Hybridization studies in Columnea L. (Gesneriaceae). 1. Jamaican species

A breeding programme using Jamaican species of Columnea demonstrated that a majority are interfertile, and made possible the synthesis of artificial hybrids between C. urbanii Stearn and C. rutilans Swartz resembling wild plants previously suspected to be of hybrid origin.
Evidence is provided to show that species in sect. Columnea are interfertile with species in sect. Pterygoloma.     

Reproductive Isolation of Hybrid Populations Driven by Genetic Incompatibilities

Despite its role in homogenizing populations, hybridization has also been proposed as a means to generate new species. The conceptual basis for this idea is that hybridization can result in novel phenotypes through recombination between the parental genomes, allowing a hybrid population to occupy ecological niches unavailable to parental species. Here we present an alternative model of the evolution of reproductive isolation in hybrid populations that occurs as a simple consequence of selection against genetic incompatibilities. Unlike previous models of hybrid speciation, our model does not incorporate inbreeding, or assume that hybrids have an ecological or reproductive fitness advantage relative to parental populations. We show that reproductive isolation between hybrids and parental species can evolve frequently and rapidly under this model, even in the presence of substantial ongoing immigration from parental species and strong selection against hybrids. An interesting prediction of our model is that replicate hybrid populations formed from the same pair of parental species can evolve reproductive isolation from each other. This non-adaptive process can therefore generate patterns of species diversity and relatedness that resemble an adaptive radiation. Intriguingly, several known hybrid species exhibit patterns of reproductive isolation consistent with the predictions of our model.     

Hybridization studies in Columnea L. (Gesneriaceae). 2. The C. querceti complex

Sherk's cytogenetic findings are outlined, as far as they relate to the possible hybrid origin of Columnea querceti Oersted. Morphological evidence is provided to show that a hybrid origin is not impossible because there is a precedent for natural hybridization within the genus.     

New evidence for a postglacial homoploid hybrid origin of the widespread Central European Scabiosa columbaria L. s. str. (Dipsacaceae)

We have tested whether Scabiosa columbaria s. str. could have originated by recent homoploid hybrid speciation because it is of intermediate morphology and is distributed in areas under glacial influence. Existing morphological and new RAPD data for 11 ingroup populations were analysed using multivariate statistics. Most methods indicated that S. columbaria s. str. could be the first example for a homoploid hybrid origin from Central Europe although a rigorous proof was missing partly because of ongoing introgression. Sequence variation was absent for 5,000 bp from 11 regions supporting a postglacial origin of S. columbaria s. str. An analysis of realized macroclimatic niches using GIS data showed that the niches of S. columbaria s. str. were partly intermediate between its putative parental species rather than extreme as often found in other homoploid hybrids. This could be due to geographical preconditions: large new habitats were available in postglacial Europe while in the case of most other homoploid hybrids only ecologically challenging habitats were not occupied by parental taxa.     

Distribution of Phoxinus eos,Phoxinus neogaeus,and Their Asexually-Reproducing Hybrids (Pisces: Cyprinidae) in Algonquin Provincial Park,Ontario

Hybrid Phoxinus are one of the few asexually reproducing vertebrates species. The distribution of hybrid Phoxinus among lakes in Algonquin Park, Ontario, was evaluated relative to the distribution of parental species and relative to physiochemical lake characteristics. No association between the distribution of hybrids and the distribution of parental species was found, suggesting that the hybrids can successfully coexist with either parental species. In addition, we found no association between hybrid distribution and the physiochemical characteristics of lakes, suggesting that the hybrids are generalists with respect to the ecological niches available in Algonquin Park. There was a difference between the physiochemical characteristics of lakes with and without the parental species P. neogaeus. The lakes containing P. neogaeus were lower elevation than the lakes containing the other parental species, P. eos. The difference in distribution between the parental species may therefore be due to different dispersal abilities, to later arrival following post-glacial dispersal, or to differences in ecology. These results suggest that asexual reproduction is a successful strategy for hybrid Phoxinus in Algonquin Park because these sperm-dependent asexual hybrids are able to survive and persist regardless of which parental species is present, and regardless of the physiochemical characteristics of their habitat.     

Influence of niche similarity on hybridization between Myriophyllum sibiricum and M. spicatum

The impact of ecological factors on natural hybridization is of widespread interest. Here, we asked whether climate niche influences hybridization between the two closely related plant species Myriophyllum sibiricum and M. spicatum. Eight microsatellite loci and two chloroplast fragments were used to investigate the occurrence of hybridization between these two species in two co‐occurring regions: north‐east China (NEC) and the Qinghai‐Tibetan Plateau (QTP). The climate niches of the species were quantified by principal component analysis with bioclimatic data, and niche comparisons were performed between the two species in each region. Reciprocal hybridization was observed, and M. sibiricum was favoured as the maternal species. Furthermore, hybrids were rare in NEC but common in the QTP. Accordingly, in NEC, the two species were climatically distinct, and hybrids only occurred in the narrow geographical or ecological transition zone, whereas in the QTP, obvious niche overlaps were found for the two species, and hybrids occurred in multiple contact zones. This association between hybridization pattern and climate niche similarity suggests that the level of hybridization was promoted by niche overlap. Compared with the parental species, similar climate niches were found for the hybrid populations in the QTP, indicating that other environmental factors rather than climate were important for hybrid persistence. Our findings highlight the significance of climate niche with respect to hybridization patterns in plants.     

A test of hybrid growth disadvantage in wild, free-ranging species pairs of threespine stickleback (Gasterosteus aculeatus) and its implications for ecological speciation

Ecological speciation is the evolution of reproductive isolation as a direct or indirect consequence of divergent natural selection. Reduced performance of hybrids in nature is thought to be an important process by which natural selection can favor the evolution of assortative mating and drive speciation. Benthic and limnetic sympatric species of threespine stickleback (Gasterosteus aculeatus) are adapted to alternative trophic niches (bottom browsing vs. open water planktivory, respectively) and reduced feeding performance of hybrids is thought to have contributed to the evolution of reproductive isolation. We tested this “hybrid‐disadvantage hypothesis” by inferring growth rates from otoliths sampled from wild, free‐ranging benthic, limnetic, and hybrid sticklebacks in two lakes. There were significant differences in growth rate between lakes, life‐history stages, and among years (maximum P = 0.02), as well as interactions between most factors, but not between hybrid and parental species sticklebacks in most comparisons. Our results provide little evidence of a growth disadvantage in hybrid sticklebacks when free‐ranging in nature. Although trophic ecology per se may contribute less to ecological speciation than envisioned, it may act in concert with other aspects of stickleback biology, such as interactions with parasites, predators, competitors, and/or sexual selection, to present strong multifarious selection against hybrids.     

Does migration of hybrids contribute to post-zygotic isolation in flycatchers?

In the face of hybridization, species integrity can only be maintained through post-zygotic isolating barriers (PIBs). PIBs need not only be intrinsic (i.e. hybrid inviability and sterility caused by developmental incompatibilities), but also can be extrinsic due to the hybrid's intermediate phenotype falling between the parental niches. For example, in migratory species, hybrid fitness might be reduced as a result of intermediate migration pathways and reaching suboptimal wintering grounds. Here, we test this idea by comparing the juvenile to adult survival probabilities as well as the wintering grounds of pied flycatchers (Ficedula hypoleuca), collared flycatchers (Ficedula albicollis) and their hybrids using stable isotope ratios of carbon (delta13C) and nitrogen (delta15N) in feathers developed at the wintering site. Our result supports earlier observations of largely segregated wintering grounds of the two parental species. The isotope signature of hybrids clustered with that of pied flycatchers. We argue that this pattern can explain the high annual survival of hybrid flycatchers. Hence, dominant expression of the traits of one of the parental species in hybrids may substantially reduce the ecological costs of hybridization.     

Genotype‐environment associations support a mosaic hybrid zone between two tidal marsh birds

Local environmental features can shape hybrid zone dynamics when hybrids are bounded by ecotones or when patchily distributed habitat types lead to a corresponding mosaic of genotypes. We investigated the role of marsh‐level characteristics in shaping a hybrid zone between two recently diverged avian taxa – Saltmarsh (Ammodramus caudacutus) and Nelson's (A. nelsoni) sparrows. These species occupy different niches where allopatric, with caudacutus restricted to coastal marshes and nelsoni found in a broader array of wetland and grassland habitats and co‐occur in tidal marshes in sympatry. We determined the influence of habitat types on the distribution of pure and hybrid sparrows and assessed the degree of overlap in the ecological niche of each taxon. To do this, we sampled and genotyped 305 sparrows from 34 marshes across the hybrid zone and from adjacent regions. We used linear regression to test for associations between marsh characteristics and the distribution of pure and admixed sparrows. We found a positive correlation between genotype and environmental variables with a patchy distribution of genotypes and habitats across the hybrid zone. Ecological niche models suggest that the hybrid niche was more similar to that of A. nelsoni and habitat suitability was influenced strongly by distance from coastline. Our results support a mosaic model of hybrid zone maintenance, suggesting a role for local environmental features in shaping the distribution and frequency of pure species and hybrids across space.     

Fine-scale environmental variation contributes to introgression in a three-species spruce hybrid complex

Hybridization is common for many forest trees, where weak barriers to reproduction obscure species boundaries. We characterized the genomic structure of Picea populations comprising three species spanning two well-known contact zones, the Picea sitchensis?×?Picea glauca and the P. engelmannii?×?P. glauca hybrid zones, using a set of 71 candidate-gene single nucleotide polymorphisms. The genetic structure of populations suggests a complex genomic architecture shaped by interspecific gene flow and strong environmental selection, with increased genetic diversity in hybrids. The presence of admixture among all three species suggests that three-way hybrids with mixed ancestry occur where species ranges overlap in transitional environments. Significant clinal variation and associations with climatic variables (including continentality, temperature, and precipitation) differ between hybrid zones, indicating that individual species and their hybrids are adapted to distinct environmental niches. Allele–environmental association analysis revealed that most of the candidate genes with evidence of selection were unique to either the Sitka?×?white or the Engelmann?×?white hybrid zones, with few shared between these zones. Management of these widespread and diverse gene pools will be best served through development of climate-based seed transfer, with recommended seed sources informed by a combination of genetic and climatic information for future climates.     

Ecologically dependent and intrinsic genetic signatures of postzygotic isolation between sympatric host races of the leaf beetle Lochmaea capreae

The fitness of hybrids might be compromised as a result of intrinsic isolation and/or because they fall between ecological niches due to their intermediate phenotypes (“extrinsic isolation”). Here, we present data from several crosses (parental crosses, F1, F2, and backcrosses) between the two host races of Lochmaea capreae on willow and birch to test for extrinsic isolation, intrinsic isolation, and environmentally dependent genetic incompatibilities. We employed a reciprocal transplant design in which offspring were raised on either host plant and their survival was recorded until adulthood. We also applied joint‐scaling analysis to determine the genetic architecture of hybrid inviability. The relative fitness of the backcrosses switched between environments; furthermore, the additive genetic–environment interaction was detected as the strongest effect in our analysis. These results provide strong evidence that divergent natural selection has played a central role in the evolution of hybrid dysfunction between host races. Joint‐scaling analysis detected significant negative epistatic effects that are most evident in the poor performance of F2‐hybrids on willow, indicating signs of intrinsic isolation. We did not find any evidence that genetic incompatibilities are manifested independently of environmental conditions. Our findings suggest the outcome of natural hybridization between these host races is mainly affected by extrinsic isolation and a weak contribution of intrinsic isolation.     

Lifetime fitness in two generations of Ipomopsis hybrids

Various models purporting to explain natural hybrid zones make different assumptions about the fitness of hybrids. One class of models assumes that hybrids have intrinsically low fitness due to genetic incompatibilities, whereas other models allow hybrid fitness to vary across natural environments. We used the intrinsic rate of increase to assess lifetime fitness of hybrids between two species of montane plants Ipomopsis aggregata and Ipomopsis tenuituba planted as seed into multiple field environments. Because fitness is predicted to depend upon genetic composition of the hybrids, we included F1 hybrids, F2 hybrids, and backcrosses in our field tests. The F2 hybrids had female fitness as high, or higher, than expected under an additive model of fitness. These results run counter to any model of hybrid zone dynamics that relies solely on intrinsic nuclear genetic incompatibilities. Instead, we found that selection was environmentally dependent. In this hybrid zone, cytoplasmic effects and genotype-by-environment interactions appear more important in lowering hybrid fitness than do intrinsic genomic incompatibilities between nuclear genes.     

Incomplete reproductive isolation between Rhododendron taxa enables hybrid formation and persistence

The evolutionary consequences of hybridization ultimately depend on the magnitude of reproductive isolation between hybrids and their parents. We evaluated the relative contributions of pre-and post-zygotic barriers to reproduction for hybrid formation, hybrid persistence and potential for reproductive isolation of hybrids formed between two Rhododendron species,R. spiciferum and R. spinuliferum. Our study established that incomplete reproductive isolation promotes hybrid formation and persistence and delays hybrid speciation.All pre-zygotic barriers to reproduction leading to hybrid formation are incomplete: parental species have overlapping flowering; they share the same pollinators;reciprocal assessments of pollen tube germination and growth do not differ among parents. The absence of post-zygotic barriers between parental taxa indicates that the persistence of hybrids is likely. Reproductive isolation was incomplete between hybrids and parents in all cases studied, although asymmetric differences in reproductive fitness were prevalent and possibly explain the genetic structure of natural hybrid swarms where hybridization is known to be bidirectional but asymmetric. Introgression, rather than speciation, is a probable evolutionary outcome of hybridization between the two Rhododendron taxa. Our study provides insights into understanding the evolutionary implications of natural hybridization in woody plants.     

An examination of hybridization between the cattail species typha latifolia and typha angustifolia using random amplified polymorphic DNA and chloroplast DNA markers

Typha glauca represents a significant portion of the biomass of the wetlands surrounding the Great Lakes, USA. It is generally accepted to be a form of hybrid between T. latifolia and T. angustifolia, which itself appears to be an exotic introduction from Europe. Based on morphological and isozyme data, conflicting theories have been proposed for the hybrid nature of T. glauca: it has been described as a hybrid swarm, a distinct hybrid species and an F1 hybrid. Therefore, we developed random amplified polymorphic DNA (RAPD) and chloroplast DNA markers, specific to the parental species, to assess hybrids. Ten RAPD primers gave 17 fragments specific to T. angustifolia and 13 fragments specific to T. latifolia. All of the interspecific hybrids contained each of the species-specific markers, indicating an F1 hybrid status. Furthermore, all hybrids tested contained the T. angustifolia chloroplast haplotype, which is consistent with differential interspecific crossing success found previously. Additional confirmation of an F1 hybrid status was gained by examining seedlings from T. glauca. These progeny were expected to be advanced-generation hybrids, as opposed to the F1 hybrid parent. Analysis of the seedlings revealed segregating marker patterns consistent with patterns observed in experimental advanced-generation hybrids, although these advanced hybrids do not appear to be a significant part of mature stands. Our data do not provide support for extensive gene flow between T. latifolia and T. angustifolia. However, our results suggest that hybridization between the native and introduced Typha species has impacted the native population through the spread of the F1 hybrid, T. glauca.     

Risk assessment by crow phenotypes in a hybrid zone

Predation is one of the most selective forces in evolution and, thus, predation may select against hybrids in narrow hybrid zones. It may be possible that parental phenotypes and hybrids differ in their responses towards predators or humans. As predation is difficult to observe I used flight-initiation distance (FID) as a metric of risk assessment. FID is a measurable outcome of the trade-off between fleeing and remaining. Here, I tested whether hybrid and parent crow phenotypes (Corvus corone, Corvus cornix) from the hybrid zone in Eastern Germany differ in their FID. Further, I measured many environmental and social variables to control statistically for their influence on FID. I sampled 154 individuals (53 hooded crows, 54 carrion crows, and 48 hybrids) in the hybrid zone in eastern Germany. I calculated a general linear model using a stepwise backward procedure to establish a minimum model containing only significant variables that explained FID in crows. The variable phenotype (hooded, carrion, hybrid) was then added to the model. There were no differences in FID between hybrids and both parental phenotypes types, suggesting similar risk assessment. This suggests that hybrids may behave similarly in their decision to flee as their parent phenotypes, which, in turn, provides no evidence for a selective disadvantage. An additional analysis focusing on pure phenotypic flocks showed that hybrids in pure hybrid flocks had a lower FID than both parental species in pure flocks. This suggests that hybrids in pure hybrid flocks may be at a disadvantage.     

Molecularmarker diversity and hybrid sterility in indica-japonica rice crosses

 The partial sterility of hybrids between the indica and japonica rice subspecies of Asian cultivated rice is a serious constraint for utilizing inter-subspecific heterosis in hybrid rice breeding. In this study, we have investigated the relationship between molecular-marker polymorphism and indica-japonica hybrid fertility using a diallel set involving 20 rice accessions including 9 indica and 11 japonica varieties. Spikelet fertility of the resulting 190 F₁s and their parents was examined in a replicated field trial. Intra-subspecific hybrids showed much higher spikelet fertility than inter-subspecific hybrids except in crosses involving wide-compatibility varieties. The parents were surveyed for DNA polymorphism using 96 RFLP and ten SSR markers, which revealed extensive genetic differentiation between indica and japonica varieties. A large number of markers detected highly significant effects on hybrid fertility. The chromosomal locations for many of the positive markers coincided well with previously identified loci for hybrid sterility. The correlation between hybrid fertility and parental distance was low in both intra- and inter-subspecific crosses. The results suggest that the genetic basis of indica-japonica hybrid sterility is complex. It is the qualitative, rather than the quantitative, difference between the parents that determines the fertility of hybrids. Received: 3 January 1997/Accepted: 17 January 1997     

四倍体大燕麦×六倍体裸燕麦的杂种F1的产生及鉴定

本研究以四倍体大燕麦 (AvenamagnaL .)做母本 ,六倍体裸燕麦 (AvenanudaL .)做父本进行杂交 ,利用幼胚拯救技术获得了杂种F1,并对其后代形态特征进行了观察 ;对杂种F1同工酶图谱和DNA指纹图谱进行了分析。杂种F1形态特征偏亲本或介于双亲之间 ;同工酶研究表明多数F1具有双亲互补酶带 ;RAPD分析不同引物扩增产物F1呈共显性或偏父、偏母。这些结果表明F1为真杂种     

四倍体大燕麦×六倍体裸燕麦的杂种F1的产生及鉴定

本研究以四倍体大燕麦（Avena magna L.）做母本,六倍体裸燕麦（Avena nuda L.）做父本进行杂交,利用幼胚拯救技术获得了杂种F1,并对其后代形态特征进行了观察;对杂种F1同工酶图谱和DNA指纹图谱进行了分析。杂种F1形态特征偏亲本或介于双亲之间;同工酶研究表明多数F1具有双亲互补酶带;RAPD分析不同引物扩增产物F1呈共显性或偏父、偏母。这些结果表明F1为真杂种。