甘草野生种群果实与种子形态特征的地理变异研究

目的：研究甘草荚果和种子形态特征地理变异规律并分析其形成的生态学机制。方法：采用样方调查的方法测定全国4省区8个旗县甘草野生种群的地上植株形态特征,采用双重筛选逐步回归分析方法探讨甘草荚果和种子形态特征的地理变异规律及其形成的生态学机制。结果：（1）不同甘草种群的荚果数、荚果腺毛长度、种室数目、每荚种子粒数和结实率5项指标的差异达到了极显著水平（P〈0．01）;荚果长度、荚果厚度、荚果腺毛密度和种子长度4项指标的差异达到了显著水平（P〈0．05）。（2）荚果厚度、种室数目等5项形态特征与经度呈显著相关,只有荚果腺毛长度1项指标与纬度显著相关。结论：甘草野生种群荚果与种子形态特征存在显著的地理变异,其变异趋势以经向变异为主,光照因子差异是导致地理变异的主要原因,其次为温度因子和水分因子,其相关关系可用多元回归模型解释。     

Photoperiodic response and heritability of the pre-flowering interval of two red clover (Trifolium pratense) populations

The reaction to 12, 14, and 16 h photoperiods of two medium-maturing red clover (Trifolium pratense) populations (cv. Kenland and Cycle 6 of a selection for long stems) under controlled environment conditions was studied. Under both 14 and 16 h photoperiods, Kenland flowered sooner and had shorter stems, more stems per plant, and greater herbage yield per plant but had a similar number of internodes compared to the long stem Cycle 6 population. A daylength between 12 and 14 h was critical for flower initiation in both populations. The critical daylength to induce stem initiation was shorter than the daylength to induce flowering in some genotypes. From half-sib analyses, the narrow–sense heritability of the pre-flowering interval in Kenland ranged from 15 to 23%. The expected response to one cycle of selection (5% selection intensity) for lengthening or shortening the pre-flower interval was 2.3 to 2.4 days at the 16 h photoperiod and 3.0 days at the 14 h photoperiod. Low correlation between pre-flowering interval, stem length, and stem number indicated that it may be possible to change one character without affecting the other two. The need for further studies of the genetic nature of the control of flowering in red clover was indicated.     

Life history variation and infraspecific heterochrony inRhinanthus glacialis (Scrophulariaceae)

Comparative cultivation experiments were made with 14 populations of the annual, hemiparasitic speciesRhinanthus glacialis from seven different habitat types. Phenological data were recorded and plants from field samples and from corresponding experimental samples were morphologically analysed.There are up to ten weeks difference in the start of flowering between means of the extreme experimental populations. Flowering time variation correlates with the number of internodes and also with most of the other morphological characters. Flowering time also shows close relationships to different grassland types and the examined populations appear to be well-adapted to the environmental conditions in natural or man-influenced grassland types as far as flowering time is concerned. The number of internodes is mostly insensitive to the applied experimental influences. The rate of internode production is constant. Trade-offs are suggested for some morphological traits (i.e., few internodes with few branches and thus few flowers as trade-off for early flowering). — Previous studies of life history variation based on population data have not been incorporated into the theory of heterochrony. The pattern of life history variation of populations ofR. glacialis together with information about morphology, ecology, and phylogeny suggests that some variants have evolved from ancestors via heterochronic processes. Earlier flowering populations from montane hay meadows, litter meadows, and dry meadows belong to paedomorphic variants. These have fewer internodes and branches as well as reduced height in comparison with the suggested ancestor. Later flowering populations from subalpine limestone grassland belong to a peramorphic variant with more internodes, more branches, and taller stems in comparison with the suggested ancestor.     

Phenotypic Variation among and within Three Peppers Species (<i>Capsicum</i>) from Mexico

The phenotypic variation and its distribution among species, morphotypes, and among and within populations was estimated in 71 populations pertaining to 15 morphotypes of three domesticated species of Capsicum from Mexico. Collections were made in the states of Sinaloa, Nayarit, Oaxaca, Tabasco, and Yucatán in two agroecosystems: Backyard-garden and monoculture. Fifteen phenotypic characteristics were analyzed through one-way variance analysis and multivariate analyses of principal components analysis (PCA) and hierarchical clustering using Ward’s method. The study was performed in a uniform greenhouse experiment. A high variation was found among and within populations in all the measured characteristics. Of the total variation, 13.0% was distributed among species, 27.9% among morphotypes, 8.1% among populations, and 51.0% within populations. Because plants grew in a uniform environment, these results indicate that the differences observed among and within species, morphotypes, and populations have a genetic basis. Univariate and multivariate analyses clearly differentiated morphotypes, suggesting that the category of morphotypes must be used to nominate the infraspecific variation in the domesticated Capsicum. The principal components analysis identified a total of 15 principal components that contributed to explain the total variation. The first two components explained 59.64% of the total variation and seven components explained more than 90% of the total variation. Among the measured characteristics, number of seed per fruit, weight of the fruit, width of the fruit, length of the fruit, stem diameter, days to flowering, and height of the plant contributed to component 1 variation, whereas width of the leaf, length of the leaf, and number of locules, number of fruits and number of seeds per plant contributed to component 2. The hierarchical clustering separated the populations and the morphotypes in two large different groups. One group consisted of populations collected in monoculture conditions and the other group corresponded to population collected from backyard-garden conditions. The monoculture populations were characterized mainly by their longer, wider, and higher weight fruits, plants were of less height, had smaller stem diameters, and lower number of fruits than the populations collected from the backyard-garden conditions. The backyard-garden populations of the Capsicum annuum and Capsicum frutescens species, considered wild or semi-domesticated, constituted a non-differentiated phenotypic group that does not allow dividing them in different species.     

Effect of ascochyta blight (Mycosphaerella pinodes) on yield components of single pea (Pisum sativum) plants under field conditions

Field studies were made in 1992 and 1993 to examine the yield components of pea inoculated with Mycosphaerella pinodes and those of healthy pea (sprayed with a mixture of flutriafol + chlorothalonil), in a split-plot design with the cv. Solara sown at different plant densities. Ascochyta blight was severe on leaves and on internodes of the basal part of the plants; pods had few lesions. The number and length of stems per plant were the same for diseased and healthy plants. The number of reproductive nodes and pods per stem were affected by disease only in 1993. In 1992 and 1993 respectively, disease caused reductions in the number of seeds per stem of 18% and 25%, and in seed size of 13.5% and 16.7%, compared with healthy plants. The harvest index and total biomass were lower in diseased than in healthy plants and seed yield was reduced by 40% in diseased plots. These results show a high relationship between the disease parameters (disease mean on stipules/nodes 8–18/ and on internodes/nodes 5–15/, percentages of stipules or internodes with a disease score 4, and percentage of stems encircled by lesions), plant density and yield reduction.     

Phenotypic plasticity of Vaccinium meridionale (Ericaceae) in wild populations of mountain forests in Colombia

Vaccinium meridionale is a promising crop for the Andean region of South America and is currently available only in the wild. Spontaneous populations of this plant are found across the Colombian mountains, but very few published records on this plant morphology are available. A zonification study of V. meridionale was conducted in four principal areas of a low mountain forest of Colombia (Provinces of Boyacá, Cundinamarca, Santander and Nari?o) in 2007. A total of 20 populations and 100 plants of V. meridionale were individually characterized and surveyed, using a list of 26 characters of morphological variables (9 quantitative and 17 qualitative characters). Our results indicated that natural populations of V. meridionale might be found in the tropical forest under a highly heterogeneous climate and microclimate conditions, at different mountain regions between 2 357 and 3 168masl. The shrubs of V. meridionale exhibited a high level of intra-population variation in several quantitative (plant height, stem diameter) and qualitative (growth habit, ramification density, presence of anthocyanins in stems) morphological characters, suggesting an environmentally induced phenotypic plasticity. Plant height, stem diameter and foliar density were the most variable morphological traits, with coefficients of variation higher than 50%. However, several quantitative characters of its reproductive potential, such as berry dimensions, rachis length and number of flowers per inflorescence, resulted with low plasticity with coefficients of variation lower than 30.2%, indicating that these characters were genetically determined. The highest correlation coefficients (p < 0.05) resulted to be between fruit length and fruit width (0.90), leaf length and leaf width (0.78), plant height and stem diameter (0.60), and inflorescence length and flowers number per inflorescence (0.57). The results suggest that an important genetic resource exists for this species in the wild. Low variation in fruit size, which constitutes a target trait for plant breeders, could be useful for selection of cultivars of V. meridionale. The results of this study could also be applied in conservation programs aimed to protect these diverse populations in the mountain forests of Colombia.     

三峡水库长期水淹条件下耐淹植物甜根子草的资源分配特征

为研究历经三峡水库长期水淹的驯化后,不同海拔高程之间相同种源的甜根子草的生物量分配特征是否发生改变,进而探讨该物种对水淹胁迫表现出的适应性进化特征,试验于2008年初选育相同种源的甜根子草同龄幼苗栽植于三峡水库消落区甜根子草种植试验示范区,并考察了2012年、2013年不同海拔高程甜根子草植株的形态和生物量特征。试验共设置3个海拔高程,即水淹高程168、172m和不受水淹对照高程176m。试验结果表明:(1)较低高程的甜根子草植株较矮小细弱,168m高程的甜根子草植株主茎长和主茎基径显著低于对照176m高程(P0.05);平均节间长度随高程的降低而缩短;与之相反,主茎长/主茎基径随高程的降低而增大。(2)甜根子草的叶片厚度、叶片长/叶片宽、叶片长/叶鞘长均随海拔高程的降低而减小;与之相反,比叶面积随高程的降低而增大。(3)水淹前,甜根子草近端成熟节间的质量密度随高程的降低而增大;水淹后,其地上存活茎段基部成熟节间的质量密度在各高程之间无显著差异(P0.05)。以上研究结果表明,甜根子草历经三峡水库长期水淹的驯化后,生物量分配特征在不同海拔高程之间发生了改变,表现出了相应的驯化特征。相较于高高程的甜根子草植株而言,低高程的植株生长缓慢,采取低株高下的高向生物量投资策略;对叶的物质投资大部分分配到叶面积的增加、叶鞘的伸长生长和叶片的直立生长上,以加强植株的光合生产。     

Rapid genetic and morphologic divergence between captive and wild populations of the endangered Leon Springs pupfish,Cyprinodon bovinus

The Leon Springs pupfish (Cyprinodon bovinus) is an endangered species currently restricted to a single desert spring and a separate captive habitat in southwestern North America. Following establishment of the captive population from wild stock in 1976, the wild population has undergone natural population size fluctuations, intentional culling to purge genetic contamination from an invasive congener (Cyprinodon variegatus) and augmentation/replacement of wild fish from the captive stock. A severe population decline following the most recent introduction of captive fish prompted us to examine whether the captive and wild populations have differentiated during the short time they have been isolated from one another. If so, the development of divergent genetic and/or morphologic traits between populations could contribute to a diminished ability of fish from one location to thrive in the other. Examination of genomewide single nucleotide polymorphisms and morphologic variation revealed no evidence of residual C. variegatus characteristics in contemporary C. bovinus samples. However, significant genetic and morphologic differentiation was detected between the wild and captive populations, some of which might reflect local adaptation. Our results indicate that genetic and physical characteristics can diverge rapidly between isolated subdivisions of managed populations, potentially compromising the value of captive stock for future supplementation efforts. In the case of C. bovinus, our findings underscore the need to periodically inoculate the captive population with wild genetic material to help mitigate genetic, and potentially morphologic, divergence between them and also highlight the utility of parallel morphologic and genomic evaluation to inform conservation management planning.     

Vertical and radial profiles in tracheid characteristics along the trunk of Douglas-fir trees with implications for water transport

The main stems of three young Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirbel) Franco) trees were dissected to obtain samples of secondary xylem from internodes axially along the trunk and radially within each internode. From these samples, measurements were obtained of tracheid diameter, length, the number of inter-tracheid pits per tracheid, and the diameter of the pit membranes. In addition, samples were obtained along the trunks of three old growth trees and also a small sample of roots for measurement of tracheid diameter. A gradient was apparent in all measured anatomical characters vertically along a sequence among the outer growth rings. These gradients arose not because of a gradient vertically along the internodes, but because of the strong gradients present at each internode among growth rings out from the pith. Tracheid characteristics were correlated: wider and longer tracheids had more numerous pits and wider pits, such that total pit area was about 6% of tracheid wall area independent of tracheid size. A stem model combining growth rings in parallel and internodes in series allowed for estimates of whole trunk conductance as a function of tree age. Conductance of the stem (xylem area specific conductivity) declined during the early growth of the trees, but appeared to approach a stable value as the trees aged.     

<Emphasis Type="Italic">In vitro</Emphasis> stem elongation of stemless carline thistle

In vitro culture of stemless carline thistle was established using immature zygotic embryos. A satisfactory bud multiplication was achieved on MS medium supplemented with BAP (1 mg L^–1) and IAA (0.2 mg L^–1). Maximum rooting of buds was induced upon short cultivation (4 or 8 days) on an auxin-supplemented medium. Highest number of roots was obtained with NAA in the medium while the longest roots developed on the IAA-supplemented medium. Plantlets that subsequently developed were in rosette form if grown in light (16/8 h light to darkness photoperiod) and with elongated stems if raised in darkness. Light grown plantlets treated with GA₃ showed dose dependent stem increase in length, reaching maximum at the concentration of 10^–4 M. This was correlated with the length and the average number of internodes. If cultivated in the presence of ancymidol, dark grown plantlets showed reduced stem length. However, the inhibitory effect of the growth retardant on stem elongation was completely overcome by the addition of GA₃.     

Differences in mate acceptance and host plant recognition between wild and laboratory-reared Busseola fusca (Fuller)

The present study was aimed at characterizing differences in mate acceptance and host plant recognition between Busseola fusca (Lep.: Noctuidae) reared for several generations under laboratory conditions and wild conspecifics, directly collected from maize stems in the field. The mating success was significantly higher in laboratory reared when compared with the wild B. fusca population. Oviposition on artificial stems was significantly higher for laboratory‐reared insects than for the wild ones. Moreover, unlike adults of the wild strain, laboratory‐reared B. fusca showed no preference to oviposit on surrogate stems impregnated with maize extracts. Long‐range attraction to the host plant was significantly lower for laboratory‐reared insects. Furthermore, the polymerase chain reaction‐restriction fragment length polymorphism (PCR‐RFLP) analysis of mitochondrial DNA showed that the wild type and laboratory populations belonged to the same genetic strain, indicating that these differences between the populations were mainly phenotypic.     

Multivariate analysis of motherwort germplasm in Iran using morphological variables and essential oil content

The morphological variation and essential oil content of 150 individuals of motherwort (Leonurus cardiaca) from six natural regions of Iran were examined biometrically based on multivariate analysis. The highest variation was observed in plant height, length of main stems, main inflorescence length, floral cycles of main stem, lateral branches length, basal leaf length, floral leaf length width, flower length, and calyx color. Results of simple correlation analysis showed the existence of significant positive and negative correlations among some important parameters. The highest correlation was observed between plant height and length of main stems and between lateral branch length and lower surface color of leaf. Populations were clustered in two groups representing subspecies cardiaca and persicus. The subspecies cardiaca populations were closely related with each other and differentiated from the subspecies persicus by pubescence stems and leaves, more number of floral cycles, higher compaction of floral cycles, lower number of lateral branches, shorter length of petioles and leaves and smaller floral leaves. Essential oil yields varied from 0.02 (for Taleghan population) to 0.053 mg/100 g dry mater (for Kerman population). The conservation of the highly diverse native populations of Iranian motherwort germplasm is recommended.     

Population structure and genetic diversity of wild <Emphasis Type="Italic">Helianthus</Emphasis> species from Mozambique

The production of sunflower suffered a major decline in Mozambique after its independence in 1975. Civil war, human activities and environmental damage subjected the species to an ecological stress contributing to reduce the number and size of wild populations. As this reduction is often related to a loss of genetic variation we estimated the genetic diversity within and among populations of wild Helianthus from five districts of Mozambique using RAPD markers. The 44 accessions studied grouped into four major clusters exhibiting structured variability with regard to geographic origin. A high level of genetic diversity (He = 0.350 and I = 0.527) was retained at the population level. The genetic variation among populations was high (59.7%), which is consistent with low gene flow (Nm = 0.338). The proportion of total genetic diversity residing among these populations should be kept in mind to devise different conservation strategies in order to preserve these populations. Currently wild Helianthus genetic resources present in Maputo and Sofala are on the edge of extinction mainly due to excessive urbanization. Therefore, conservation of what remains of this plant genetic diversity is essential for sustainable utilization and can be useful for breeding programs.     

Geographic variation in primary sex allocation per flower within and among 12 species of Pedicularis (Orobanchaceae): Proportional male investment increases with elevation

? Premise of the study: The study of geographic variation in ecologically important traits within and among taxa is a first step toward understanding the environmental factors that contribute to population differentiation and species divergence. This study examines variation in mean sex allocation per flower (androecium mass/gynoecium mass) among 49 wild populations representing 12 Pedicularis species across an elevation gradient on the eastern Tibetan Plateau. ? Methods: We used population means to evaluate sources of variation in per-flower sex allocation within and across species. In particular, we evaluate the relative influence of intrinsic (i.e., plant size, estimated as aboveground stem biomass) vs. extrinsic factors affecting mean sex allocation among populations. ? Key results: Mean sex allocation per flower (the relative investment in male floral organs) is negatively correlated with mean plant size; populations of large plants produce relatively female-biased flowers. This relationship between mean plant size and mean sex allocation is not statistically significant, however, when the effect of elevation is controlled statistically. Among populations within and across species, mean sex allocation increases with elevation. This relationship persists even when the effect of mean plant size is controlled statistically. Factors associated with increasing elevation appear to favor genotypes and/or taxa with male-biased flowers. ? Conclusion: Extrinsic environmental conditions may be more important than intrinsic resource status in determining patterns of geographic variation in mean sex allocation among populations or species of Pedicularis. We cannot conclude whether the effect of elevation on mean sex allocation is the result of environmentally induced plasticity, genetically based adaptation, or species sorting, but it is only partly mediated by mean plant size.     

Adaptive variation in physiological traits underpinning stem elongation responses among nodally-rooting stoloniferous herbs

For nodally-rooting stoloniferous herbs a close physiological link exists between the growth of nodal roots and development of shoot tissues of phytomers distal to them. Furthermore, in Trifolium, presence of nodal roots at phytomers proximal to the youngest nodal root on a stem influences the allocation of carbon along the stem when that stem encounters a patch preventing further formation of nodal roots. In this study six further phylogenetically and morphologically diverse stoloniferous species, selected so that they varied along the phalanx–guerrilla continuum (Acaena (Rosaceae), Leptinella (Asteraceae), Mentha (Lamiaceae), Vinca (Apocynaceae), Lamiastrum (Lamiaceae) and Glechoma (Lamiaceae)), were used to test the null hypothesis: that in all stoloniferous species the proportional allocation of carbon to the youngest nodal root decreases if proximally located roots are present when their stems extend into an environment preventing further nodal rooting. As Glechoma and Lamiastrum failed to change their proportional allocation to the youngest nodal root whilst the other five species varied their proportional allocation, with the species with shortest internodes (Acaena and Leptinella) changing the most, we reject this hypothesis. In addition, upon the transition to a dry environment, internode lengths in species with longer internodes (Glechoma and Lamiastrum) were markedly increased (doubled) whilst those in species with shorter internodes showed no response. We suggest this response is driven by apical bud tissues sensing changes in the humidity of their micro-environment and then modifying the length of internodes developing proximal to them. The extent of both these physiological responses correlated linearly with internode length of species (or position along the phalanx–guerrilla continuum), suggesting that these three traits were co-evolving, possibly in response to variation in grain of environmental heterogeneity, in such a way that the trade-off between the explorative and exploitive components of growth was underpinned and the performance of species in particular niches optimised as a result.     

Phenotypic Characterization of <i>Oryza nivara</i> (Sharma <i>et</i> Shastry) Collected from Different Ecological Niches of Sri Lanka

Information on the genetic diversity of wild rice species in Sri Lanka is relatively meagre, though it plays a key role in crop improvement programs of cultivated rice (Oryza sativa L.). The present study was carried out to identify the morphological variation pattern of the wild populations of O. nivara in Sri Lanka. Seven populations (P1 to P7) collected from different agro-ecological regions were characterized in a common garden based on nine morphological traits. The findings revealed a high level of phenotypic variation between populations when compared to within a population. The most variable traits were the flag leaf panicle neck length (FLPNL) and flag leaf angle (FLA), whereas the least variable trait was the flag leaf length (FLL). Box plots clearly illustrated the large differentiation of phenotypic traits in the entire distribution of wild rice populations. The cumulative values of the two principal components, i.e., FLPNL and FLA, explained 58.7% of the total variance. Populations from similar natural habitats clustered together. The P7 was adapted to intercept more sunlight by increasing flag leaf width (FLW) and FLA to compete with weeds and other shrubs. P2 and P5 were the most closely related populations representing approximately similar ecological conditions of the dry zone. The P3 population from the intermediate zone showed a vigorous plant growth with the highest plant height, culm girth and awn length (P < 0.05). Knowledge of such morphological diversity would facilitate designing conservation strategies and basic information for the proper utilization of wild resources in rice genetic improvement.     

Genome‐wide single nucleotide polymorphisms reveal population history and adaptive divergence in wild guppies

Adaptation of guppies (Poecilia reticulata) to contrasting upland and lowland habitats has been extensively studied with respect to behaviour, morphology and life history traits. Yet population history has not been studied at the whole‐genome level. Although single nucleotide polymorphisms (SNPs) are the most abundant form of variation in many genomes and consequently very informative for a genome‐wide picture of standing natural variation in populations, genome‐wide SNP data are rarely available for wild vertebrates. Here we use genetically mapped SNP markers to comprehensively survey genetic variation within and among naturally occurring guppy populations from a wide geographic range in Trinidad and Venezuela. Results from three different clustering methods, Neighbor‐net, principal component analysis (PCA) and Bayesian analysis show that the population substructure agrees with geographic separation and largely with previously hypothesized patterns of historical colonization. Within major drainages (Caroni, Oropouche and Northern), populations are genetically similar, but those in different geographic regions are highly divergent from one another, with some indications of ancient shared polymorphisms. Clear genomic signatures of a previous introduction experiment were seen, and we detected additional potential admixture events. Headwater populations were significantly less heterozygous than downstream populations. Pairwise F_ST values revealed marked differences in allele frequencies among populations from different regions, and also among populations within the same region. F_ST outlier methods indicated some regions of the genome as being under directional selection. Overall, this study demonstrates the power of a genome‐wide SNP data set to inform for studies on natural variation, adaptation and evolution of wild populations     

Genetic diversity of Ustilago maydis strains

Thirty wild isolates belonging to five different locations in Mexico plus two laboratory strains of Ustilago maydis were characterized by restriction fragment length polymorphism (RFLP) analysis using 23 different clones as probes derived from a PstI library and two restriction enzymes. All loci analysed presented a high level of polymorphism, including one locus with thirty one different alleles. Geographical grouping of the populations was based on Nei's genetic distance and there was no correlation between genetic and geographic distances among these isolates. Our results suggest that DNA fingerprinting is a useful method for detecting genetic variation in populations of U. maydis. This work demonstrated that considerable genetic variation may be present within field populations of U. maydis.     

A study of the diversity and geographical variation in numbers of leg‐bearing segments in centipedes (Chilopoda: Geophilomorpha) in north‐western Europe

Five species of geophilomorphs, Strigamia maritima (Leach, 1817), Geophilus flavus (De Geer, 1778), Geophilus truncorum Bergsøe and Meinert, 1866, Geophilus proximus C. L. Koch, 1847, and Pachymerium ferrugineum (C. L. Koch, 1835), from various sample sites in north‐western Europe were examined for numbers of leg‐bearing segments. Where data was adequate, mean segment numbers were correlated with latitudinal gradients for each species. Solely in S. maritima and some populations of P. ferrugineum did the results show a highly significant geographic pattern towards more leg‐bearing segments in southern populations of both sexes. As concerns G. proximus, we presented for the first time a remarkable geographic shift towards an increased number of pairs of legs in northern populations. Contrary to the conventional geographic pattern, we found that G. flavus and G. truncorum did not exhibit a north–south increase or decrease in segment number. Although there was no general/universal evidence supporting the occurrence of a significant regression slope between mean segment number and latitude/temperature, more information shows that the overall region‐to‐region segment variation was highly significant in both sexes. In S. maritima and P. ferrugineum mean segment number was correlated with the north–south temperature cline. The same was not observed in G. proximus. Parthenogenesis in G. proximus, and a series of ecological characteristics such as habitat preferences, spatial distribution, and fragmented populations could explain the presence or absence of a geographic patterning of segment number variation along a latitudinal cline. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 899–909.