Ecological divergence and speciation in common bottlenose dolphins in the western South Atlantic

Coastal and offshore ecotypes of common bottlenose dolphins have been recognized in the western South Atlantic, and it is possible that trophic niche divergence associated with social interactions is leading them to genetic and phenotypic differentiation. The significant morphological differentiation observed between these ecotypes suggests they represent two different subspecies. However, there is still a need to investigate whether there is congruence between morphological and genetic data to rule out the possibility of ecophenotypic variation accompanied by gene flow. Mitochondrial DNA (mtDNA) control region sequence data and 10 microsatellite loci collected from stranded and biopsied dolphins sampled in coastal and offshore waters of Brazil as well as 106 skulls for morphological analyses were used to determine whether the morphological differentiation was supported by genetic differentiation. There was congruence among the data sets, reinforcing the presence of two distinct ecotypes. The divergence may be relatively recent, however, given the moderate values of mtDNA nucleotide divergence (dA = 0.008), presence of one shared mtDNA haplotype and possibly low levels of gene flow (around 1% of migrants per generation). Results suggest the ecotypes may be in the process of speciation and reinforce they are best described as two different subspecies until the degree of nuclear genetic divergence is thoroughly evaluated: Tursiops truncatus gephyreus (coastal ecotype) and T. t. truncatus (offshore ecotype). The endemic distribution of T. t. gephyreus in the western South Atlantic and number of anthropogenic threats in the area reinforces the importance of protecting this ecotype and its habitat.     

Isotopic niche width differentiation between common bottlenose dolphin ecotypes and sperm whales in the Gulf of California

World populations or stock distinction of Tursiops truncatus has been difficult to assess, because of large variations in morphology, habitat, feeding habits, and social structure among areas, which may reflect phylogenetic segregation or ecological plasticity. In the Gulf of California, Mexico, two common bottlenose dolphin ecotypes (inshore and offshore) have been reported. The offshore ecotype is frequently observed in association with sperm whales (Physeter macrocephalus) but the reason for this is still unknown. To explore the degree of resource partitioning/overlap between these species and stocks, we used skin stable isotope values (δ¹³C, δ¹⁵N) to estimate quantitative metrics of isotopic niche width (Bayesian standard ellipse areas, SEA_B) and estimated their diet composition using Bayesian isotopic mixing models. The inshore ecotype in different regions (north, central, and south) of the Gulf of California exhibited distinct δ¹⁵N values and SEA_B, suggesting a latitudinal gradient in nitrogen sources of coastal localities. The SEA_B of inshore and offshore bottlenose dolphin ecotypes was completely distinct, indicating resource partitioning. Associated offshore ecotype and sperm whales had overlapping SEA_B. The isotopic mixing model indicates that a considerable proportion of both species’ diet is large Humbolt squid. Our results suggest that resource partitioning and species association are two strategies that bottlenose dolphin ecotypes use in this zone.     

Genetic differentiation among coastal and offshore common bottlenose dolphins,Tursiops truncatus,in the eastern North Pacific Ocean

Common bottlenose dolphins (Tursiops truncatus) are found worldwide in temperate and tropical regions, often occurring as distinct coastal and offshore ecotypes. Along the west coast of the United States, two stocks are recognized for management based on morphological and photo‐identification studies: a California coastal stock, estimated at 450–500 individuals, and a California/Oregon/Washington offshore stock of about 1,000 animals. This study is the first to analyze genetic differentiation between these stocks. We examined both the hypervariable portion of the mitochondrial DNA (mtDNA) control region and fifteen microsatellite markers for coastal (n = 64) and offshore (n = 69) dolphins. Significant genetic differentiation was found between the two stocks for mtDNA (Φ_ST = 0.30, P < 0.001; F_ST = 0.14, P < 0.001) and microsatellite loci (F_ST = 0.19, P < 0.001). Coastal dolphins had less genetic diversity than offshore dolphins. Further substructuring within the offshore stock was not detected. The level of genetic differentiation between the coastal and offshore dolphins is consistent with long‐term separation and reinforces recognizing them as separate stocks. These findings are particularly important for management of the smaller, less genetically diverse, coastal stock that is vulnerable to a variety of anthropogenic threats.     

Occurrence,site fidelity,and associations of oceanic common bottlenose dolphins (Tursiops truncatus) off northeastern New Zealand

Two ecotypes of the common bottlenose dolphin (Tursiops truncatus) occur in New Zealand waters: a widely studied Nationally Endangered coastal ecotype and a little-known oceanic ecotype. Site fidelity and association patterns of the oceanic ecotype, and home range overlap with the coastal ecotype, are examined from photo-identification records collected off northeastern New Zealand between 2005 and 2016. The oceanic ecotype occurs widely in the study area: distance from shore ranged from <1 to ~150 km and home ranges of the two ecotypes overlap in some areas. Forty-nine percent of the 478 identified distinctive or very distinctive individuals were sighted during more than 1 year and resightings spanned over 10 years and 650 km. All individuals were linked by association in a single, albeit clustered, social network. Unlike the coastal ecotype, interspecific associations with false killer (Pseudorca crassidens) and southern long-finned pilot whales (Globicephala melas edwardii) were frequent, occurring during 84% of encounters. Only one oceanic individual matched any of the individuals from the coastal ecotype photo-identification catalogues throughout the study area, suggesting that the two ecotypes co-occur parapatrically. We recommend that the two ecotypes be considered independent management units for conservation purposes due to their divergent ecologies.     

Cranial variation in bottlenose dolphins<Emphasis Type="Italic">Tursiops spp.</Emphasis> from the Indian andwestern Pacific Oceans: additional evidence for two species

We examined the skulls of 72 bottlenose dolphins from the Indian and western Pacific Oceans to clarify the systematics in genusTursiops. We also examined type skulls ofTursiops aduncus (Ehrenberg, 1832),T. eurynome (Gray, 1846),T. catalania (Gray, 1862),Delphinus (Steno) gadamu Owen 1866,T. dawsoni Lydekker 1908, andT. truncatus (Montagu, 1821). The specimens were divided into two groups, A and B, based on rostrum length and characters of the basal part of the rostrum. The rostrum of group A was longer and tapered abruptly near the base, whereas that of group B was shorter and tapered more gradually throughout its length. Group A contained specimens from western, central, and eastern Asia, Africa, and Australasia. Group B contained specimens from central and eastern Asia and Australasia. The six type skulls were also clearly divided into two groups; the type skulls ofT. aduncus, T. catalania, andDelphinus (Steno) gadamu were included in group A, whereas the type skulls ofT. truncatus, T. eurynome, andT. dawsoni were included in group B. This grouping was consistent with the genetic status. Therefore, we identified group A specimens asT. aduncus and group B specimensT. truncatus, the oldest scientific names related to the two groups.     

A comparative study of allozyme variation of peripheral and central populations of Silene nutans L. (Caryophyllaceae) from Western Europe: implications for conservation

In Belgium, at the western border of its continental distribution range, the perennial herb S. nutans has evolved two parapatric ecotypes (calcicolous or silicicolous), which show contrasted allozyme, morphological and adaptive patterns, and isolating reproductive barriers. We examined allozyme variation in 21 populations close to Belgium (from France and Luxemburg) in relation to their peripheral or central geographical position, and investigated their genetic relationships with the Belgian ecotypes. Both peripheral and central populations showed high genetic variation. They were differentiated from each other, but not in relation to the substrate (calcareous or siliceous). The peripheral and central populations were related to the Belgian silicicolous and calcicolous ecotype, respectively. This suggests an ancient differentiation and different past histories for the Belgian ecotypes, and parapatric speciation processes. This study exemplifies the high evolutionary potential of populations at range peripheries and the need for considering them in conservation strategies.     

Habitat‐driven population structure of bottlenose dolphins,Tursiops truncatus,in the North‐East Atlantic

Despite no obvious barrier to gene flow, historical environmental processes and ecological specializations can lead to genetic differentiation in highly mobile animals. Ecotypes emerged in several large mammal species as a result of niche specializations and/or social organization. In the North‐West Atlantic, two distinct bottlenose dolphin (Tursiops truncatus) ecotypes (i.e. ‘coastal’ and ‘pelagic’) have been identified. Here, we investigated the genetic population structure of North‐East Atlantic (NEA) bottlenose dolphins on a large scale through the analysis of 381 biopsy‐sampled or stranded animals using 25 microsatellites and a 682‐bp portion of the mitochondrial control region. We shed light on the likely origin of stranded animals using a carcass drift prediction model. We showed, for the first time, that coastal and pelagic bottlenose dolphins were highly differentiated in the NEA. Finer‐scale population structure was found within the two groups. We suggest that distinct founding events followed by parallel adaptation may have occurred independently from a large Atlantic pelagic population in the two sides of the basin. Divergence could be maintained by philopatry possibly as a result of foraging specializations and social organization. As coastal environments are under increasing anthropogenic pressures, small and isolated populations might be at risk and require appropriate conservation policies to preserve their habitats. While genetics can be a powerful first step to delineate ecotypes in protected and difficult to access taxa, ecotype distinction should be further documented through diet studies and the examination of cranial skull features associated with feeding.     

Evidence for the genetic basis and epistatic interactions underlying ocean‐ and river‐maturing ecotypes of Pacific Lamprey (Entosphenus tridentatus) returning to the Klamath River,California

Surveys of genomic variation have improved our understanding of the relationship between fitness‐related phenotypes and their underlying genetic basis. In some cases, single large‐effect genes have been found to underlie important traits; however, complex traits are expected to be under polygenic control and elucidation of multiple gene interactions may be required to fully understand the genetic basis of the trait. In this study, we investigated the genetic basis of the ocean‐ and river‐maturing ecotypes in anadromous Pacific lamprey (Entosphenus tridentatus). In Pacific lamprey, the ocean‐maturing ecotype is distinguished by advanced maturity of females (e.g., large egg mass) at the onset of freshwater migration relative to immature females of the river‐maturing ecotype. We examined a total of 219 adult Pacific lamprey that were collected at‐entry to the Klamath River over a 12‐month period. Each individual was genotyped at 308 SNPs representing known neutral and adaptive loci and measured at morphological traits, including egg mass as an indicator of ocean‐ and river‐maturing ecotype for females. The two ecotypes did not exhibit genetic structure at 148 neutral loci, indicating that ecotypic diversity exists within a single population. In contrast, we identified the genetic basis of maturation ecotypes in Pacific lamprey as polygenic, involving two unlinked gene regions that have a complex epistatic relationship. Importantly, these gene regions appear to show stronger effects when considered in gene interaction models than if just considered additive, illustrating the importance of considering epistatic effects and gene networks when researching the genetic basis of complex traits in Pacific lamprey and other species.     

Genetic divergence between two phenotypically distinct bottlenose dolphin ecotypes suggests separate evolutionary trajectories

Due to their worldwide distribution and occupancy of different types of environments, bottlenose dolphins display considerable morphological variation. Despite limited understanding about the taxonomic identity of such forms and connectivity among them at global scale, coastal (or inshore) and offshore (or oceanic) ecotypes have been widely recognized in several ocean regions. In the Southwest Atlantic Ocean (SWA), however, there are scarce records of bottlenose dolphins differing in external morphology according to habitat preferences that resemble the coastal‐offshore pattern observed elsewhere. The main aim of this study was to analyze the genetic variability, and test for population structure between coastal (n = 127) and offshore (n = 45) bottlenose dolphins sampled in the SWA to assess whether their external morphological distinction is consistent with genetic differentiation. We used a combination of mtDNA control region sequences and microsatellite genotypes to infer population structure and levels of genetic diversity. Our results from both molecular marker types were congruent and revealed strong levels of structuring (microsatellites F_ST = 0.385, p < .001; mtDNA F_ST =  0.183, p < .001; Φ_ST = 0.385, p < .001) and much lower genetic diversity in the coastal than the offshore ecotype, supporting patterns found in previous studies elsewhere. Despite the opportunity for gene flow in potential “contact zones”, we found minimal current and historical connectivity between ecotypes, suggesting they are following discrete evolutionary trajectories. Based on our molecular findings, which seem to be consistent with morphological differentiations recently described for bottlenose dolphins in our study area, we recommend recognizing the offshore bottlenose dolphin ecotype as an additional Evolutionarily Significant Unit (ESU) in the SWA. Implications of these results for the conservation of bottlenose dolphins in SWA are also discussed.     

Size selection by a gape‐limited predator of a marine snail: Insights into magic traits for speciation

The intertidal snail Littorina saxatilis has repeatedly evolved two parallel ecotypes assumed to be wave adapted and predatory shore crab adapted, but the magnitude and targets of predator‐driven selection are unknown. In Spain, a small, wave ecotype with a large aperture from the lower shore and a large, thick‐shelled crab ecotype from the upper shore meet in the mid‐shore and show partial size‐assortative mating. We performed complementary field tethering and laboratory predation experiments; the first set compared the survival of two different size‐classes of the crab ecotype while the second compared the same size‐class of the two ecotypes. In the first set, the large size‐class of the crab ecotype survived significantly better than the small size‐class both on the upper shore and in the laboratory. In the second set, the small size‐class of the crab ecotype survived substantially better than that of the wave ecotype both on the upper shore and in the laboratory. Shell‐breaking predation on tethered snails was almost absent within the lower shore. In the laboratory shore crabs (Pachygrapsus marmoratus) with larger claw heights selected most strongly against the small size‐class of the crab ecotype, whereas those with medium claw heights selected most strongly against the thin‐shelled wave ecotype. Sexual maturity occurred at a much larger size in the crab ecotype than in the wave ecotype. Our results showed that selection on the upper shore for rapid attainment of a size refuge from this gape‐limited predator favors large size, thick shells, and late maturity. Model parameterization showed that size‐selective predation restricted to the upper shore resulted in the evolution of the crab ecotype despite gene flow from the wave ecotype snails living on the lower shore. These results on gape‐limited predation and previous ones showing size‐assortative mating between ecotypes suggest that size may represent a magic trait for the thick‐shelled ecotype.     

MULTIVARIATE ANALYSIS OF SELECTION IN EXPERIMENTAL POPULATIONS DERIVED FROM HYBRIDIZATION OF TWO ECOTYPES OF THE ANNUAL PLANT DIODIA TERES W. (RUBIACEAE)

Morphologically variable F₂ genotypes derived from hybridization of coastal and inland ecotypes of the annual plant Diodia teres were used to identify selection on morphological traits in the natural habitat of each ecotype. These ecotypes occur in very different habitats, and have evolved pronounced morphological differentiation. Selection analysis can suggest whether present patterns of selection can explain morphological differences between ecotypes. F₂ genotypes were characterized morphologically, clonally replicated, and transplanted into the habitat of each ecotype. Selection was measured on six morphological traits. Directional and stabilizing selection occurred on many traits; direction and strength of selection varied sharply at different stages of growth, as revealed by a path-analysis approach that divided selection into a set of independent components. Directional selection favored traits of the native population at the coastal habitat, but less so at the inland habitat. Selection was of sufficient strength to create the observed morphological differences between ecotypes in 25–100 generations, given constant selection and sufficient genetic variation. In effects on fitness, most traits were neither independent nor consistently interactive with other traits. Rather, many traits entered into strong but evanescent interactions affecting particular components of fitness. Observed interactions did not support the hypothesis that the morphology of each ecotype was functionally integrated to a high degree.     

Lung size and thoracic morphology in shallow‐ and deep‐diving cetaceans

Shallow‐diving, coastal bottlenose dolphins (Tursiops truncatus) and deep‐diving, pelagic pygmy and dwarf sperm whales (Kogia breviceps and K. sima) will experience vastly different ambient pressures at depth, which will influence the volume of air within their lungs and potentially the degree of thoracic collapse they experience. This study tested the hypotheses that lung size will be reduced and/or thoracic mobility will be enhanced in deeper divers. Lung mass (T. truncatus, n = 106; kogiids, n = 18) and lung volume (T. truncatus, n = 5; kogiids, n = 4), relative to total body mass, were compared. One T. truncatus and one K. sima were cross‐sectioned to calculate lung, thoracic vasculature, and other organ volumes. Excised thoraxes (T. truncatus, n = 3; kogiids, n = 4) were mechanically manipulated to compare changes in thoracic cavity shape and volume. Kogiid lungs were half the mass and one‐fifth the volume of those of similarly sized T. truncatus. The lungs occupied only 15% of the total thoracic cavity volume in K. sima and 37% in T. truncatus. The kogiid and dolphin thoraxes underwent similar changes in shape and volume, although the width of the thoracic inlet was relatively constrained in kogiids. A broader phylogenetic comparison demonstrated that the ratio of lung mass to total body mass in kogiids, physeterids, and ziphiids was similar to that of terrestrial mammals, while delphinids and phocoenids possessed relatively large lungs. Thus, small lung size in deep‐diving odontocetes may be a plesiomorphic character. The relatively large lung size of delphinids and phocoenids appears to be a derived condition that may permit the lung to function as a site of respiratory gas exchange throughout a dive in these rapid breathing, short‐duration, shallow divers. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Common bottlenose dolphins (Tursiops truncatus) in California waters: Cranial differentiation of coastal and offshore ecotypes

Coastal and offshore bottlenose dolphins in California waters are currently assessed and managed as separate stocks. Recent molecular studies (of mtDNA haplotypes and microsatellites) have shown the two populations to be genetically differentiated. This study investigated cranial osteological differentiation of the forms. The sample analyzed included 139 skulls from live captures, direct takes, fishery bycatch, and strandings; the skulls were assigned to form based on collection locality or mtDNA haplotype. The coastal form differs from the offshore form mainly in features associated with feeding: larger and fewer teeth, more robust rostrum, larger mandibular condyle, and larger temporal fossa. This suggests that it may feed on larger and tougher prey than the offshore form. Differences between the forms in other features of the skull may reflect differences in diving behavior and sound production. Approximately 86% of the stranded specimens were estimated to be of coastal origin; based on relative estimated sizes of the two populations and assuming similar mortality rates, this suggests that a coastal carcass is about 50 times more likely to beach than an offshore one. The morphological differences between the two ecotypes indicate evolutionary adaptation to different environments and emphasize the importance of conserving the relatively small coastal population and its habitat.     

Genetic isolation between coastal and fishery‐impacted,offshore bottlenose dolphin (Tursiops spp.) populations

The identification of species and population boundaries is important in both evolutionary and conservation biology. In recent years, new population genetic and computational methods for estimating population parameters and testing hypotheses in a quantitative manner have emerged. Using a Bayesian framework and a quantitative model‐testing approach, we evaluated the species status and genetic connectedness of bottlenose dolphin (Tursiops spp.) populations off remote northwestern Australia, with a focus on pelagic ‘offshore’ dolphins subject to incidental capture in a trawl fishery. We analysed 71 dolphin samples from three sites beyond the 50 m depth contour (the inshore boundary of the fishery) and up to 170 km offshore, including incidentally caught and free‐ranging individuals associating with trawl vessels, and 273 dolphins sampled at 12 coastal sites inshore of the 50 m depth contour and within 10 km of the coast. Results from 19 nuclear microsatellite markers showed significant population structure between dolphins from within the fishery and coastal sites, but also among dolphins from coastal sites, identifying three coastal populations. Moreover, we found no current or historic gene flow into the offshore population in the region of the fishery, indicating a complete lack of recruitment from coastal sites. Mitochondrial DNA corroborated our findings of genetic isolation between dolphins from the offshore population and coastal sites. Most offshore individuals formed a monophyletic clade with common bottlenose dolphins (T. truncatus), while all 273 individuals sampled coastally formed a well‐supported clade of Indo‐Pacific bottlenose dolphins (T. aduncus). By including a quantitative modelling approach, our study explicitly took evolutionary processes into account for informing the conservation and management of protected species. As such, it may serve as a template for other, similarly inaccessible study populations.     

IMPROVING MANAGEMENT OF OVERLAPPING BOTTLENOSE DOLPHIN ECOTYPES THROUGH SPATIAL ANALYSIS AND GENETICS

In the Northwest Atlantic the distribution of coastal bottlenose dolphins ( Tursiops truncatus ) overlaps with that of the offshore ecotype. We hypothesized that the distribution of the two ecotypes could be delineated by depth and/or distance from shore, facilitating their identification during surveys. We obtained 304 skin biopsy samples and identified each as either coastal or offshore using analysis of mitochondrial DNA. We then interpreted the spatial distribution of coastal and offshore forms using spatial analysis. Using a Classification and Regression Tree (CART) analysis, we found a statistically significant break in ecotype distribution at 34 km from shore. In waters beyond 34 km from shore and deeper than 34 m, all bottlenose dolphins were of the offshore ecotype. Within 7.5 km of shore, all 65 samples were of the coastal ecotype. Between these two areas only nine samples were collected, so the genetic composition of bottlenose dolphins in this area remains poorly known. To enhance our understanding of the spatial distribution of the two ecotypes, future research should obtain more biopsy samples in this zone. Nevertheless, our results indicate that a conservative abundance estimate for the coastal ecotype could be generated from surveys of bottlenose dolphins within 7.5 km of shore.     

THE DIETS OF MODERN AND HISTORIC BOTTLENOSE DOLPHIN POPULATIONS REFLECTED THROUGH STABLE ISOTOPES

The δ¹³C and δ¹⁵N compositions of teeth used in combination with existing data provide dietary information for different populations of western North Atlantic bottlenose dolphins (Tursiops truncatus). The dental isotopic signatures of bottlenose dolphins collected during the 1980s significantly differ for coastal and offshore ecotypes and are consistent with reports that coastal forms feed primarily on fish whereas offshore individuals consume more squid. In a second study, the isotopic compositions of teeth from bottlenose dolphins that span a 100-yr period and data from published stomach content analyses as well as field observations made during the past 100 yr provide evidence that coastal bottlenose dolphins from the 1880s, 1920s, and 1980s had similar diets.     

Mismatch in the distribution of floral ecotypes and pollinators: insights into the evolution of sexually deceptive orchids

Plants are predicted to show floral adaptation to geographic variation in the most effective pollinator, potentially leading to reproductive isolation and genetic divergence. Many sexually deceptive orchids attract just a single pollinator species, limiting opportunities to experimentally investigate pollinator switching. Here, we investigate Drakaea concolor, which attracts two pollinator species. Using pollinator choice tests, we detected two morphologically similar ecotypes within D. concolor. The common ecotype only attracted Zaspilothynnus gilesi, whereas the rare ecotype also attracted an undescribed species of Pogonothynnus. The rare ecotype occurred at populations nested within the distribution of the common ecotype, with no evidence of ecotypes occurring sympatrically. Surveying for pollinators at over 100 sites revealed that ecotype identity was not correlated with wasp availability, with most orchid populations only attracting the rare Z. gilesi. Using microsatellite markers, genetic differentiation among populations was very low (G_ST = 0.011) regardless of ecotype, suggestive of frequent gene flow. Taken together, these results may indicate that the ability to attract Pogonothynnus has evolved recently, but this ecotype is yet to spread. The nested distribution of ecotypes, rather than the more typical formation of ecotypes in allopatry, illustrates that in sexually deceptive orchids, pollinator switching could occur throughout a species' range, resulting from multiple potentially suitable but unexploited pollinators occurring in sympatry. This unusual case of sympatric pollinators highlights D. concolor as a promising study system for further understanding the process of pollinator switching from ecological, chemical and genetic perspectives.     

Symbiont‐conferred reproduction and fitness benefits can favour their host occurrence

Double infections of Wolbachia and Spiroplasma are frequent in natural populations of Tetranychus truncatus, a polyphagous mite species that has been a dominant species in China since 2009. However, little is known about the causes and ecological importance of such coexistences. In this study, we established T. truncatus strains with different infection types and then inferred the impact of the two endosymbionts on host reproduction and fitness. Double infection induced cytoplasmic incompatibility, which was demonstrated by reduction in egg hatchability of incompatible crosses. However, doubly infected females produced more eggs relative to other strains. Wolbachia and Spiroplasma did not affect host survival, whereas doubly infected females and males developed faster than other strains. Such reproduction and fitness benefits provided by double infections may be associated with the lower densities of each symbiont, and the quantitative results also confirmed competition between Wolbachia and Spiroplasma in doubly infected females. These symbiont‐conferred beneficial effects maintain stable prevalence of the symbionts and also help drive T. truncatus outbreaks in combination with other environmental factors.     

Skull morphology of bottlenose dolphins from different ocean populations with emphasis on South America

The bottlenose dolphin, genus Tursiops, is cosmopolitan occurring in tropical and temperate regions, with morphological variation between and within different oceans. Since the genus' taxonomy has been under discussion for a long time, this work aimed at analyzing the cranial variability of T. truncatus from different regions of the world. Geometric Morphometrics analyses were performed in 201 skulls of adult specimens, on dorsal, ventral, and lateral views, from the Eastern North Pacific, Eastern North Atlantic, Eastern South Atlantic, and Western South Atlantic oceans. The results indicate differences between individuals that inhabit the Atlantic and Pacific oceans. Within the Atlantic Ocean, there is an evident longitudinal differentiation of specimens from the eastern and western regions. A latitudinal separation was also observed, considering specimens from the North and South Atlantic Ocean. In the Western South Atlantic statistical differences were found between two morphological groups, identified as T. gephyreus (sensu Lahille, 1908) and T. truncatus, and the cross-validation presented 98% as minimum confidence for correct classification of these two groups. The present study provides strong morphological support to consider these two lineages as separate species.     

Hybridization,polyploidy and clonality influence geographic patterns of diversity and salt tolerance in the model halophyte seashore paspalum (Paspalum vaginatum)

In plant species, variation in levels of clonality, ploidy and interspecific hybridization can interact to influence geographic patterns of genetic diversity. These factors commonly vary in plants that specialize on saline habitats (halophytes) and may play a role in how they adapt to salinity variation across their range. One such halophyte is the turfgrass and emerging genomic model system seashore paspalum (Paspalum vaginatum Swartz). To investigate how clonal propagation, ploidy variation, and interspecific hybridization vary across ecotypes and local salinity levels in wild P. vaginatum, we employed genotyping‐by‐sequencing, cpDNA sequencing and flow cytometry in 218 accessions representing > 170 wild collections from throughout the coastal southern United States plus USDA germplasm. We found that the two morphologically distinct ecotypes of P. vaginatum differ in their adaptive strategies. The fine‐textured ecotype is diploid and appears to reproduce in the wild both sexually and by clonal propagation; in contrast, the coarse‐textured ecotype consists largely of clonally‐propagating triploid and diploid genotypes. The coarse‐textured ecotype appears to be derived from hybridization between fine‐textured P. vaginatum and an unidentified Paspalum species. These clonally propagating hybrid genotypes are more broadly distributed than clonal fine‐textured genotypes and may represent a transition to a more generalist adaptive strategy. Additionally, the triploid genotypes vary in whether they carry one or two copies of the P. vaginatum subgenome, indicating multiple evolutionary origins. This variation in subgenome composition shows associations with local ocean salinity levels across the sampled populations and may play a role in local adaptation.