The Genus Cerion (Gastropoda: Cerionidae) in the Florida Keys

The systematic relationships and phylogeography of Cerion incanum, the only species of Cerion native to the Florida Keys, are reviewed based on partial sequences of the mitochondrial COI and 16S genes derived from 18 populations spanning the range of this species and including the type localities of all four described subspecies. Our samples included specimens of Cerion casablancae, a species introduced to Indian Key in 1912, and a population of C. incanum x C. casablancae hybrids descended from a population of C. casablancae introduced onto Bahia Honda Key in the same year. Molecular data did not support the partition of C. incanum into subspecies, nor could populations be apportioned reliably into subspecies based on morphological features used to define the subspecies. Phylogenetic analyses affirmed the derived relationship of C. incanum relative to other cerionids, and indicated a Bahamian origin for the Cerion fauna of southern Florida. Relationships among the populations throughout the Keys indicate that the northernmost populations, closest to the Tomeu paleoislands that had been inhabited by Cerion petuchi during the Calabrian Pleistocene, are the oldest. The range of Cerion incanum expanded as the archipelago that is the Florida Keys was formed since the lower Tarantian Pleistocene by extension from the northeast to the southwest, with new islands populated as they were formed. The faunas of the High Coral Keys in the northeast and the Oölite Keys in the southwest, both with large islands that host multiple discontinuous populations of Cerion, are each composed of well supported clades that are characterized by distinctive haplotypes. In contrast, the fauna of the intervening Low Coral Keys consist of a heterogeneous series of populations, some with haplotypes derived from the High Coral Keys, others from the Oölite Keys. Individuals from the C. incanum x C. casablancae hybrid population inhabiting the southeastern coast of Bahia Honda Key were readily segregated based on their mitogenome lineage, grouping either with C. incanum or with C. casablancae from Indian Key. Hybrids with C. casablancae mitogenomes had haplotypes that were more divergent from their parent mitogenome than were hybrids with C. incanum mitogenomes.     

PATTERNS OF HYBRIDIZATION IN THE PIRIQUETA CAROLINIANA COMPLEX IN CENTRAL FLORIDA: EVIDENCE FOR AN EXPANDING HYBRID ZONE

We have identified a broad zone of hybridization between two morphologically and ecologically distinct herbaceous perennial taxa (morphotypes) within the Piriqueta caroliniana complex, which extends more than 300 km across the central Florida peninsula. Phylogeographic analyses indicate that the caroliniana morphotype has been present in north and central Florida since the early Pleistocene and that the viridis morphotype has immigrated into southern Florida much more recently. We examine the distribution of diagnostic morphological characters and nuclear genetic markers to assess the extent and patterns of introgression in this system. The results are consistent with the hypothesis that this hybrid zone has expanded north in recent history as viridis alleles have introgressed into regions that were previously occupied by populations of caroliniana. Genetic markers diagnostic for caroliniana have consistently high frequencies across the hybrid zone, whereas markers for viridis are extremely variable among populations with frequency reversals in adjacent populations. The latter pattern is probably the result of the combined stochastic effects of dispersal and drift on viridis alleles as they introgressed northward. Additional evidence for the recent expansion of this hybrid zone comes from patterns of variation for morphological and genetic markers. As expected for an expanding hybrid zone, within-population morphological variation was greatest toward the advancing front of introgression and levels of genetic variation for neutral diagnostic markers were greatest in the region of initial contact and lower in areas of recent expansion. The observed patterns of variation suggest that at least some hybrid genotypes have high fitnesses, which has led to the expansion of the hybrid zone via the displacement of parental genotypes in central Florida.     

Hybridization and asymmetric introgression between Cypripedium tibeticum and C. yunnanense in Shangrila County,Yunnan Province,China

Hybridization and introgression are thought to be important for speciation and adaptation in many plants. However, little is known about the hybridization and introgression among Cypripedium species. To investigate the evidence for hybridization and the pattern of introgression between Cypripedium yunnanense and C. tibeticum in Shangrila County, Yunnan Province, China, morphological characters and amplified fragment length polymorphism (AFLP) data for both the species and their putative hybrids were studied. Hand pollination was also performed to verify the crossability of the putative parents. Principal coordinate analysis based on morphological characters and the AFLP data suggested that the putative hybrids were true hybrids of these two Cypripedium species. Analysis with the NewHybrids software indicated that the putative hybrids were F1 generation individuals and backcrosses to C. yunnanese, but no F2 generation was found. Analysis with the Structure software demonstrated asymmetric introgression from C. tibeticum to C. yunnanense. We conclude that natural hybridization and introgression can occur between these two species and that in situ conservation of the parental species is required before fully assessing the evolutionary potential of hybrids.     

SIGNIFICANT ROLE FOR HISTORICAL EFFECTS IN THE EVOLUTION OF REPRODUCTIVE ISOLATION: EVIDENCE FROM PATTERNS OF INTROGRESSION BETWEEN THE CYPRINID FISHES,LUXILUS CORNUTUS AND LUXILUS CHRYSOCEPHALUS

Samples of Luxilus cornutus, Luxilus chrysocephalus, and their hybrids were collected along hypothesized routes of dispersal from Pleistocene refugia to examine the significance of geographic variation in patterns of introgression between these species. Patterns of allozyme and mitochondrial DNA (mtDNA) variation were generally consistent with those from previous studies. Tests of Hardy-Weinberg equilibrium revealed significant deficiencies of heterozygotes in all samples, indicating some form of reproductive isolation. Mitochondrial DNAs of each species were not equally represented in F₁ hybrids; however, this bias was eliminated when the two largest samples were excluded from the analysis. Backcross hybrids exhibited biased mtDNA introgression, as samples from Lake Erie (eastern) and Lake Michigan (western) drainages showed significant excesses of mtDNAs from L. chrysocephalus and L. cornutus, respectively, relative to frequencies of diagnostic allozyme markers. The extent and direction of allozyme and mtDNA introgression was quantified by calculating isolation index values from morphologically “pure” individuals of each species from each locality. Analysis of variance of these measures identified limited introgression of allozyme variants with no geographic pattern, but significant differences in direction of mtDNA introgression between drainages (i.e., postglacial dispersal route). Association between patterns of mtDNA introgression and dispersal route across the latitudinal width of the contact zone is best explained by genetic divergence during past isolation of ancestral populations from these drainages. These results identify a significant role for historical effects in the evolution of reproductive isolation and the process of speciation.     

Extensive hybridization in hawksbill turtles (Eretmochelys imbricata) nesting in Brazil revealed by mtDNA analyses

Bahia state hosts over 90% of hawksbill (Eretmochelys imbricata) nests registered in the main nesting sites monitored by Projeto Tamar-IBAMA in Brazil. The genetic diversity of this hawksbill population (n=119) was assayed through the analyses of 752 bp of the mitochondrial DNA control region in nesting females. Seven distinct haplotypes, defined by 125 polymorphic sites, were found. Most of the individuals (n=67) display four typical hawksbill haplotypes, 50 individuals display two haplotypes characteristic of the loggerhead turtle (Caretta caretta) and two individuals had a haplotype affiliated with the olive ridley (Lepidochelys olivacea). These results demonstrate hybridization between the hawksbills and two species that nest along the Bahia coast. Of special interest is the high occurrence of loggerhead × hawksbill hybrids (42%), which display loggerhead mtDNA haplotypes but are characterized morphologically as hawksbills. The true hawksbill haplotypes present only three variable sites and low genetic diversity values (h=0.358±0.069; π=0.0005±0.0001). The occurrence of several nesting individuals with identical mtDNA from another species may also suggest a long history of introgression between species producing likely F2 or further generation hybrids. Marine turtle hybrids have been previously reported, but the high frequency observed in Bahia is unprecedented. Such introgression may influence evolutionary pathways for all three species, or may introduce novel morphotypes that develop apart from the parental species. The presence of a unique hybrid swarm has profound conservation implications and will significantly influence the development and implementation of appropriate management strategies for these species.     

Genetic evidence for natural hybridization and apparent introgression between freshwater snail species (Viviparus ater and V. contectus)

We examined six allozyme markers and shell morphology to study gene flow between the naturally sympatric freshwater snail species Viviparus ater and V. contectus in Lake Garda, Italy, using offspring from experimental crosses and snails collected from natural populations. Hybrid offspring obtained from experimental crosses and hybrids collected from natural populations were heterozygous at four or five out of five diagnostic loci. Shell morphology was a poor predictor of the hybrid status of individual snails. Backcrossed offspring from experimentally bred hybrids and either of the parental species morphologically resembled the parental species. There was no evidence of segregation distortion at the six loci. Linkage analysis revealed one pair of linked loci (GPI and PNP). These cross experiments indicated a Mendelian type hybrid system in which gene introgression may occur. F₁ hybrids were found at four out of six sampling sites of Lake Garda, Italy. Local frequencies of F₁ hybrids ranged from 0% to 1.6% (estimated average = 0.74%) of the total population including both species. Alleles typical of V. ater were found at low frequencies in V. contectus at all six sampling sites. Alleles typical of V. contectus were found at low frequencies in V. ater at three out of six sampling sites. This is consistent with the hypothesis of introgression in both directions.     

A CYTOLOGICAL CONSPECTUS OF THE GENUS CALYCADENIA (ASTERACEAE): AN EXAMPLE OF CONTRASTING MODES OF EVOLUTION

The chromosome numbers of 12 taxa of Calycadenia are documented by determinations in 77 widespread populations of the genus. Results of biosystematic studies indicate that hybrids among C. multiglandulosa, C. ciliosa, C. pauciflora, C. hispida, C. spicata, and C. oppositifolia characteristically have multiple associations of chromosomes producing complex meiotic configurations, whereas hybrids among C. villosa, C. mollis, C. truncata, and C. tenella generally yield few or no bivalents and a predominance of univalents during meiosis. Hybrids between C. multiglandulosa and C. hispida exhibit strict bivalent pairing and high pollen stainability. It is demonstrated that C. pauciflora and C. ciliosa comprise several sibling taxa that are morphologically cryptic, yet highly differentiated cytologically. Conversely, it is concluded that C. multiglandulosa and C. hispida comprise at least five morphologically distinct taxa that so far as known are highly interfertile (as judged by pollen stainability of hybrids) and cytologically uniform. Reproductive biology, chromosome mutation rates, habitats, flowering time, and genetic recombination mechanisms of the C. pauciflora-C. ciliosa and C. multiglandulosa-C. hispida complexes are compared in an attempt to account for the different modes of evolution observed in these closely related groups of Calycadenia. Lacking other plausible explanations, it is suggested that differential methods for limiting recombination of critical adaptive gene complexes might account for the divergent evolutionary patterns encountered in Calycadenia.     

A BIOSYSTEMATIC STUDY OF THE POLYGONUM HYDROPIPEROIDES (POLYGONACEAE) COMPLEX

The Polygonum hydropiperoides complex includes P. hydropiperoides Michx., P. opelousanum Riddell ex Small, P. setaceum Baldwin ex Ell., and P. hisutum Walter. Studies of morphology, cytology, and crossing compatibility were conducted with specimens from North Carolina, South Carolina, Georgia, and Florida. A multivariate cluster analysis using 212 specimens and 34 characters indicated that P. hirsutum and P. setaceum are each morphologically distinct while P. hydropiperoides and P. opelousanum are morphologically indistinct from one another. The chromosome counts of 2n = 20 for P. setaceum and P. hirsutum, and 2n = 40 for P. hydropiperoides and P. opelousanum are the first reported for the respective species. Experimental hybridizations produced the fertile hybrids P. hydropiperoides x opelousanum and P. setaceum × hirsutum. Results of these studies and population structure, habitat, and pollination studies suggest that P. hydropiperoides, P. setaceum, and P. hirsutum are each distinct species while P. opelousanum is indistinct and should be merged with P. hydropiperoides.     

THE ORIGIN OF AGROPYRON ALBICANS

Previous suggestions of introgression between Agropyron spicatum (Pursh) Scribn. & Smith, 2n = 14 & 28, and Agropyron dasystachyum (Hook.) Scribn., 2n = 28, were confirmed. Fertile, meiotically regular, 28-chromosome plants morphologically identical to Agropyron albicans Scribn. & Smith, 2n = 28, occurred in first- and second-generation open-pollination progenies of diploid A. spicatum × A. dasystachyum hybrids, presumably by backcrossing to A. dasystachyum. These A. albicans-like derivatives were fully cross-compatible with naturally occurring A. albicans. First and second generation open-pollination progeny of tetraploid A. spicatum × A. dasystachyum F₁'s contained approximately 5% A. albicans-like plants; but none was tetraploid, cytologically stable, and fertile. Although introgression occurs freely between tetraploid A. spicatum and A. dasystachyum, derivation of fertile true-breeding A. albicans from their early-generation progeny seems unlikely. Agropyron griffithsii Scribn. & Smith ex. Piper, the glabrous counterpart of A. albicans, probably originated from hybrids between diploid A. spicatum and Agropyron riparium Scribn. & Smith, the glabrous form of A. dasystachyum. Genome formulas of diploid A. spicatum, A. dasystachyum (riparium), and A. albicans (griffithsii) may be written as S₁S₁, S₂S₂XX, and S_1-2 S_1-2XX, respectively. The relationship between A. albicans and A. dasystachyum is so close that A. albicans should be regarded as no more than a subspecies of A. dasystachyum.     

ESSENTIAL OILS AS TAXONOMIC AIDS IN THE CLASSIFICATION OF DICHANTHIUM PARVIFLORUM

Essential oil components and gross morphological characters are closely correlated in Dichanthium parviflorum (R. Br.) de Wet et Harlan (Gramineae) and related species. Different species, varieties, and geographical races, as well as hybrids between them, can be identified on the basis of absence or presence and quantity of essential oil components. The morphologically variable D. parviflorum was subdivided into four varieties: var. parviflorum, var. capilliflorum (Steud.) de Wet et Harlan comb. nov., var. mutispiculum (Ohwi) de Wet et Harlan comb. nov., and var. spicigerum (S. T. Blake) de Wet et Harlan comb. nov. These varieties differ from each other morphologically in having respectively racemes with 1-4 and awned, 3-5 and awned, 1-2 and awnlass, and 4-10 and awned spikelet pairs per raceme.     

History as a cause of area effects: an illustration from Cerion on Great Inagua, Bahamas

The two parts of this paper work towards the common aim of setting contexts for and documenting explanations based on historical contingencies. The first part is a review of area effects in Cepaea. We discuss the original definitions and explanations, emphasizing the debate of adaptationist vsstochastic approaches, but arguing that the contrast of historical contingency vs. selective fit to environment forms a more fruitful and fundamental context in discussing the origin of area effects. We argue that contingencies of bottlenecks and opening of formerly unsuited habitats may explain the classic area effects of Cepaea better than selectionist accounts originally proposed. The second part is a documentation of an area effect within Cerion columnaon the northern coast of Great Inagua, Bahamas. Historical explanations are often plagued by insufficiency of preserved information, but the Inagua example provides an unusual density of data, with several independent criteria all pointing to the same conclusion. Shells in the area effect are squat and flat-topped in contrast with typical populations of long, thin, tapering shells living both east and west of the area effect. The flat-topped area effect is a result of introgression with a propagule of the C. dimidiatum stock (living on nearby Cuba, and most apically flattened of all Cerion). Fossils of this propagule were found fully cemented into highly indurated fossil soil crusts within the region of the current area effect. Multivariate morphometry, based on complex patterns of covariation, not just intermediacy in single characters, identifies the area effect samples as hybrids between this propagule and typical C. columna. Genetic analysis has identified three unexpected alleles in area effect samples only, and in no other snails of any other Cerion taxon anywhere else on Inagua. We hypothesize that the flattopped area effect did not arise as a selective response to local environments within C. columna, but by introgression from a fortuitously introduced propagule of the C. dimidiaium complex. The unexpected alleles therefore represent genetic phantoms of C. dimidiatum's former presence or are hybrizymes—novel alleles produced by interspecific hybridization     

NATURAL AND ARTIFICIAL HYBRIDS OF HELIANTHUS MAXIMILIANI X H. GROSSESERRATUS

Long , Robert W. (Ohio Wesleyan U., Delaware.) Natural and artificial hybrids of Helianthus Maximiliani × H. grosseserratus. Amer. Jour. Bot. 46(10): 687–692. Illus. 1959.—An investigation of the occurrence of natural hybridization in two perennial sunflowers, Helianthus Maximiliani and H. grosseserratus, was begun in 1950. Subsequently, artificial F₁, F₂, and first and second backcross generations were produced. Fertility and vigor were high in all these plants, but F₁ plants appeared to excel the others in these characteristics. Observations in the experimental garden were supplemented by examination of chromosomes in pollen mother cells, comparisons of herbarium collections, and study of wild populations. Evidence pointed to close genetic relationship of the species and to the occurrence of natural hybridization in areas of distributional overlap. In 1957 and 1958, field work in these areas resulted in the scoring of 18 natural populations, 3 of which consisted of both parental species plus putative F₁ hybrids. Two explanations are offered to account for the seeming absence of introgression. The results support the conclusion that natural hybridization leads to the establishment of F₁ hybrids and that introgression does not occur to any significant extent. Although both species display a high degree of interfertility, they are distinct morphologically. For this reason, it is advisable to maintain them as separate species.     

Interspecific hybridization between Vigna radiata (L.) Wilczek and V. glabrescens

Summary Interspecific hybrids of the mungbean, Vigna radiata (L.) Wilczek (2n=22) and V. glabrescens (2n=44) were generated with the aid of embryo culture. V. glabrescens x V. radiata hybrids were recovered via germination of the immature embryos. Reciprocal hybrids were obtained via shoot formation from embryonic callus. The authenticity of the hybrids was determined by morphological characteristics, chromosome number, and isozyme patterns. The hybrids were highly sterile upon selfing, but backcrossing to the diploid parent yielded viable seeds. Some of the plants resembled the diploid parent morphologically while others resembled neither parent. The backcross plants were sufficiently fertile to give a large number of mature, selfed seeds. Plants obtained differed morphologically and in their isozyme patterns from either parent, indicating introgression. These progeny populations will be used as bridging materials to transfer pest resistance from the wild tetraploid to the cultivated mungbean.     

Evidence for introgression in azaleas (Rhododendron; Ericaceae): Chloroplast DNA and morphological variation in a hybrid swarm on Stone Mountain,Georgia

Hybridization and introgression have been suggested by previous workers as important factors in the evolution of the southeastern azaleas. However, because of high levels of intraspecific variation and lack of intensive sampling, documentation of this phenomenon has been difficult. We investigated a population of azaleas on Stone Mountain, Georgia, composed of Rhododendron flammeum, R. canescens, and their interspecific hybrids. After analysis of chloroplast DNA, we found evidence of extensive localized cytoplasmic introgression into R. flammeum. Many individuals in the sample population that are morphologically indistinguishable from R. flammeum possess the chloroplast genome of R. canescens. This suggests that some of the observed variation in some populations of R. flammeum may be due to past introgression from R. canescens.     

Molecular and morphological evidence of hybridization between newts Triturus vulgaris and T. montandoni (Caudata: Salamandridae) in Slovakia

Natural hybridization and introgression is recorded between the newt species Triturus vulgaris and T. montandoni in Slovakia. To confirm a hybrid status of two putative hybrids, morphological and Randomly Amplified Polymorphic DNA (RAPD) markers specific for T. vulgaris and T. montandoni were used. The individuals found in Zbojská (Veporské vrchy Mts, Slovakia; analyzed morphologically and genetically) and Zubrohlava (Oravská kotlina basin, Slovakia; analyzed only morphologically) possessed markers of both species. Segregation of RAPD markers together with a model-based Bayesian analysis revealed that the specimen from Zbojská belonged to later generation hybrids.     

THE ORIGIN AND RELATIONSHIPS OF LASTHENIA BURKEI (COMPOSITAE)

Lasthenia burkei (Compositae) is a narrowly restricted California endemic closely related to L. conjugens and L. fremontii. These three species differ from each other by pappus and phyllary characters and in geographical distribution. All are freely intercrossable, but L. fremontii forms rather sterile artificial hybrids with its two relatives which, in turn, form fairly fertile artificial hybrids with each other. Lasthenia burkei and L. conjugens have homologous chromosomes, four of which are homologous with four of those of L. fremontii. The remaining two chromosomes probably have reciprocal translocations which lead to multivalent formation during meiosis in interspecific hybrids. Pollen viability is restored in most F₂ generations, suggesting a close genetic relationship among the three species. The evolutionary relationship among these species may be a linear one with L. burkei occupying an intermediate position between L. fremontii and L. conjugens, although the direction of this linear phylogeny is not certain, or it may be one in which L. burkei has been derived from hybridization between its two relatives. Support for the latter hypothesis comes from the appearance of some individuals in F₁ progenies of L. conjugens × L. fremontii that are morphologically indistinguishable from L. burkei (although fairly sterile). The apparently rather simple genetic basis for the morphological characteristics of each of the species in this trio suggests that the morphologically heterogeneous genus Lasthenia may be considerably more homogeneous genetically than might be suspected. Because of the diverse kinds of relationships among these three Lasthenias, possible alternative taxonomies for the group are dependent upon those relationships that a taxonomist wishes to communicate. Nevertheless, the patterns of diversification in this group have led to reaffirmation of an earlier decision that three species should be recognized.     

NATURAL HYBRIDIZATION BETWEEN THE SYMPATRIC HAWAIIAN SPECIES DROSOPHILA SILVESTRIS AND DROSOPHILA HETERONEURA

Two newly formed, morphologically distinct species of Drosophila from the island of Hawaii have been found to form fertile hybrids in two areas of sympatry. Both F₁ and backcross hybrids have been recognized in nature; in one case, the hybridization events extended over three years. Original hybridizations involved one or more D. silvestris females mating with D. heteroneura males. Female F₁ hybrids from this cross have participated in backcrosses to D. silvestris. In any one locality, less than 2% hybrids have been found in nature. A hybrid swarm was not formed; selection appears to favor a strict maintenence of morphologies characteristic of the separate species. This result is attributed to pervasive sexual selection, which serves to preserve the syndromes of sexual characteristics that arose during past allopatric divergence. Populations of D. silvestris both within and outside the present range of D. heteroneura often display heritable variation in color patterns involving the abdomen, pleurae, legs, and wings. Genes effecting variation in these characters may be derived from genes involved in a past introgression from D. heteroneura. Independent evidence for past hybridization between these species comes from study of mitochondrial DNA. Although the inferred direction of the cross is the opposite of that observed in the recent case described here, both reciprocal crosses have been obtained experimentally in the laboratory. Accordingly, we suggest that these species may have been open to hybridization since their first sympatic encounters following their inception in allopatry. That they remain as strictly recognizable morphological entities is due both to their current partial allopatry and to the action of sexual selection in maintaining two separate major modes of efficient reproduction. There is no reason to invoke specific reinforcing selection that has imposed reproductive isolation.     

OBSERVATIONS ON INTROGRESSION OF TRIPSACUM INTO MAIZE

Derivatives of a cross between diploid Zea mays L. and Tripsacum dactyloides (L.) L. (2n = 72) were compared cytologically and morphologically. The objective of this study was to detect introgression from Tripsacum to maize that might have occurred during seven backcross generations with maize. Thirty-three morphological characters were used to analyze variation among aneuploid (20Zm + 2Td), 20-chromosome recovered maize, and the recurrent maize parent plants. Aneuploid and maize checks were extreme types, with 20-chromosome hybrid derivatives being morphologically intermediate. Several recovered maizes clustered with aneuploid plants and these hybrid derivatives have the greatest chance of Tripsacum introgression. Many traits such as endosperm abnormalities, tassel seed, albinos, tunicate glumes, tassel-tipped ears, fasciated and branched ear, and male spikelets between rows of kernels were observed. Although the genetic basis of many traits is unknown, mutations, epistatic effects or expression of Tripsacum chromatin are possible causes. The number of abnormal and tripsacoid traits observed in 20-chromosome recovered maizes indicates genetic transfer from Tripsacum to the maize genome.     

Morphological and molecular evidence of natural hybridization in Shorea (Dipterocarpaceae)

Shorea (Dipterocarpaceae) is a large genus in which many closely related species often grow together in Southeast Asian lowland tropical rain forests. Many Shorea species share common pollinators, and earlier studies suggested occurrence of interspecific hybridization and introgression. Here, we show morphological and molecular evidence of hybridization between Shorea species. In the census of all the trees of Shorea curtisii, Shorea leprosula, and Shorea parvifolia (>30 cm dbh) within the 164-ha area of Bukit Timah Nature Reserve in Singapore, we found 21 morphologically recognizable hybrid individuals. All of the putative hybrids could be distinguished obviously from the parental species on the basis of vegetative characters. Population genetic analysis of DNA sequences of two nuclear (GapC and PgiC) and chloroplast (trnL-trnF) regions demonstrated that each of the three species had several species-specific mutations. The nuclear sequences of the putative hybrids were heterozygote at all the species-specific sites between two parental species. Hybrid between S. curtisii and S. leprosula was found most, while S. curtisii × S. parvifolia and S. leprosula × S. parvifolia hybrids were also found. Almost no shared polymorphism between populations of the parental species suggests rarity of introgression. The study indicated that natural hybridization between sympatric Shorea species should not be uncommon, but all of the hybrid individuals were F₁, and the post-F₁ hybrids were considerably rare.     

HYBRIDIZATION AND INTROGRESSION IN THE GRASS GENUS ELYMUS

Brown , W. V., and G. A. Pratt . (U. Texas, Austin.) Hybridization and introgression in the grass genus Elymus. Amer. Jour. Bot. 47 (8) : 669–676. Illus. 1960.—The production of artificial F₁ hybrids that produce some seed between E. virginicus and E. canadensis, E. virginicus and E. interruptus, and E. canadensis and E. interruptus proves that similar plants found in nature are true hybrids. The artificial hybrids constituted the basis of comparison in an analysis of natural local populations in the Austin region. Analysis by scatter diagrams indicates that there is introgression of genes of some species into the genomes of others through the development of hybrid swarms. In the area studied the great variability of virginicus is due to the introgression of canadensis and, possibly, interruptus genes. Such hybridization and subsequent introgression have occurred many times and in many places and are still taking place. It is likely that the named varieties of canadensis and virginicus may be introgressant types that are the results of hybridizations between those species and others that occur sympatrically in Eastern United States.