CLUTCH SIZES AND BREEDING STRATEGIES AMONG BERMUDAN AND NORTH AMERICAN PASSERINES

Evidence for reduction of clutch size in insular passerines is presented. Average clutch sizes for the Catbird, Cardinal and White-eyed Vireo were reduced 14.0, 18.2 and 30.7%, respectively as compared with coastal North American populations from similar latitudes. Differences are highly significant. Clutch size in the Eastern Bluebird was not reduced. Trends in latitudinal and longitudinal variations of clutch size in North America are examined. With the exception of a slight latitudinal gradient in coastal populations, no consistent trends existed. There is no evidence for reduced variability in clutch size or increased egg size in the Bermudan populations, but both these parameters may show some response to combined effects of latitude and longitude in continental populations.
Reduction of clutch size in insular nidicolous birds is consistent with both the Lack-Ashmole resource limitation hypothesis and Cody's energy allocation hypothesis. In the absence of experimental evidence for life history optimization in response to K-selection, the modified Lack hypothesis seems sufficient.     

MOLECULAR EVOLUTIONARY DIVERGENCE AMONG NORTH AMERICAN CAVE CRICKETS. II. DNA-DNA HYBRIDIZATION

Single-copy DNA divergence among 23 populations of cave crickets belonging to two genera (Euhadenoecus and Hadenoecus) has been determined by DNA-DNA hybridization employing the TEACL method. These same populations have been studied for allozyme variation (Caccone and Sbordoni, 1987). In addition, a European relative (Dolichopoda laetitiae) has been included as an outgroup for rooting the phylogeny. One of the most remarkable findings is the large degree of DNA divergence among these species and populations. A ΔT_m of up to 5°C has been found between populations of the same species; even further divergence is indicated by a lowered normalized percentage of reassociation. A phylogeny was constructed and tested for synchrony of rates, i.e., a molecular clock. Statistically, we could not reject the clock hypothesis. Attempts to calibrate the clock led to the conclusion that these insects are among the fastest evolving (with respect to single-copy DNA) groups yet studied—at least as fast as Drosophila and sea urchins—where a ΔT_m of 1°C indicates 0.5 to 1.5 MY since the last common ancestor. In general, the phylogeny derived from the DNA data agrees with that derived from isozymes. Nei's D and ΔT_m are correlated; in this group a D of 0.1 corresponds to a ΔT_m of about 1.5°C. This indicates that, relative to total single-copy DNA, the protein-coding regions of the genome are slowly evolving.     

MOLECULAR EVOLUTIONARY DIVERGENCE AMONG NORTH AMERICAN CAVE CRICKETS. I. ALLOZYME VARIATION

Forty-nine populations of nine species of North American cave crickets (genera Euhadenoecus and Hadenoecus) have been studied for genetic variation at 41 loci by electrophoresis. Wright's F_ST, Slatkin's Nm^* gene-flow estimator, and Nei's genetic distances (D) have been used to compare closely related species that have different ecological requirements (cave vs. forest species), distribution patterns, and/or different degrees of geographic isolation among populations. Cave and epigean (noncave) species differ greatly in their levels of genetic differentiation. Cave species have lower rates of gene exchange (low Nm, high D, and F_ST) than epigean species. Within cave species the degree of genetic differentiation among populations is correlated with the limestone structure of the area where the species occur. Species or groups of populations inhabiting areas where the limestone is continuous and highly fissured (e.g., H. subterraneus populations in the Mammoth Cave region) are genetically less differentiated than are populations occurring in regions where the limestone distribution is more fragmented, such as the Appalachian Ridge where E. fragilis occurs; this effect is more extreme in Central Tennessee where genetically differentiated E. insolitus populations occur only a few kilometers apart. This suggests that epigean dispersal through forest habitat in cave-dwelling species is negligable. For forest species, the data indicate relatively recent radiation with ongoing gene exchange among populations. For cave species, the distribution of protein polymorphisms is apparently more a function of historical patterns of gene exchange rather than current gene exchange. Phylogenetic relationships were studied using cluster analyses (UPGMA and Wagner algorithms) of Nei's and Edwards' genetic distances and multivariate analysis (correspondence analysis) of the raw allele frequencies. Different algorithms result in branching patterns that are similar but not entirely concordant with one another or with the phylogeny based on morphology.     

BIOCHEMICAL HETEROPHYLLY AND FLAVONOID EVOLUTION IN NORTH AMERICAN POTAMOGETON (POTAMOGETONACEAE)

Morphologically heterophyllous species of Potamogeton also commonly display biochemical heterophylly with respect to flavonoid compounds. Generally, floating leaves contain an assortment of flavonoids, whereas submersed leaves often exhibit reduced flavonoid profiles. In strictly submersed (homophyllous) species, two patterns occur. Linear-leaved species have few flavonoids and their biochemical profiles resemble those of submersed leaves of heterophyllous species. Broad-leaved homophyllous species possess flavonoid profiles more similar to those of the floating leaves of heterophyllous species. Numerical analysis of these chemical data is consistent with phylogenetic relationships within the genus derived independently on the basis of morphological and chromosomal data. Glycoflavones, which are probably maintained in floating leaves because of their UV filtering ability, exhibit the most pronounced biochemical heterophylly in Potamogeton. The lack of glycoflavones in submersed leaves of heterophyllous species and in linear-leaved homophyllous species is attributable to the ability of naturally colored water to significantly absorb harmful UV radiation. These observations provide strong support for earlier hypotheses suggesting the importance of flavonoid evolution in the conquest of exposed terrestrial habitats by plants.     

CHROMOSOME NUMBERS FOR NORTH AMERICAN EUPHORBIACEAE

Chromosome number determinations are reported for 101 collections distributed among 54 species of the North American Euphorbiaceae. First counts are presented for 44 species. When appropriate, systematic discussions regarding chromosomal information are included. Camera lucida drawings are published for previously undocumented chromosome reports.     

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.     

ECOLOGICAL DIVERSITY IN NORTH AMERICAN PINES

Ecological groups were identified from 34 North American species of pine using multivariate analysis of 18 ecological traits. Five adaptive modes are described: 1) fire-resistant species that are large, thick-barked, and have large cones and long needles; 2) tall, fast-growing mesophytic species with moderately high shade tolerance; 3) stress-tolerant species with animal-dispersed seeds, occurring mainly on cold or dry sites where fire is infrequent; 4) fire-resilient species that are precocious reproducers with small seeds, often in serotinous cones; and 5) species of southern mesic sites with fast growth, strong, heavy wood and short persistence of needles. Intermediates between these modes exist. Convergent evolution has occasionally occurred, as shown by high ecological similarity of species in different taxonomic sections within Pinus. However, the analogies between species are imperfect, suggesting the importance of constraint by shared ancestry and divergence produced by a diversity of environments.     

PHYLOGEOGRAPHIC PATTERNS IN MITOCHONDRIAL DNA OF THE DESERT TORTOISE (XEROBATES AGASSIZI), AND EVOLUTIONARY RELATIONSHIPS AMONG THE NORTH AMERICAN GOPHER TORTOISES

Restriction-fragment polymorphisms in mitochondrial DNA (mtDNA) were used to evaluate population-genetic structure in the desert tortoise Xerobates agassizi and to clarify evolutionary affinities among species of the gopher tortoise complex. Fourteen informative endonucleases were employed to assay mtDNAs from 56 X. agassizi representing 22 locations throughout the species' range. The mtDNA genotypes observed were readily partitioned into three major phylogenetic assemblages, each with striking geographic orientation. Overall, the X. agassizi mtDNA genotypes typify a common phylogeographic pattern, in which broad genetic uniformity of populations is interrupted by geographic features that presumably have functioned as dispersal barriers. The geologic history of the Colorado River area, which includes extensive marine incursions, may account for the marked mtDNA divergence between eastern and western X. agassizi assemblages. In mtDNA comparisons among the four species of the gopher tortoise complex, both UPGMA and Wagner parsimony analysis strongly support the recognition of two distinct species groups previously suggested by traditional systematic approaches. Furthermore, the mtDNA data identify the eastern X. agassizi assemblage as the probable inceptive lineage of X. berlandieri. Results from both intra- and interspecific comparisons illustrate how clues to historical events may be present in the geographic structure of mtDNA phylogenies.     

KARYOTYPIC DATA ON NORTH AND CENTRAL AMERICAN ZAMIACEAE (CYCADALES) AND THEIR PHYLOGENETIC IMPLICATIONS

Chromosome numbers and karyotypes of species from four American Zamiaceae (Cycadales) are reported. Zamia shows interspecific and intraspecific chromosome variation, whereas Microcycas, Ceratozamia, and Dioon have constant karyotypes within each genus. In Zamia, all karyotypes have the same number of submetacentric and acrocentric chromosomes, but they differ in the number of metacentric and telocentric chromosomes. Centric fission of metacentric chromosomes is proposed to explain the karyotypic variation in this genus. Zamia shows karyological relationships with Microcycas and Ceratozamia, whereas Dioon appears very distinct from the other American cycad genera. Affinity among Zamia, Ceratozamia, and Microcycas karyotypes and distinctiveness of Dioon karyotypes are supported by comparative analysis of phenotypic characters in the four genera.