Speciation in the Cladophorales species flock of ancient Lake Baikal

<p>Understanding speciation is one of the great challenges in evolutionary biology as many of the processes involved in speciation, as well as the forces leading to morphological and genetic differentiation, are not fully understood. Three main modes of speciation have been described: allopatric, parapatric and sympatric. Sympatric speciation is the most enigmatic mode because in the absence of physical barriers, disruptive selection, assortative mating and hybridization play central roles in reproductive isolation. Although it is accepted that sympatric speciation is possible, only a few examples of this process exist to date. Another common method of speciation in plants and algae is via polyploidization. Recently, a promising system to study speciation in sympatry was discovered: the endemic Cladophorales species flock in ancient Lake Baikal, Russia. The flock consists of sixteen taxa grouped in four genera: Chaetocladiella, Chaetomorpha, Cladophora and Gemmiphora. In spite of their morphological diversity, recent molecular analyses have shown that this is a monophyletic group with low genetic variation and nested within the morphologically simple genus Rhizoclonium. Due to their high number of species, endemism and sympatric distribution, many interesting questions have arisen such as what processes are involved in speciation, and whether this group might be a novel example of sympatric speciation. In this study, we analysed the population genetics of the endemic Baikalian Cladophorales to infer the processes shaping the evolution of the group. First, a set of microsatellites was designed using high-throughput sequencing data. Second, species delimitation methods based on genetic clustering were performed. Third, the population genetics of three widely distributed species was analysed looking for evidence of panmixia, a common criteria to support sympatric speciation. A total of 11 microsatellites that mostly cross-amplify between most species were obtained. The genotyping revealed that most loci had more than two alleles per individual indicating polyploidy. As such, the analyses required a different approach which consisted in coding the genotypes as ‘allelic phenotypes’, allowing the use of individuals of different ploidy levels in the same data set. The species delimitation of 15 operative morphotaxa and 727 individuals supported reproductive isolation of five morphotaxa and two hypotheses of conspecificity. However, some morphotaxa showed unclear assignments revealing the need of further research to clarify their reproductive limits. Finally, the population genetics of Chaetomorpha moniliformis, Cladophora compacta and Cl. kursanovii revealed patterns of genetic variation and structure that suggest different reproductive strategies and dispersal abilities. This demonstrates that contrasting biological characteristics may arise in closely related lineages: Chaetomorpha moniliformis with dominant asexual reproduction and long dispersal abilities; Cladophora compacta with high genetic diversity, no population structure and likely to reproduce sexually; Cl. kursanovii with a structure congruent with geographic distribution and more restricted dispersal. The results suggest that polyploidy, rather than speciation with gene flow, is the force driving the reproductive isolation and evolution of this flock. Although many questions remain to be studied, this research provides the first insights into the diversification of this Cladophorales species flock and contributes to the understanding of speciation in freshwater algae.</p>

Speciation in the Cladophorales species flock of ancient Lake Baikal

<p>Understanding speciation is one of the great challenges in evolutionary biology as many of the processes involved in speciation, as well as the forces leading to morphological and genetic differentiation, are not fully understood. Three main modes of speciation have been described: allopatric, parapatric and sympatric. Sympatric speciation is the most enigmatic mode because in the absence of physical barriers, disruptive selection, assortative mating and hybridization play central roles in reproductive isolation. Although it is accepted that sympatric speciation is possible, only a few examples of this process exist to date. Another common method of speciation in plants and algae is via polyploidization. Recently, a promising system to study speciation in sympatry was discovered: the endemic Cladophorales species flock in ancient Lake Baikal, Russia. The flock consists of sixteen taxa grouped in four genera: Chaetocladiella, Chaetomorpha, Cladophora and Gemmiphora. In spite of their morphological diversity, recent molecular analyses have shown that this is a monophyletic group with low genetic variation and nested within the morphologically simple genus Rhizoclonium. Due to their high number of species, endemism and sympatric distribution, many interesting questions have arisen such as what processes are involved in speciation, and whether this group might be a novel example of sympatric speciation. In this study, we analysed the population genetics of the endemic Baikalian Cladophorales to infer the processes shaping the evolution of the group. First, a set of microsatellites was designed using high-throughput sequencing data. Second, species delimitation methods based on genetic clustering were performed. Third, the population genetics of three widely distributed species was analysed looking for evidence of panmixia, a common criteria to support sympatric speciation. A total of 11 microsatellites that mostly cross-amplify between most species were obtained. The genotyping revealed that most loci had more than two alleles per individual indicating polyploidy. As such, the analyses required a different approach which consisted in coding the genotypes as ‘allelic phenotypes’, allowing the use of individuals of different ploidy levels in the same data set. The species delimitation of 15 operative morphotaxa and 727 individuals supported reproductive isolation of five morphotaxa and two hypotheses of conspecificity. However, some morphotaxa showed unclear assignments revealing the need of further research to clarify their reproductive limits. Finally, the population genetics of Chaetomorpha moniliformis, Cladophora compacta and Cl. kursanovii revealed patterns of genetic variation and structure that suggest different reproductive strategies and dispersal abilities. This demonstrates that contrasting biological characteristics may arise in closely related lineages: Chaetomorpha moniliformis with dominant asexual reproduction and long dispersal abilities; Cladophora compacta with high genetic diversity, no population structure and likely to reproduce sexually; Cl. kursanovii with a structure congruent with geographic distribution and more restricted dispersal. The results suggest that polyploidy, rather than speciation with gene flow, is the force driving the reproductive isolation and evolution of this flock. Although many questions remain to be studied, this research provides the first insights into the diversification of this Cladophorales species flock and contributes to the understanding of speciation in freshwater algae.</p>

Context-specific behavior serves as a mechanism of interspecific cohesion in mixed-species flocks

We investigated whether context-specific behavior is responsible for the cohesion of mixed-species flocks of antshrikes and antwrens in Amazonian Peru. Antshrikes perform a behavior while spatially repositioning, to which antwrens respond by approaching. The cohesion of interspecific associations requires communication, although the mechanisms often are unexplored. In monospecific groups, cohesion among individuals is maintained with actions or vocalizations given in a certain context. Dusky-throated Antshrikes (Thamnomanes ardesiacus) vocalize while in flight and the number of times they vocalize covaries with the flight distance. We refer to this pairing of flight and vocalization as repositioning behavior. Furthermore, antshrikes pair a different call type with perching, which we refer to as perching behavior. We followed Long-winged (Myrmotherula longipennis) and White-flanked Antwrens (M. axillaris) and recorded their response following natural vocalizations (no playback used) given by the antshrikes. Long-winged Antwrens, but not White-flanked, flew toward an antshrike significantly sooner and were more likely to approach the antshrikes after the repositioning behavior than after perching behavior. In addition, Long-winged Antwrens, but not White-flanked, flew toward an antshrike sooner after a longer series of repositioning calls than after a shorter series. We did not distinguish between the Long-winged Antwrens’ response as a function of movement vs. vocalizations of the antshrikes, although we argue that vocalizations are likely a more important communication component of repositioning behavior than movement. It remains unclear whether the antshrikes are deliberately signaling the Long-winged Antwrens or the antwrens are taking advantage of the repositioning behavior; active signaling is possible since antshrikes benefit from antwrens. White-flanked Antwrens may be less responsive to the antshrikes since they have a lower propensity to occur in a mixed-species flock. The results indicate that the repositioning behavior of Dusky-throated Antshrikes is a key mechanism of interspecific cohesion of Amazonian mixed-species flocks of the understory.

The Hyalella species flock of Lake Titicaca (Crustacea: Amphipoda): perspectives and drawbacks of dna-based identification

Lake Titicaca, in the High Andes of Perú and Bolivia, harbours the world’s third most speciose ancient-lake amphipod radiation on record. A minimum of nineteen species of Hyalella derived from at least five independent colonization episodes concentrate in this high altitude water body, although the actual species number present has not yet been established and could be much higher. Herein, we take advantage of the description of three new species (H. krolli, H. gonzalezi, and H. hirsuta) and the re-description of other two (H. solida and H. nefrens) to assess the feasibility of adopting a dna-based identification approach to resolve the magnitude of this highly speciose amphipod assemblage. A Bayesian phylogenetic analysis of the evolutionary relationships among South American Hyalella cox1 haplotypes, including those of four out of the five species dealt with herein, shows a great disagreement between taxonomic units delimited under morphological and genetic data, hampering species identification exclusively based on cox1 dna barcode sequences.

Disturbed Amazonian forests support diminished breeding bird communities

Twenty percent of the Brazilian Amazon has now been deforested, and deforestation rates are increasing. This compels us to evaluate the conservation potential of human-modified landscapes; yet the ecological value of regenerating and fragmented Amazonian forests remains poorly understood. To date, most faunal studies in disturbed forests have examined metrics derived from presence or abundance. Although valuable, these data cannot tell us how old-growth species are using disturbed forests. In this study, we complement presence data with breeding observations to assess more comprehensively the habitat quality of disturbed forests in central Amazonia. We synthesized breeding evidence from standardized passive mist-netting, mixed-species flock-following, and opportunistic nest-searching across a full annual cycle in 30–35-year-old secondary forest, 10-ha fragments, and undisturbed forest. We then used multistate occupancy models to estimate the number of undetected breeding species in each forest type, which illustrated that fewer species bred in secondary forest (–43%) and 10-ha fragments (–17%) than in undisturbed forest. Both these losses are larger than the associated decrease in species richness alone (–17% and –10%, respectively). Notably, we confirmed breeding by only three terrestrial and near-ground insectivores in fragments and secondary forest combined (of the 9 species found in undisturbed forest). Disturbed forests also supported fewer breeding individuals (–35% to –50%) and, in secondary forest, fewer successful breeding attempts (–24%). Encouragingly, however, some forest-dependent birds are breeding and producing fledglings in disturbed forests, including representatives from almost every guild. This was especially apparent for mixed-species flocks and army-ant followers, two guilds that have historically been considered vulnerable to anthropogenic disturbance. Therefore, despite a loss of breeding habitat in disturbed forests, these data suggest that landscapes composed of regenerating forest and small fragments have conservation potential for forest bird populations.

Effects of foraging site distances on the intestinal bacterial community compositions of the sympatric wintering Hooded Crane (Grus monacha) and Domestic Duck (Anas platyrhynchos domesticus)

Background The composition of intestinal microflora in animals is affected by cross-species transmission. In a nature reserve, the foraging sites of waterbirds are relatively fixed, but frequently close to residential areas and can also be visited by domestic fowls. It is easy to result in the trans-species-flock dispersal of gut microbes between the wild birds and domestic fowls. The effects of the variable foraging site distances on the gut microbe structures of the waterbirds and the sympatric domestic fowls are currently unclear, and further research is required to evaluate the impacts of geographic location on cross-infection. Methods Illumina high-throughput sequencing and bioinformatics analysis software were utilized to compare and analyze the composition of gut microbes from the fecal samples of Hooded Cranes (HC; Grus monacha) and two groups of Domestic Ducks (Anas platyrhynchos domesticus) that foraged at 1 km (ducks in near areas, D-N), and 4 km (ducks in far areas, D-F) away from the habitats of the Hooded Cranes at Shengjin Lake, China. Results The results showed that there were significant differences in the alpha-diversity of the gut bacteria in the HC, D-N, and D-F samples under the interspecific distance factor. The dominant bacterial phyla, Cyanobacteria and Proteobacteria, showed correlations with distance for each host. The D-N group had more diverse intestinal flora than the D-F, as they were physically closer to the HC and had more indirect contact and cross-transmission of their gut microbes. More potentially pathogenic bacterial sequences, and Operational Taxonomic Units (OTUs) were found in the D-N than in HC and D-F. Conclusions Hooded Cranes and the Domestic Duck populations at variable distances from the cranes showed significant differences in their intestinal bacteria and potentially pathogenic bacteria. The closer the foraging sites were, the easier the intestinal flora spread across species. The results provide a basis for determining the safe distance between wild birds and domestic fowls in a nature reserve.

The genetic population structure of Lake Tanganyika's Lates species flock, an endemic radiation of pelagic top predators

Life history traits are important factors in shaping gene flow within species, and these traits can thus be determinants of whether a species exhibits genetic homogeneity across its range, or considerable population structure. Furthermore, understanding genetic connectivity plays crucially into species conservation decisions, and genetic connectivity is an important component of modern fisheries management in fishes exploited for human consumption. In this study, we investigated the four endemic Lates species of Lake Tanganyika (Lates stappersii, L. microlepis, L. mariae and L. angustifrons), sampled along the Tanzanian shoreline, using reduced-representation genomic sequencing methods. Based on previous studies, we predicted little genetic population structure in the entirely pelagic L. stappersii and the predominantly pelagic L. microlepis. In contrast, we expected the highest genetic differentiation in populations of L. mariae, which is the most resident and for which females aggregate over spawning grounds. We predicted that L. angustifrons would show intermediate structure. We indeed find the most strongly differentiated genetic clusters in L. mariae. However, contrary to our predictions, we find evidence for genetically distinct (albeit weakly differentiated) groups within the pelagic L. stappersii and L. microlepis, and no genetic structure in L. angustifrons. We call for management approaches accounting for genetically differentiated populations of L. stappersii, L. microlepis, and L. mariae, which are commercially important species in the region.

DNA barcoding in recognition of Gammarusflock diversity and distribution in the ancient Lake Ohrid

Lake Ohrid, located on the Balkan Peninsula at the Albanian-Macedonian border, is the oldest European lake (1.3-1.9 My old) and one of the world’s smallest ancient lakes. Taking into account the size of the lake and its biodiversity, it harbors the highest level of endemism, especially within amphipod crustaceans (ca. 90%) with the endemic Gammarus species flock. Our previous studies upon this flock have shown a substantial decoupling between molecular and morphological diversity, existence of cryptic species and puzzling speciation history. In order to explore sources of observed diversity, in the current study we are investigating ecological preferences of the species within the flock, based on their distribution in depth gradient, in relation to molecular diversity based on DNA barcoding. In the study over 200 barcodes were generated and combined with 173 previously published. The specimens were collected from all depth ranges of Lake Ohrid as well as from springs located on or near the banks of the lake. Within the species flock, 13 BIN’s were identified, 12 previously known and one newly recognized, representing separate lineage and putatively a new species. Two of the flock species were found only in the springs: G. sketi and G. cryptosalemaai. G. sketi, previously found only in springs on the southern banks of Lake Ohrid, has now also been discovered in springs in its north-eastern part. Both species show low haplotypic diversity. All remaining species were recorded from the depth between 20 and 60 meters, that is characterized by the highest ecological diversity with different types of substrates: stones, macrophytes, abundant Dreissena shells as well as sand and silt. Among them G. sywulai, G. macedonicus, G. cryptoparechiniformis, G. lychnidensis, G. ochridensis, G. parechinifromis were found exclusively within this depth range. The three latter species represent single BIN and share haplotypes, at the same time this BIN has the highest number of haplotypes in comparison to others. The remaining species found on this depth represent separate BINs with different levels of haplotype diversity. Only G. lychnidensis, G. stankokaramani and G. solidus were found below the depth of 60 meters, in a quite homogenous environment dominated by silt. In the deepest parts of the lake, between 260 and 290 meters, only G. solidus was found. This species is represented only by three haplotypes while G. stankokaramani is characterized by multiple haplotypes partially shared with G. lychnidensis. The shared haplotype represents the only G. lychnidensis occurrences on the depths below 60 meters. Summarizing, the highest abundance of BINs, species and haplotypes was recovered from the most ecologically diversified depth range of the lake (20 to 60 meters). This suggests that ecological heterogeneity could be the main driver of Gammarus species flock diversification in the ancient Lake Ohrid. Due to the complex pattern of morphological diversity, DNA barcoding proved to be the best if not the only method in identification of the species flock diversity.