The ontogeny of sexual dimorphism in the cranium of Bornean orang-utans (Pongo pygmaeus pygmaeus) as detected by principal-components analysis

The ontogeny of cranial sexual dimorphism in the Bornean orang-utan (Pongo pygmaeus pygmaeus) is examined by means of principal-components analysis (PCA). Normalized first components are called allometry vectors or vectors of relative growth and show that sexual dimorphism is present at all stages of growth. Two patterns of sexual dimorphism are present: (1) sexual differences at age groups 2 and 3 are the result primarily of differences in principal component II scores, reflecting mainly shape-related differences, and (2) age groups 5, 6, and 7 show a trend of stronger size-related shape differences with increasing age in the allometry vector along with decreasing differences in principal component II scores, reflecting an increase in size-related shape differences between the sexes. Age group 4 shows a combination of both patterns. Our results support Shea's hypothesis (1985a) that when using multigroup PCAs in closely related taxa, the allometry vector will generally estimate the shape variation resulting from the extension of common growth allometry patterns (ontogenetic scaling). The second and subsequent components summarize shape variation from slope and intercept differences between the groups, provided that ontogenetic scaling is not solely responsible for all the shape differences present. Subanalyses of those dimensions previously found to show ontogenetic scaling and acceleration follow this pattern well. The total sample provides a pattern whereby ontogenetically scaled dimensions possess a stronger influence over accelerated dimensions but still generally follow Shea's hypothesis. Finally, variously derived coefficients provided several interesting findings: (1) strong evidence was found against multivariate isometry for both the pooled and the separate samples, (2) multivariate allometric coefficients for both sexes follow the general growth pattern of negative scaling in neurocranial dimensions and positive scaling in the viscerocranium, and (3) multivariate slopes have a very high correlation with bivariate slopes relative to the same independent X variable, thereby lending further support to Jolicoeur's (1963a, b) allometry generalization.     

同域分布3种啄木鸟冬季取食的生态位差异

为了掌握黑啄木鸟、三趾啄木鸟和大斑啄木鸟的冬季取食行为特征,特别是三者之间取食生态位的差异,于2016年1月5-13日,在黑龙江省凉水国家级自然保护区以样线法结合样方法对3种啄木鸟的取食生境和取食行为进行了系统调查,收集了15个生境和行为特征变量数据。共布设45条样线,484个对照样方,收集312组啄木鸟取食数据,其中黑啄木鸟73组,三趾啄木鸟97组,大斑啄木鸟142组。多变量回归树和多分类逻辑斯谛分析结果显示,3种啄木鸟在所调查的15项特征上存在显著分异。采用基于利用-可利用方法的Bailey''s方法和双因子方差分析,分别对3种啄木鸟的生境选择和行为特征进行分析,结果显示：黑啄木鸟和三趾啄木鸟偏好在郁闭度较高的原始云、冷杉林中取食,而大斑啄木鸟则随机地在各种林型、生境中取食。黑啄木鸟、三趾啄木鸟多在树干取食,黑啄木鸟更常在倒木上取食,而大斑啄木鸟则多在树冠层取食。黑啄木鸟基本只在主干上凿洞,其他两种特别是大斑啄木鸟则可以在侧枝上取食。与黑啄木鸟和大斑啄木鸟凿洞取食昆虫不同,三趾啄木鸟多通过扒去树皮获得食物。黑啄木鸟的取食树基本为死树,单树取食时间最长,大斑啄木鸟多在活树上取食,单树取食时间最短,经常更换取食树,而三趾啄木鸟的取食树则死活参半,单树取食时间也较长。黑啄木鸟的冬季取食行为节律表现为双峰形,日出后和日落前各有一个活动高峰,其他两种则于白天持续取食。3种啄木鸟取食生境和行为生态位的差异,使它们能够更有效地利用有限的食物资源,共存于同一森林。     

Pre-Pleistocene Refugia and Differentiation between Populations of the Caucasian Salamander (Mertensiella caucasica)

A 350-bp fragment of the mitochondrial cytochrome-b gene was sequenced in the Caucasian salamander, Mertensiella caucasica, representing 10 populations from across its range along the Black Sea coast. Five haplotypes were discovered among 65 fragments analyzed, differing at 2–50 positions. The highest differentiation between haplotypes was observed in animals from the eastern part of the species' range (Borjomi) compared to those from the remainder of the species' range. Randomly amplified nuclear DNA revealed a pattern of spatial genetic variation similar to that of the mitochondrial genome. M. caucasica, as currently known, represents two evolutionary lineages that evolved independently, perhaps since the lower Pliocene. These lineages represent taxa, possibly to be described as species, distributed in the Borjomi area in central Georgia and in southwestern Georgia and northeastern Turkey. The multivariate analysis of morphological data did not reveal significant differences between the taxa. However, substantial morphological differentiation was observed within both lineages, showing parallel patterns in body proportions and coloration patterns. This variation is possibly associated with extant ecological conditions. Salamanders with reduced pigmentation from southwestern Georgia were not genetically distinguishable from neighboring populations.     

A multivariate analysis of geographic variation in morphology inPersoonia mollis (Proteaceae)

This study highlights the utility and limitations of multivariate procedures for the analysis and taxonomic interpretation of geographic variation within a complex species. Morphological data from herbarium specimens ofPersoonia mollis (Proteceae) were analyzed by two ordination procedures, multidimensional scaling (MDS) and canonical variate analysis (CVA). Support for the earlier recognition of nine infraspecific taxa was found. However, plots of canonical variate scores against latitute or longitude across parapatric subspecies boundaries revealed unexpected phenetic homogeneity within subspecies with sharp transitions between subspecies. The geographic variation in morphology withinP. mollis can be best described as a mosaic of nine recognisably distinct and internally homogeneous allopatric or parapatric taxa, where neighbouring taxa share surprisingly narrow zones of morphological transition at ecotones. The utility of these procedures was highlighted by the recovery of unexpected patterns of variation. However, questions of rank remain unresolved by phenetic analysis alone.     

天然和人工固沙灌木林蜘蛛和甲虫分布与环境因子的关系

干旱区人工植被恢复驱动的土地利用变化强烈影响了地表和土栖的节肢动物群落结构及多样性。然而,我们对地表节肢动物群落关键类群-蜘蛛和甲虫对固沙植被恢复的响应及与环境变化关系的认识还很有限。以天然固沙灌木林和2种人工固沙灌木林为研究对象,运用方差分析和多变量分析等方法定量研究了干旱区天然和人工固沙植被区地表蜘蛛和甲虫分布特征及影响要素。结果表明,天然灌木林与人工梭梭、柽柳林地表蜘蛛和甲虫群落组成明显不同,人工梭梭、柽柳林地表蜘蛛活动密度和甲虫多样性均显著高于天然灌木林,而地表甲虫密度和蜘蛛多样性变化与之相反。两种人工固沙灌木林之间蜘蛛和甲虫群落组成也存在一定差异,人工柽柳林地表蜘蛛活动密度、多样性和甲虫物种丰富度均显著高于人工梭梭林。进一步分析发现,蜘蛛群落中狼蛛科、平腹蛛科、皿蛛亚科和球蛛科与甲虫群落中拟步甲科、步甲科和象甲科等一些甲虫种属对3种生境的选择模式不同决定了蜘蛛和甲虫群落聚集结构。植被、土壤环境因子与蜘蛛和甲虫pRDA和pCCA结果表明,草本生物量、凋落物量、土壤含砂量、电导率和灌木盖度是影响蜘蛛分布的主要环境因子,它们解释了82.1%的蜘蛛群落变异;灌木盖度、草本生物量、土壤pH和砂含量是影响甲虫群落分布的主要环境因子,它们解释了60.6%的甲虫群落变异。总之,人工固沙灌木恢复影响了植被和土壤环境,它们相互作用改变了荒漠-绿洲过渡区蜘蛛和甲虫等地表节肢动物的分布格局。     

Spatial and Temporal Dynamics of Exotic Earthworm Communities Along Invasion Fronts in a Temperate Hardwood Forest in South-Central New York (USA)

The invasion of North American forests by exotic earthworms is producing profound ecosystem changes, such as alterations in soil nutrient cycling, and redistribution and loss of soil organic matter. However, the present and future extent of these invasions is difficult to evaluate without a better understanding of the factors that control the distribution and abundance of earthworms in previously non-invaded habitats. In this study, the species composition and short-term dynamics of three exotic earthworm invasion fronts were studied at a northern hardwood forest in south-central New York State (USA). Belt transects were established at each of the three locations to sample from earthworm-invaded areas through transition zones and into invasion front areas. Lumbricus rubellus, L. terrestrisandOctolasion tyrtaeum were the most common species, but their distribution was not homogeneous along the transects. Whereas, L. rubellus was the only species with relatively high adult densities at transition zones and invasion fronts, L. terrestris and O. tyrtaeum occurred mostly in the heavily earthworm-invaded areas and were rare at the invasion fronts. The density of earthworms along the transects decreased by 60–87 from June 2001 to October 2002 and then recovered in 2003 to values similar to those of 2001. This decrease was apparently caused by reduced recruitment of immature earthworms, probably related to the severe drought periods that the study area experienced in 2001 and 2002. Our data suggest that climate and topography, through their effects on soil moisture patterns, can be critical factors controlling the distribution and spread of exotic earthworms in previously non-invaded habitats.     

Predation and life-history variation in Poecilia reliculata (Cyprinodontiformes: Poeciliidae)

Synopsis Reznick and Endler investigated natural variation in life-history traits of populations of Trinidad guppies exposed to one of three intensities of predation: (i) high predation directed primarily at adults, (ii) moderate predation directed primarily at juveniles, and (iii) low predation. They were able to document significant interpopulational differences in life-history traits associated with this differential predation on a trait-by-trait basis. However, the present extended multivariate analysis indicates that (1) life-history traits do not differ significantly between populations exposed to moderate versus low predation, although both differ greatly from high-predation populations; (2) life-history variation is strongly unifactorial; and (3) despite the importance of predation effects, approximately 17% of the variation in life-history variables cannot be accounted for by predation intensity. Residual variation has no obvious geographical patterns, but instead seems to reflect local environmental variability. Life-history differences between predation regimes are consistent with residual patterns of variation within regimes, suggesting that local variation provides the raw material for extrapolation in response to predation, but also that it influences the direction of correlated change in life-history traits.     

A biometrical analysis of the South AmericanElymus glaucescens complex (Poaceae: Triticeae)

Based on a simple multivariate analysis of the morphology of theAgropyron pubiflorum complex—A. patagonicum, A. antarcticum, A. fuegianum, andA. magellanicum—the variation pattern is shown to be clinal, and all taxa are included in a single, variable species. Currently the majority of species ofAgropyron is included inElymus and the correct name of this species isE. glaucescens.     

Aspects of population dynamics in Halimione portulacoides communities

Summary Studies on sample plots inHalimione portulacoides communities show that environmental disturbances, either natural or induced by man, start a sequence of partly overlapping density maxima inSuaeda maritima, Aster tripolium andPuccinellia maritima successively, before the originalHalimione community totally recovers. When succession time before recovering is long enough, there are tendencies in redundancy of this sequence stressing the unilinear character of the succession. Minor environmental impacts induce a longer time-lag period of theSuaeda density maximum, suggesting threshold values of these impacts for the species to maintain minimal population densities or to become locally extinct. This sequence of interim species starting after an environmental disturbance, suggests also a gradient character in various biological attributes, for instance in life-time, propagation, nutrient and genetic plasticity strategies. The mechanism described can therefore be interpreted as a complex of mostly well-adapted and well-integrated inherent species strategies capable of absorbing environmental shocks. It is suggested that in the salt-marsh ccosystem the pattern of spatial variation in densities and that of temporal variation in fluctuations of the three species populations under natural conditions reflect corresponding patterns of environmental disturbances in the vegetation taking into account a timelag associated with the magnitude of the impact concerned.Contribution to the Symposium on Plant species and plant communities held at Nijmegen, 11–12 November 1976, on the occasion of the 60th birthday of Professor Victor Westhoff.Nomenclature follows Heukels-van Ooststroom. Flora van Nederland, 18e druk, 1975. Wolters-Noodhoff, Groningen.The authors are greatly indebted to Dr K.F. Vaas (Yerseke) for reviewing the English text.Communication Nr. 160.     

Patterns of morphometric and allozyme variation in Aedes albopictus

A survey of microgeographic variation using morphometric and allozyme analyses was conducted on 19 US populations of Aedes albopictus (Skuse) (Diptera: Culicidae), a mosquito that was recently introduced into the US. There was considerable variation within and among populations both in morphometric traits and allele frequencies. A multivariate discriminant analysis enabled the separation of populations into distinct groups; separation among the populations in the morphometric analysis was incomplete with an average of 70% of the individuals being correctly classified. In the allozyme analysis, the discrimination was complete. The populations from Texas were placed close together in the morphometric analysis, whereas in the allozyme analysis a geographic clustering of populations could not be detected. A test of association between the distance matrices derived from the morphometric and allozyme analyses was statistically nonsignificant. The results are discussed in the context of the colonization of the US by A. albopictus. The possible factors underlying the differences in the patterns of variation derived from morphometric and allozyme analyses are also discussed.     

Variation in the flash pattern of the firefly,Photuris versicolor quadrifulgens (Coleoptera: Lampyridae)

We sampled a population of signalingPhoturis versicolor quadrifulgens fireflies to quantify the variation in flash patterns emitted by males. Males produced five distinct flash patterns during their mate-searching flights. Four of the patterns consisted of two to five equal-intensity pulses and the fifth pattern type was a flicker, a group of rapid modulations in intensity. We found that the proportions of each pattern remained relatively constant from night to night throughout the season. The different flash patterns produced varied significantly with time of night; patterns having fewer pulses occurred earlier in the evening. Local density, an estimate of competition, did not significantly correlate with flash pattern type. On consecutive emissions, individuals changed their flash types with a mean probability of 0.12 (over all males), and they usually switched between patterns differing by a single pulse (from a two- to a three-pulse pattern, from a three- to a two-pulse pattern, etc.). The nightly temporal changes in flash patterns may be related to tradeoffs between female availability and energetic costs of signaling or the changes may be related to increased predation risk from visual predators. Photuris versicolor quadrifulgens was originally described by Barber (1951) as a subspecies ofP. versicolor. The genusPhoturis is currently under revision by Dr. James E. Lloyd, and this firefly will be given species status.     

Mating pattern variability among western toad (Bufo boreas) populations

Summary Breeding ecology and mating patterns of the western toad, Bufo boreas, were examined in three large explosively breeding populations in the Oregon Cascade Mountains. Two mating patterns occurred variably within and among the three populations. First, a large male mating advantage was observed in two of the three populations when data from all the days of data collection were combined. When each day of breeding was analyzed separately, there was a large male mating advantage on 3 of 5 days at one population, and mating was random on all days at the two other populations. The second mating pattern, positive assortative mating by size, was observed at two of the three populations. This pattern was found on separate days of breeding as well as when data from all days were combined at one population, and on only one day of breeding at the second population.In a survey of anuran amphibian mating patterns, intraspecific variation was found in 13 of 15 species, including the present study of the western toad. Intrapopulation variation in mating patterns among breeding years has been observed in 5 of 8 anurans, whereas within-site, within-year mating pattern variation has only been reported for the western toad. These results strongly suggest that anuran mating patterns are frequently neither species-specific nor population-specific attributes. Variable mating patterns were most commonly observed in explosively-breeding anurans. Explosive breeders may be susceptible to variable mating patterns because they may be more sensitive to fluctuations in environmental conditions, demographic parameters, and the intensities of intrasexual competition and mate choice.     

燕麦属颖果微形态特征及其分类学意义

为探讨燕麦属颖果微形态特征的分类学意义,采用体视显微镜和扫描电子显微镜观察燕麦属27种颖果的微形态特征,结合分子系统发育证据分析其分类学意义。燕麦属颖果有纺锤形、倒披针形、椭圆形3种形状,条纹、棱纹、网纹3种纹饰。燕麦属颖果形状、纹饰和花柱基宿存模式具有有限的属下分类学意义,颖果大小和表面大毛密度具有种间鉴定价值,而颖果腹面形态、压扁方式、胚比不具有种间鉴定价值。大穗燕麦(Avena macrostachya Balansa ex Coss.Durieu)颖果纺锤形,条纹纹饰,隶属于燕麦属颖果微形态特征的变异范围。大粒裸燕麦(A.nuda L.)与普通栽培燕麦(A.sativa L.)颖果大小、形状及纹饰特征的差异支持将大粒裸燕麦作为独立种处理。燕麦属颖果大小、表面大毛密度、胚比变异幅度大,推测与分布区广幅的气候变异相适应,凹腹面颖果体积相对缩减,有利于颖果快速发育、成熟,推测与燕麦属植物在温带、寒带分布区适生期较短相适应。     

The scale of demographic heterogeneity in a population of Peromyscus leucopus

Summary The patterns of spatial heterogeneity in density and demography were studied in a population of Peromyscus leucopus inhabiting a deciduous forest in west-central Indiana. A series of 9 live-trapping grids sampled densities from 3 spatial scales: 3 ha, 80 ha and 350 km². We found high levels of variation within all three spatial scales. There was as much variation within a single, large grid as within the entire Sugar Creek Valley (350 km²). The patterns of density variation were not temporally stable and thus represent variation in demographic processes rather than simply differences in habitat quality. We suggest that the Sugar Creek Valley represents a metapopulation of Peromyscus leucopus composed of a number of temporally unstable subpopulations. This structure provides considerable demographic variation that dispersing animals can exploit.     

Microgeographic edaphic differentiation in allozyme polymorphisms of wild barley (Hordeum spontaneum,Poaceae)

Allozymic variation in proteins encoded by 22 loci was analyzed electrophoretically in 278 individual plants of wild barley,Hordeum spontaneum, the progenitor of cultivated barley, in four 100 meter transects, in Israel, each equally subdivided into basalt and terra rossa soil types. Significant differentiation according to soil was found in 9 alleles. Our results suggest that allozyme polymorphisms in wild barley are at least partly adaptive, and differentiate by edaphic natural selection rather than by stochastic processes, and/or neutrality of allozymic variants.     

Atrazine-resistant cytoplasmic male-sterile-nigra broccoli obtained by protoplast fusion between cytoplasmic male-sterile Brassica oleracea and atrazine-resistant Brassica campestris

Summary By using restriction endonuclease digestion patterns, the degree of intraspecific polymorphism of mitochondrial DNA in four diploid species of wheat and Aegilops, Ae. speltoides, Ae. longissima, Ae. squarrosa, and Triticum monococcum, was assessed. The outbreeding Ae. speltoides was found to possess the highest degree of variability, the mean number of nucleotide substitutions among conspecific individuals being 0.027 substitutions per nucleotide site. A very low degree of mtDNA variation was detected among Ae. longissima accessions, with most of the enzyme-probe combinations exhibiting uniform hybridization patterns. The mean number of substitutions among Ae. longissima individuals was 0.001 substitutions per nucleotide site. The domesticated diploid wheat T. monococcum var. monococcum and its conspecific variant T. monococcum var. boeoticum seem to lack mitochondrial DNA variability altogether. Thus, the restriction fragment pattern can be used as a characteristic identifier of the T. monococcum cytoplasmic genome. Similarly, Ae. squarrosa accessions were found to be genetically uniform. A higher degree of variation among accessions is observed when noncoding sequences are used as probes then when adjacent coding regions are used. Thus, while noncoding regions may contain regulatory functions, they are subject to less stringent functional constraints than protein-coding regions. Intraspecific variation in mitochondrial DNA correlates perfectly with the nuclear variability detected by using protein electrophoretic characters. This correlation indicates that both types of variation are selectively neutral and are affected only by the effective population size.     

The structure of sculpin populations along a stream size gradient

Synopsis Data on spatial variation of sculpin density, growth and fecundity support the hypothesis that populations of stream fish are structured by changes in risk of predation and prey availability along a gradient in stream size. Cottus bairdi in warm streams and C. cognatus in cold streams exhibit similar patterns. Sculpins in large streams have faster individual growth rates and higher fecundities than those in small streams, but occur at lower density. The patterns appear to be persistent and suggest that predation reduces sculpin density in larger streams. Competitive release, variations in prey productivity, and local factors probably contribute to the variation in sculpin growth.     

Age-related differences in the performance, diffusion, and maintenance of stone handling, a behavioral tradition in Japanese macaques

Identifying the sources of behavioral diversity in non-human primates is vital to understanding the evolution of human behavior. Stone handling (SH, hereafter) is a form of object play consisting of the manipulation of stones by performing various behavioral patterns. This behavior is socially transmitted from generation to generation in Japanese macaques (Macaca fuscata), as a behavioral tradition. SH behavior in particular may reflect on the origin and evolution of stone-tool material culture. The objective of this study was to assess how group size, age structure, and age- and sex-related differences may account for the substantial intra- and inter-troop variations in SH reported in ten troops of Japanese macaques. Our results supported the hypothesis that patterns of variation in SH across troops reflected variability in group size and composition in age classes. We found that troop size was correlated with the proportion of troop members exhibiting SH simultaneously. The effect of troop size on the synchronized performance of SH may reveal the contagious nature of play. Our results suggest that the age structure of the group may affect the diffusion of SH. As predicted by the surplus energy hypothesis, a major functional hypothesis about play, intra-group variation in SH reflected more age- than sex-related differences. SH mainly occurred and was more frequent in younger than in older individuals, whereas no significant sex differences were found. SH episodes were shorter, more vigorous, and SH patterns were more diverse and less complex in immature than in mature individuals. The present findings reveal that age-related factors and group size may constrain the performance, diffusion, and maintenance of SH within a troop. Contrary to most other socially transmitted stone-tool using behaviors in non-human primates and early hominids, there is no optimal SH pattern. Provided some form of social learning, the non-adaptive nature of SH may allow particular SH pattern preferences to emerge at the group level. Our findings urge the use of an inter-populational comparative approach based on multivariate analyses when addressing the question of the evolution of behavioral traditions in primate and human populations.     

Codon usage and base composition inRickettsia prowazekii

Codon usage and base composition in sequences from the A + T-rich genome ofRickettsia prowazekii, a member of the alpha Proteobacteria, have been investigated. Synonymous codon usage patterns are roughly similar among genes, even though the data set includes genes expected to be expressed at very different levels, indicating that translational selection has been ineffective in this species. However, multivariate statistical analysis differentiates genes according to their G + C contents at the first two codon positions. To study this variation, we have compared the amino acid composition patterns of 21R. prowazekii proteins with that of a homologous set of proteins fromEscherichia coli. The analysis shows that individual genes have been affected by biased mutation rates to very different extents: genes encoding proteins highly conserved among other species being the least affected. Overall, protein coding and intergenic spacer regions have G + C content values of 32.5% and 21.4%, respectively. Extrapolation from these values suggests thatR. prowazekii has around 800 genes and that 60–70% of the genome may be coding. Correspondence to: S.G.E. Andersson