Variable floral phenology: temporal resource heterogeneity and its implication for flower visitors

I investigated within- and between-year patterns of flowering phenology of entomophilous plants in an isolated old field in northern Michigan, USA. The number of open flowers was censused at 5-d intervals from June-September of 1984–1986, and the data analyzed by detrended correspondence analysis. DCA axis 1 reflected within-year phenology, was highly correlated with date of censusing, and implied substantial constancy in flowering sequences. However, DCA axis 2 revealed significant between-year variation in abundances of flowers; years differed significantly in their scores on axis 2, and these differences were driven by variable plant species abundances during late summer. Within-year patterns showed that the floral community consisted of three more-or-less distinct temporal "guilds" of plants, and that the rate of turnover in community composition throughout the season was not uniform. The appearance of these guilds was not synchronous across years.
Insects foraging on these species throughout the floral season can expect constancy in the relative timing of blooming, but are exposed to considerable temporal heterogeneity in flower resource abundance both between- and within-years. Opportunities for coevolutionary specialization are therefore likely to be constrained.     

中卫山羊核心产地植物群落的数量分类与排序

应用数量分类(TWINSPAN)和DCA排序方法,对我国特有裘皮用山羊(中卫山羊)核心产地——宁夏香山地区植物群落进行多元分析,根据TWINSPAN分类结果,并结合生态特征将所调查的28个样地植被分为6组,代表了该山地在海拔、地形、土壤、山坡坡度等环境作用下植被空间的分布格局。TWINSPAN分类结果与DCA排序结果较一致,DCA排序第一轴结果主要体现出海拔和山坡坡度对植被分布的作用,第二轴结果主要体现了地形(坡向)和土壤基质(沙地、石质山坡、土质山坡)对植被分布的作用。影响该干旱山地植被分布的主要环境因子有海拔和山坡的坡度,另外长期过度放牧对植被的空间分布影响较大,导致了该山地系统植被空间结构紊乱,并干扰了群落数量分析结果的准确性。     

Chydorid assemblages in the sedimentary sequence of Lake La Cruz (Spain) subject to water level changes

Changes among chydorid assemblages in Lake La Cruzsediments, from a core of 178 cm length, have beenstudied using community structure indices andmultivariate methods (cluster analysis, principalcomponents analysis).The results revealed the existence of two main sourcesof variation in these assemblages. One (axis I ofPCA1) is associated with the trophic level of thelake, which is hypothetised to be greater in thebottom and upper part of the core. Both zones arecharacterised by an increase in the relative frequencyof Chydorus sphaericus, accompanied by a markedreduction in the relative frequency of the mostabundant species throughout the history of the lake,Acroperus neglectus.The other source of variation (axis II of PCA1) may beinterpreted as the alternation of periods of dryer and wetter weather. Ouranalyses show the separation of two assemblages whichalternatively prevail in the sedimentary sequence, oneconstituted by Pleuroxus laevis, Alona guttata,Graptoleberis testudinaria and, sometimes, Alonarectangula, and another dominated by Chydorussphaericus, Alona quadrangularis, Alona affinis andsometimes Leydigia species. According to theecological preferences of these species, the dominanceof the first chydorid assemblage can be related toepisodes of higher temperature and with very low andfluctuating water level, or with a high water levelbut a very reduced extent of shallow water in thelake. Prevalence of the second group corresponds toepisodes with a colder environment and permanentwaters, the level of which could be variable but insuch a way that a benthic sublittoral zone maydevelop. The structure of the community is stronglyaffected by changes in lake level. High indices offluctuation in community structure and small mean bodysize of the chydorid assemblage marked the period withmore extreme dry conditions, characterised by Alona rectangula maxima. The peaks of C. sphaericuscoincided with peaks of the index offluctuation D₀ and prevalence of smaller speciesof the chydorid assemblage.     

Craniodental variation in the African macaque, with reference to various Asian species

Based on twenty-seven craniodental measurements and ratios derived from them, the relationship between the African macaque (M. sylvanus) and the others in Asia were examined with principal components analyses (PCA) and Euclidean distance analysis based upon prior discriminant function analyses (DFA). Results based on analyses of raw measurements indicate that the variation between species lies in the first axis of PCA; the species are dispersed according to their differences in size. The variation between sexes (sexual dimorphism) lies in the second axis. In the analyses of ratio variables, though these two patterns of separation remain orthogonal, they lie at approximately forty-five degrees to each axis. Variables relating to anterior teeth were found to play an important role in variation analysis, and this may be related to the special food preferences of these monkeys: more frequent usage of the incisor teeth for processing frugivorous diets than in other primates that are mainly folivorous. The results from Euclidean distance analyses indicate that the average distance of species within the Asian group is shorter than that between Asian and African groups regardless of sex and variable type. In addition the variation between African and Asian groups is larger than that within Asian group. Thus, it is reasonable to suggest that the African macaque has a range of measurements and ratios quite distinct from the species found in Asia (though the greatest separations result from the analyses of ratio data). These results therefore support the view that M. sylvanus may be regarded as an independent species group in the genus Macaca as proposed by Delson [1980].     

The dorsoventral axis is specified prior to first cleavage in the direct developing sea urchin Heliocidaris erythrogramma

Previous fate mapping studies as well as the culture of isolated blastomeres have revealed that the dorsoventral axis is specified as early as the 2-cell stage in the embryos of the direct developing echinoid, Heliocidaris erythrogramma. Normally, the first cleavage plane includes the animal-vegetal axis and bisects the embryo between future dorsal and ventral halves. Experiments were performed to establish whether the dorsoventral axis is set up prior to the first cleavage division in H. erythrogramma. Eggs were elongated and fertilized in silicone tubes of a small diameter in order to orient the cleavage spindle and thus the first plane of cell division. Following first cleavage, one of the two resulting blastomeres was then microinjected with a fluorescent cell lineage tracer dye. Fate maps were made after culturing these embryos to larval stages. The results indicate that the first cleavage division can be made to occur at virtually any angle relative to the animal-vegetal and dorsoventral axes. Therefore, the dorsoventral axis is specified prior to first cleavage. We argue that this axis resides in the unfertilized oocyte rather than being set up as a consequence of fertilization.     

Electrical Sizing of Particles in Suspensions: III. Rigid Spheroids and Red Blood Cells

The processes involved during the passage of a suspended particle through a small cylindrical orifice across which exists an electric field are investigated experimentally for an approximate prolate spheroid in the form of two tangent, rigid spheres (ragweed pollen particles) and for fresh, human red blood cells. Oscillograms of current pulses produced by both types of particles are presented and discussed in terms of particle shape and orientation and the effects of the hydrodynamic field. It is concluded that all the particles enter the orifice with their major axes aligned parallel to the orifice axis (electric field), but that during their passage some are rotated by the hydrodynamic field. Cells with their equatorial plane perpendicular to a radius of the orifice change their orientation with respect to the electric field as they are rotated, the others do not; only in the former case is there any deformation. It is shown that the bimodal or skewed size distributions can be explained on this basis, and that size (shape factor × volume) is actually a normally distributed variable (P > 95%). The average size of samples from 10 healthy adults was found to be 102.7 μ³ with a coefficient of variation of 1.8%. For a volume of 87 μ³, this corresponds to a shape factor of 1.18, an axial ratio (assuming a perfect oblate spheroid) of 0.26, and an equivalent major axis of 8.6 μ. The effect of high electric fields on red cell size distributions is mentioned.     

The oral-aboral axis of a sea urchin embryo is specified by first cleavage

Several lines of evidence suggest that the oral-aboral axis in Strongylocentrotus purpuratus embryos is specified at or before the 8-cell stage. Were the oral-aboral axis specified independently of the first cleavage plane, then a random association of this plane with the blastomeres of the four embryo quadrants in the oral-aboral plane (viz. oral, aboral, right and left) would be expected. Lineage tracer dye injection into one blastomere at the 2-cell stage and observation of the resultant labeling patterns demonstrates instead a strongly nonrandom association. In at least ninety percent of cases, the progeny of the aboral blastomeres are associated with those of the left lateral blastomeres and the progeny of the oral blastomeres with the right lateral ones, respectively. Thus, ninety percent of the time the oral pole of the future oral-aboral axis lies 45 degrees clockwise from the first cleavage plane as viewed from the animal pole. The nonrandom association of blastomeres after labeling of the 2-cell stage implies that there is a mechanistic relation between axis specification and the positioning of the first cleavage plane.     

Estimation of Sheet Thickness Distributions from Linear and Plane Sections

A biological “membrane” or a “barrier” can often be modelled by a sheet, namely a portion of space comprised between two smooth, quasi-parallel faces, such that sheet thickness is in general variable but very small in relation to face extent. Sheet thickness measurements are often of interest to the morphometrist, the physiologist and the pathologist. In the present paper, some stereological techniques are developed for estimating true sheet thickness distributions, and their moments, from the apparent thickness or intercept length measurements obtained, respectively, via random plane or linear sections. An important premise for the models to work is that of isotropic random (‘non-preferential’) sheet orientation relative to the sectioning probe.     

Head‐turning morphologies: Evolution of shape diversity in the mammalian atlas–axis complex

Mammals flex, extend, and rotate their spines as they perform behaviors critical for survival, such as foraging, consuming prey, locomoting, and interacting with conspecifics or predators. The atlas–axis complex is a mammalian innovation that allows precise head movements during these behaviors. Although morphological variation in other vertebral regions has been linked to ecological differences in mammals, less is known about morphological specialization in the cervical vertebrae, which are developmentally constrained in number but highly variable in size and shape. Here, we present the first phylogenetic comparative study of the atlas–axis complex across mammals. We used spherical harmonics to quantify 3D shape variation of the atlas and axis across a diverse sample of species, and performed phylogenetic analyses to investigate if vertebral shape is associated with body size, locomotion, and diet. We found that differences in atlas and axis shape are partly explained by phylogeny, and that mammalian subclades differ in morphological disparity. Atlas and axis shape diversity is associated with differences in body size and locomotion; large terrestrial mammals have craniocaudally elongated vertebrae, whereas smaller mammals and aquatic mammals have more compressed vertebrae. These results provide a foundation for investigating functional hypotheses underlying the evolution of neck morphologies across mammals.     

山西高原植被与气候的关系分析及植被数量区划的研究

本文利用去势典范对应分析和数量区划的方法,研究了山西高原植被与气候之间的关系,并进行了数量区划。排序的结果表明：DCCA的第—轴代表山西高原植被和气候梯度的纬向性,热量梯度是决定植被分布最主要的气候因子,水分梯度中的年降水量也对第—轴有较大的影响,由于山西高原南北跨度大,植被与气候因子表现出明显的纬向性;DCCA第二轴代表山西高原植被和气候梯度的经向性,与DCCA第二轴相关性较大的是水分因子中的年降水量、年蒸发量,由于山西高原东西跨度不大,而且大部分地区处在吕梁山脉和太行山脉之间,东西向的气候变化幅度不大,所以植被与气候梯度的经向性不明显。植被数量区划的结果表明：山西高原可划分为17个植被区,用图示的方法确定山西高原大致有三个极点和—个中心。     

Analysis of the clinal variation pattern inSilene latifolia (Caryophyllaceae) pollen morphology

Reticulate and microechinate pollen forms withinSilene latifolia (S. alba, S. pratensis) were first described from N. America, where the species is an introduced weed. A previous study showed that the two forms also exist in Europe and intergrade along a zone roughly congruent with intergradation zones in seed morphology and flavone glycosylation genotypes. The present survey of pollen from herbarium specimens is more extensive and covers localities from nearly the whole of the species' native Eurasian and North African range. The first axis of a detrended correspondence analysis (DCA) of qualitative characterstate data for 11 pollen morphological characters shows a gradient between the two extreme exine types, reticulate and microechinate. When mapped, the co-ordinates of pollen samples on this axis indicate a broad zone of intergradation which coincides approximately with the somewhat sharper transition between low- and high-tubercle seeds in C. and N. Europe, but becomes diffuse in Italy and the Mediterranean region and diverges from the seed transition zone in SW. Asia and the Middle East. The biological significance of the pollen morphs is unknown; the variation pattern in pollen morphology, unlike that in seeds, is not consistently correlated with macroclimate.     

Spatial autocorrelation in biology 1. Methodology

Spatial autocorrelation analysis tests whether the observed value of a nominal, ordinal, or interval variable at one locality is independent of values of the variable at neighbouring localities. The computation of autocorrelation coefficients for nominal, ordinal, and for interval data is illustrated, together with appropriate significance tests. The method is extended to include the computation of correlograms for spatial autocorrelation. These show the autocorrelation coefficient as a function of distance between pairs of localities being considered, and summarize the patterns of geographic variation exhibited by the response surface of any given variable.
Autocorrelation analysis is applied to microgeographic variation of allozyme frequencies in the snail Helix aspersa. Differences in variational patterns in two city blocks are interpreted.
The inferences that can be drawn from correlograms are discussed and illustrated with the aid of some artificially generated patterns. Computational formulae, expected values and standard errors are furnished in two appendices.     

Genetic similarities within and between human populations

The proportion of human genetic variation due to differences between populations is modest, and individuals from different populations can be genetically more similar than individuals from the same population. Yet sufficient genetic data can permit accurate classification of individuals into populations. Both findings can be obtained from the same data set, using the same number of polymorphic loci. This article explains why. Our analysis focuses on the frequency, omega, with which a pair of random individuals from two different populations is genetically more similar than a pair of individuals randomly selected from any single population. We compare omega to the error rates of several classification methods, using data sets that vary in number of loci, average allele frequency, populations sampled, and polymorphism ascertainment strategy. We demonstrate that classification methods achieve higher discriminatory power than omega because of their use of aggregate properties of populations. The number of loci analyzed is the most critical variable: with 100 polymorphisms, accurate classification is possible, but omega remains sizable, even when using populations as distinct as sub-Saharan Africans and Europeans. Phenotypes controlled by a dozen or fewer loci can therefore be expected to show substantial overlap between human populations. This provides empirical justification for caution when using population labels in biomedical settings, with broad implications for personalized medicine, pharmacogenetics, and the meaning of race.     

Variation in occlusal dental wear of two Chalcolithic populations in the southern Levant

Occlusal wear rate and wear plane in two Chalcolithic ( approximately 6500-5500 BP) samples from the southern Levant were compared, using paired first and second mandibular molars. Though food staples in both societies were derived from agro-pastoralism, they were located in distinct environmental regions: Wadi (W.) Makkukh in the Judean desert, and Peqi'in in the Upper Galilee. Accordingly, it was predicted that variation in wear should occur due to their location in distinct environments. Jaw size and tooth size were measured to estimate the possible impact of these variables on wear scores. Molar occlusal surfaces were divided into four quadrants, and wear scores were recorded for each quadrant. Principal axis analysis was then performed between total wear scores of paired, adjacent first and second molars to assess wear rates. Principal axis analysis was also used to analyze the change in occlusal wear plane in each sample by comparing between-buccal-cusps wear scores of the first molar with lingual wear scores of the second molar. The results indicate that the occlusal wear plane was similar in both samples but that wear tended to be more rapid in W. Makkukh. Since both samples were similar in jaw/tooth size, it is argued that the results reflect less refined food-processing methods as well as the unintentional ingestion of sand by individuals interred in the Judean desert.     

Stand characteristics,understorey associates and environmental correlates ofEucalyptus tetrodonta F. Muell. forests on Gunn Point,northern Australia

An ordination study of 20Eucalyptus tetrodonta forest stands growing on deep earths in monsoonal Australia revealed two major gradients in understorey vegetation type. The first axis reflected both floristic and structural understorey variation, where litter and shrub cover were inversely related to grass cover. This axis is thought to reflect a complex fire-vegetation type interaction, where vegetation is primarily determined by the saturation of the soil profile in the wet season, as measured by the colour of the iron rich soils. On the second axis of the ordination, floristic composition but not vegetation structure, and stand height were found to vary with the intercorrelated measures of soil gravel and moisture supply.E. tetrodonta is able to regenerate in the absence of fire, but firing appears to stimulate regeneration. All stands contained some advance growth, which occurs in distinct clumps, probably reflecting these plants clonal origin. Sapling presence in the stands is variable and the recruitment of advance growth into this size class appears to be related to over-wood competition. The size class distribution of trees was found to be similar amongst the stands, therefore stand structure appeared to be independent of understorey type. In comparison to general models of temperate eucalypt regeneration processes the tropical eucalypts have evolved different regeneration strategies, possibly in response to the severe annual drought.Nomenclature: Chippendale (1971), unless otherwise indicated. Structural classification: Specht (1970).I acknowledge the help of Mr. Clyde Dunlop who identified plant specimens and provided stimulating discussion, and Dr. Peter Minchin for permission to use the computer package ECOPACK to prepare and handle the floristic data, and his invaluable assistance with the ordination analysis. Mr. Lee Belbin and CSIRO Division of Water and Land Resources kindly gave their permission to use the Numerical Taxonomy Package (NTP).     

Community composition and soil properties in northern Bolivian savanna vegetation

Abstract. Direct gradient analysis (Canonical Correspondence Analysis) of northern Bolivian savanna vegetation revealed correlations between the composition of plant communities and physical and chemical soil properties. Cover/abundance values for 193 species from 27 sites were related to data on eight soil factors. A water-regime variable and the percentage of sand and silt were correlated with the first axis of the species-environment biplot and explained most of the variation in community composition. Along this axis, species and sites of flood-plain vegetation were separated from sites not affected by flooding rivers. The second axis of the biplot was correlated with soil-chemical variables, namely extract-able phosphate, base saturation, organic carbon, pH, and effective cation exchange capacity. Part of the variation in community composition can be explained by these soil nutrient variables. Grassland communities were separated from woody vegetation along the soil nutrient gradient, and floodplain communities of white-water rivers from those of clear-water rivers. The results of the gradient analysis indicate that the soil texture-moisture gradient is the prime factor determining the variation in the floristic composition of the savanna communities examined, and that, in addition, the soil nutrient gradient accounts for some of the variation.     

Evolutionary dissociation between cleavage, cell lineage and embryonic axes in sea urchin embryos.

Using vital dye staining and the microinjection of fluorescent cell lineage-autonomous tracers, the relationship between the first cleavage plane and the prospective larval dorsoventral axis was examined in several sea urchin species, including: Strongylocentrotus purpuratus, S. droebachiensis, Lytechinus pictus, Clypeaster rosaceus, Heliocidaris tuberculata and H. erythrogramma. The results indicate that there is no single relationship between the early cleavage pattern and the dorsoventral axis for all sea urchins; however, specific relationships exist for individual species. In S. purpuratus the first cleavage plane occurs at an angle 45 degrees clockwise with respect to the prospective dorsoventral axis in most cases, as viewed from the animal pole. On the other hand, in S. droebachiensis, L. pictus and H. tuberculata, the first cleavage plane generally corresponds with the plane of bilateral symmetry. There does not appear to be a predominant relationship between the first cleavage plane and the dorsoventral axis in C. rosaceus. In the direct-developing sea urchin H. erythrogramma the first cleavage plane bisects the dorsoventral axis through the frontal plane. Clearly, evolutionary differences have arisen in the relationship between cleavage pattern and developmental axes. Therefore, the mechanism of cell determination is not necessarily tied to any particular pattern of cell cleavage, but to an underlying framework of axial systems resident within sea urchin eggs and embryos.     

Evolutionary divergence in directions of high phenotypic variance in the ostracode genus poseidonamicus

Trait variation and covariation are understood to influence the response of populations to natural selection on generational time scales, but their role, if any, in shaping long-term macroevolutionary divergence is still unclear. The present study uses the rich fossil record of the ostracode genus Poseidonamicus to reconstruct in great detail the evolutionary history of a set of landmark-based morphometric characters. This reconstruction included two kinds of evolutionary inferences: ancestor-descendant transitions among populations repeatedly sampled at the same location and divergence between lineages measured as independent contrasts on a phylogeny. This reconstructed history was then used to test if evolutionary changes were concentrated in directions (traits or combinations of traits) with high phenotypic variance. Two different statistics of association between evolution and variation tested the null hypothesis that evolutionary changes occur in random directions with respect to trait variability. The first of these measured the similarity between the directions of evolutionary change and the axis of maximum variance, and the second measured the degree to which evolutionary changes were concentrated in directions of high phenotypic variation. Randomization tests indicated that both kinds of evolutionary inferences (ancestor-descendant and phylogenetic contrasts) occurred preferentially in directions of high phenotypic variance (and close to the axis of maximal variation), suggesting that within-population variation can structure long-term divergence. This effect decayed after a few million years, but at least for one metric, never disappeared completely. These results are consistent with Schluter's genetic constraints model in which evolutionary trajectories on adaptive landscapes are deflected by variation within and covariation among traits.     

The geography of crushing: Variation in claw performance of the invasive crab Carcinus maenas

The major claws of predatory, durophagous decapods are specialized structures that are routinely used to crush the armor of their prey. This task requires the generation of extremely strong forces, among the strongest forces measured for any animal in any activity. Laboratory studies have shown that claw strength in crabs can respond plastically to, and thereby potentially match, the strength of their prey's defensive armor. These results suggest that claw strength may be variable among natural populations of crabs. However, very few studies have investigated spatial variation in claw strength and related morphometric traits in crabs. Using three geographically separate populations of the invasive green crab in the Gulf of Maine, we demonstrate, for the first time, geographic variation in directly measured claw crushing forces in a brachyuran. Despite variation in mean claw strength however, the scaling of claw crushing force with claw size was consistent among populations. We found that measurements of crushing force were obtained with low error and were highly repeatable for individual crabs. We also show that claw mass, independent of a linear measure of claw size, and carapace color, which is an indicator of time spent in the intermoult, were important predictors of claw crushing force.     

Effects of sample distribution along gradients on eigenvector ordination

In general, disproportionately heavy sampling of the ends of a gradient increases the interpretability of eigenvector ordinations. More specifically, correspondence analysis (CA) and detrended correspondence analysis (DCA) best reproduce the original positions of samples in simulated coenoclines when samples are clustered toward the ends of the axis. Principal components analysis (PCA) reproduces the original sample positions less well than either CA or DCA and shows no improvement as samples are increasingly clustered toward the ends of the axis. PCA and CA show less curvature of one dimensional data into the second axis when sampling favors the ends of the axis.