Hybridization between Senecio smallii and S. tomentosus (Compositae) on the granitic flatrocks of the Southeastern United States

Senecio smallii andS. tomentosus occasionally hybridize on the granitic flatrocks of the Piedmont of the Southeastern United States. These hybrids are intersectional crosses between sectionsAurei andTomentosi. The parental species have different chromosome numbers withn = 22 inS. smallii andn = 23 inS. tomentosus. Meiosis in the hybrids is abnormal with 22ns and li or 2iis and 3is, and the percentage of stainable pollen is greatly reduced over that of the parents. The hybrid pollen grains are structurally abortive as demonstrated by the scanning electron microscope. Origin of the natural hybrids was documented by the production of artificial hybrids including F1’s, F2’s, and BC’s. Very few of the F2 seeds germinated, and all of the F2 seedlings later died, indicating hybrid breakdown. Seeds of the F1 and BC generations germinated well, and the seedlings were vigorous. Population samples indicate that there is some variation within the natural population systems; however, no evidence of introgression was detected. The isolating mechanisms and possible evolutionary effects of this natural hybridization are discussed.     

Apomixis,hybridization, and taxonomic complexity in eastern north AmericanAmelanchier (Rosaceae)

Apomixis and hybridization together contribute to taxonomic complexity inAmelanchier. Hybridization combines genetically divergent genomes and spawns new forms that apomixis perpetuates. Apomixis is aposporous, facultative, and pseudogamous in the genus, and apomicts are generally polyploid, pollen fertile, and pollinated by generalists. That gene flow actually occurs is empirically evident. As apomixis is genetically dominant over sexuality, hybrids involving at least one apomictic parent are apomictic. Clonal reproduction may thus perpetuate F₁ individuals and generate agamospecies. Alternatively hybrids may interbreed and backcross to create hybrid swarms or cross with species other than the parents. In eastern North America, the abundance of published names and general taxonomic confusion in the genus doubtless result at least in part from this interplay of apomixis and hybridization. The roles of apomixis and hybridization in diversification withinAmelanchier are examined in light of new data about breeding system of an apomictic, hybrid microspecies, informally namedA. “erecta” and its formation of a hybrid swarm with anotherAmelanchier apomict,A. laevis.     

How do biological control and hybridization affect enemy escape?

Two mechanisms often linked with plant invasions are escape from enemies and hybridization. Classical biological control aims to reverse enemy escape and impose top-down population control. However, hybridization has the potential to alter interactions with enemies and thus affect biological control. We examined how introductions of biological control agents affect enemy escape by comparing specialist enemy loads between the native and introduced ranges of two noxious weeds (spotted and diffuse knapweed; Centaurea stoebe L. and C. diffusa Lam.) that have been the targets of an extensive biological control program. Hybrids between spotted and diffuse knapweed are often found within diffuse knapweed sites in North America, so we also compared enemy loads on plants that appeared morphologically like diffuse knapweed and hybrids. Finally, we tested the preference for diffuse knapweed, hybrids, and spotted knapweed of one of the agents thought to be instrumental in control of diffuse knapweed (Larinus minutus; Curculionidae). In North America spotted knapweed has largely escaped its root herbivores, while seedhead herbivore loads are comparable in the introduced and native ranges. Diffuse knapweed exhibited seedhead herbivore loads five times higher in the introduced compared to native range. While this pattern of seedhead herbivory is expected with successful biological control, increased loads of specialist insect herbivores in the introduced range have rarely been reported in the literature. This finding may partially explain the better population control of diffuse vs. spotted knapweed. Within North American diffuse knapweed sites, typical diffuse knapweed and hybrid plants carried similar herbivore loads. However, in paired feedings trials, the specialist L. minutus demonstrated a preference for newly created artificial hybrids over North American diffuse knapweed and for European diploid spotted knapweed over North American tetraploid spotted knapweed. Overall though, hybridization does not appear to disrupt biological control in this system.     

Taxonomic notes on Senecio tomentosus and its allies (Asteraceae)

Distribution maps, specimen citations and taxonomic commentary are provided for 14 species ofSenecio in temperate North America allied withS. tomentosus (the “Tomentosi”). These data support the author’s treatment of the group inNorth American Flora.     

Biology and host specificity ofChamaesphecia hungarica andCh. Astatiformis (Lep.: Sesiidae) two candidates for the biological control of leafy spurge,Euphorbia esula (Euphorbiaceae) in North America

Leafy spurge (Euphorbia esula (s.1.)) is an herbaceous perennial and serious weed of Eurasian origin that has been accidentally introduced into North America. The two European root-boring mothsChamaesphecia hungarica andCh. astatiformis are univoltine and overwinter as mature larvae. Both species have a lower survival rate on leafy spurge than on their field hosts, and thus are not optimal candidates for the biological control of leafy spurge. However, the rate of larval development and larval growth on the target weed and on the two field hosts is nearly the same. The experimental host range of both species is restricted to a few species in the subgenusEsula within the genusEuphorbia. The two species occupy different habitats in the steppe biome and are targeted for similar leafy spurge habitats in North America.     

Tricholoma fulvimarginatum (Tricholomataceae): A new species from North America associated with cottonwood

Tricholoma fulvimarginatum is described as a new species of Agaricales from eastern North America. This fungus belongs in subgenusTricholoma sectionGenuina due to the coloration of the pileus surface. The orange-brown pileus, marginate lamellae, and presumed mycorrhizal association withPopulus deltoides distinguish this species from others in the section.     

Population genetic structure of Picea engelmannii, P. glauca and their previously unrecognized hybrids in the central Rocky Mountains

Areas of geographic overlap between potentially hybridizing species provide the opportunity to study interspecific gene flow and reproductive barriers. Here we identified hybrids between Picea engelmannii and P. glauca by their genetic composition at 17 microsatellite markers, and determined the broad-scale geographic distribution of hybrids in the central Rocky Mountains of North America, a geographic region where hybrids and isolation between species had not previously been studied. Parameter estimates from admixture models revealed considerable variation in ancestry within and among collection sites, suggesting that within this area of geographic overlap, the interaction of the two species varies extensively. The results document a previously unrecognized patchy distribution of hybrids between P. engelmannii and P. glauca, including locations where hybrids were not known or expected to exist. Further, the ancestry of many hybrids was consistent with multiple generations of hybridization, with probable directional backcrossing to P. engelmannii, suggesting a relatively porous species boundary. The identification and characterization of hybridization between these spruce in this region raises the question of what factors maintain barriers to gene flow in these long-lived forest trees. The current research lays the groundwork for future study of the ecological and evolutionary contexts of their hybridization, as well as of differential introgression and permeability of species boundaries.     

Presence du genre Simoedosaurus (Reptilia,Choristodera) en Amerique du Nord

The genus Simoedosaurus, whose distinction from Champsosaurus has recently been proven in Europe, exists also in the Clarkforkian of North America: four specimens in the Collection of Princeton University provide the evidence. This genus can be added to the already long list of forms common to the two continents at the beginning of the Eocene, which attests to the ease of migration from one to the other at this time.     

Social behaviour of a subtropical paper wasp,Ropalidia fasciata (F.): Field observations during founding stage

Field studies showed that the percentage of foundation by multiple foundresses was remarkably high (34.5%) inRopalidia fasciata in Okinawa. Although one of the foundresses of the association usually remained on the nest and others tended to do extranidal activities, the dominance-subordination relation among cofoundresses seemed to be far milder than that reported inPolistes spp. of Europe and North America. Dominance relation sometimes reversed. All the foundresses could receive a share of food and feed larvae. It was suggested that a large risk of parasitism and predation should favour the trait to join foundress associations inR. fasciata.     

Genetic characterization of an arctic zooplankter: insights into geographic polyploidy

Species of Bosmina from the temperate regions of North America and Europe are diploid and reproduce by cyclical parthenogenesis. By contrast, this study provides evidence that the dominant bosminid taxon in High Arctic lakes reproduces by obligate parthenogenesis and is a polyploid derived from interspecific hybridization. Sinobosmina liederi, a species common in temperate North America, is likely to have been one parent of these hybrids, but the other parent is unknown. As neither parent was detected in the Arctic, it seems unlikely that the hybrid clones that now occupy arctic lakes were synthesized locally. Most habitats contained only one or two clones, despite a total of 38 clones in the region, suggesting that priority effects have been important in restricting diversity within single lakes. The high regional diversity of arctic bosminids could reflect either repeated hybridization between the parent taxa or the genetic instability of newly formed polyploid lineages. These processes would produce hybrid polyploids that are considerably more diverse than their sexual parent taxa, and this difference in genetic diversity may confer an advantage to the polyploid biotype. As many zooplankton taxa from the arctic possess genetic characteristics similar to those of bosminids, these processes may provide a general explanation for the widespread occurrence of polyploids in the Arctic.     

Meiotic anomalies induced by x-rays inCapsicum annuum L.

Various types of meiotic anomalies in Mi generation such as multivalents, fragments, bridges, mioronuclei, polyads and in particular the multispindle formation, were observed after seed X-ray irradiation inCapsicum annuum L. As the dose increased there was a gradual increase in the number of aberrations.     

Wood and stem anatomy of Misodendraceae: Systematic and ecological conclusions

The wood anatomy of the monogeneric South American family of epiparasites, Misodendraceae, contains numerous peculiarities. In the four species of the subgenusAngelopogon the cambium is virtually inactive in ray areas. In three of those species, a second ring of fascicular areas (“bundles”) forms in the pith only well after the initiation of the outer ring. Fibers occur in fascicular areas in paired strands (sheathed by crystalliferous parenchyma), as the background tissue, or may be absent, depending on the species; rays consist wholly of fibers or are composed of thin-walled cells, depending on the species. In the subgenusMisodendron (seven species), the cambium is active in fascicular areas and interfascicular areas equally; it contributes fibers to potential ray areas and to fascicular areas. If occurrence of these fibers is a criterion, the subgenusMisodendron may be considered rayless (except forM. gayanum, in which fibers occur only in rays). Growth rings are present in all species. Vessel elements are extremely short and narrow, numerous per mm², and pores are markedly grouped; all these are highly xeromorphic features appropriate to a parasite in an extreme climatic zone. In subgenusAngelopogon, lateral wall pitting of vessels consists of laterally much-widened pits, an appearance falsely resembling helical bands of primary xylem vessels. In subgenusMisodendron, pits on vessels are alternate or scalariformlike. The difference in vessel pitting between the two subgenera may relate to wood parenchymatization. Perforation plates are simple. Pits have wide apertures inM. angulatum andM. recurvum. Pith is sclerenchymatous in subgenusMisodendron but composed of thin-walled cells in which vessel strands occur in subgenusAngelopogon. Because of the numerous specific characteristics which correspond to the taxonomic system for the family, a key to species based on wood anatomy is given. New names must be employed for both subgenera for nomenclatural reasons; thus the combinationM. subgenusAngelopogon (=M. subgenusEumyzodendron) is coined. Wood anatomy of Misodendraceae is similar to that ofPsittacanthus and other Loranthaceae, disregarding the unique features found inMisodendron.     

ABNORMAL REDUCTIONAL AND NONREDUCTIONAL MEIOSIS IN CERATOPTERIS: ALTERNATIVES TO HOMOZYGOSITY AND HYBRID STERILITY IN HOMOSPOROUS FERNS

Natural and synthesized hybrids of Ceratopteris were investigated to determine the effect of hybridization on the genetic system. Studies indicated that the hybrids exhibited massive spore abortion and pairing abnormalities at meiotic prophase, characteristic of “sterile diploids and triploids” reported in hybridization studies of other fern genera. However, a small percentage of viable spores also was produced by the hybrids. Cytological investigations indicated the presence of previously unreported meiotic adaptations that allowed the production of unreduced spores and reduced spores exhibiting chromatid heterozygosity. The reduced spores allow haploid gametophytes to form heterozygous zygotes in spite of intragametophytic selfing. The unreduced spores were shown to be responsible for the fertility of the “sterile” hybrid and allowed the subsequent production of up to three generations of sporophytes. The literature suggests that these meiotic adaptations are present in other fern genera and may play a significant role in evolution through hybridization.     

Chromosome numbers,polyploidy, and hybridization in castilleja (scrophulariaceae ) of the Great Basin and Rocky Mountains

Chromosome numbers are reported for 190 collections ofCastilleja of the Great Basin-Rocky Mountains and adjacent regions in western North America. Polyploidy has played a major role in evolu tion of this genus. Of the 35 taxa represented, 14 are exclusively diploid (n =12). The remaining 21 taxa exhibit polyploidy: eight are tetraploid only, two are octopIoid only, while 11 of the more wide-ranging species have two or more levels of polyploidy. BothC. rhexifolia andC. sulphured have diploid, tetraploid, and octoploid races while the widespreadC. miniata has four (2x, 4x, 8x, and 10x). Where appropriate, the role of polyploidy and hybridization inCastilleja evolution and their relevance to taxonomy are discussed for individual species.     

Seedling morphology ofLespedeza (Leguminosae)

Seedling morphology was studied in 22 species ofLespedeza, of which six belong to subgenusMacroles-pedeza and 16 to subgenusLespedeza. Two seedling types were recognized: 1) those with opposite leaves at the first node; 2) those having the first and subsequent leaves alternate. The two types are distinguished in the number of leaf primordia in the plumule of the seed: the opposite type has two leaf primordia, but the alternate type has only one primordium. Most species exhibited one of the two types, but rarely both types were observed in several species. In species having two types, one type always far outnumbers the other. The opposite type was common in Asian species, while the alternate type was common in North American ones. Because seedlings are of the opposite type inKummerowia which is the most closely related genus withLespdeza, the alternate type is considered to be apomorphic inLespedeza. The apomorphic seedling morphology is, there-fore, dominantly occurred in North American species ofLespedeza. This fact may be an evidence which suggests a monophyletic origin for North American species from an Asian ancesfor.     

Global distribution of cryptic native,introduced and hybrid lineages in the widespread estuarine amphipod Ampithoe valida

Biological invasions can pose a severe threat to coastal ecosystems, but are difficult to track due to inaccurate species identifications and cryptic diversity. Here, we clarified the cryptic diversity and introduction history of the marine amphipod Ampithoe valida by sequencing a mtDNA locus from 683 individuals and genotyping 10,295 single-nucleotide polymorphisms (SNPs) for 349 individuals from Japan, North America and Argentina. The species complex consists of three cryptic lineages: two native Pacific and one native Atlantic mitochondrial lineage. It is likely that the complex originated in the North Pacific and dispersed to the north Atlantic via a trans-arctic exchange approximately 3 MYA. Non-native A. valida in Argentina have both Atlantic mitochondrial and nuclear genotypes, strongly indicating an introduction from eastern North America. In two eastern Pacific estuaries, San Francisco Bay and Humboldt Bay, California, genetic data indicate human-mediated hybridization of Atlantic and Pacific sources, and possible adaptive introgression of mitochondrial loci, nuclear loci, or both. The San Francisco Bay hybrid population periodically undergoes population outbreaks and profoundly damages eelgrass Zostera marina thalli via direct consumption, and these ecological impacts have not been documented elsewhere. We speculate that novel combinations of Atlantic and Pacific lineages could play a role in A. valida’s unique ecology in San Francisco Bay. Our results reinforce the notion that we can over-estimate the number of non-native invasions when there is cryptic native structure. Moreover, inference of demographic and evolutionary history from mitochondrial loci may be misleading without simultaneous survey of the nuclear genome.

     

Population studies in Goodyera (Orchidaceae) with emphasis on the hybrid origin of G. Tesselata

Morphological, cytological, and paper Chromatographic studies of populations from northern Michigan and examination of herbarium specimens from throughout North America were used to clarify the relationships ofGoodyera oblongifolia, G. repens var.ophioides, andG. tesselata. A canonical analysis of morphological data from mixed populations of these three species depictsG. tesselata as intermediate betweenG. oblongifolia andG. repens var.ophioides. The latter two species are diploid (2n = 30) andG. tesselata is tetraploid (2n = 60). Triploids (2n = ca. 45) were found in two mixed-species populations in northern Michigan.Goodyera tesselata produces three phenolic compounds present inG. oblongifolia and five different compounds present inG. repens var.ophioides. The range ofG. tesselata is confined to glaciated territory (except for two stations) in northeastern North America where the postglacially produced ranges ofG. oblongifolia andG. repens var.ophioides overlap. However,G. tesselata is quite abundant in areas outside the region of sympatry of the other two species. Based on this evidence, it is postulated thatG. tesselata is an allotetraploid species which resulted from hybridization betweenG. oblongifolia andG. repens var.ophioides during early post-Pleistocene. The slightly earlier blooming season ofG. tesselata may have been selected for to provide a measure of reproductive isolation between the tetraploid and its parents and to adapt the new species to the rather short growing season of northeastern North America.     

Patterns of hybridization in plants

Hybridization plays an important role in the evolution of many taxonomic groups, but large-scale phylogenetic patterns of hybridization are poorly known. Here, we investigate patterns of hybridization in vascular plants. Our dataset included 282 families, 3212 genera and ≈37,000 species accounts from eight regional floras covering continental Europe, two island regions, and parts of North America and Australia. Interspecific hybrids were common in the wild, occurring in 40% of families and 16% of genera, with an overall frequency of 0.09 hybrids per nonhybrid species. Taxon species richness explained a large amount of variation in the number of hybrids, but taxon bias (study effort) did not. We accounted for species richness in calculating hybridization propensities, and found that both families and genera differed in hybridization propensity. Hybridization propensity of a given group was generally consistent across regions (with the exception of Hawaii), suggesting that hybridization behavior may be determined more by intrinsic properties of a group than by environmental conditions. We found evidence of a strong phylogenetic signal (λ=0.93) in hybridization propensity as hybrids were not uniformly distributed across orders of vascular plants. Characterization of the hybridization behavior of groups should lead to increased predictive power regarding their traits and evolutionary trajectories, and will allow comparative tests of the traits driving differences in hybridization propensity.     

Apoptose et ségrégation méiotique dans les spermatozoïdes d'hommes porteurs de translocations

Men with a chromosomal translocation produce a significant percentage of unbalanced spermatozoa. In order to determine a correlation between chromosomal anomalies and apoptosis in human sperm, we analysed DNA fragmentation and meiotic segregation in sperm from men with a (13;14) Robertsonian translocation. We studied sperm from 12 (13;14) translocation carriers and 9 proven fertile men with a normal karyotype. Meiotic segregation of chromosomes 13 and 14 was analysed using dual-colour fluorescencein situ hybridization with locus-specific probes for chromosomes 13 and 14. Apoptosis in spermatozoa was measured byin situ TUNEL assay. The meiotic segregation study showed a significantly increased frequency of unbalanced spermatozoa for chromosomes 13 and 14 in (13;14) carriers (15.9%) compared to the control population (1.3%) (p=0.00016). The study of apoptosis showed an increase of DNA fragmentation in (13;14) carriers (34.9%) compared to the control population (13.8%) (p=0.0036). This increased apoptosis was observed in spermatozoa presenting an increase of unbalanced chromosomal anomalies concerning chromosomes 13 and 14, but with a predominance of balanced spermatozoa compared to the theoretical risk of meiotic segregation. These results suggest that apoptosis could be involved as a regulatory mechanism to eliminate unbalanced chromosomal spermatozoa in men with a (13;14) Robertsonian translocation.     

The systematics of Rigidella (Iridaceae)

Rigidella and the four species contained therein are defined primarily on characters related to, and a direct consequence of, adaptation to hummingbird pollination. The genus is probably derived from either the “multiflora” complex inTigridia subgenusHydrotaenia or is an evolutionary line parallel toTigridia. Illustrations, distribution maps, and a key to the species are included. The rediscovery ofRigidella flammea Lindl. is reported, and a new species,R. inusitata, is described.