GENETIC STRUCTURE OF THE AUSTRALIAN CYCAD,MACROZAMIA COMMUNIS (ZAMIACEAE)

Cycads have been the subject of a wide variety of botanical studies. However, nothing is known of the genetic structure of their populations. Seeds were collected from five populations of the common Australian cycad Macrozamia communis along a north to south transect through its 500-km-long range in eastern New South Wales. Leaf samples from seedlings grown from those seeds were the subject of isozyme analysis. Nine of the 18 isozyme loci studied were polymorphic. Levels of polymorphism varied among loci and among populations. The three southern-most populations were genetically similar to one another but well differentiated from the other two. The levels of species-wide genetic diversity observed for M. communis were relatively low compared with those of other gymnosperms and angiosperms. On the other hand, the levels of spatial differentiation were relatively high, especially compared with other gymnosperms.     

INSECT POLLINATION IN THE CYCAD ZAMIA PUMILA (ZAMIACEAE)

A wind and insect exclusion pollination experiment was conducted in a wild population of the cycad Zamia pumila L. in Florida. Cones from which insects but not wind were excluded produced no viable seeds, while cones from which wind but not insects were excluded produced abundant viable seeds. Two beetle species have been identified which may be effecting pollination. Adults and larvae of both beetles are found abundantly on the male cones, and adults of both species and larvae of one species are found occasionally on female cones. Z. pumila produces sugar and amino acid-rich micropyle droplets which may serve as pollinator rewards. Mimicry of resource-rich male cones by female cones and the use of cones as refuges between mating and feeding bouts may account for beetle movement from male to female cones.     

PATTERNS OF ALLOZYME DIVERSITY IN THE WEST INDIES CYCAD ZAMIA PUMILA (ZAMIACEAE)

The West Indies cycad, Zamia pumila, is restricted to the Greater Antilles, northern Bahama Islands, Florida, and the southeastern coast of Georgia. An electrophoretic study based on nine enzymes compared 21 accessions from throughout the range of the species. Lower levels of intrapopulation variation than those reported for ferns, other gymnosperms, and angiosperms were discovered for the two more extensively sampled populations. However, this variation was similar to that found in other island taxa and in the endemic Australian cycad, Macrozamia communis. In contrast, allozyme divergence among accessions of Z. pumila appeared relatively high, mostly as a result of rare alleles restricted in geographic distribution. The age and biogeography of Z. pumila may gave contributed to population differentiation. Also, mean number of alleles per locus was low for the species (1.75). Finally, the time-since-divergence value (10.8 million years ago) between Z. pumila and its closest extant relative, Z. splendens, was much smaller than the age of Z. pumila suggested by the fossil record and historical geology of the Caribbean (30–60 million years ago). Together, these data indicate that biochemical evolution within this species, and perhaps in all cycads, is slow when compared to that of noncycad seed plants.     

GROUP SIZE FACTORS AND GEOGRAPHIC VARIATION OF MORPHOMETRIC CORRELATION

Within-locality correlations among eight morphological traits of the fundatrix and among 15 morphological traits of the alate fundatrigeniae of the aphid Pemphigus populicaulis Fitch were examined for among-locality variation. A jackknife procedure revealed highly significant differences among correlation matrices representing 34 local samples from eastern North America. Most bivariate correlations also differed significantly among samples. Although very low correlations cannot differ, for moderate or strong correlations the average magnitude of correlation is not a good predictor of the degree of interlocality difference. Nearly half of the variation among localities is in the level of “overall correlation,” which is positively correlated with intralocality size variation. Despite significant differences among localities, spatial autocorrelation tests failed to reveal any significant geographic pattern in correlations. Monte Carlo experiments suggest that if geographic patterns in the correlations were of the same magnitude as those for the means of these traits, some of these patterns would have been detected. Factor analyses of the pooled within-locality correlation matrices, after oblique rotation to simple structure, reveal appendage and body-size factors for both alates and stem mothers. In general, correlations between traits with high loadings on the same factor differ among localities, whereas substantial correlations that do not vary geographically are between traits that are not well resolved by the factor analyses. It is hypothesized that the apparently random geographic differences in correlation are caused by chance differences in the mode of response to short-term selection. Geographic patterns in trait means are established by selection, but, because local population sizes are finite, similar response to selection occurs by diverse physiological or developmental mechanisms in different populations, just as replicate lines in artificial selection experiments may achieve the same response by diverse mechanisms. These diverse mechanisms of response will have diverse effects on genetic variance and covariance, causing correlation patterns to vary geographically. Several forces will oppose continued divergence of these patterns.     

VOCAL DIALECTS AND THE STRUCTURE OF GEOGRAPHIC VARIATION IN MORPHOLOGICAL AND ALLOZYMIC CHARACTERS IN THE RUFOUS-COLLARED SPARROW,ZONOTRICHIA CAPENSIS

The geographical patterns of variation shown at 20 allozyme and non-enzymatic protein-coding loci, in 8 external, and in 12 skeletal morphological characters in the rufous-collared sparrow, Zonotrichia capensis, were analyzed in order to test the local (genetic) adaptation hypothesis regarding the origin and maintenance of vocal dialects in birds. Approximately 20 males were collected from each of four sites within each of six different dialect zones. There was significant variability in both external and skeletal morphology among all 24 sites and among dialect groups. Average Wright's corrected fixation coefficient (F_ST) was 0.118, indicating significant genetic differentiation among all sites, regardless of dialect. Hierarchical F statistics indicated that only 50% of among site variability was due to a dialect effect. Puna dialect sites were highly differentiated from all other sites with respect to both morphology (external and skeletal measures) and allozyme frequencies. Heterogeneity at the PGM-1 locus among puna scrub sites was the major cause of the high average F_ST across all sites, and within the puna scrub dialect. Average genetic differentiation among non-puna sites (F_ST = 0.018) was similar to differentiation among sites within each of the five non-puna dialect groups (mean F_ST = 0.0132 ± 0.0069). Hierarchical F statistics indicated that none of the among-site differentiation in this subset of samples was due to a dialect effect. These observations are not consistent with the local adaptation hypothesis. All significant genetic heterogeneity occurred among sites in mountainous habitats, and we suggest that topography and patchiness of habitat may have been major factors involved in population differentiation, rather than vocal dialects.     

BERGMANN'S RULE,SEASONALITY, AND GEOGRAPHIC VARIATION IN BODY SIZE OF HOUSE SPARROWS

Body size of House Sparrows is positively correlated with seasonality (annual temperature range) in both North America and Europe. Previously used climatic variables gave inconsistent or contrasting results for the two continental samples. The physiological advantage of large size in seasonal environments appears to be increased fasting ability, but this may be offset somewhat, particularly for females, by other factors, e.g., competitive interactions that influence rates of energy intake or ability to initiate breeding in the spring. It is proposed that minimization of maintenance energy costs on a per individual basis, and thus small size, is advantageous in equable (and predictable) environments.     

COMPARATIVE METHODS AT THE SPECIES LEVEL: GEOGRAPHIC VARIATION IN MORPHOLOGY AND GROUP SIZE IN GREY-CROWNED BABBLERS (POMATOSTOMUS TEMPORALIS)

We show that a new comparative method that sheds light on evolutionary trends among species may also illuminate trends within species. This finding comes from a phylogenetic autocorrelation analysis of morphological traits among individuals sampled from ten populations of a cooperatively breeding songbird, the Grey-crowned Babbler (Pomatostomus temporalis). Highly variable mitochondrial DNA (mtDNA) sequences from both the eastern (Pomatostomus temporalis temporalis) and western (Pomatostomus temporalis rubeculus) lineages were used to define genetic distances among 120 individuals and to estimate correlations among individuals in wing length, tarsus length, and body weight via an intraspecific weighting matrix. The autoregressive model effectively removed intraspecific correlations for all three morphological variables, and the proportion of the total phenotypic variance due to genealogical relationships varied from 0.68 (weight) to 0.23 (tarsus). The analysis revealed correlations among the specific components of traits, in which none were previously detected (type-I error) and diminished correlations that appeared strong when phylogeny was ignored. Group size was the only trait for which the autoregressive model failed to remove intraspecific correlations, a result likely due to the plasticity, convergence, and clinal variation in this trait in both the eastern and western lineages. The autocorrelation analysis weakened significant negative correlations between group size and total values for wing length and body weight, but the interpretation of this result depends on the adaptive significance ascribed to the “phylogenetic component” of trait values removed by the analysis. Comparative methods employing distance matrices are one useful way of summarizing the pattern of nonhierarchical relationships among conspecific individuals (“tokogenetic” relationships, sensu Hennig).     

SEX-RATIO AND REPRODUCTIVE VARIATION IN THE MISTLETOE PHORADENDRON JUNIPERINUM (VISCACEAE)

Despite the fact that many parasitic and hemiparasitic plant species such as mistletoes are dioecious and occur in both the new and the old world, few data exist on variation in the sex ratio and allocation to reproduction in these taxa. We investigated 1) the sex-ratio of the xylem-tapping mistletoe Phoradendron juniperinum in relation to its age and position within the canopy of its host tree Juniperus osteosperma, and 2) reproductive effort in relation to the gender and age of mistletoe plants. Our surveys showed that P. juniperinum has a male-biased sex ratio. Despite this predominance of male individuals, females lived longer and had a greater reproductive effort than did males. A statistical analysis of the age distribution data indicated that the peak in the frequency of reproductively mature individuals was later in females than in males. These gender-specific distributions may have resulted 1) from sequential hermaphroditism (age-specific sex switching), or 2) because the average age of peak reproduction is later in female individuals. Because sex is genetically determined in a closely related genus of mistletoe and because we have no data to indicate sex switching in this species, we feel that our data support the interpretation that female individuals, on average, show a peak in reproductive vigor at an older age relative to males. While delayed reproduction in females may be favored because reproductive effort and success appear to be age-dependent in females of this species, both sexes can become reproductively mature relatively early in life. Further, because 1) allocation to reproduction as a function of age increases more rapidly for females of this species relative to males, and 2) because there may be a higher resource cost associated with reproduction in females, we hypothesized that female individuals would be more abundant in the best quality locations within the host tree so as to maximize the opportunity to meet those costs. In spite of the association between gender and some host characteristics, there was no indication that female plants were located in sites most favorable to either their carbon or water balance. We discuss reasons why this may be the case.     

POPULATION STRUCTURE IN ZAMIA DEBILIS (ZAMIACEAE) I. SIZE CLASSES,LEAF PHENOLOGY,AND LEAF TURNOVER

Population structure, leaf phenology and leaf turnover were followed over a 29-month period in Zamia debilis L.f. ex Aiton (Zamiaceae), an understory species in the Cambalache Forest in northern Puerto Rico. It was not possible to determine plant age or to measure the subterranean stems; size classes based on leaf number and leaf × leaflet number indices were used to determine population structure. Despite seasonal and year to year fluctuations in leaf number at the individual and population level, population profiles remained relatively constant. At any one time, over 50% of the population was composed of unbranched individuals with one or two leaves. Only 7% of the plants were branched. Plants with seven or more leaves comprised at a maximum 8% of the population, but accounted for 28% of the total foliage. Size classes based on leaf number and on a leaf × leaflet index gave approximately reverse J-shaped curves typical of trees with shade tolerant seedlings and saplings. New leaves emerged throughout the year, with a peak at the beginning of the rainy season in May or June and lowest production during the dry months of February through April. Average leaf life expectancy was approximately 2.3 years. Leaf death occurred over an extended period of time by the loss of individual leaflets. Patterns in leaf production and loss differed between few- and many-leaved plants. On the average, as the number of mature leaves on a plant increased, time between emergence of new leaves decreased. In many-leaved plants more than one event of new leaf emergence per year was common. Individuals with one to three mature leaves and individuals with four or more mature leaves differed in their response to water stress: few-leaved plants generally reduced the rate of new leaf production and retained old leaves longer. Plants with more than three leaves continued to produce new leaves, but the rate of leaf mortality increased so that most had a net leaf loss. There was no evidence that leaf emergence or retention were affected by cone production or seed maturation.     

LEAF DEMOGRAPHY OF A NEOTROPICAL RAIN FOREST CYCAD,ZAMIA SKINNERI (ZAMIACEAE)

Tropical rain forest understories are extremely light-limited environments. Little is known about the demography of leaves in these habitats. To determine leaf longevity and causes of mortality for a species in the understory of a neotropical rain forest, we studied 634 leaves on 173 individuals of the cycad shrub Zamia skinneri for 10 yr at the La Selva Biological Station, Costa Rica. Five different seasonal cohorts of leaves were followed. No significant differences among cohorts were found in median leaf longevity, but the cohorts did differ in the proportion of leaves dying in annual intervals from <1 to ≥6 yr. Median minimum time alive for all leaves was 4.6 yr. Epiphyll coverage increased from 0% in the first year of life to 75%-100% by year 4. Leaf mortality was well described by two separate exponential functions, approximately 10% mortality/yr until year 4, then abruptly increasing to ca. 51%/yr. Of 625 leaf deaths, 77.0% were due to unexplained causes, 16.5% to physical damage by falling litter, 4.8% to attacks by larvae of the lycaenid butterfly Eumaeus minyas, and 1.8% to other causes. Leaf longevity was positively correlated with leaf size. Individuals of Z. skinneri frequently survive more than a century in very dark understory conditions. We suggest that leaf longevity in these circumstances is determined by the inevitable deterioration in leaf carbon balance due to leaf aging and increasing epiphyll coverage, and by loss due to physical damage and attacks by the specialist herbivore.     

GEOGRAPHIC VARIATION OF SEXUAL DIMORPHISM IN SIZE OF SAVANNAH SPARROWS (PASSERCULUS SANDWICHENSIS): A TEST OF HYPOTHESES

The Savannah Sparrow (Passerculus sandwichensis) is a widespread and common North American bird that shows both geographic variation and sexual dimorphism in size. I used information from 24 measurements on 1,791 individuals from 51 populations to test two hypotheses (sexual-selection and niche-partitioning) about the evolution of sexual dimorphism. Throughout their range male Savannah Sparrows are larger, on average, than females. This doubtless reflects Darwinian sexual selection, for territorial fights usually involve males, many of whom fail to obtain mates. In some parts of their range, Savannah Sparrows are commonly polygynous, whereas in others they are characteristically monogamous. Among species of American sparrows (subfamily Emberizinae) sexual size dimorphism is generally greater in polygynous species than in monogamous ones. However, I did not find a similar trend among populations of Savannah Sparrows. The amount of dimorphism in all populations of Savannah Sparrows is equivalent in magnitude to that of other species of sparrows that are commonly or regularly polygynous, and it is greater than that of other sparrow species that are characteristically monogamous. The amount of sexual dimorphism, either in overall size or in bill size, does not correlate with species diversity and does not differ between island and mainland populations. These results do not support the niche-variation hypothesis. Size dimorphism is relatively great in populations of Savannah Sparrows that are resident in southwestern salt marshes, and these birds are the only sparrow-like birds that generally breed in these marshes. Dimorphism is, in general, relatively great in marsh-dwelling species in the family Emberizidae. These species are commonly, but not always, polygynous; the mating systems of the salt-marsh Savannah Sparrows are not known. There are no significant differences in the extent of dimorphism among populations of salt-marsh sparrows, and there are few among the non-salt-marsh ones, probably reflecting conservatism in the evolution of size dimorphism.     

GEOGRAPHIC VARIATION AND HETEROCHRONY IN TWO SPECIES OF COWRIES (GENUS CYPRAEA)

Geographic variation in the marine, Indo-Pacific cowry, Cypraea caputserpentis, involves clinal variations that parallel the ontogenetic development of adult shell characteristics. Cypraea caputdraconis, a closely related species endemic to Easter Island and Sala y Gómez, is morphologically similar to juvenile C. caputserpentis. Using multivariate measures of size and shape, I examine these patterns as a possible outcome of heterochrony, or changes in the timing of developmental events in ontogeny. Whorl-expansion rates of juvenile shells are significantly higher in C. caputdraconis when compared to C. caputserpentis and are negatively correlated with surface seawater temperatures among populations of C caputserpentis. High expansion rates, often associated with slow growth, result in a delay in the onset of lateral callus development and subsequent paedomorphosis. Ontogenetic trajectories calculated from growth series of adult and preadult shells indicate that paedomorphosis results from the combined effects of neoteny and post-displacement. Paedomorphosis among cowries may result from the advantages of larger body size relative to shell size under reduced predation intensities and associated increases in fecundity.     

GEOGRAPHIC VARIATION IN THE LIFE HISTORY OF MASTOCARPUS PAPILLATUS (RHODOPHYTA)1

Variation in the geographic distribution of the life histories of Mastocarpus papillatus was investigated. Carpospores were isolated from 377 female gametophytes collected from eight localities on the Pacific coast of Baja California, Mexico, and California, U.S.A., and grown in laboratory culture. All carpospores from a single female gave rise either to basal discs with gametophyte-like uprights or crustose plants formerly referred to the genus Petrocelis. Early stages in the development of each type of germling were observed, and environmental factors affecting development were suggested. Based on carpospore germlings, females from each location were scored as having either the 1) sexual life history (crustose germlings) or 2) direct-development life history (discoid germlings with uprights). All females from the two southernmost locations in Baja California exhibited the sexual life history. In the three locations from the central-southern California coast, 70-95% of the females exhibited the sexual life history and the remainder exhibited the direct-development life history. In two of the three populations from the central-northern California coast, 70-90% of the females exhibited the direct-development life history and the remainder the sexual life history. In the third location from the central-northern California coast, the northernmost location sampled in the current study, 60% of the females exhibited the sexual life history and 40% the direct-development life history. The relative ecological advantages and disadvantages of the life histories are unknown as are the environmental factors that produced the ratios of sexual to direct-development females observed at each location.     

黄鼬(Mustela sibirica)体型大小的性二型及地理变异

本文报道黄鼬体型大小的性二型及其与地理变异的关系。个体大小的性二型现象很可能与繁殖行为有关,许多在繁殖季节有争偶行为的一雄多雌兽类,雄性都有较大的个体。     

中国西部麦红吸浆虫(双翅目:瘿蚊科)酯酶同工酶与种群遗传结构研究(英文)

本文用PAGE方法测定了我国西部麦红吸浆虫Sitodiplosismosellana (G啨ｈｉｎ) 3个地理种群酯酶同工酶 ,分析了其遗传规律及种群的遗传结构。采样地点有春麦区的皋兰县 (36.3°N ,1 0 3.9°E)和武威市(37.9°N ,1 0 2 .6°E)及冬麦区的长安县 (34.1°N ,1 0 8.9°E)。结果分析认为该酶受两个基因位点控制 ,命名为Est 1和Est 2。Est 1可能为质体基因 ,其酶谱为所有带中移动最快的一条 ,每个个体都有。Est 2为重复基因位点 ,共观察到 8个等位基因 ,在所有种群中表现为 8条带 ,依其迁移速度命名为a、b、c、d、e、f、g、h。Est 2在每个个体中表现 1 - 4条带 ,且为一多态位点 ,因此该等位酶可作为一个稳定的遗传标记。在所有研究种群的个体中 ,共观察到 1 9种酶谱类型。在冬麦区长安种群中个体酶谱类型多达1 7个 ;春麦区个体酶谱类型少 ,在皋兰种群内有 5个、武威种群内有 4个。在所有的种群中 ,以d、e、g 3个等位基因的频率最高。冬麦区长安种群内有 8个等位基因、而春麦区种群仅有 3个。通过对Est 2单个基因位点的遗传一致度和聚类分析表明 ,春麦区两种群亲缘关系较近 ,且都与冬麦区种群较远。上述所有分析显示麦红吸浆虫有一定程度的种下分化 ,遗传漂移可能是引起该分化的主要原因。不同种群间有可能发生过     

ESTERASE VARIATION AND GENETIC STRUCTURE IN THREE GEOGRAPHIC POPULATIONS OF SITODIPLOSIS MOSELLANA (GEHIN) (DIPTERA: CECIDOMYIIDAE) IN WESTERN CHINA*

Abstract This paper investigates the esterase variation and genetic structure in three geographic populations of Sitodiplosis mosellana (Géhin) in western China by PAGE. The localities surveyed are Gaolan (36.3°N, 103.9°E) and Wuwei (37.9°N, 102.6°E) in spring wheat region and Chang'an (34. 1°N, 108.9° E) in winter wheat region. The results suggest that the esterase is coded by two loci: Est‐1 and Est‐2. Est‐1 is coded by a plastogene producing only one band that is the fastest on the gel among all bands. The Est‐2 is duplicated loci with 8 alleles, namely, a, b, c, d, e, f, g, h, which produce altogether 8 bands in all the populations and 1–4 bands in individual samples. There are 19 zymogram types observed in the three geographic populations. Seventeen zymogram types emerge in Chang'an population, but 5 and 4 zymogram types are found respectively in Gaolan and Wuwei populations. II₂ zymogram type is the commonest in all the populations. The alleles that had the highest frequencies in all the populations are d, e, g. All 8 alleles at the Est2 were observed in Chang'an population, but only total 3 alleles‐d, e, g at the Est‐2 appeared in Gaolan and Wuwei populations. The analysis of genetic identity and cluster (UPGMA) on the alloenzyme indicates that the relationship between the two populations of spring wheat region seems to be closer, as compared with the relationship between spring wheat population and winter wheat population. It is evident that there exists some infraspecific variation caused mainly by genetic drift in S. mosellana and the gene flow among the populations possibly took place to some extent.