, only five sub-tribes from tribe Areceae (Basseliniinae, Linospadicinae, Oncospermatinae, Rhopalostylidinae and Verschaffeltiinae) failed to receive support. Three major higher level clades were strongly supported: (1) the RRC clade (Roystoneeae, Reinhardtieae and Cocoseae), (2) the POS clade (Podococceae, Oranieae and Sclerospermeae) and (3) the core arecoid clade (Areceae, Euterpeae, Geonomateae, Leopoldinieae, Manicarieae and Pelagodoxeae). However, new data sources are required to elucidate ambiguities that remain in phylogenetic relationships among and within the major groups of Arecoideae, as well as within the Areceae, the largest tribe in the palm family.     

Existence of excitation waves for a collection of cardiomyocytes electrically coupled to fibroblasts

We consider mathematical models of a collection of cardiomyocytes (myocardial tissue) coupled to a varying number of fibroblasts. Our aim is to understand how conductivity (δ) and fibroblast density (η) affect the stability of the collection. We provide mathematical and computational arguments indicating that there is a region of instability in the η-δ space. Mathematical arguments, based on a simplified model of the coupled myocyte-fibroblast system, show that for certain parameter choices, a stationary solution cannot exist. Numerical experiments (1D, 2D) are based on a recently developed model of electro-chemical coupling between a human atrial myocyte and a number of associated atrial fibroblasts. The numerical experiments demonstrate that there is a region of instability of the form observed in the simplified model analysis.     

Are spring mass migrations of bumblebees and wasps driven by vole cyclicity?

In search for factors which could drive the spectacular spring migrations of pre-reproductive queens of bumblebees and social wasps, we made a literature search of potential mortality factors of individuals and colonies. Although the role of parasitism cannot be ruled out, we highlight the potential of vole cyclicity in explaining the migrations. Because the spatial scale of vole-cycle asynchrony is located at the same level (γ scale) as the range of migratory movements, spring flights could rid bumblebees and wasps of strong nest-site competition among colony-founding queens, and of severe predation by voles in high vole-density regions, to take advantage of abundant, empty vole burrows as nest sites in low-predation, relatively vole-free areas. As a corollary of the migratory movements, relaxed parasite pressure is expected.     

The geographical pattern of speciation and floral diversification in the neotropics: the tribe sinningieae (gesneriaceae) as a case study

The geographical pattern of speciation and the relationship between floral variation and species ranges were investigated in the tribe Sinningieae (Gesneriaceae), which is found mainly in the Atlantic forests of Brazil. Geographical distribution data recorded on a grid system of 0.5 x 0.5 degree intervals and a near-complete species-level phylogenetic tree of Sinningieae inferred from a simultaneous analysis of seven DNA regions were used to address the role of geographical isolation in speciation. Geographical range overlaps between sister lineages were measured across all nodes in the phylogenetic tree and analyzed in relation to relative ages estimated from branch lengths. Although there are several cases of species sympatry in Sinningieae, patterns of sympatry between sister taxa support the predominance of allopatric speciation. The pattern of sympatry between sister taxa is consistent with range shifts following allopatric speciation, except in one clade, in which the overlapping distribution of recent sister species indicates speciation within a restricted geographical area and involving changes in pollinators and habitats. The relationship between floral divergence and regional sympatry was also examined by analyzing floral contrasts, phenological overlap, and the degree of sympatry between sister clades. Morphological contrast between flowers is not increased in sympatry and phenological divergence is more apparent between allopatric clades than between sympatric clades. Therefore, our results failed to indicate a tendency for sympatric taxa to minimize morphological and phenological overlap (geographic exclusion and/or character displacement hypotheses). Instead, they point toward adaptation in phenology to local conditions and buildup of sympatries at random with respect to flower morphology. Additional studies at a lower geographical scale are needed to identify truely coexisting species and the components of their reproductive isolation.