Geographic independence and phylogenetic diversity of red shiner introductions

Identifying areas at risk of invasion can be difficult when the distribution of a non-native species encompasses geographically disjunct regions. Understanding genealogical relationships among native and non-native populations can clarify the origins of fragmented distributions, which in turn can clarify how fast and far a non-native species may spread. We evaluated genetic variation across the native and invasive ranges of red shiner (Cyprinella lutrensis), a minnow known to displace and hybridize with native species, to reconstruct invasion pathways across the United States (USA). Examination of mitochondrial cytochrome-b variation found that native range populations of red shiner fall into four highly divergent lineages that likely warrant species recognition. Introduced red shiner populations in the eastern and western USA are derived from only two of these lineages. Western USA populations originate from the mid-western and western genetic lineages, whereas eastern introductions derive only from the mid-western lineage. Western USA invasive populations exhibit fewer, but more diverse haplotypes compared to eastern USA invasive populations. We also recovered an undescribed, divergent lineage of Cyprinella that has been cryptically introduced into the western USA, which raises the possibility that hybridization has proceeded following secondary contact between previously allopatric lineages. Approximate Bayesian Computation modeling suggests that the disjunct distribution of red shiner across North America is an agglomeration of independent regional invasions with distinct origins, rather than stepwise advance of an invasion front or secondary introductions across regions. Thus localized control may be effective in managing non-native red shiner, including further spread to areas of conservation concern.     

Dispersal and selection mediate hybridization between a native and invasive species

Hybridization between native and non-native species has serious biological consequences, but our understanding of how dispersal and selection interact to influence invasive hybridization is limited. Here, we document the spread of genetic introgression between a native (Oncorhynchus clarkii) and invasive (Oncorhynchus mykiss) trout, and identify the mechanisms influencing genetic admixture. In two populations inhabiting contrasting environments, non-native admixture increased rapidly from 1984 to 2007 and was driven by surprisingly consistent processes. Individual admixture was related to two phenotypic traits associated with fitness: size at spawning and age of juvenile emigration. Fish with higher non-native admixture were larger and tended to emigrate at a younger age―relationships that are expected to confer fitness advantages to hybrid individuals. However, strong selection against non-native admixture was evident across streams and cohorts (mean selection coefficient against genotypes with non-native alleles (s) = 0.60; s.e. = 0.10). Nevertheless, hybridization was promoted in both streams by the continuous immigration of individuals with high levels of non-native admixture from other hybrid source populations. Thus, antagonistic relationships between dispersal and selection are mediating invasive hybridization between these fish, emphasizing that data on dispersal and natural selection are needed to fully understand the dynamics of introgression between native and non-native species.     

Hybridization rapidly reduces fitness of a native trout in the wild

Human-mediated hybridization is a leading cause of biodiversity loss worldwide. How hybridization affects fitness and what level of hybridization is permissible pose difficult conservation questions with little empirical information to guide policy and management decisions. This is particularly true for salmonids, where widespread introgression among non-native and native taxa has often created hybrid swarms over extensive geographical areas resulting in genomic extinction. Here, we used parentage analysis with multilocus microsatellite markers to measure how varying levels of genetic introgression with non-native rainbow trout (Oncorhynchus mykiss) affect reproductive success (number of offspring per adult) of native westslope cutthroat trout (Oncorhynchus clarkii lewisi) in the wild. Small amounts of hybridization markedly reduced fitness of male and female trout, with reproductive success sharply declining by approximately 50 per cent, with only 20 per cent admixture. Despite apparent fitness costs, our data suggest that hybridization may spread due to relatively high reproductive success of first-generation hybrids and high reproductive success of a few males with high levels of admixture. This outbreeding depression suggests that even low levels of admixture may have negative effects on fitness in the wild and that policies protecting hybridized populations may need reconsideration.     

A molecular approach to detect hybridisation between crucian carp (Carassius carassius) and non-indigenous carp species (Carassius spp. and Cyprinus carpio)

1. Releases of non‐native fish into the wild is an increasing problem posing considerable ecological and genetic threats through direct competition and hybridisation. 2. We employed six microsatellite markers to identify first generation hybrids and backcrosses between native crucian carp (Carassius carassius) and introduced goldfish (C. auratus) and common carp (Cyprinus carpio) in the U.K. We also investigated the genetic characteristics of the taxonomically controversial gibel carp (Carassius spp.) from sites across Europe. 3. Natural hybridisation between goldfish and crucian carp occurs frequently, although hybrids between all other species pairs were observed. Only 62% of British crucian carp populations (n = 21) consisted exclusively of pure crucian carp. In some populations hybrids were so frequent, that no pure crucian carp were caught, indicating a high competitive ability of hybrids. 4. Most hybrids belonged to the F1 generation but backcrossing was evident at a low frequency in goldfish × crucian carp hybrids and goldfish × common carp hybrids. Furthermore, some local populations had high frequencies of backcrosses, raising the opportunity for introgression. 5. Gibel carp from Germany and Italy belonged to two triploid clonal lineages that were genetically closely related to goldfish, whereas all individuals identified from British populations proved to be crucian carp × goldfish hybrids. 6. Our study suggests that the release of closely related exotic cyprinids not only poses a threat to the genetic integrity and associated local adaptations of native species, but may also contribute to shifts in community structure through competitive interactions.     

Condition status and parasite infection of Neogobius kessleri and N. melanostomus (Gobiidae) in their native and non-native area of distribution of the Danube River

The success of introduced species is often facilitated by escape from the effects of natural predators and parasites. Introduced species can profit from this favourable situation, attaining higher population densities and greater individual sizes in novel areas. In this study, somatic condition and parasite infection were compared between native and non-native populations of Neogobius kessleri Günther; introduced only within the interconnected Danube and Rhine River system, and N. melanostomus (Pallas); widely introduced throughout several river systems in Europe and North America. Higher values of Fulton’s condition factor were observed in non-native populations of both goby species. Neogobius melanostomus attained higher gonadosomatic index values in non-native populations, indicating potential increased investment in reproduction in its new area. A lower splenosomatic index was observed in non-native populations, especially in N. melanostomus. Parasite infracommunity richness and mean abundance were higher in N. kessleri in both native and non-native populations, suggesting higher susceptibility of N. kessleri to these parasites. Non-native populations of both hosts showed higher infra-community richness as a result of acquiring parasites native to the new area, but lower parasite abundance. Differences in success of the introduction and establishment in new areas between the two fish species may be associated with a relatively low parasite infection rate and a higher gonadosomatic index in non-native populations of N. melanostomus in comparison to N. kessleri.     

Red shiner invasion and hybridization with blacktail shiner in the upper Coosa River,USA

Human disturbance increases the invasibility of lotic ecosystems and the likelihood of hybridization between invasive and native species. We investigated whether disturbance contributed to the invasion of red shiner (Cyprinella lutrensis) and their hybridization with native blacktail shiner (C. venusta stigmatura) in the Upper Coosa River System (UCRS). Historical records indicated that red shiners and hybrids rapidly dispersed in the UCRS via large, mainstem rivers since the mid to late 1990s. We measured the occurrence and abundance of parental species and hybrids near tributary-mainstem confluences and characterized populations at these incipient contact zones by examining variation across morphological traits and molecular markers. Red shiners represented only 1.2% of total catch in tributaries yet introgression was widespread with hybrids accounting for 34% of total catch. Occurrence of red shiners and hybrids was highly correlated with occurrence of blacktail shiners, indicating that streams with native populations are preferentially colonized early in the invasion and that hybridization is a key process in the establishment of red shiners and their genome in new habitats. Tributary invasion was driven by post-F1 hybrids with proportionately greater genomic contributions from blacktail shiner. Occurrence of red shiners and hybrids and the relative abundance of hybrids significantly increased with measures of human disturbance including turbidity, catchment agricultural land use, and low dissolved oxygen concentration. Red shiners are a significant threat to Southeast Cyprinella diversity, given that 41% of these species hybridize with red shiner, that five southeastern drainages are invaded, and that these drainages are increasingly disturbed by urbanization.     

Behavioural and genetic interactions between an endangered and a recently-arrived hummingbird

The invasion or expansion of non-native species into new geographic areas can pose a major threat to the conservation of biodiversity. These threats are augmented when the newly-arrived species interacts with native species that are already threatened by other ecological or anthropogenic processes. Potential interactions can include both competition for scarce resources and reproductive interference, including hybridisation. Understanding the dynamics of these interactions forms a crucial component of conservation management strategies. A recent contact zone occurs in the north of Chile between the endangered Chilean woodstar (Eulidia yarrellii) and the closely-related and recently-arrived Peruvian sheartail (Thaumastura cora), which expanded its range from Peru into Chile during the 1970s. We characterised the interactions between the species by combining population size estimates with molecular, morphological and behavioural data. We show that a low degree of hybridisation, but not introgression, is occurring between the two species. Despite interspecific morphological similarities, behavioural observations indicate that food niche overlap between the species is relatively low, and that the dietary breadth of sheartails is larger, which may have aided the species’ range expansion. Finally, woodstars dominate the sheartails in male–male territorial interactions. However, potentially increased energetic costs for woodstars associated with frequent territorial chases and courtship displaying with sheartails may exacerbate the effects of other threats on woodstar viability, such as human-induced habitat modification. This study highlights the value of implementing multidisciplinary approaches in conservation biology to gain a more complete understanding of interactions between recently-arrived and endangered species.     

Human disturbance causes the formation of a hybrid swarm between two naturally sympatric fish species

Most evidence for hybrid swarm formation stemming from anthropogenic habitat disturbance comes from the breakdown of reproductive isolation between incipient species, or introgression between allopatric species following secondary contact. Human impacts on hybridization between divergent species that naturally occur in sympatry have received considerably less attention. Theory predicts that reinforcement should act to preserve reproductive isolation under such circumstances, potentially making reproductive barriers resistant to human habitat alteration. Using 15 microsatellites, we examined hybridization between sympatric populations of alewife (Alosa pseudoharengus) and blueback herring (A. aestivalis) to test whether the frequency of hybridization and pattern of introgression have been impacted by the construction of a dam that isolated formerly anadromous populations of both species in a landlocked freshwater reservoir. The frequency of hybridization and pattern of introgression differed markedly between anadromous and landlocked populations. The rangewide frequency of hybridization among anadromous populations was generally 0–8%, whereas all landlocked individuals were hybrids. Although neutral introgression was observed among anadromous hybrids, directional introgression leading to increased prevalence of alewife genotypes was detected among landlocked hybrids. We demonstrate that habitat alteration can lead to hybrid swarm formation between divergent species that naturally occur sympatrically, and provide empirical evidence that reinforcement does not always sustain reproductive isolation under such circumstances.     

Population-specific responses to an invasive species

Predicting the impacts of non-native species remains a challenge. As populations of a species are genetically and phenotypically variable, the impact of non-native species on local taxa could crucially depend on population-specific traits and adaptations of both native and non-native species. Bitterling fishes are brood parasites of unionid mussels and unionid mussels produce larvae that parasitize fishes. We used common garden experiments to measure three key elements in the bitterling–mussel association among two populations of an invasive mussel (Anodonta woodiana) and four populations of European bitterling (Rhodeus amarus). The impact of the invasive mussel varied between geographically distinct R. amarus lineages and between local populations within lineages. The capacity of parasitic larvae of the invasive mussel to exploit R. amarus was higher in a Danubian than in a Baltic R. amarus lineage and in allopatric than in sympatric R. amarus populations. Maladaptive oviposition by R. amarus into A. woodiana varied among populations, with significant population-specific consequences for R. amarus recruitment. We suggest that variation in coevolutionary states may predispose different populations to divergent responses. Given that coevolutionary relationships are ubiquitous, population-specific attributes of invasive and native populations may play a critical role in the outcome of invasion. We argue for a shift from a species-centred to population-centred perspective of the impacts of invasions.     

Investigating Differences across Host Species and Scales to Explain the Distribution of the Amphibian Pathogen Batrachochytrium dendrobatidis

Many pathogens infect more than one host species, and clarifying how these different hosts contribute to pathogen dynamics can facilitate the management of pathogens and can lend insight into the functioning of pathogens in ecosystems. In this study, we investigated a suite of native and non-native amphibian hosts of the pathogen Batrachochytrium dendrobatidis (Bd) across multiple scales to identify potential mechanisms that may drive infection patterns in the Colorado study system. Specifically, we aimed to determine if: 1) amphibian populations vary in Bd infection across the landscape, 2) amphibian community composition predicts infection (e.g., does the presence or abundance of any particular species influence infection in others?), 3) amphibian species vary in their ability to produce infectious zoospores in a laboratory infection, 4) heterogeneity in host ability observed in the laboratory scales to predict patterns of Bd prevalence in the landscape. We found that non-native North American bullfrogs (Lithobates catesbeianus) are widespread and have the highest prevalence of Bd infection relative to the other native species in the landscape. Additionally, infection in some native species appears to be related to the density of sympatric L. catesbeianus populations. At the smaller host scale, we found that L. catesbeianus produces more of the infective zoospore stage relative to some native species, but that this zoospore output does not scale to predict infection in sympatric wild populations of native species. Rather, landscape level infection relates most strongly to density of hosts at a wetland as well as abiotic factors. While non-native L. catesbeianus have high levels of Bd infection in the Colorado Front Range system, we also identified Bd infection in a number of native amphibian populations allopatric with L. catesbeianus, suggesting that multiple host species are important contributors to the dynamics of the Bd pathogen in this landscape.     

Hybridisation in European ungulates: an overview of the current status,causes, and consequences

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Hybridisation among groupers (genus <Emphasis Type="Italic">Cephalopholis</Emphasis>) at the eastern Indian Ocean suture zone: taxonomic and evolutionary implications

Hybridisation is a significant evolutionary process that until recently was considered rare in the marine environment. A suture zone in the eastern Indian Ocean is home to numerous hybridising sister species, providing an ideal opportunity to determine how hybridisation affects speciation and biodiversity in coral reef fishes. At this location, hybridisation between two grouper (Epinephelidae) species: Cephalopholis urodeta (Pacific Ocean) and C. nigripinnis (Indian Ocean) was investigated to determine the genetic basis of hybridisation and to compare the ecology and life history of hybrids and their parent species. This approach aimed to provide insights into the taxonomic and evolutionary consequences of hybridisation. Despite clear phenotypic differences, multiple molecular markers revealed hybrids, and their parent species were genetically homogenous within and (thousands of kilometres) outside of the hybrid zone. Hybrids were at least as fit as their parent species (in terms of growth, reproduction, and abundance) and were observed in a broad range of intermediate phenotypes. The two species appear to be interbreeding at Christmas Island due to inherent biological and ecological compatibilities, and the lack of genetic structure may be explained by three potential scenarios: (1) hybridisation and introgression; (2) discordance between morphology and genetics; and (3) incomplete lineage sorting. Further molecular analyses are necessary to discriminate these scenarios. Regardless of which applies, C. urodeta and C. nigripinnis are unlikely to evolve in reproductive isolation as they cohabit where they are common (Christmas Island) and will source congeneric mates where they are rare (Cocos Keeling Islands). Our results add to the growing body of evidence that hybridisation among coral reef fishes is a dynamic evolutionary factor.     

Assessment of hybridisation between the endangered Chatham Island black robin (<Emphasis Type="Italic">Petroica traversi</Emphasis>) and the Chatham Island tomtit (<Emphasis Type="Italic">Petroica macrocephala chathamensis</Emphasis>)

Hybridisation between an endangered species and a more common species can facilitate population decline and extinction of the endangered species due to wasted reproductive effort, outbreeding depression and/or swamping of alleles due to widespread or complete admixture. The Chatham Island black robin (Petroica traversi) is an endangered songbird species, which was reduced to only five individuals in 1980. Intensive cross-fostering, whereby black robin offspring were placed into nests of the closely related Chatham Island tomtit (Petroica macrocephala chathamensis) to increase reproductive output, contributed to the rapid recovery of the species within 10 years. Several hybridisation events occurred and although those hybrids were successfully eliminated from the population, concerns remained for the possibility of introgression between the two species that may have gone unnoticed. In this study, we genotyped seven microsatellite loci in both species from the two islands where they coexist, to assess the level of hybridisation and the extent of introgression between the two species. The two species shared no alleles at five of the seven loci genotyped, and cluster analysis, AMOVA and admixture analysis of a total of 174 black robins and 78 Chatham Island tomtits showed no evidence of hybridisation or introgression on either of the two islands where they co-exist. As a result, there is no evidence that black robins are currently in any danger of population decline or extinction through hybridisation with tomtits, although small population size and skewed sex ratio, particularly in the smaller of the two populations, may facilitate future hybridisation events.     

A review of growth and life-history traits of native and non-native European populations of black bullhead <Emphasis Type="Italic">Ameiurus melas</Emphasis>

North American black bullhead, Ameiurus melas, which were introduced to Europe in the nineteenth and twentieth centuries, have received relatively little study. With focus on growth and reproduction, this extensive review, which includes new European data, aims to inform the risk analysis process concerning this non-native species in Europe. Surprisingly, the new data for Europe were more comprehensive than for native populations, with data available mainly from Oklahoma, and North and South Dakota (USA). In terms of relative growth, juvenile A. melas were found to have a relatively uniform body shape regardless of the population’s origin, whereas adults developed different phenotypes depending upon location. Overall growth trajectory was significantly faster for native than for non-native populations. Growth index values decreased significantly with increasing latitude in non-native but not native populations—the latter decreasing weakly with increasing altitude in the populations located at latitudes <40°. Mean general condition (slope ‘b’), mean sex ratio and mean egg diameter did not differ significantly between native and non-native populations. Absolute fecundity was slightly (but not significantly) higher in non-native than native populations. GSI data, which were very scarce for native populations, suggest gonad production may be slightly higher in native than in non-native populations. Precise data on age at maturity (AaM) are lacking for the native range, where 2–5 years is reported. Whereas, in the introduced range the greatest AaM was 3.5 years, and AaM decreases with increasing juvenile growth (TL at age 3). The populations with fastest juvenile growth tended to be from warmer water bodies where they are considered to be invasive. The great growth and life-history plasticity of black bullhead affords the species great potential to invade and establish viable populations in new areas.     

Ancient and current gene flow between two distantly related Mediterranean oak species, Quercus suber and Q. ilex

Background and Aims

Quercus suber and Q. ilex are distantly related and their distributions partially overlap. They hybridize occasionally, but the complete replacement of Q. suber chloroplast DNA (cpDNA) by that of Q. ilex was identified in two specific geographical areas. The objective of this study was to determine whether the contrasting situation reflected current or recent geographical interspecies gene flow variation or was the result of ancient introgression.

Methods

cpDNA PCR-RFLPs (restriction fragment length polymorphisms) and variation at ten nuclear microsatellite loci were analysed in populations of each species, in 16 morphologically intermediate individuals and the progeny of several of them. Interspecies nuclear introgression was based on individual admixture rates using a Bayesian approach with no a priori species assignment, and on a maximum-likelihood (ML) method, using allele frequencies in the allopatric populations of each species as controls. Gene flow was compared specifically between populations located within and outside the specific areas.

Key Results

High interspecies nuclear genetic differentiation was observed, with twice the number of alleles in Q. ilex than in Q. suber. According to Bayesian assignment, approx. 1 % of individuals had a high probability of being F₁ hybrids, and bidirectional nuclear introgression affected approx. 4 % of individuals in each species. Hybrid and introgressed individuals were identified predominantly in mixed stands and may have a recent origin. Higher proportions including allospecific genes recovered from past hybridization were obtained using the ML method. Similar rates of hybridization and of nuclear introgression, partially independent of cpDNA interspecies transfer suggestive of gene filtering, were obtained in the populations located within and outside the areas of complete cpDNA replacement.

Conclusions

The results did not provide evidence for geographical variation in interspecies gene flow. In contrast, historical introgression is supported by palynological records and constitutes the more reliable origin of cpDNA replacement in specific regions.Key words: cpDNA PCR-RFLPs, nuclear microsatellite (nSSR) variation, hybridization, interspecies genetic introgression, Quercus suber, Quercus ilex     

A biogeographic genetic approach for testing the role of reinforcement: the case of Drosophila pseudoobscura and D. persimilis

Abstract.— The role of reinforcement in speciation can be explained by two distinct models. In model I, two diverged populations hybridize and produce fertile hybrids that successfully backcross (hybridization with gene flow). In model II, two populations hybridize but succeeding backcrosses are unproductive (hybridization without gene flow). Using Drosophila persimilis and D. pseudoobscura , we have tested model I by comparing the extent of heterospecific introgression in sympatric versus allopatric populations. We show that certain expectations of this particular model of reinforcement, which is based on hybridization and gene flow between divergent populations after secondary contact, are not realized in these two species. The evidence consists of the similarity of genetic distances as well as proportions of unique/rare alleles between sympatric and allopatric heterospecific populations and a negative correlation between genetic distance and geographical distance between heterospecific populations, which suggests ecological differentiation. This approach in quantifying differential gene flow has important consequences to studies that compare sympatric and allopatric isolation using genetic distance. Following model I, one would expect a pattern of higher prezygotic isolation in sympatric species compared to allopatric species of the same genetic distance simply as a result of an underestimation of genetic distance due to introgression between sympatric populations. We suggest more parsimonious explanations such as reinforcement without genetic exchange (model II) and ecological differentiation, which require high levels of preexisting reproductive isolation between populations.     

Trophic dynamics within a hybrid zone – interactions between an abundant cyprinid hybrid and sympatric parental species

1. Recent proliferation of hybridisation in response to anthropogenic ecosystem change, coupled with increasing evidence of the importance of ancient hybridisation events in the formation of many species, has moved hybridisation to the forefront of evolutionary theory. 2. In spite of this, the mechanisms (e.g. differences in trophic ecology) by which hybrids co‐exist with parental taxa are poorly understood. A unique hybrid zone exists in Irish freshwater systems, whereby hybrid offspring off two non‐native cyprinid fishes often outnumber both parental species. 3. Using stable isotope and gut content analyses, we determined the trophic interactions between sympatric populations of roach (Rutilus rutilus), bream (Abramis brama) and their hybrid in lacustrine habitats. 4. The diet of all three groups displayed little variation across the study systems, and dietary overlap was observed between both parental species and hybrids. Hybrids displayed diet, niche breadth and trophic position that were intermediate between the two parental species while also exhibiting greater flexibility in diet across systems.     

Large-scale asymmetric introgression of cytoplasmic DNA reveals Holocene range displacement in a North American boreal pine complex

Jack pine (Pinus banksiana) and lodgepole pine (Pinus contorta var. latifolia) are two North American boreal hard pines that hybridize in their zone of contact in western Canada. The main objective of this study was to characterize their patterns of introgression resulting from past and recent gene flow, using cytoplasmic markers having maternal or paternal inheritance. Mitochondrial DNA (mtDNA) and chloroplast DNA (cpDNA) diversity was assessed in allopatric populations of each species and in stands from the current zone of contact containing morphological hybrids. Cluster analyses were used to identify genetic discontinuities among groups of populations. A canonical analysis was also conducted to detect putative associations among cytoplasmic DNA variation, tree morphology, and site ecological features. MtDNA introgression was extensive and asymmetric: it was detected in P. banksiana populations from the hybrid zone and from allopatric areas, but not in P. contorta populations. Very weak cpDNA introgression was observed, and only in P. banksiana populations. The mtDNA introgression pattern indicated that central Canada was first colonized by migrants from a P. contorta glacial population located west of the Rocky Mountains, before being replaced by P. banksiana migrating westward during the Holocene. In contrast, extensive pollen gene flow would have erased the cpDNA traces of this ancient presence of P. contorta. Additional evidence for this process was provided by the results of canonical analysis, which indicated that the current cpDNA background of trees reflected recent pollen gene flow from the surrounding dominant species rather than historical events that took place during the postglacial colonization.     

Morphological differences between native and non-native pumpkinseed in traits associated with locomotion

Adaptive strategies in morphology can significantly influence the successful invasion and establishment of non-native species. Since its introduction, the pumpkinseed (Lepomis gibbosus), a sunfish of North American origin, has spread throughout most of Europe, including the Iberian Peninsula. We hypothesized that 12 morphological traits, functionally significant for locomotion, would differ according to geographic origin (native/non-native) and habitat type (fluvial/lacustrine). Using flow-through raceways, we simultaneously reared F₁ young-of-the-year pumpkinseed from two native and two non-native populations, produced from adults kept in a common environment. Morphometric measurements were recorded at the beginning and end of the 90-day rearing period. Median-fin size and placement differed significantly between native and non-native populations, whereas paired fin size differed between fluvial and lacustrine populations. Other functionally significant traits, such as body width, also differed between native and non-native populations. Spanish populations were considered to have acquired these adaptive external morphologies through successive generations, following the species’ range expansion through the variable environments of the Iberian Peninsula.     

Evolutionary History of the Grey-Faced Sengi,Rhynchocyon udzungwensis,from Tanzania: A Molecular and Species Distribution Modelling Approach

Rhynchocyon udzungwensis is a recently described and poorly understood sengi (giant elephant-shrew) endemic to two small montane forests in Southern Tanzania, and surrounded in lower forests by R. cirnei reichardi. In this study, we investigate the molecular genetic relationship between R. udzungwensis and R. c. reichardi, and the possible role that shifting species distributions in response to climate fluctuations may have played in shaping their evolutionary history. Rhynchocyon udzungwensis and R. c. reichardi individuals were sampled from five localities for genetic analyses. Three mitochondrial and two nuclear loci were used to construct species trees for delimitation and to determine whether introgression was detectable either from ancient or ongoing hybridization. All species-tree results show R. udzungwensis and R. c. reichardi as distinct lineages, though mtDNA shows evidence of introgression in some populations. Nuclear loci of each species were monophyletic, implying introgression is exclusively historical. Because we found evidence of introgression, we used distribution data and species distribution modelling for present, glacial, and interglacial climate cycles to predict how shifting species distributions may have facilitated hybridization in some populations. Though interpretations are affected by the limited range of these species, a likely scenario is that the mtDNA introgression found in eastern mid-elevation populations was facilitated by low numbers of R. udzungwensis that expanded into lowland heavily occupied R. c. reichardi areas during interglacial climate cycles. These results imply that relationships within the genus Rhynchocyon may be confounded by porous species boundaries and introgression, even if species are not currently sympatric.