Variation and correlation in functional traits of main woody plants in the Cyclobalanopsis glauca community in the karst hills of Guilin,southwest China

研究植物功能性状变异以及不同性状之间的关系, 有助于了解植物对环境的适应策略, 对揭示群落构建和生物多样性维持机制具有重要意义。本研究以桂林岩溶石山青冈群落18种主要木本植物为研究对象, 采用单因素方差分析、混合线性模型以及Pearson相关分析和主成分分析等方法探讨了不同生长型和生活型物种的9个枝、叶功能性状(叶绿素含量、叶片厚度、叶面积、叶干质量、比叶面积、叶干物质含量、叶组织密度、小枝干物质含量和小枝组织密度)的变异与关联。结果表明: (1) 9个功能性状的变异程度不同, 叶面积和叶干质量的种内和种间变异系数最大, 小枝干物质含量和小枝组织密度的种内和种间变异系数最小。(2)在生长型水平上, 乔木、灌木和木质藤本的绝大部分功能性状差异显著。(3)对于不同生活型植物, 除叶面积和比叶面积为落叶植物显著大于常绿植物外, 其余7个功能性状皆为常绿植物显著大于落叶植物。(4)不同生长型和生活型植物功能性状的种内和种间变异大小存在差异, 除部分植物功能性状在一些功能型上表现为种内变异高于种间变异, 其余功能性状的种间变异皆高于种内变异。(5)各功能性状之间关系在个体水平和物种水平大致相同, 但是个体水平上的显著相关比例高于物种水平。研究发现, 植物功能性状的种间变异基本高于种内变异, 但种内变异不容忽略。此外, 不同生长型和生活型物种采取不同的生态策略来适应喀斯特生境。今后的研究应基于个体水平采样并结合环境因子从生长型和生活型等不同功能型角度上深入探究植物功能性状在种内和种间等不同尺度上的变异与关联。     

亚热带常绿落叶阔叶混交林植物功能性状的种间和种内变异

不同物种间的功能性状差异是自然生态系统中物种共存的基础, 而物种内个体间的性状变异对物种的共存和分布同样具有重要作用。本文以湖北星斗山自然保护区亚热带常绿落叶阔叶混交林内28种主要树种(通过物种多度排序获得, 其中常绿和落叶树种各14种)为研究对象, 探讨不同叶习性树种的4种功能性状(比叶面积、叶干物质含量、叶面积和比茎密度)在种间和种内的差异程度。结果表明: (1)常绿和落叶树种在4种功能性状上均存在显著差异, 常绿树种的比叶面积和叶面积显著低于落叶树种, 但叶干物质含量和比茎密度则显著高于落叶树种; (2)比叶面积的变化主要来源于叶习性(57.49%), 叶面积变化主要来源于种间(66.80%)和种内变异(27.52%), 叶干物质含量的变化主要来源于种间(38.12%)和种内(33.88%)变异, 但比茎密度的变化主要来源于种内变异(51.50%), 其次为种间变异(32.52%); (3)常绿和落叶树种种间水平的性状相关性可能掩盖各功能性状之间的相关性。种内变异能够显著影响群落间的植物功能性状差异, 但不同功能性状的种内变异程度存在差异。     

桂林岩溶石山青冈群落主要木本植物功能性状变异与关联

研究植物功能性状变异以及不同性状之间的关系, 有助于了解植物对环境的适应策略, 对揭示群落构建和生物多样性维持机制具有重要意义。本研究以桂林岩溶石山青冈群落18种主要木本植物为研究对象, 采用单因素方差分析、混合线性模型以及Pearson相关分析和主成分分析等方法探讨了不同生长型和生活型物种的9个枝、叶功能性状(叶绿素含量、叶片厚度、叶面积、叶干质量、比叶面积、叶干物质含量、叶组织密度、小枝干物质含量和小枝组织密度)的变异与关联。结果表明: (1) 9个功能性状的变异程度不同, 叶面积和叶干质量的种内和种间变异系数最大, 小枝干物质含量和小枝组织密度的种内和种间变异系数最小。(2)在生长型水平上, 乔木、灌木和木质藤本的绝大部分功能性状差异显著。(3)对于不同生活型植物, 除叶面积和比叶面积为落叶植物显著大于常绿植物外, 其余7个功能性状皆为常绿植物显著大于落叶植物。(4)不同生长型和生活型植物功能性状的种内和种间变异大小存在差异, 除部分植物功能性状在一些功能型上表现为种内变异高于种间变异, 其余功能性状的种间变异皆高于种内变异。(5)各功能性状之间关系在个体水平和物种水平大致相同, 但是个体水平上的显著相关比例高于物种水平。研究发现, 植物功能性状的种间变异基本高于种内变异, 但种内变异不容忽略。此外, 不同生长型和生活型物种采取不同的生态策略来适应喀斯特生境。今后的研究应基于个体水平采样并结合环境因子从生长型和生活型等不同功能型角度上深入探究植物功能性状在种内和种间等不同尺度上的变异与关联。     

Evolution and functional significance of tendon ossification in woodcreepers (Aves: Passeriformes: Dendrocolaptinae)

The woodcreepers, a clade of scansorial, neotropical birds, are distinctive among passerines in having extensive tendon ossification. Dissection of 42 of the 50 species indicates that such ossification in the hindlimb is limited almost entirely to tendons of insertion of the crural muscles. Most crural muscles have ossifications, and in all but one the ossified tendons are long and thin. Preliminary dissection revealed a similar pattern among ossified wing tendons. Phylogenetic analysis suggests that extensive tendon ossification is a synapomorphy of the woodcreepers. The species of Dendrocincla, which form a clade, show secondary reduction of ossification in some tendons, which may be correlated with increased intraspecific variation and with an expansion of foraging habits and postures to include nonscansorial behaviors. In contrast, the larger woodcreepers, other than Drymornis bridgesii and Nasica longirostris, form a clade with virtually no loss in ossification or evidence of intraspecific variation, even in large series of two species. Phylogenetic losses do not occur for the primary flexor of the ankle (M. tibialis cranialis), whereas two extensors (Mm. fibularis longus and gastrocnemius pars lateralis) show a complex pattern of derivation and loss. Previous biomechanical studies demonstrate that ossification increases the stiffness of tendons, making them stretch less under a given force. These structural and phylogenetic patterns are consistent with the view that hindlimb tendon ossification in woodcreepers is an adaptation to resist increased forces that act to extend the limb during vertical climbing. © 1993 Wiley-Liss, Inc.     

Feeding biology of sunfishes: patterns of variation in the feeding mechanism

Fishes are generally believed to differ in their ability to alter (modulate) their feeding behaviour in response to different prey. We investigated modulation quantitatively in four species of sunfishes (Centrarchidae) by evaluating the variation in 11 electromyographic variables measured from recordings of electrical activity in head muscles during feeding on three prey types. The experimental design used allowed us to partition variation between species, among individuals within species, among prey types, and among feedings. Duration of activity of the sternohyoideus muscle was the only variable significantly different among the four species. All variables showed significant differences among individuals within species. The overall range of activity of each muscle activity variable was about the same for all four sunfishes. However, three species showed a significant ability to modulate most muscle variables while a fourth did not change its feeding response with respect to prey type. The results indicate that: (1) intraspecific variation is an important source of variability in functional attributes that should be accounted for in comparisons between species; (2) the ability of some species to modulate appears to be independent of the potential variability in muscle activity possessed by each species; (3) closely related species can differ considerably in their ability to alter muscle electrical activity patterns during feeding; and (4) a quantitative assessment of variation in electromyographic patterns is a powerful approach for asking questions about differences in feeding behaviour.     

鼎湖山南亚热带常绿阔叶林叶功能性状沿群落垂直层次的种内变异

探究功能性状沿着环境梯度如何变化一直以来是基于性状的群落生态学的核心问题之一。尽管功能性状存在种内和种间变异, 但种内变异沿环境梯度如何变化仍有待探究。本文以鼎湖山南亚热带常绿阔叶林1.44 ha塔吊样地内16个树种的2,820个个体为研究对象, 探究4种叶功能性状(比叶面积、叶干物质含量、叶厚度和叶面积)沿群落垂直层次的种内变异。首先, 利用随机效应线性模型量化塔吊样地内的种内变异和种间变异; 其次, 利用Kmeans函数将森林的垂直层次划分为灌木层、亚冠层和林冠层, 并通过构建回归模型探究叶功能性状在群落垂直层次中的种内变异格局。最后, 应用混合线性模型和单因素方差分析的方法探究叶功能性状沿垂直层次的种内变异是否具有物种依赖性。结果表明: 在局域群落中, 并非所有叶功能性状的种内变异都低于种间变异; 叶功能性状在不同垂直层次的种内变异格局存在显著差异, 且种内变异与垂直范围呈正相关; 叶功能性状的种内变异具有较强的物种依赖性, 因此树种差异相对于小环境解释了更多的性状变异; 此外, 不同叶功能性状的种内变异沿垂直层次的变化趋势并不一致。本研究发现种内变异对于物种共存具有重要作用。     

Timing of ossification in duck, quail, and zebra finch: intraspecific variation, heterochronies, and life history evolution

Skeletogenic heterochronies have gained much attention in comparative developmental biology. The temporal appearance of mineralized individual bones in a species - the species ossification sequence - is an excellent marker in this kind of study. Several publications describe interspecific variation, but only very few detail intraspecific variation. In this study, we describe and analyze the temporal order of ossification of skeletal elements in the zebra finch, Taeniopygia guttata, the Japanese quail, Coturnix coturnix japonica, and the White Pekin duck, a domestic race of the mallard Anas platyrhynchos, and explore patterns of intraspecific variation in these events. The overall sequences were found to be conserved. In the duck, variability is present in the relative timing of ossification in the occipital, the basisphenoid and the otic regions of the skull and the phalanges in the postcranium. This variation appears generally in close temporal proximity. Comparison with previously published data shows differences in ossification sequence in the skull, the feet, and the pelvis in the duck, and especially the pelvis in the quail. This clearly documents variability among different breeds.     

Individual variation in two host plants of the ladybird beetle,Epilachna pustulosa (Coleoptera: Coccinellidae)

Individual variation in two species of host plants (thistle,Cirsium kamtschaticum, and blue cohosh,Caulophyllum robustum) of the herbivorous ladybird beetleEpilachna pustulosa was examined under laboratory conditions for their acceptability to adult beetles as a food resource, for adult preference and for larval performance. When clones of these plants were subjected to non-choice tests using posthibernating female beetles, there was found to be significant intraspecific variation among clones in terms of their acceptability, but interspecific variation was not detected. Significant intraspecific as well as interspecific variation were frequently detected in the two host plants when clones of these plants were subjected to choice tests using posthibernating female beetles; the magnitude of interspecific plant variation for beetle preference is not necessarily larger than that of intraspecific plant variation. Individual variation across plant species with respect to beetle larval performance was also significant. A positive correlation between adult preference and larval performance is suggested across the two taxonomically remote host plant species, thistle and blue cohosh, although this needs further investigation.     

The community and ecosystem consequences of intraspecific diversity: a meta‐analysis

Understanding the relationships between biodiversity and ecosystem functioning has major implications. Biodiversity–ecosystem functioning relationships are generally investigated at the interspecific level, although intraspecific diversity (i.e. within‐species diversity) is increasingly perceived as an important ecological facet of biodiversity. Here, we provide a quantitative and integrative synthesis testing, across diverse plant and animal species, whether intraspecific diversity is a major driver of community dynamics and ecosystem functioning. We specifically tested (i) whether the number of genotypes/phenotypes (i.e. intraspecific richness) or the specific identity of genotypes/phenotypes (i.e. intraspecific variation) in populations modulate the structure of communities and the functioning of ecosystems, (ii) whether the ecological effects of intraspecific richness and variation are strong in magnitude, and (iii) whether these effects vary among taxonomic groups and ecological responses. We found a non‐linear relationship between intraspecific richness and community and ecosystem dynamics that follows a saturating curve shape, as observed for biodiversity–function relationships measured at the interspecific level. Importantly, intraspecific richness modulated ecological dynamics with a magnitude that was equal to that previously reported for interspecific richness. Our results further confirm, based on a database containing more than 50 species, that intraspecific variation also has substantial effects on ecological dynamics. We demonstrated that the effects of intraspecific variation are twice as high as expected by chance, and that they might have been underestimated previously. Finally, we found that the ecological effects of intraspecific variation are not homogeneous and are actually stronger when intraspecific variation is manipulated in primary producers than in consumer species, and when they are measured at the ecosystem rather than at the community level. Overall, we demonstrated that the two facets of intraspecific diversity (richness and variation) can both strongly affect community and ecosystem dynamics, which reveals the pivotal role of within‐species biodiversity for understanding ecological dynamics.     

Disentangling the relative importance of species occurrence,abundance and intraspecific variability in community assembly: a trait‐based approach at the whole‐plant level in Mediterranean forests

Understanding which factors and rules govern the process of assembly in communities constitutes one of the main challenges of plant community ecology. The presence of certain functional strategies along broad environmental gradients can help to understand the patterns observed in community assembly and the filtering mechanisms that take place. We used a trait‐based approach, quantifying variations in aboveground (leaf and stem) and belowground (root) functional traits along environmental gradients in Mediterranean forest communities (south Spain). We proposed a new practical method to quantify the relative importance of species turnover (distinguishing between species occurrence and abundance) versus intraspecific variation, which allowed us to better understand the assemblage rules of these plant communities along environmental gradients. Our results showed that the functional structure of the studied plant communities was highly determined by soil environment. Results from our modelling approach based on maximum likelihood estimators showed a predominant influence of soil water storage on most of the community functional traits. We found that changes in community functional structure along environmental gradients were mainly promoted by species turnover rather than by intraspecific variability. Specifically, our new method of variance decomposition demonstrated that between‐site trait variation was the result of changes in species occurrence rather than in the abundance of certain dominant species. In conclusion, this study showed that water availability promoted the predominance of specific trait values (both in above and belowground fractions) associated to a resource acquisition or conservation strategy. In addition, we provided evidence that changes on community functional structure along the environmental gradient were mainly promoted by a process of species replacement, which represent a crucial step towards a more general understanding of the relative importance of intraspecific versus interspecific trait variation in these woody Mediterranean communities.     

Prey abundance and male parental behaviour in Acrocephalus warblers

This study examined the prey abundance at nesting sites of five different Acrocephalus warbler species. Intraspecific variations in prey density were analysed to determine the relative significance of this ecological factor for different mating systems and components of male parental behaviour. The Great Reed Warbler Acrocephalus arundinaceus, a facultatively polygynous species, nested in areas with the highest insect abundance. Its intraspecific variation in prey density (between different territories) was the highest while the male: female offspring feeding ratio was the lowest. At the other extreme, the monogamous Moustached Warbler Acrocephalus melanopogon and Reed Warbler Acrocephalus scirpaceus nested in areas with the lowest prey abundance and lowest intraspecific variation. Males of these species had the highest involvement in feeding young. In the Moustached Warbler and Marsh Warbler Acrocephalus palustris, an intraspecific comparison revealed a negative correlation between insect abundance and the male feeding component; a tendency in this direction was found for the Reed Warbler, while no correlation was evident in the polygynous Great Reed Warbler. These data demonstrate a general pattern relating prey abundance, mating systems and male parental investment on an interspecific level. However, intraspecific correlations revealed a net of complex interdependence. One of the factors which may be important is resource predictability. We found a significant correlation between certain vegetation types and insect abundance. The vegetation composition of territories is further influenced by interspecific dominance relationships whereby the dominant Great Reed Warbler occupies territories with the vegetation type which correlates with high prey abundance while the overlapping Moustached Warbler and Reed Warbler are restricted to areas with lower insect density.     

An experimental study of intraspecific variation, developmental timing, and heterochrony in fishes

Heterochrony is widely regarded as an important evolutionary mechanism, one that may underlie most, if not all, morphological evolution, yet relatively few studies have examined variation in the sequence of development. Even fewer studies have been designed so that intraspecific variation in the relative sequence of developmental events can be assessed, although this variation must be the basis for evolutionary change. Intraspecific variation in developmental ossification sequences was documented from the zebrafish (Danio rerio) by Cubbage and Mabee (1996) and from the Siamese fighting fish (Betta splendens) by Mabee and Trendler (1996), but a quantitative analysis of the patterns within this variation was not made. Here, we quantify the effect of rearing temperature on the sequence of ossification and characterize the levels and patterns of intraspecific variation in these fishes. For Danio, there were no temperature effects on the sequence of bone development across the cranium, cranial region development, cartilage versus dermal bones, or lateral line bone versus nonassociated bones. Likewise the level of variation in relative sequence (position) of ossification was low, about two ranks, across temperatures. At higher temperatures, we found higher levels of variation in iterated cranial bones and less in bones forming early in the sequence. No temperature effects on variation were found among regions, between lateral line-associated bones and nonassociated bones, between median and paired bones, or across the entire sequence, indicating concordant variability among the three temperatures. Individual bones with the highest levels of variability were not consistent among temperatures. Baseline patterns of intraspecific variation in Danio were compared to those of Betta. For both species, the level of intraspecific variation in sequence position was low and the variability of cranial bones was concordant. Individual bones with the highest levels of variability were not consistent between species. In both species, variation was widespread (distributed evenly across the sequence). We used comparisons (among regions, between dermal and cartilage bones, between lateral line-associated and other bones, between median and paired bones, between iterated and noniterated bones, between feeding-associated bones and others) to see which subsets were most variable and thus potentially useful in predicting high levels of evolutionary change. The only subset of bones that was significantly more variable than others was cartilage bones. If interspecific patterns are parallel to these intraspecific differences, cartilage bones would be expected to show higher levels of heterochrony. Although concordance across the cranial ossification sequence and among regions in Danio, Betta, and two other teleosts, Oryzias and Barbus, suggests an evolutionarily conserved pattern of ossification, identity in sequence position across taxa was not observed for any bone. Thus, variation existed in sequence position across temperatures and species. Intraspecific variation of this sort may influence the morphological outcome and evolutionary trajectories of species.     

Genome size and genome evolution in diploid Triticeae species.

One of the intriguing issues concerning the dynamics of plant genomes is the occurrence of intraspecific variation in nuclear DNA amount. The aim of this work was to assess the ranges of intraspecific, interspecific, and intergeneric variation in nuclear DNA content of diploid species of the tribe Triticeae (Poaceae) and to examine the relation between life form or habitat and genome size. Altogether, 438 plants representing 272 lines that belong to 22 species were analyzed. Nuclear DNA content was estimated by flow cytometry. Very small intraspecific variation in DNA amount was found between lines of Triticeae diploid species collected from different habitats or between different morphs. In contrast to the constancy in nuclear DNA amount at the intraspecific level, there are significant differences in genome size between the various diploid species. Within the genus Aegilops, the 1C DNA amount ranged from 4.84 pg in A. caudata to 7.52 pg in A. sharonensis; among genera, the 1C DNA amount ranged from 4.18 pg in Heteranthelium piliferum to 9.45 pg in Secale montanum. No evidence was found for a smaller genome size in annual, self-pollinating species relative to perennial, cross-pollinating ones. Diploids that grow in the southern part of the group's distribution have larger genomes than those growing in other parts of the distribution. The contrast between the low variation at the intraspecific level and the high variation at the interspecific one suggests that changes in genome size originated in close temporal proximity to the speciation event, i.e., before, during, or immediately after it. The possible effects of sudden changes in genome size on speciation processes are discussed.     

Temperature-dependent plasticity of segment number in an arthropod species: the centipede Strigamia maritima

SUMMARY The evolution of arthropod segment number provides us with a paradox, because, whereas there is more than 20‐fold variation in this character overall, most classes and orders of arthropods are composed of species that lack any variation in the number of segments. So, what is the origin of the higher‐level variation? The centipede order Geophilomorpha is unusual because, with the exception of one of its families, all species exhibit intraspecific variation in segment number. Hence it provides an opportunity to investigate how segment number may change in a microevolutionary context. Here, we show that segment number can be directly altered by an environmental factor (temperature)—this is the first such demonstration for any arthropod. The direction of the effect is such that higher temperature during embryogenesis produces more segments. This potentially explains an intraspecific cline in the species concerned, Strigamia maritima, but it does not explain how such a cline is translated into the parallel interspecific pattern of lower‐latitude species having more segments. Given the plastic nature of the intraspecific variation, its link with interspecific differences may lie in selection acting on developmental reaction norms.     

桂林岩溶石山青冈群落植物功能性状的种间和种内变异研究

研究植物功能性状的种间和种内变异,有助于揭示植物对环境的适应策略及其群落构建机制研究。以桂林岩溶石山青冈群落主要木本植物为研究对象,选取叶干物质含量（LDMC）、叶厚度（LT）、比叶面积（SLA）和木材密度（WD）4个功能性状,采用混合线性模型结合方差分解方法对其在种内和种间尺度的变异程度和相对贡献进行分析,同时运用零模型方法探讨驱动岩溶石山青冈群落构建的潜在机制。结果表明：（1） LDMC,LT,SLA和WD这4个性状在种内和种间水平上均存在不同程度的变异幅度,种间变异大于种内变异,但种内变异亦不容忽略。（2）零模型检验表明,将种内变异纳入考虑的环境过滤检测得到改善,基于个体性状值的尺度考虑群落构建机制研究是必要的。（3）对于桂林岩溶石山青冈群落,环境过滤作用确实是群落构建的重要驱动机制,与相似性限制作用和随机作用共同塑造了以青冈为优势物种的群落构建。     

Ontogenetic Sequence Analysis: using parsimony to characterize developmental sequences and sequence polymorphism

Ontogenetic sequences are a pervasive aspect of development and are used extensively by biologists for intra- and interspecific comparisons. A tacit assumption behind most such analyses is that sequence is largely invariant within a species. However, recent embryological and experimental work emphasizes that ontogenetic sequences can be variable and that sequence polymorphism may be far more prevalent than is generally realized. We present a method that uses parsimony algorithms to map hierarchic developmental patterns that capture variability within a sample. This technique for discovering and formalizing sequences is called the "Ontogenetic Sequence Analysis" (OSA). Results of OSA include formalized diagrams of reticulating networks, describe all most parsimonious sequences, and can be used to develop statistics and metrics for comparison of both intraspecific and interspecific sequence variation. The method is tested with examples of human postnatal skeletal ossification, comprising a time-calibrated data set of human hand and wrist epiphyseal unions, and a longitudinal data set of human wrist ossification. Results illustrate the validity of the method for discovering sequence patterns and for predicting morphologies not represented in analytic samples. OSA demonstrates the potential and challenges of incorporating ontogenetic sequences of morphological information into evolutionary analyses.     

Intraspecific Variation in the Vocalizations and Hand Pad Morphology of Southern Lesser Bush Babies (Galago moholi): A Comparison with G. senegalensis

Previous studies have shown the taxonomic value of vocal repertoires and hand (volar) pad characteristics in the classification of cryptic nocturnal primates such as bush babies. However, no study included quantitative comparisons within the geographical range of any one species. We investigated levels of intraspecific variation in calls and hand pad characteristics of the southern lesser bush baby (Galago moholi), using the northern lesser bush baby (Galago senegalensis) for interspecific comparisons. Examination of calls recorded from different regions along a transect of 1500 km across southern Africa revealed low levels of intraspecific variation in Galago moholi, whereas comparisons with homologous call-types in G. senegalensis revealed them to be significantly different. Volar pad measurements across the ranges of both species also showed low levels of intraspecific variation and relatively high interspecific variation. These findings demonstrate that vocal and volar pad characteristics can be used as consistent measures of difference between species that look almost identical. These methods provide a practical means of distinguishing between cryptic species, whether in the field, in captivity, or, in the case of volar pads, of preserved specimens.     

Extensive intraspecific chloroplast DNA (cpDNA) variation in the alpine Draba aizoides L. (Brassicaceae): haplotype relationships and population structure

Chloroplast DNA (cpDNA) sequence variation is currently the most widely used tool for the inference of phylogenetic relationships among plants at all taxonomic levels. Generally, noncoding regions tend to evolve faster than coding sequences and have recently been applied to the study of phylogenetic relationships among closely related taxa. An implicit assumption of many of these studies is that intraspecific cpDNA variation is either absent or low and therefore will not interfere with the reconstruction of interspecific relationships. A survey of cpDNA sequence variation in the common alpine plant species Draba aizoides L. was undertaken to assess levels of intraspecific cpDNA sequence variation. These levels were compared to levels of interspecific sequence divergence between D. aizoides and related alpine Draba species. Intraspecific cpDNA sequence divergence was extensive in D. aizoides, and intraspecific differences were often larger than interspecific differences. cpDNA haplotype relationships were explored using a maximum parsimony approach and minimum-spanning networks. Results from both methods were largely congruent but comparisons provided interesting insights into the presumed evolutionary history of cpDNA haplotypes. A combined effect of cpDNA introgression and complex lineage sorting was inferred to explain the pattern of cpDNA variation found in D. aizoides. Our results suggest that intraspecific cpDNA variation can be extensive and that intraspecific variation needs to be taken into account when inferring phylogenetic relationships among closely related taxa.     

Ecogeographical rules: elements of a synthesis

The development of a more synthetic approach to understanding spatial patterns in biogeography, particularly of the way in which these patterns interact, constitutes a major challenge for the field. Here we propose some key elements of such a synthesis for what can broadly be termed 'ecogeographical rules', that is spatial patterns in biological traits. These include understanding: (1) the different kinds of patterns (intraspecific, interspecific and assemblage), and the distinctions between them; (2) the unifying role that geographical ranges play in linking the patterns together; (3) that this unification can be obscured by the methodological assumptions made in documenting some patterns (e.g. assuming that intraspecific variation does not significantly influence interspecific and assemblage patterns in traits); (4) the implications of other methodological issues for the nature of observed patterns (e.g. how ranges are located on positional or environmental axes for interspecific patterns); (5) the need for further development of models linking different types of traits; (6) the nature of the generality of documented patterns at all levels, and particularly the difference between the frequency with which patterns are documented in the literature and the variety of extant species; and (7) the constraints that the form of intraspecific patterns place on interspecific and assemblage patterns, and that interspecific patterns place on assemblage patterns.