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.     

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.     

SIZE STRUCTURE AND DYNAMICS IN A POPULATION OF SARGASSUM MUTICUM (PHAEOPHYCEAE)

Sargassum muticum (Yendo) Fensholt is an introduced brown seaweed with a very distinctive seasonal growth cycle on European shores. The present study links the dynamics of a population of S. muticum with the seasonal growth cycle of the species and the density-dependent processes operating throughout this cycle. Results indicate that both growth cycle and intraspecific competition influenced the structure and population dynamics. Size inequality increased during the slow growth phase (autumn–winter) of the 2-year study. Mechanisms generating inequality of size could be the existence of asymmetric competition and the inherent differences in growth rates between old (regenerated) and new thalli (recruits). Inequality of size distributions decreased progressively during the last months of the growth phase (spring–summer) and could be related to a process of self-thinning. There was a negative biomass–density relationship (as a measure of biomass accumulation-driven mortality) that confirms the importance of self-thinning as a major demographic factor in the S. muticum population.     

GEOGRAPHIC VARIATION IN REPRODUCTIVE BEHAVIOR AND SIZE STRUCTURE OF THE AUSTRALIAN CYCAD MACROZAMIA COMMUNIS (ZAMIACEAE)

Macrozamia communis is a common cycad that occurs in eastern New South Wales, Australia, where it forms an understory in various forest or woodland communities. Leaf number of reproductive individuals as well as of nonreproductive ones, taken as an indication of size class, varies clinally from north to south, with reproductive individuals in southern populations having mean leaf numbers about 12 times greater than those at the northern end of the range of this species. Female cone size and mature ovule weight appear to vary also along this gradient. Using leaf number as a predictor of sexual maturity, from 37 to 100% of the individuals present in a population were judged to be mature. The percent of mature individuals participating in the 1987 coning episode ranged from none in three southern populations to 68% elsewhere. In 9 of 10 samples more males than females participated in the 1987 coning episode, with a collective sex ratio slightly male biased for all populations sampled. In three population samples average cone numbers for male and female plants were equal. In two samples males produced more cones per plant than females. Leaf numbers of males and females in individual populations were equal. Few individuals producing cones in 1987 showed evidence of recent prior cone production, suggesting that successive cone production by both sexes is separated by an interval of several years. The periodicity with which fire is used as a fire suppression method may affect the frequency of cone production in some populations, perhaps preventing it entirely. The apparent absence of recruitment in some populations may result from long-term seed collection by humans for horticultural purposes as well as the very poor seed dispersal that appears to characterize this cycad throughout its range. The weevil Tranes internatus was observed occupying the internal tissues of female cones; this insect may be involved in the pollination of M. communis.     

DEVELOPMENT OF THE POLLEN TUBE OF ZAMIA FURFURACEA (ZAMIACEAE) AND ITS EVOLUTIONARY IMPLICATIONS

Compared with pollen tubes of conifers, gnetophytes, and angiosperms, the pollen tube of cycads is exclusively a vegetative structure, uninvolved with the siphonogamous conduction of sperm to an egg. The cycad pollen tube appears to function primarily to obtain nutrients for the extensive growth and development of the male gametophyte. Previous workers have suggested that, similar to an haustorial fungus, the cycad pollen tube penetrates the reproductive tissues of the sporophyte by enzymatically destroying nucellar cells. These earlier studies did not document the precise structural relationship between the growing male gametophyte and its “host” tissue, the nucellus. Pollen tube growth, and its relation to the nucellus, was examined in Zamia furfuracea with light and transmission electron microscopy. Following germination, the pollen tube of Zamia furfuracea grows intercellularly through the subepidermal layers of the micropylar apex of the nucellus. Electron micrographs clearly show additional localized outgrowths of the pollen tube penetrating the walls of individual nucellar cells. Intracellular haustorial growth ultimately leads to the complete destruction of each penetrated cell, and appears to induce the degeneration of proximal unpenetrated nucellar cells. This pattern of intracellular penetration of the sporophyte by the male gametophyte in Zamia furfuracea is fundamentally different from what has been described in any other major group of seed plants (where intercellular growth of the male gametophyte is the rule), and suggests that the heterotrophic and tissue-specific relationships that male gametophytes of seed plants have with their host sporophytes are substantially more diverse than had previously been known.     

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.     

佛坪自然保护区羚牛的种群数量与结构特征

１９９６年对佛坪自然保护区内的羚牛种群进行了数量调查,结果表明,保护区内羚牛的数量为４３５～５２７只,种群密度达到１．２９～１．５６只／ｋｍ２。羚牛种群的成体雌雄个体之比约为１∶０．５,明显地偏向雌性。在羚牛种群中幼仔占１２．２４％,亚成体占３５．０３％,成年雄性个体占１７．３３％,成年雌性个体占３５．４０％。该羚牛种群在１９９１年至１９９６年间的年均增长率为１８．４８％,明显高于１９８４至１９９１年间羚牛种群的年均增长率。保护区周边地区的生境不断丧失,迫使羚牛向保护区境内迁移,是保护区内羚牛种群数量在短期内迅速升高的原因。     

川西平原大足鼠的种群生态学Ⅰ.种群动态和个体大小

我们利用标志重捕、夹捕解剖和笼养等三种方法于１９８９年１月至１９９５年６月对川西平原的大足鼠的种群密度、个体大小、生长和生物量进行了研究。种群密度最高１１．３只／ｈａ,最低０．７只／ｈａ,平均３．６只／ｈａ,并显示出与农作物相同的季节变动特征。雌雄性比由幼年组和亚成体组约１：１到成体组１：０．７８到老年组１：０．５３。雌雄个体的平均质量分别为１３６ｇ和１１４ｇ,两性的个体质量和后足长有明显差异,体长和耳长差异不明显。饲养的幼体及亚成体个体质量和尾长均持续增长,并可分别用Ｌｏｇｉｓｔｉｃ和对数方程较好地拟合。生物量则显示出比种群密度和个体质量更稳定的特征。     

INSECT-CYCAD SYMBIOSIS AND ITS RELATION TO THE POLLINATION OF ZAMIA FURFURACEA (ZAMIACEAE) BY RHOPALOTRIA MOLLIS (CURCULIONIDAE)

Zamia furfuracea, a cycad of Mexican origin, in cultivation is pollinated by the snout weevil, Rhopalotria mollis, also of Mexican origin. This weevil apparently is host specific, swarming upon male cones of the cycad, where mating, feeding, and oviposition occur. Sporophylls of male cones are rich in starch; those of female cones are poor in starch, and weevils feed upon male cones and are visitors to but not feeders upon or within female cones. Pollen transport to female cones occurs during such visitation. All stages of metamorphosis of R. mollis occur within male cones; larvae feed exclusively on parenchyma of microsporophylls, pupate within stalks of microsporophylls, and emerge as adults from the outer ends of microsporophylls. They do not feed on pollen and do not damage microsporangia or pollen. Toward the end of the breeding season of the weevil (and the cycad), some larvae enter diapause in thick-walled pupal cases within microsporangial stalks of pollen-spent cones. These remain in diapause until the next reproductive season of the cycad.     

北京山区辽东栎林的径级结构、种群起源及生态学意义

 北京山区的辽东栎(Quercus wutaishanica)林径级结构的调查分析结果表明,辽东栎林是稳定性的群落类型,其更新主要通过萌生来完成。能够与辽东栎长期共存的伴生种有色木(Acer mono)、大果榆(Ulmus macrocarpa)、大叶白蜡(Fraxinus rhynchophylla)等。萌生在辽东栎种群中十分普遍,除辽东栎伐桩、枯木和大树根际有较多的萌生茎干外,多年生实生苗也有萌生现象发生。辽东栎林的更新在辽东栎总体种群的幼树阶段(径级2)存在着明显的瓶颈现象,但萌生特性使辽东栎林在更新时或受到干扰后形成较大的萌生种群,使辽东栎的大幼树能够得到迅速的补充,顺利地通过瓶颈进行更新,保持群落的相对稳定。一定数量的辽东栎实生种群的存在对于维持辽东栎的遗传多样性、种群整体活力和群落的稳定,提高辽东栎种群对环境变化的反应能力等方面有着十分重要的生态学意义。     

GENETIC DIVERSITY AND POPULATION STRUCTURE IN CAMELLIA JAPONICA L. (THEACEAE)

Camellia japonica is a widespread and morphologically diverse tree native to parts of Japan and adjacent islands. Starch gel electrophoresis was used to score allelic variation at 20 loci in seeds collected from 60 populations distributed throughout the species range. In comparison with other plant species, the level of genetic diversity within C. japonica populations is very high: 66.2% of loci were polymorphic on average per population, with a mean number of 2.16 alleles per locus; the mean observed and panmictic heterozygosities were 0.230 and 0.265, respectively. Genotypic proportions at most loci in most populations fit Hardy-Weinberg expectations. However, small heterozygote deficiencies were commonly observed (mean population fixation index = 0.129). It is suggested that the most likely cause of the observed deficiencies is population subdivision into genetically divergent subpopulations. The overall level of population differentiation is greater than is typically observed in out-breeders: The mean genetic distance and identity (Nei's D and I) between pairs of populations were 0.073 and 0.930, respectively, and Wright's Fst was 0.144. Differences among populations appeared to be manifested as variation in gene frequencies at many loci rather than variation in allelic composition per se. However, the patterns of variation were not random. Reciprocal clinal variation of gene frequencies was observed for allele pairs at six loci. In addition, principal components analysis revealed that populations tended to genetically cluster into four regions representing the geographic areas Kyushu, Shikoku, western Honshu, and eastern Honshu. There was a significant relationship between genetic and geographic distance (r = 0.61; P < 0.01). Analysis of variance on allozyme frequencies showed that there was approximately four times as much differentiation among populations within regions, as among regions. It is likely that the observed patterns of population relationships result from the balance between genetic drift in small subpopulations and gene flow between them.     

GENETIC DIVERSITY AND POPULATION STRUCTURE IN TRADESCANTIA HIRSUTICAULIS (COMMELINACEAE)

Tradescantia hirsuticaulis, the hairy-stemmed spiderwort, is an insect-pollinated perennial plant species found primarily on rock outcrops in Georgia, South Carolina, and Alabama. Although populations of T. hirsuticaulis are rare and scattered, local populations are frequently large. Levels of genetic variation were assessed for 13 populations representing the species' range in these three states. Despite the disjunct distribution of this habitat specialist and apparent lack of specialized seed and pollen dispersal mechanisms, exceptionally high levels of genetic variation are maintained within the species, with a moderate level of variation (18%) found among populations. Twenty-nine of the 33 loci resolved (88%) were polymorphic within the species; the mean number of loci polymorphic within populations was 54%. The mean number of alleles per polymorphic locus was 3.24 across all populations and averaged 2.37 within populations. Genetic diversity was 0.206 for the species, whereas mean population genetic diversity was 0.157, both much higher than the average for other short-lived herbaceous perennials. Estimated levels of gene flow were moderate (Nm = 0.95), and a significant association between geographic distance and genetic distance between populations was found (r = 0.68; P < 0.0001). Habitat destruction is the major threat to this genetically diverse species. Since gene flow among its highly dispersed populations is limited, diminution or extinction of local populations could jeopardize the long-term evolutionary potential of this species.     

EFFECTS OF DISPLAY SIZE,FLOWERING PHENOLOGY,AND NECTAR AVAILABILITY ON EFFECTIVE VISITATION FREQUENCY IN COMPARETTIA FALCATA (ORCHIDACEAE)

Comparettia falcata is an epiphytic, neotropical orchid that produces nectar as a pollinator reward. In Puerto Rico, C. falcata is allogamous and pollinated by the endemic hummingbird Chlorostilbon maugaeus. Autogamous pollinations are possible, but may result in reduced fruit set. For the 1989 and 1990 flowering seasons, the probability of pollinarium removals and natural pollinations increased with individual inflorescence display size. However, the frequency of effective pollinator visits was independent of flowering phenology in both years. A positive correlation between inflorescence size and reproductive success occurred in 1990 but not in 1989. In 1990 plants produced longer spurs, a higher standing crop of nectar, and a more concentrated nectar than in 1989. There was no relation between nectar availability (= standing crop of nectar) and sugar concentration in either year. Nectar availability and sugar concentration did not vary among the first four flowers of an inflorescence in either season. Nectar availability was not a good predictor of effective visitation. Comparettia falcata has a higher natural fruit set than tropical deceptive orchids, suggesting that pollinator visitation may be enhanced by nectar reward. The small, dilute nectar volumes secreted by C. falcata may benefit the plant by increasing interplant pollinator movement and pollen dispersal.     

EFFECTIVE POPULATION SIZE AND GENETIC DRIFT IN TRISTYLOUS EICHHORNIA PANICULATA (PONTEDERIACEAE)

Populations of the tristylous, annual Eichhornia paniculata are markedly differentiated with respect to frequency of mating types. This variation is associated with evolutionary changes in mating system, from predominant outcrossing to high self-fertilization. To assess the potential influence of genetic drift acting on this variation, we estimated effective population size in 10 populations from northeastern Brazil using genetic and demographic methods. Effective size (N_e) was inferred from temporal changes in allele frequency at two to eight isozyme loci and also calculated using five demographic variables: 1) the number of flowering individuals (N); 2) temporal fluctuations in N; 3) variance in flower number; 4) frequency of mating types; and 5) selfing rate. Average N_e based on isozyme data was 15.8, range 3.4–70.6, and represented a fraction (mean N_e/N = 0.106) of the census number of individuals (mean N = 762.8; range: 30.5–5,040). Temporal variation in N and variance in flower number each reduced N_e to about a half of N whereas mating type frequencies and selfing rate caused only small reductions in N_e relative to N. All estimates of N_e based on demographic variables were considerably larger than those obtained from genetic data. The two kinds of estimates were in general agreement, however, when all demographic variables were combined into a single measure. Monte Carlo simulations indicated that effective size must be fewer than about 40 for drift to overcome the frequency-dependent selection that maintains the polymorphism for mating type. Applying the average N_e/N value to 167 populations censused in northeastern Brazil indicated that 72% had effective sizes below this number. This suggests that genetic drift is likely to play a dominant role in natural populations of E. paniculata.     

POPULATION STRUCTURE,BIOMASS ALLOCATION,AND PHENOTYPIC PLASTICITY IN MURDANNIA KEISAK (COMMELINACEAE)

Field and glasshouse studies were used to explain differences in plant biomass, shoot length, and reproductive effort in four populations of a wetland annual herb, Murdannia keisak. Populations were chosen from continually thermally disturbed, intermittently thermally disturbed, revegetating, and undisturbed portions of a floodplain forest in South Carolina, USA. Plants in the two thermally disturbed areas were shorter, flowered earlier in the autumn, and produced more and smaller seeds than plants in revegetating and undisturbed sites. Reproductive effort was higher in populations from undisturbed and revegetating sites than in the two thermally disturbed sites. Generally, differences observed in the field were not expressed in the glasshouse plants. Glasshouse experiments suggested that most of the observed among-population differences in size and reproductive effort in the field study were a result of a plastic response to water depth and light. The combination of field and glasshouse data showed that this wetland weed adjusts readily to newly disturbed habitats, thus spreading rapidly and maintaining local dominance.     

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.     

SEPARATING POPULATION STRUCTURE FROM POPULATION HISTORY IN ASCOPHYLLUM NODOSUM (FUCALES)

Ascophyllum nodosum is dominant seaweed along many rocky intertidal shores throughout the North Atlantic. Next to the kelps, fucalean taxa such as Ascophyllum are the largest macrophytes and provide important habitat for invertebrates. Understanding the underlying genetic structure of natural populations over a range of spatial scales can reveal how the causes of structure may change with scales. Separating population history from population structure may also be elucidated. The analysis is based on six polymorphic microsatellite loci and> 1000 individuals. Strong genetic structure at small spatial scale was found and is consistent with demographic models based on long-lived individuals, low recruitment and many sib matings. At large spatial scales only weak population differentiation was found. This is consistent with recent recolonization of the North Atlantic following the last glacial maximum.