Hybridization and speciation

Hybridization has many and varied impacts on the process of speciation. Hybridization may slow or reverse differentiation by allowing gene flow and recombination. It may accelerate speciation via adaptive introgression or cause near‐instantaneous speciation by allopolyploidization. It may have multiple effects at different stages and in different spatial contexts within a single speciation event. We offer a perspective on the context and evolutionary significance of hybridization during speciation, highlighting issues of current interest and debate. In secondary contact zones, it is uncertain if barriers to gene flow will be strengthened or broken down due to recombination and gene flow. Theory and empirical evidence suggest the latter is more likely, except within and around strongly selected genomic regions. Hybridization may contribute to speciation through the formation of new hybrid taxa, whereas introgression of a few loci may promote adaptive divergence and so facilitate speciation. Gene regulatory networks, epigenetic effects and the evolution of selfish genetic material in the genome suggest that the Dobzhansky–Muller model of hybrid incompatibilities requires a broader interpretation. Finally, although the incidence of reinforcement remains uncertain, this and other interactions in areas of sympatry may have knock‐on effects on speciation both within and outside regions of hybridization.     

长白山垂直自然带弹尾纲物种概况

以长白山弹尾纲为研究对象, 基于1758‒2022年度发表的分类学和生态学文献资料, 编制了长白山垂直自然带弹尾纲物种名录, 分析了首次发表物种的学者贡献、出版物、发表时间和各属沿海拔梯度的分布情况。本研究共梳理弹尾纲物种92种, 隶属于3目11科46属, 长角䖴目、原䖴目和愈腹䖴目的物种分别占比61%、31%和8%, 其中长角䖴目的等节䖴科(该类群物种数占总体比例, 26%)、长角䖴科(16%)和鳞䖴科(16%)、原䖴目的棘䖴科(26%)是该地区的主要类群。统计的92种长白山弹尾纲物种由36位学者首次定名和报道, 中国学者占主要贡献(33%), 其次为波兰(11%)、瑞典(11%)、俄罗斯(8%)和美国(8%); 相关论文多发表在国际期刊上, Zootaxa是发表的主要期刊。基于海拔梯度的统计发现, 鳞䖴属(Tomocerus)和符䖴属(Folsomia)的物种沿海拔梯度的分布范围最广(海拔800‒1,700 m), 分布在海拔800 m (15个物种)、1,100 m (20个物种)和1,400 m (14个物种)的物种最多。基于以上结果, 本文讨论了长白山弹尾纲分类学现状和发展前景。     

Hybridization among sympatric species of Rhododendron (Ericaceae) in Turkey: morphological and molecular evidence

Rhododendron (Ericaceae) is a large genus in which barriers to hybridization are especially weak, but many species are maintained in sympatry. Hybridization among four species of Rhododendron subsect. Pontica, which occur in sympatry in Turkey, was investigated. Material of R. ponticum, R. smirnovii, R. ungernii, and R. caucasicum and their putative hybrids was collected from the wild. Based on morphology, chloroplast DNA and nuclear ribosomal DNA restriction fragment length polymorphism (RFLP) profiles, each accession was identified as a species or hybrid combination. Five of the six possible hybrid combinations among the four species were detected. Rhododendron ponticum × R. smirnovii was represented by a single individual and R. caucasicum × R. smirnovii by one small group of hybrid plants. The combinations R. ponticum × R. ungernii and R. ungernii × R. smirnovii showed evidence of frequent backcrossing, while R. ponticum × R. caucasicum appeared unusual in that an intermediate hybrid type was abundant, whereas hybrids with phenotypes approaching either parent were rare. Possible explanations of this latter situation are discussed. The results suggest that natural hybridization among Rhododendron species is common and that ecological factors are important in maintaining integrity when species occur in sympatry.     

Hybridization between tetraploid species of Alopecurus L. (Poaceae): morphological studies of natural and artificial hybrids

The morphology of individuals of mixed natural populations of three species of Alopecurus, A. pratensis, A. geniculars and A. arundinaceus , is described. It is suggested that plants morphologically intermediate between the parental species are hybrids or of hybrid origin. Hybridization occurs more readily between A. pratensis and A. arundinaceus than between A. pratensis and A. geniculate , since intermediate forms were recovered from all mixed populations of A. pratensis and A. arundinaceus. The intermediate forms occur only in a few populations of A. pratensis and A. geniculars where an appropriate habitat is available. Evidence of backcrossing and therefore introgression comes from the continuum of intermediate forms that exists between the species. Scatter diagrams for individual populations show distinct differences in the degree of introgression. Observations on artificially produced hybrids are similar to those on naturally occurring hybrids.     

Hybridization between two insectivorous bird species and the effect on prey-handling efficiency

Gene flow between closely related species is not always impeded by hybrid inviability or sterility but may be limited by behavioural or ecological factors. European common redstarts, Phoenicurus phoenicurus, and black redstarts, P. ochruros, are insectivorous passerine bird species which regularly interbreed in the wild and produce viable and fertile progeny. Yet, the isolation barriers, which prevent extensive gene flow between these distinct species, are unknown. I studied prey-handling time and efficiency in both species and the F₁-hybrids in captivity. All birds changed prey-handling mode from a single pick to more complex behaviour with increasing prey length and their handling time was a positive function of prey length. Common redstarts tended to handle their prey quicker than black redstarts. Hybrids appeared to be intermediate. However, individual differences were significant, but group membership was not. Handling efficiency decreased with increasing prey length and was determined by individual and family effects. Hybrid females handled their prey as efficient as hybrid males. Small differences in beak morphology could not explain individual differences in prey-handling behaviour. The results of this study suggest that prey-handling in F₁-hybrids is not a postzygotic barrier which prevent gene flow between common and black redstarts in the wild.     

Sharp acoustic boundaries across an altitudinal avian hybrid zone despite asymmetric introgression

Birdsong is a sexually selected trait that could play an important evolutionary role when related taxa come into secondary contact. Many songbird species, however, learn their songs through copying one or more tutors, which complicates the evolutionary outcome of such contact. Two subspecies of a presumed vocal learner, the grey‐breasted wood‐wren (Henicorhina leucophrys), replace each other altitudinally across the western slope of the Ecuadorian Andes. These subspecies are morphologically very similar, but show striking differences in their song. We examined variation in acoustic traits and genetic composition across the altitudinal range covered by both subspecies and between two allopatric populations. The acoustic boundary between the subspecies was found to be highly abrupt across a narrow elevational range with virtually no evidence of song convergence. Mixed singing and use of hetero‐subspecific song occurred in the contact zone and was biased towards the use of leucophrys song types. Hetero‐subspecific song copying by hilaris and not by leucophrys reflected a previously found asymmetric pattern of response to song playback. Using amplified fragment length polymorphisms (AFLP) markers, we detected hybridization in the contact zone and asymmetric introgression in parapatric populations, with more leucophrys alleles present in hilaris populations than vice versa. This pattern may be a trail of introgression due to upslope displacement of leucophrys by hilaris. Our data suggest that song learning may impact speciation and hybridization in contrasting ways at different spatial scales: although learning may speed up population divergence in songs, thereby enhancing assortative mating and reducing gene flow, it may at a local level also lead to the copying of heterospecific songs, therefore allowing some level of hybridization and introgression.     

基于形态特征探究鼻狂蝇属两近缘物种鉴别

鼻狂蝇属物种是有蹄类动物鼻咽腔内常见的专性寄生蝇,是畜牧兽医、农林生产、动物保护领域重点关注的寄生虫.本研究通过系统比较紫鼻狂蝇Rhinoestrus purpureus和亚非鼻狂蝇Rhinoestrus usbekistanicus这两种寄生于马科动物的近缘物种的成虫外部形态及雌雄尾器特征,明确了二者关键鉴别特征,并绘制了形态特征图;此外,还总结了两近缘种在中国的寄主、分布地信息,不仅为鼻狂蝇的快速识别和精准鉴定提供了科学数据,还为其流行病学调查及防控提供了基础资料.     

运用稀疏法分析物种丰富度的海拔梯度分布格局:以样方实测乔木种数据为例

植物物种丰富度随山地海拔梯度的变化格局是生物多样性研究的热点之一.基于种-面积关系的任何模型对群落物种数目所作估计,其精度都依赖于样本的代表性、抽样尺度以及所涉及的分类群.作者以秦岭南坡森林群落样方实测的乔木种数据为例,借鉴群落最小面积(minimum area,MA)的概念及其确定方式,利用稀疏法(rarefacti...     

Vascular plant species richness along latitudinal and altitudinal gradients: a contribution from New Zealand temperate rainforests

Vascular plant species richness is known to often decrease with both increasing latitude and increasing altitude. However, a number of studies have shown the reverse trend and the primary cause of these gradients remains unknown. In the present work, generalized linear models were used to assess the relative importance of latitude and altitude as well as of a number of other factors (mean annual precipita-tion, slope, substrate and forest type) on species richness in temperate rainforests of New Zealand. The effect of Southern beech ( Nothofagus spp.) as dominant canopy species on total species richness was shown to be much smaller than postulated in most previous studies. Within the region studied, altitude had by far the strongest effect on species richness. This effect was independent of latitude and was significant for woody but not for herbaceous vegetation.     

Reproductive isolation among two interfertile Rhododendron species: low frequency of post-F1 hybrid genotypes in alpine hybrid zones

Hybrids between the acid-loving species Rhododendron ferrugineum and the basic soil species Rhododendron hirsutum occur on soils of intermediate pH in the European Alps. Material from two hybrid zones ~500 m apart, and also nearby populations of each parent species, was surveyed for presence/absence of 31 random amplified polymorphic DNA markers that distinguish parents. Based on morphological assessment, the material comprised 51 putative hybrids, 18 putative R. ferrugineum individuals and 26 putative R. hirsutum plants. RAPD data were analysed using a Bayesian approach implemented by the program newhybrids , and also by principal coordinates analysis. The identity of all R. ferrugineum plants examined was confirmed; however, of the putative R. hirsutum individuals examined, two were certainly and 11 possibly hybrid derivatives. Among all hybrid derivatives examined, about half were designated as F₁s or a similar class, otherwise backcrosses to R. hirsutum appeared to be common whereas other hybrid classes were rare and backcrosses to R. ferrugineum possibly absent. Despite this, artificially generated seed of F₂ class and backcrosses in each direction showed greater viability than one parent ( R. hirsutum ). Introgression from R. ferrugineum was also detected in a population that from morphology appeared to contain only R. hirsutum . Hence, the direction of backcrossing might be highly asymmetric within hybrid zones, causing unidirectional gene flow from R. ferrugineum into R. hirsutum . Conversely, the rarity of backcrosses to R. ferrugineum , F₂s and later hybrid generations, which might be due to phenology effects and habitat-mediated selection, could play a part in restricting gene flow towards R. ferrugineum .     

Genetic,morphological, and niche variation in the widely hybridizing Rhus integrifolia-Rhus ovata species complex

Hybridization and introgression are common processes among numerous plant species that present both challenges and opportunities for studies of species delimitation, phylogenetics, taxonomy and adaptation. Rhus integrifolia and R. ovata are two ecologically important shrubs native to the southwestern USA and Mexico, and are known to hybridize frequently, but the morphological, genetic and ecological implications of hybridization in these species are poorly studied on a broad geographic scale. Analyses were conducted using leaf morphology, genetic variation of plastid and nuclear loci, and species distribution models for both species and their putative hybrid introgressants across 19 localities in California and Arizona, USA. These analyses revealed evidence for morphological and genetic distinction among localities comprising putative parental species, but a high degree of morpho-genetic intermediacy among localities with putative hybrids. Comparison of morphological and genetic population structure among localities revealed evidence for putative local adaptation or widespread phenotypic plasticity. Multiple regression models identified a weak but statistically significant negative association between leaf area and precipitation. Finally, species distribution modeling inferred northward range shifts over time, with both species predicted to occupy more coastal regions in the future, possibly increasing the frequency of hybridization among them. These findings underscore the importance of integrative assessment of multiple data sources in the study of hybridizing species and highlight the R. integrifolia-ovata complex as a powerful model for investigating the adaptive implications of hybridization.     

Hybridization and sex pheromone responses in two closely related mealybug species (Homoptera: Pseudococcidae)

Abstract. Reciprocal crosses were made between the closely related species Planococcus citri and Planococcus ficus by using mating inducers. The pheromone responses of the hybrid offspring were bioassayed. The sex ratio of the offspring showed a striking shift in favour of females in comparison with homospecific matings mainly because of high mortality of male nymphs during the first instar. Hybrid males gave positive reactions only to pheromone extracts from the maternal species and showed no response to extracts from the paternal species. This result may be explained by the lecanoid chromosome system known for mealybug males, in which the paternal set of chromosomes becomes heterochro-matic (condensed and genetically inactive) starting from early embryogeny. Further evidence is, however, presented that (at least in certain tissues) the paternal set reverts to the euchromatic state and to genetical activity.     

From hybridization to introgression between two closely related sympatric ant species

Interspecific hybridization is becoming more frequent worldwide due to increasing global changes and translocations of organisms. For individual organisms, the most significant negative consequences are sterility or inviability of hybrid offspring. However, hybridization sometimes leads to fertile offspring, promoting introgression from one species into another. In such situations, hybridization can play a key role in evolution and speciation. Combining hypervariable DNA (microsatellites) and mitochondrial DNA markers with the use of several modeling methods allow an efficient detection of hybridization processes. The present study therefore investigates hybridization between two ant species, Tetramorium immigrans and T. caespitum, using multiple methods, and systematically comparing results with simulated data to ensure accurate identification of hybrids. Introgression was revealed both by backcross detection based on 14 nuclear microsatellite loci and by mitochondrial‐nuclear discordance based on comparison with mitochondrial DNA (cytochrome c oxidase subunit I). Results were spatially consistent, with hybrids located at latitudes where parental species are sympatric. The causes and consequences of hybridization and introgression between T. caespitum and T. immigrans remain to be further investigated, especially because T. immigrans could be an invasive species in France.     

Recombination rate variation in closely related species

Despite their importance to successful meiosis and various evolutionary processes, meiotic recombination rates sometimes vary within species or between closely related species. For example, humans and chimpanzees share virtually no recombination hotspot locations in the surveyed portion of the genomes. However, conservation of recombination rates between closely related species has also been documented, raising an apparent contradiction. Here, we evaluate how and why conflicting patterns of recombination rate conservation and divergence may be observed, with particular emphasis on features that affect recombination, and the scale and method with which recombination is surveyed. Additionally, we review recent studies identifying features influencing fine-scale and broad-scale recombination patterns and informing how quickly recombination rates evolve, how changes in recombination impact selection and evolution in natural populations, and more broadly, which forces influence genome evolution.     

Strong divergent selection at multiple loci in two closely related species of ragworts adapted to high and low elevations on Mount Etna

Recently diverged species present particularly informative systems for studying speciation and maintenance of genetic divergence in the face of gene flow. We investigated speciation in two closely related Senecio species, S. aethnensis and S. chrysanthemifolius, which grow at high and low elevations, respectively, on Mount Etna, Sicily and form a hybrid zone at intermediate elevations. We used a newly generated genome‐wide single nucleotide polymorphism (SNP) dataset from 192 individuals collected over 18 localities along an elevational gradient to reconstruct the likely history of speciation, identify highly differentiated SNPs, and estimate the strength of divergent selection. We found that speciation in this system involved heterogeneous and bidirectional gene flow along the genome, and species experienced marked population size changes in the past. Furthermore, we identified highly‐differentiated SNPs between the species, some of which are located in genes potentially involved in ecological differences between species (such as photosynthesis and UV response). We analysed the shape of these SNPs’ allele frequency clines along the elevational gradient. These clines show significantly variable coincidence and concordance, indicative of the presence of multifarious selective forces. Selection against hybrids is estimated to be very strong (0.16–0.78) and one of the highest reported in literature. The combination of strong cumulative selection across the genome and previously identified intrinsic incompatibilities probably work together to maintain the genetic and phenotypic differentiation between these species – pointing to the importance of considering both intrinsic and extrinsic factors when studying divergence and speciation.     

Avian life history variation along altitudinal gradients: an example with cardueline finches

Elevation has long been considered a major influence on the evolution of life-history traits. Most elevation-induced variation in life history traits has been attributed to changes in climate, duration of breeding season, predation, and food limitation. I use a phylogenetic approach to show that life histories are closely associated with breeding elevation in extant cardueline finches. Finches at high elevations had smaller clutches, fewer broods, and longer incubation periods. Neither food limitation nor nest predation appear to readily account for this strong elevational variation in cardueline life histories. However, juvenile survival may be greater at higher elevations as a result of prolonged parental care and shorter natal dispersal and can potentially compensate for reduced fecundity in high-elevation finches. Received: 28 September 1996 / Accepted: 24 March 1997     

Sperm-related phenotypes implicated in both maintenance and breakdown of a natural species barrier in the house mouse

The house mouse hybrid zone (HMHZ) is a species barrier thought to be maintained by a balance between dispersal and natural selection against hybrids. While the HMHZ is characterized by frequency discontinuities for some sex chromosome markers, there is an unexpected large-scale regional introgression of a Y chromosome across the barrier, in defiance of Haldane''s rule. Recent work suggests that a major force maintaining the species barrier acts through sperm traits. Here, we test whether the Y chromosome penetration of the species barrier acts through sperm traits by assessing sperm characteristics of wild-caught males directly in a field laboratory set up in a Y introgression region of the HMHZ, later calculating the hybrid index of each male using 1401 diagnostic single nucleotide polymorphisms (SNPs). We found that both sperm count (SC) and sperm velocity were significantly reduced across the natural spectrum of hybrids. However, SC was more than rescued in the presence of the invading Y. Our results imply an asymmetric advantage for Y chromosome introgression consistent with the observed large-scale introgression. We suggest that selection on sperm-related traits probably explains a large component of patterns observed in the natural hybrid zone, including the Y chromosome penetration.     

A new morphometric model in distinguishing two closely related extinct brachiopod species

Pseudoatrypa (Atrypida, Brachiopoda) are extinct Paleozoic brachiopods that are the closest relatives of the modern brachiopods, rhynchonellids. Two Paleozoic brachiopod species were qualitatively distinguished based on close examination of phenotypic characters and traditional morphometrics. These species were recovered for the first time from the Givetian localities of Traverse Group of Michigan. The efficacy of geometric morphometric analyses in discriminating species has been proven for various organisms, but rarely studies have been performed in distinguishing brachiopod species. The advanced morphometric model developed here using the Procrustes-based landmark method is based on the analysis of valve shape in two extinct brachiopod species: Pseudoatrypa lineata and Pseudoatrypa devoniana. Results from quantitative analyses reinforce the qualitative identifications of the two species. The application of this method on four different orientations (dorsal, ventral, posterior and anterior regions) of the brachiopods permits the two species to be clearly separated, and this study proposes that the model can be effectively used for future identification and distinction of fossil species. Additionally, this study will aid biologists, paleontologists and neontologists in making appropriate advancement in systematic studies and further assist with gauging biodiversity congruously.     

Hybridization and the inheritance of female colour polymorphism in two ischnurid damselflies (Odonata: Coenagrionidae)

Female‐limited polychromatism is frequent in many species of Odonata. Ischnura elegans has three colour morphs: one male‐like coloured (androchrome) and two additional gynochrome brown morphs (infuscans and rufescens‐obsoleta morphs). A total of 19 progenies obtained from once‐mated females were reared in the laboratory in three generations. Results indicate that the colour morphs are controlled by the same genetic system as previously described for I. graellsii, i.e. an autosomal locus with female‐limited expression and with three alleles with a hierarchy of dominance (p^a > pⁱ > p°). Five interspecific crossings between female I. graellsii and male I. elegans, five crossings between hybrid females and male I. elegans and one crossing between female I. graellsii and a hybrid male further confirmed that the genetic system is the same in both species. A survey of morph frequencies in north‐west Spain revealed that I. elegans shows high variability in androchrome frequency (4–91%) between nearby populations, whereas in I. graellsii androchromes never are the majority morph (5–40%). The highest androchrome frequency in I. graellsii was found in populations closest to a locality where both species have hybridized, and that now has the highest androchrome frequency of I. elegans. We hypothesize that I. elegans genes have been incorporated into the genome of I. graellsii resulting in increased androchrome frequency in the latter species. Low androchrome frequency in I. elegans seems also related to the influence of I. graellsii genes. Therefore, we suggest that hybridization between both taxa is contributing to the temporal maintenance of contrasting androchrome frequencies in nearby populations. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 85 , 471–481.     

Intraspecific variation in reproductive barriers between two closely related Arabidopsis sister species

Reproductive isolation (RI) is a critical component of speciation and varies strongly in timing and strength among different sister taxa, depending on, for example the geography of speciation and divergence time. However, these factors may also produce variation in timing and strength among populations within species. Here we tested for variation in the expression of RI among replicate population pairs between the sister taxa Arabidopsis lyrata subsp. lyrata and A. arenicola. While the former is predominantly outcrossing, the latter is predominantly selfing. We focused on intrinsic prezygotic and postzygotic RI as both species occur largely in allopatry. We assessed RI by performing within-population crosses and interspecific between-population crosses, and by raising offspring. RI was generally high between all interspecific population pairs, but it varied in timing and strength depending on population history. Prezygotic isolation was strongest between the closest-related population pair, while early postzygotic isolation was high for all other population pairs. Furthermore, the timing and strength of RI depended strongly on cross direction. Our study provides empirical support that reproductive barriers between species are highly variable among population pairs and asymmetric within population pairs, and this variation seems to follow patterns typically described across species pairs.