Early Stages Of Diversification Research Articles

Phylogenomics of Neogastropoda: the backbone hidden in the bush.

The molluscan order Neogastropoda encompasses over 15,000 almost exclusively marine species playing important roles in benthic communities and in the economies of coastal countries. Neogastropoda underwent intensive cladogenesis in early stages of diversification, generating a 'bush' at the base of their evolutionary tree, that has been hard to resolve even with high throughput molecular data. In the present study to resolve the bush, we use a variety of phylogenetic inference methods and a comprehensive exon capture dataset of 1,817 loci (79.6% data occupancy) comprising 112 taxa of 48 out of 60 Neogastropoda families. Our results show consistent topologies and high support in all analyses at (super)family level, supporting monophyly of Muricoidea, Mitroidea, Conoidea, and, with some reservations, Olivoidea and Buccinoidea. Volutoidea and Turbinelloidea as currently circumscribed are clearly paraphyletic. Despite our analyses consistently resolving most backbone nodes, three prove problematic: First, uncertain placement of Cancellariidae, as the sister group to either a Ficoidea-Tonnoidea clade, or to the rest of Neogastropoda, leaves monophyly of Neogastropoda unresolved. Second, relationships are contradictory at the base of the major 'core Neogastropoda' grouping. Third, coalescence-based analyses reject monophyly of the Buccinoidea in relation to Vasidae. We analysed phylogenetic signal of targeted loci in relation to potential biases, and we propose most probable resolutions in the latter two recalcitrant nodes. The uncertain placement of Cancellariidae may be explained by orthology violations due to differential paralog loss shortly after the whole genome duplication, which should be resolved with a curated set of longer loci.

Environmental selection, rather than neutral processes, best explain regional patterns of diversity in a tropical rainforest fish

To conserve the high functional and genetic variation in hotspots such as tropical rainforests, it is essential to understand the forces driving and maintaining biodiversity. We asked to what extent environmental gradients and terrain structure affect morphological and genomic variation across the wet tropical distribution of an Australian rainbowfish, Melanotaenia splendida splendida. We used an integrative riverscape genomics and morphometrics framework to assess the influence of these factors on both putative adaptive and non-adaptive spatial divergence. We found that neutral genetic population structure was largely explainable by restricted gene flow among drainages. However, environmental associations revealed that ecological variables had a similar power to explain overall genetic variation, and greater power to explain body shape variation, than the included neutral covariables. Hydrological and thermal variables were the strongest environmental predictors and were correlated with traits previously linked to heritable habitat-associated dimorphism in rainbowfishes. In addition, climate-associated genetic variation was significantly associated with morphology, supporting heritability of shape variation. These results support the inference of evolved functional differences among localities, and the importance of hydroclimate in early stages of diversification. We expect that substantial evolutionary responses will be required in tropical rainforest endemics to mitigate local fitness losses due to changing climates.

탄소 배출에 따른 제조업 집적의 비선형 영향에 관한 연구

Purpose : This study examined the non-linear effect of manufacturing clusters on carbon emissions using the semi-parametric spatial autoregressive model. Research design, data and methodology : This study calculated the manufacturing specialization and diversification agglomeration index according to the region in Korea for 17 regions and 25 manufacturing fields in Korea from 2013 to 2019. Considering the impact of manufacturing agglomeration on carbon emissions, the non-linear effect of manufacturing agglomeration on carbon emissions was examined using a semi-parametric model for the effect of manufacturing agglomeration on carbon emissions. In addition, considering the spatial fluidity of carbon emissions and policy imitation of neighboring regions, this study constructed a semi-parametric spatial autoregressive model considering the spatial interactions between individuals. Results : An analysis of the carbon emission reduction effect according to the manufacturing cluster type showed that, the effect of a specialized cluster on carbon emission was an N-shaped structure. In the early stages of specialization integration, the action of promoting carbon emission is weakened as the level of integration increases and gradually suppresses carbon emission. The effect of the diversification of the manufacturing industry on carbon emissions was found to be a distinct inverted U-shaped structure. In the early stages of diversification and integration, carbon emissions increased strongly, and carbon emissions were gradually suppressed as the level of integration increased, Diversification agglomeration in the manufacturing industry had a greater effect on suppressing carbon emissions than specialization agglomeration. Conclusions : Attention should be paid to the relationship between the expansion of production scale and carbon emission in large and relatively mature manufacturing specialization clusters. The expansion of the production scale mat weaken the eco friendly effect of technological progress. In the early stages of diversification and agglomeration in the manufacturing industry, attention should be paid to improving resource use efficiency and reducing carbon emissions. While diversifying the agglomeration areas, it is necessary to promote the rational use of existing resources by improving public infrastructure within the industrial complex. Comprehensively, policies should be set to maintain the functions of various related industries, enhance the energy efficiency, and lower carbon intensity through process improvement.

Demographic consequences of dispersal-related trait shift in two recently diverged taxa of montane grasshoppers.

Although the pervasiveness of intraspecific wing-size polymorphism and transitions to flightlessness have long captivated biologists, the demographic outcomes of shifts in dispersal ability are not yet well understood and have been seldom studied at early stages of diversification. Here, we use genomic data to infer the consequences of dispersal-related trait variation in the taxonomically controversial short-winged (Chorthippus corsicus corsicus) and long-winged (Chorthippus corsicus pascuorum) Corsican grasshoppers. Our analyses revealed lack of contemporary hybridization between sympatric long- and short-winged forms and phylogenomic reconstructions supported their taxonomic distinctiveness, rejecting the hypothesis of intraspecific wing polymorphism. Statistical evaluation of alternative models of speciation strongly supported a scenario of Pleistocene divergence (<1.5 Ma) with ancestral gene flow. According to neutral expectations from differences in dispersal capacity, historical effective migration rates from the long- to the short-winged taxon were threefold higher than in the opposite direction. Although populations of the two taxa present a marked genetic structure and have experienced parallel demographic histories, our coalescent-based analyses suggest that reduced dispersal has fueled diversification in the short-winged C. c. corsicus. Collectively, our study illustrates how dispersal reduction can speed up geographical diversification and increase the opportunity for allopatric speciation in topographically complex landscapes.

Phylogenomics of the genus Tursiops and closely related Delphininae reveals extensive reticulation among lineages and provides inference about eco-evolutionary drivers

Phylogeographic inference has provided extensive insight into the relative roles of geographical isolation and ecological processes during evolutionary radiations. However, the importance of cross-lineage admixture in facilitating adaptive radiations is increasingly being recognised, and suggested as a main cause of phylogenetic uncertainty. In this study, we used a double digest RADseq protocol to provide a high resolution (~4 Million bp) nuclear phylogeny of the Delphininae. Phylogenetic resolution of this group has been especially intractable, likely because it has experienced a recent species radiation. We carried out cross-lineage reticulation analyses, and tested for several sources of potential bias in determining phylogenies from genome sampling data. We assessed the divergence time and historical demography of T. truncatus and T. aduncus by sequencing the T. aduncus genome and comparing it with the T. truncatus reference genome. Our results suggest monophyly for the genus Tursiops, with the recently proposed T. australis species falling within the T. aduncus lineage. We also show the presence of extensive cross-lineage gene flow between pelagic and European coastal ecotypes of T. truncatus, as well as in the early stages of diversification between spotted (Stenella frontalis; Stenella attenuata), spinner (Stenella longirostris), striped (Stenella coeruleoalba), common (Delphinus delphis), and Fraser’s (Lagenodelphis hosei) dolphins. Our study suggests that cross-lineage gene flow in this group has been more extensive and complex than previously thought. In the context of biogeography and local habitat dependence, these results improve our understanding of the evolutionary processes determining the history of this lineage.

Bridging the Process-Pattern Divide to Understand the Origins and Early Stages of Adaptive Radiation: A Review of Approaches With Insights From Studies of Anolis Lizards.

Understanding the origins and early stages of diversification is one of the most elusive tasks in adaptive radiation research. Classical approaches, which aim to infer past processes from present-day patterns of biological diversity, are fraught with difficulties and assumptions. An alternative approach has been to study young clades of relatively few species, which may represent the putative early stages of adaptive radiation. However, it is difficult to predict whether those groups will ever reach the ecological and morphological disparity observed in the sorts of clades usually referred to as adaptive radiations, thereby making their utility in informing the early stages of such radiations uncertain. Caribbean Anolis lizards are a textbook example of an adaptive radiation; anoles have diversified independently on each of the 4 islands in the Greater Antilles, producing replicated radiations of phenotypically diverse species. However, the underlying processes that drove these radiations occurred 30-65 million years ago and so are unobservable, rendering major questions about how these radiations came to be difficult to tackle. What did the ancestral species of the anole radiation look like? How did new species arise? What processes drove adaptive diversification? Here, we review what we have learned about the cryptic early stages of adaptive radiation from studies of Anolis lizards, and how these studies have attempted to bridge the process-pattern divide of adaptive radiation research. Despite decades of research, however, fundamental questions linking eco-evolutionary processes to macroevolutionary patterns in anoles remain difficult to answer.

Diversity partitioning in Phanerozoic benthic marine communities

Biotic interactions such as competition, predation, and niche construction are fundamental drivers of biodiversity at the local scale, yet their long-term effect during earth history remains controversial. To test their role and explore potential limits to biodiversity, we determine within-habitat (alpha), between-habitat (beta), and overall (gamma) diversity of benthic marine invertebrates for Phanerozoic geological formations. We show that an increase in gamma diversity is consistently generated by an increase in alpha diversity throughout the Phanerozoic. Beta diversity drives gamma diversity only at early stages of diversification but remains stationary once a certain gamma level is reached. This mode is prevalent during early- to mid-Paleozoic periods, whereas coupling of beta and gamma diversity becomes increasingly weak toward the recent. Generally, increases in overall biodiversity were accomplished by adding more species to local habitats, and apparently this process never reached saturation during the Phanerozoic. Our results provide general support for an ecological model in which diversification occurs in successive phases of progressing levels of biotic interactions.

Variation in the magnitude of morphological and dietary differences between individuals among populations of small benthic Arctic charr in relation to ecological factors.

The early stages of intraspecific diversity are important for the evolution of diversification and speciation. Early stages of diversification can be seen in individual specialization, where individuals consume only a portion of the diet of the population as a whole, and how such specialization is related to phenotypic diversity within populations. Here, we study the strength of the relationship between morphological and dietary distances among individuals in eighteen populations of Icelandic small benthic charr. We furthermore studied if the strength of the relationship could be related to variation in local ecological factors these populations inhabit. In all the populations studied, there was a clear relationship between morphological and dietary distances, indicating that fish that had similar morphology were at the same time‐consuming similar food items. Our findings show a systematic variation in the relationship between morphology and diet at early stages of diversification in a highly specialized small benthic charr morph. The results show the importance of fine scale comparisons within populations and furthermore the value that systematic comparisons among populations under parallel evolution can contribute toward our increased understanding of evolutionary and ecological processes.

Positive selection on sperm ion channels in a brooding brittle star: consequence of life-history traits evolution.

Closely related species are key models to investigate mechanisms leading to reproductive isolation and early stages of diversification, also at the genomic level. The brittle star cryptic species complex Ophioderma longicauda encompasses the sympatric broadcast-spawning species C3 and the internal brooding species C5. Here, we used de novo transcriptome sequencing and assembly in two closely related species displaying contrasting reproductive modes to compare their genetic diversity and to investigate the role of natural selection in reproductive isolation. We reconstructed 20146 and 22123 genes for C3 and C5, respectively, and characterized a set of 12229 orthologs. Genetic diversity was 1.5-2 times higher in C3 compared to C5, confirming that species with low parental investment display higher levels of genetic diversity. Forty-eight genes were the targets of positive diversifying selection during the evolution of the two species. Notably, two genes (NHE and TetraKCNG) are sperm-specific ion channels involved in sperm motility. Ancestral sequence reconstructions show that natural selection targeted the two genes in the brooding species. This may result from an adaptation to the novel environmental conditions surrounding sperm in the brooding species, either directly affecting sperm or via an increase in male/female conflict. This phenomenon could have promoted prezygotic reproductive isolation between C3 and C5. Finally, the sperm receptors to egg chemoattractants differed between C3 and C5 in the ligand-binding region. We propose that mechanisms of species-specific gamete recognition in brittle stars occur during sperm chemotaxis (sperm attraction towards the eggs), contrary to other marine invertebrates where prezygotic barriers to interspecific hybridization typically occur before sperm-egg fusion.

The importance (or lack thereof) of niche divergence to the maintenance of a northern species complex: the case of the long-toed salamander (Ambystoma macrodactylum Baird).

The relative importance of ecological vs. non-ecological factors for the origin and maintenance of species is an open question in evolutionary biology. Young lineages--such as the distinct genetic groups that make up the ranges of many northern species--represent an opportunity to study the importance of ecological divergence during the early stages of diversification. Yet, few studies have examined the extent of niche divergence between lineages in previously glaciated regions and the role of ecology in maintaining the contact zones between them. In this study, we used tests of niche overlap in combination with ecological niche models to explore the extent of niche divergence between lineages of the long-toed salamander (Ambystoma macrodactylum Baird) species complex and to determine whether contact zones correspond to (divergent) niche limits. We found limited evidence for niche divergence between the different long-toed salamander lineages, substantial overlap in the predicted distribution of suitable climatic space for all lineages and range limits that are independent of niche limits. These results raise questions as to the importance of ecological divergence to the development of this widespread species complex and highlight the potential for non-ecological factors to play a more important role in the maintenance of northern taxa.

Stratigraphy and paleoenvironments of the early to middle Holocene Chipalamawamba Beds (Malawi Basin, Africa)

Abstract. We describe the Chipalamawamba Beds, early to middle Holocene deposits at the southern margin of long-lived Lake Malawi. The beds are exposed because of downcutting of the upper Shire River. The Chipalamawamba sediments are medium to coarse, yellow to brown sands deposited in lenses varying in horizontal extent from a few meters to several hundreds of meters. Four units are recognized; the first three mainly contain lacustrine sediments deposited during lake high stands about 10.6–9.7 cal ka BP (Unit 1), 7.6–6.5 cal ka BP (Unit 2) and 5.9–5.3 cal ka BP (Unit 3). Sediments of Unit 4 overlay Units 1 to 3, are coarser and display regular foresets and oblique-bedding, suggesting deposition in riverine environments after installation of the Shire River (~ 5.5–5.0 ka BP). Freshwater mollusk assemblages and bioturbation regularly occur in the lacustrine sediments, but are largely absent from Unit 4. Diverse and often contradicting hypotheses on the lake levels of Lake Malawi have been proposed for the early and middle Holocene. The Chipalamawamba Beds allow straightforward recognition of water levels and provide strong evidence for oscillating lake levels during this period, rather than continuous high or low levels. Sedimentation rates have been high and individual shell beds have typically been deposited during a few decades. Because the Chipalamawamba Beds contain a sequence of mollusk assemblages with intervals between subsequent shell beds ranging from a century to a few millennia, they enable paleontological analysis of the fauna with an unusually high temporal resolution. That some mollusk lineages inhabiting Lake Malawi are in the early stages of diversification and radiation increases the paleobiological relevance of these beds.

Eco-Evolutionary Community Dynamics: Covariation between Diversity and Invasibility across Temperature Gradients

Understanding biodiversity gradients is a long-standing challenge, and progress requires theory unifying ecology and evolution. Here, we unify concepts related to the speed of evolution, the influence of species richness on diversification, and niche-based coexistence. We focus on the dynamics, through evolutionary time, of community invasibility and species richness across a broad thermal gradient. In our framework, the evolution of body size influences the ecological structure and dynamics of a trophic network, and organismal metabolism ties temperature to eco-evolutionary processes. The framework distinguishes ecological invasibility (governed by ecological interactions) from evolutionary invasibility (governed by local ecology and constraints imposed by small phenotypic effects of mutation). The model yields four primary predictions: (1) ecological invasibility declines through time and with increasing temperature; (2) average evolutionary invasibility across communities increases and then decreases through time as the richness-temperature gradient flattens; (3) in the early stages of diversification, richness and evolutionary invasibility both increase with increasing temperature; and (4) at equilibrium, richness does not vary with temperature, yet evolutionary invasibility decreases with increasing temperature. These predictions emerge from the "evolutionary-speed" hypothesis, which attempts to account for latitudinal species richness gradients by invoking faster biological rates in warmer, tropical regions. The model contrasts with predictions from other richness-gradient hypotheses, such as "niche conservatism" and "species energy." Empirically testing our model's predictions should help distinguish among these hypotheses.

Distribution of Hordoindoline genes in the genus Hordeum

Hordoindoline (Hin) genes, which are known to comprise Hina, Hinb-1, and Hinb-2, are associated with grain hardness in barley. However, the interspecific variation in the Hin genes in the genus Hordeum has not been studied in detail. We examined the variation in Hin genes and used it to infer the phylogenetic relationships between the genes found in two H. vulgare subspecies (cultivated barley and H. vulgare subsp. spontaneum) and 10 wild relatives (H. bogdanii, H. brachyantherum, H. bulbosum, H. chilense, H. comosum, H. marinum, H. murinum, H. patagonicum, H. pusillum, and H. roshevitzii). The Hina and Hinb genes of these species were amplified by PCR. We found two Hinb genes in three wild species (H. bogdanii, H. brachyantherum, and H. roshevitzii) and preliminarily named them Hinb-A and Hinb-B. Cluster analysis showed that the 17 Hinb genes present in Hordeum formed two distinct clusters (named A and B). Seven Hinb genes were included in Cluster-A, and 10 Hinb genes were included in Cluster-B. All Hinb-A genes were included in Cluster-A, while all of the Hinb-B genes were included in Cluster-B. In contrast, the Hinb-1 and Hinb-2 genes in H. vulgare were included in Cluster-B. These results suggest that the Hinb genes duplicated during the early stages of diversification in the genus Hordeum. On the other hand, the Hinb-1 and Hinb-2 genes in H. vulgare seem to have been generated by a duplication of the Hinb gene after the split of the lineages leading to H. vulgare and H. bulbosum.