KARYOTYPIC VARIATION OF EASTERN NORTH AMERICAN CLAYTONIA CHEMICAL RACES

Considerable karyotypic differentiation has occurred within the group of taxa comprising the eastern North American members of the genus Claytonia. Patterns of karyotypic differentiation are congruent with evolutionary groupings based on flavonoid chemistry, particularly at the diploid level. The 2n = 16 diploid chemotype found in both C. caroliniana and C. virginica possesses a karyotype composed entirely of metacentric chromosomes, while acrocentric chromosomes predominate in the karyotypes of the 2n = 12 and 2n = 14 diploid chemotypes of C. virginica. The 2n = 16 diploid also has a karyotype significantly larger than those of the other diploid cytotypes. Polyploid karyotypes of both species show varying degrees of divergence from their presumed diploid progenitors.     

ELEVATIONAL VARIATION IN FOLIAR FLAVONOIDS OF QUERCUS RUBRA L. (FAGACEAE)

Foliar extracts of high elevation Quercus rubra contain 20 major flavonol-3-0-glycosides based primarily on the aglycones myricetin, quercetin, and kaempferol. These compounds display an altitudinal cline in the Appalachian mountains, with quercetin-rich low elevation chemotypes intergrading into myricetin-rich high elevation forms. These chemotypes appear to be correlated with previously documented variation in foliar morphology. The degree of interpopulation variation also differs with elevation. At higher elevations flavonoids differ only quantitatively between sites, while below 3,000 ft there is site-to-site variability in the hydroxylation level of the aglycones.     

KARYOTYPES OF EASTERN NORTH AMERICAN PHLOX

The karyotypes of 32 Phlox taxa were studied in many plants representative of populations found throughout eastern North America. Most plants were diploid with 2n = 14, but a few tetraploids and plants with B-chromosomes were also observed. Some species were virtually indistinguishable karyotypically, while others differed. Within and between the Subsections Speciosae, Divaricatae, and Subulatae, strikingly similar karyotypes were observed. Other Subsections differed significantly, with the Ovatae showing strong intrasubsectional variation. Increased karyotype asymmetry is correlated with morphological specialization. Where polyploidy was observed, discordant chromosome sets indicated allopolyploidy.     

NATURAL INTERSPECIFIC HYBRIDIZATION IN EASTERN NORTH AMERICAN CLAYTONIA

The eastern North American spring ephemeral Claytonia virginica (Portulacaceae) is well known for its great variation in chromosome number. The origin and significance of this cytological diversity has been a source of some controversy over the last two decades; in particular it has been suggested that one major source of variation could have arisen by allopolyploidy following hybridization between C. virginica and the second eastern North American representative of the genus, C. caroliniana. However, there has been no rigorous demonstration of natural hybridization between these taxa, and attempts to document hybridity have been hampered by a paucity of distinguishing morphological characters. Nuclear ribosomal gene markers are, however, able to distinguish between the two species and provide a means of identifying hybrid plants. We have found a locality in central New York state where the two species cooccur; in this population, hybrid individuals with additive ribosomal gene patterns are found in a zone of overlap between the two parental species. These hybrids are morphologically intermediate and have reduced pollen stainabilities, and appear to be F₁ individuals.     

SPECIATION IN NORTH AMERICAN CHICKADEES: I. PATTERNS OF mtDNA GENETIC DIVERGENCE

We surveyed mitochondrial DNA haplotype divergence within and between populations of six species of North American chickadees (Parus, Subgenus Poecile) with the following results. (1) Genotype diversities (range 0.3 to 0.7) and low nucleotide diversities (range 3 to 27 × 10^?4) within populations were typical of known vertebrates. (2) The two widespread, northern species (atricapillus and hudsonicus) exhibit little mtDNA genetic differentiation throughout their previously glaciated continental distributions, most likely because of recent, postglacial range expansions. (3) Newfoundland populations of atricapillus and maritime province (Newfoundland plus Nova Scotia) populations of hudsonicus have distinct mtDNA haplotypes which differ from continental haplotypes by single restriction site changes. (4) Haplotypes of the southeastern U.S. species P. carolinensis divide into eastern and western sets which have diverged by three percent. This heretofore unrecognized, divided population structure may correspond to the Tombigbee River/ Mobile Bay disjunction known in some other vertebrate taxa. (5) Allopatric populations of the southwestern species sclateri and gambeli exhibit divergences of one and three percent respectively. (6) Prevailing interspecific divergence distances of three to seven percent suggest speciation early in the Pleistocene rather than during late (e.g., Wisconsin) glaciations. (7) Phylogenetic analyses suggest that North American taxa include two clades, hudsonicus-rufescens-sclateri versus carolinensis-atricapillus-gambeli and that carolinensis and atricapillus are not sister species.     

PATTERNS OF ISOZYME VARIATION IN WESTERN NORTH AMERICAN ANTENNARIA (ASTERACEAE: INULEAE) II. DIPLOID AND POLYPLOID SPECIES OF SECTION ALPINAE

Antennaria is a genus of dioecious, perennial, herbaceous Composites that are especially diverse in the cordillera of western North America. Section Alpinae consists of about nine taxa, among them A. aromatica, A. densifolia, A. media, A. pulchella, and A. umbrinella. Although diploids are morphologically distinct, the polyploid derivatives of the diploids obscure the morphological distinctness of the groups. A survey of 19 putative isozyme loci indicates that the diploids have diverged only moderately from one another with respect to biochemical genetics (I = 0.838 to 0.961). Additionally, only moderate amounts of genetic diversity were detected. Isozyme data are supportive of a hypothesis of a rapid mode of speciation in Antennaria, where morphological differentiation has been accompanied by small amounts of allozyme divergence. Polyploids have significantly higher amounts of heterozygosity than diploids and tetrasomic inheritance is inferred. Evidence from morphology and biochemical genetics suggests that the polyploids represent a continuum between interracial autopolyploids and segmental allopolyploids. In light of the relatively low degree of genetic and morphological divergence among many Antennaria species, taxonomic conservatism is advocated.     

FOLIAR VENATION PATTERNS IN THE RUTACEAE

Dede , Roland A. (California Concordia Coll., Oakland.) Foliar venation patterns in the Rutaceae . Amer. Jour. Bot. 49(5): 490–497. Illus. 1962.—A study of herbarium specimens selected from 80 genera of the Rutaceae revealed a series of 7 venation types. The series is based on the spatial relationship of the venation system to the secretory cavities. The hypothesis is advanced that the uncorrelated venation pattern is primitive and that the more complex types represent end products involving a branching pattern of phylogenetic sequences. A key to the illustrated representatives of the 7 types is included.     

PATTERNS OF VARIATION IN LEVELS OF HOMOPLASY

Patterns of variation in levels of homoplasy were explored through statistical analyses of standardized consistency indexes. Data were obtained from 60 recent cladistic analyses of a wide variety of organisms based on several different kinds of characters. Consistency index is highly correlated with the number of taxa included in an analysis, with homoplasy increasing as the number of taxa increases. This observation is compatible with a simple model of character evolution in which 1) the probability of character-state change increases with the total number of branches in a tree and 2) the number of possible states of a character is limited. Consistency index does not show a significant relationship to the number of characters utilized in an analysis or to the taxonomic rank of the terminal taxa. When the relationship between consistency index and number of taxa is taken into account, there is no significant difference between plant and animal data sets in the amount of homoplasy. Likewise, the level of homoplasy in morphological and molecular data sets does not appear to differ significantly, although there are still too few molecular studies to be confident of this result. Future comparisons of consistency indexes, including studies along the lines established here, must take into account the influence of the number of taxa on homoplasy.     

FOLIAR TRICHOMES OF PLEUROTHALLIDINAE (ORCHIDACEAE): FUNCTIONAL SIGNIFICANCE

Foliar surfaces of nine pleurothallid orchids (Cryptophoranthus lepidotus L. O. Wms., Octomeria sp., Pleurothallis pidax Luer, P. poeppigii Lindl., P. revoluta (Ruiz & Pav.) Garay, Restrepiella ophiocephala (Lindl.) Garay & Dunsterv., Restrepia muscifera Rchb. f. ex Lindl., Scaphosepalum rapax Luer, Stelis endresii Rchb. f.), four nonpleurothallid orchids (Cyrtopodium punctatum (L.) Lindl., Encyclia cochleata (L.) Lemee, E. tampensis (Lindl.) Small, Paphiope-dilum hybrid) and five tillandsioid bromeliads (Catopsis nutans (Sw.) Griseb., Tillandsia fas-ciculata Sw., T. streptophylla Scheidw., T. stricta Soland., T. tectorum E. Morr.) were assayed for permeability to Ca, S and P ions. Compared to leaves of the atmospheric bromeliads, those of the orchids proved less permeable to Ca and S. Moisture exchange profiles were determined for two of the bromeliads and three orchids; bromeliad leaves were able to rehydrate completely, orchid leaves only partially. Absorptive capacities of trichomes borne by five taxa were tested by autoradiographic analysis. Unlike bromeliad trichomes, those of the orchids exhibited no capacity to accumulate ³H-leucine. These results and other considerations described here suggest that, unlike tillandsioid bromeliads, neither the pleurothallid nor the nonpleurothallid orchids examined in this survey rely heavily on shoots for mineral and moisture procurement.     

GEOGRAPHIC PATTERNS OF VARIATION IN OFFSHORE SPOTTED DOLPHINS (STENELLA ATTENUATA) OF THE EASTERN TROPICAL PACIFIC OCEAN

The spotted dolphin (Stenella attenuata), a species commonly associated with the tuna fishery, is widely distributed in the eastern tropical Pacific Ocean (ETP). Geographic patterns in morphological characteristics of this dolphin were evaluated for 36 skull measurements of 574 museum specimens allocated to 5° latitude-longitude blocks. A strong subdivision between northern and southern offshore populations was demonstrated using principal components, canonical variates and cluster (UPGMA and function-point) analyses of block means derived from standardized, sex-adjusted measurements. Classification functions derived from a stepwise multiple discriminant analysis correctly identified more than 87 percent of the specimens as being from either a northern or southern locality. These results support recognition of southern spotted dolphins as a separate management stock. Dolphins from a sample representing a far-western locality show similarities with those from the south. Patterns of geographic variation in morphology were evaluated using Mantel tests and matrix correlations; 17 of the 36 characters showed regional patterning and 18 exhibited local patterning. Morphological trends were also assessed with respect to 14 oceanographic measures that characterize environmental variability in the ETP. In general, clinal variation in morphology mirrors a north-south trend found in several environmental variables. Overall, the most striking associations between morphological and environmental variables involved Solar Insolation (Jan.). Sea surface temperatures also exhibited a strong association with morphological variables; the highest correspondence was found between width of the temporal fossa and sea surface temperature in July. In addition, the thickness of the oxygen minimum layer exhibited a pattern of variation that was statistically associated with 58 percent of the skull characters. The geographic patterns found for cranial morphology and environmental measures are strongly clinal and in general congruent, suggesting that oceanographic conditions in part determine trends in morphological characteristics of spotted dolphins.     

POLLEN MORPHOLOGY IN NORTH AMERICAN SPECIES OF TRILLIUM

The pollen morphology and tapetal membranes of 17 species of Trillium from North America have been examined by scanning and transmission electron microscopy. The pollen grains of the genus are consistently spherical and inaperturate. The exine is unstratified and ultra-thin. Five types of exine structure are recognized, namely, echinate-, corrugate-, granulate-, verrucate- and spinulate-type, based on the variations in the components of the exine. The granulate-type is the most common and is divided into six subtypes. The features of tapetal membranes also vary in relation with the variation of the exine structures. This study demonstrates the potential of palynological evidence in assessment of taxonomic relationships within the genus.     

TAXONOMIC STATUS AND GEOGRAPHICAL CRANIAL VARIATION OF COMMON DOLPHINS (DELPHINUS) IN THE EASTERN NORTH ATLANTIC

The common dolphin has a widespread distribution and is relatively abundant in the temperate to subtropical waters of the eastern North Atlantic. However, it is not known whether different species, subspecies, or populations occur in this region. We examined 393 common dolphin skulls obtained from both stranded and bycaught individuals collected between 1901 and 2005. The series included skulls of 152 females and 199 males, from animals ranging in body length from 93 to 230 cm and 105 to 244 cm, respectively. The ranges of total body length, skull size, RL/ZGW ratio and maximum upper alveolar (tooth) count of common dolphins in the eastern North Atlantic overlapped with those of both short- ( D. delphis ) and long-beaked ( D. capensis ) species found off the Californian coast. However, in the absence of additional data, the common dolphin in the eastern North Atlantic is regarded here as a large form of Delphinus delphis . Sexual dimorphism and possible sex-linked characters were identified within the sample. Results of the current study indicate some population differentiation within the eastern North Atlantic, with common dolphins off Portugal showing segregation in morphometric characteristics from common dolphins in other areas.     

GENETIC VARIATION IN BROMUS TECTORUM (POACEAE): POPULATION DIFFERENTIATION IN ITS NORTH AMERICAN RANGE

Allelic variation in seedlings from 60 North American populations of the alien annual grass Bromus tectorum was determined at 25 loci using starch gel electrophoresis. Populations were collected from four regions; east of the Rocky Mountains, Nevada and California, the Intermountain West, and British Columbia. Compared to other diploid seed plants, genetic variation within these populations of B. tectorum is low: 4.60% of loci are polymorphic per population, with an average of 1.05 alleles per locus and a mean expected heterozygosity of 0.012. Although 2,141 individuals were analyzed, no heterozygous individuals were detected, and consequently, mean observed heterozygosity is 0.000. Extensive deviations from Hardy-Weinberg expectations were observed at every polymorphic locus due to heterozygote deficiencies. The mean genetic identity (Nei's I) between population pairs was 0.980 and indicates a high level of overall genetic similarity among populations. The among-population component of the total gene diversity is high (G_ST ⁼ 0.478), indicating substantial genetic differentiation among populations. These results are consistent with previous reports for highly self-pollinating plants of low genetic variation and substantial genetic differentiation among populations. Despite the lack of genetic variation as measured by enzyme electrophoresis, this weedy grass has become exceedingly abundant in a diverse array of arid environments throughout much of western North America, perhaps due to phenotypic plasticity.     

ORIGINS OF GENOTYPIC VARIATION IN NORTH AMERICAN DANDELIONS INFERRED FROM RIBOSOMAL DNA AND CHLOROPLAST DNA RESTRICTION ENZYME ANALYSIS

Restriction enzyme analysis of ribosomal DNA (rDNA) and chloroplast DNA (cpDNA) is used to assess the relative contribution of hybridization and mutation as sources of genotypic variation in weedy asexual dandelions, with focus on the dandelion flora of North America. Of 318 North American dandelions surveyed, 145 rDNA-cpDNA clones are detected. The combined rDNA-cpDNA genotypes show that most of the polymorphic rDNA and cpDNA restriction sites or lengths in these plants are also present in weedy asexual dandelions collected from natural populations in Europe and in asexual and diploid taxa (microspecies) chosen to represent diverse Eurasian members of the genus. However, of 222 combined rDNA-cpDNA genotypes found in 427 asexual plants surveyed, only 9 genotypes are found in both North American and Eurasian dandelions. Two rDNA and three cpDNA characters are unique to individual plants in North America and are consistent with mutational origins of genotypic variation in asexual lineages. But the array of genotypic diversity, characterized by different combinations of the rDNA and cpDNA characters, show that multiple hybridization events are a more important source of genotypic variation than mutation in the asexual polyploids. The rDNA and cpDNA data also indicate polyphyletic origin of several asexual Taraxacum taxa.