Components of variation in seedling potential relative growth rate: phylogenetically independent contrasts

Variation between species in seedling potential relative growth rate (RGR) is among the most important spectra of plant adaptation. Investigations are reported into the components responsible for this variation, using phylogenetically independent contrasts (PICs). The two species for each PIC were selected to diverge in seed mass at least four-fold, seed mass being a known correlate of RGR. Consistent with previous reports, the main influence on RGR differences between species was leaf area per unit leaf mass (SLA), rather than net assimilation rate per leaf area (NAR_a). The PIC design showed that SLA differences both underpinned old RGR divergences between orders and families, and also were repeatedly responsible for more recent RGR divergences between genera and species.     

Polytomies and phylogenetically independent contrasts: examination of the bounded degrees of freedom approach

We examined the effect of soft polytomies on the performance (Type I error rate and bias) of Felsenstein's (1985; Am. Nat. 125:1-15) method of phylogenetically independent contrasts for estimating a bivariate correlation. We specifically tested the adequacy of bounding degrees of freedom, as suggested by Purvis and Garland (1993; Syst. Biol. 42:569-575). We simulated bivariate character evolution under Brownian motion (assumed by independent contrasts) and eight other models on five phylogenetic trees. For non-Brownian motion simulations, the adequacy of branch-length standardization was checked with a simple diagnostic (Garland et al., 1992; Syst. Biol. 41:18-32), and transformations were applied as indicated. Surprisingly, soft polytomies tended to have negligible effects on Type I error rates when models other than Brownian motion were used. Overall, and irrespective of evolutionary model, degrees of freedom were appropriately bounded for hypothesis testing, and unbiased estimates of the correlation coefficient were obtained. Our results, along with those of previous simulation studies, suggest that independent contrasts can reliably be applied to real data, even with phylogenetic uncertainty.     

Modelling extinction risk in multispecies data sets: phylogenetically independent contrasts versus decision trees

Many recent studies of extinction risk have attempted to determine what differences exist between threatened and non-threatened species. One potential problem in such studies is that species-level data may contain phylogenetic non-independence. However, the use of phylogenetic comparative methods (PCM) to account for non-independence remains controversial, and some recent studies of extinction have recommended other methods that do not account for phylogenetic non-independence, notably decision trees (DTs). Here we perform a systematic comparison of techniques, comparing the performance of PCM regression models with corresponding non-phylogenetic regressions and DTs over different clades and response variables. We found that predictions were broadly consistent among techniques, but that predictive precision varied across techniques with PCM regression and DTs performing best. Additionally, despite their inability to account for phylogenetic non-independence, DTs were useful in highlighting interaction terms for inclusion in the PCM regression models. We discuss the implications of these findings for future comparative studies of extinction risk.     

High seedling relative growth rate and specific leaf area are traits of invasive species: phylogenetically independent contrasts of woody angiosperms

Understanding causal factors of exotic species invasions is important not only for prevention and prioritizing control efforts, but also for providing valuable insights into the underlying biology of contrasting life-history strategies. In seedling growth analyses, invasive woody species were compared with less-invasive woody species commonly cultivated in California using phylogenetically corrected procedures (12 phylogenetically independent contrasts). Invasive species were hypothesized to have higher seedling relative growth rates (RGRs) and specific leaf areas (SLAs) than did related less-invasive species. In phylogenetically independent contrasts conducted among taxa within families, high seedling RGRs and SLAs have significant positive associations with woody plant invasiveness. For contrasts containing species invasive in mediterranean regions, invasive species had significantly larger root biomass allocation than did less-invasive species. Optimization of fast seedling growth (high RGR) associated with opportunistic resource acquisition (high SLA) and increased root allocation to survive summer drought may be critical for the success of plant invaders in regions with mediterranean climates.     

Increased seed dispersal potential towards geographic range limits in a Pacific coast dune plant

Dispersal may be favoured at geographic range edges by unstable population and metapopulation dynamics. However, dispersal may also evolve in response to geographic variation in other life-history traits, especially the mating system. Here, increased dispersal at range margins was tested for with a range-wide analysis of seed dispersal and mating system traits in Abronia umbellata, a plant endemic to Pacific coastal dunes of North America. Seeds disperse within winged anthocarps. Anthocarps from 34 populations varied widely in wing size (mass-corrected wing index). Wing index correlated negatively with threshold wind velocity for dispersal in wind tunnel tests, suggesting that wings facilitate tumbling over open sandy substrate. As predicted, wing index increased and threshold velocity decreased towards both range limits. Flower size, herkogamy and self-incompatibility declined towards range limits, indicating a shift to self-fertilization, and flower size and wing index correlated negatively. However, the increase in wing index towards range limits remained after statistically controlling flower size. These results are consistent with selection favouring dispersal at range margins. The evolutionary lability of dispersal across the range may affect the interaction between selection and gene flow in the establishment and maintenance of geographic range limits.     

Seedling root anatomy and morphology: an examination of ecological differentiation with rainfall using phylogenetically independent contrasts

We examined patterns of seedling root architecture, morphology and anatomy in Australian perennial plants chosen as phylogenetically independent contrasts (PICs) for rainfall in the areas they inhabit. Our objective was to assess whether there are consistent evolutionary patterns in structure of seedling root systems in species from different rainfall environments when examined across multiple evolutionary lineages. Seedlings were grown to a standardised developmental stage under controlled conditions. We found that seedling root systems of species restricted to low rainfall environments are characterised by greater proportional allocation to main root axis and have proportionally smaller main root axis diameter and areas of stele and xylem. Species of low rainfall environments also had higher specific root length (SRL) of the main axis, but lower SRL when the entire root system was considered. Seedling root system elongation rates were higher in species of high rainfall relative to those of low rainfall environments, paralleling expected differences in relative growth rate. The higher root system elongation rates in species of high rainfall environments were associated with greater numbers of growing tips in the root system, but not with differences in elongation rates of individual tips, relative to species of low rainfall environments.     

Body size establishes the scaling of avian postnatal metabolic rate: an interspecific analysis using phylogenetically independent contrasts

The avian postnatal metabolic rate literature is reviewed using power equations, Y = aM^b, to describe the relation between postnatal resting metabolic rate (RMR) and chick body mass (M) for 25 species. In altricial species, the relation between RMR and M from hatching to fledging can be described by a single power equation, whereas in most nonaltricial species two such equations are needed, one for chicks weighing less than about 25% of mature mass ( M _a) and a second for larger chicks. For altricial chicks and larger nonaltricial chicks, the body-mass exponent, b, of 25 intraspecific power equations ranged from 0.25 to 1.67 and varied inversely with M _a. The scaling of postnatal RMR is thus unlike that of either adult or hatchling metabolism in that it is size dependent. We examined the relationship between intraspecific b and M _a using Felsenstein's independent contrasts method to control for statistical complications due to the hierarchical nature of phylogenetic relationships. This "phylogenetic regression" technique yielded the relation b = 1.6 M _a^-015, in which mature mass explained 38% of the variation in b. The mass exponent of this equation (-0.15) did not differ significantly from that determined by nonphylogenetic methods (-0.17).
In altricial chicks and larger nonaltricial chicks, the scaling coefficient, a, of the interspecific power equations varied with adult mass according to the phylogenetically determined relation a (kj/h) = 0.0052M_a^0.65and was higher in fed than in fasted chicks. Equations derived in this analysis permit one to estimate the RMR of a growing chick from its mass and adult body mass and provide a basis for evolutionary and ecological comparisons.     

Cryptostylis species (Orchidaceae) from a broad geographic and habitat range associate with a phylogenetically narrow lineage of Tulasnellaceae fungi

While many Australian terrestrial orchids have highly specialized mycorrhizal associations, we tested the hypothesis that the geographically widespread orchid genus Cryptostylis associates with a diversity of fungal species. Using fungal isolation and molecular approaches, we investigated the mycorrhizal associations of five Australian Cryptostylis species (27 sites sampled) and included limited sampling from three Asiatic Cryptostylis species (two sites). Like related orchid genera, Tulasnellaceae formed the main fungal associations of the Cryptostylis species we sampled, although some ectomycorrhizal, ericoid and saprotrophic fungi were detected infrequently. Each species of Australian Cryptostylis associated with three to seven Tulasnella Operational Taxonomic Units (OTUs), except for C. hunteriana where only one Tulasnella OTU was detected. In total, eleven Tulasnella OTUs associated with Australian Cryptostylis. The Asiatic Cryptostylis associated with four different Tulasnella OTUs belonging to the same lineage as the Australian species. While five Tulasnella OTUs (T. australiensis, T. prima, T. warcupii, T. densa, and T. punctata) were used by multiple species of Australian Cryptostylis, the most commonly used OTU differed between orchid species. The association with different Tulasnella fungi by Cryptostylis species co-occurring at the same site suggests that in any given environmental condition, Cryptostylis species may intrinsically favour different fungal OTUs.     

Macaranga ant-plants hide food from intruders: correlation of food presentation and presence of wax barriers analysed using phylogenetically independent contrasts

Many tropical ant-plants provide specialized ant partners with food, which may attract foreign ants parasitizing the mutualism. We present evidence for the ant-plant genus Macaranga , showing that ant competition has forced host plants to hide food resources and restrict access to the mutualists. In Macaranga myrmecophytes, the influence of ant competition strongly depends on the presence of slippery 'wax barriers'. Of all Macaranga ant-plant species, 50% have waxy stems that can be climbed only by the specific ant partners and not by other ant species. We compared the presentation of food (food bodies and extrafloral nectar) between waxy and non-waxy Macaranga host plants using traditional and phylogenetic comparative methods. Consistent with the hypothesized effect of ant competition, wax-free Macaranga host species had fewer extrafloral nectaries and more often produced food bodies under recurved or tubular stipules inaccessible to other ants; closed stipules were less persistent in waxy hosts. Several traits showed phylogenetic signal, but our finding of a more promiscuous food presentation in waxy Macaranga hosts was still supported by phylogenetic comparative analyses. We conclude that competition among ants is an important factor in the evolution of myrmecophytism, and that it has given rise to traits acting as protective filter mechanisms.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 177–193.     

Wind dispersal of alpine plant species: A comparison with lowland species

Question: The prominent role of wind dispersal in alpine habitats has been recognized early but has rarely been quantified. The aim of this study is to compare wind dispersal under alpine and lowland conditions and to analyse whether differences are caused by species traits, e.g. terminal velocity of seeds (V_term) or weather conditions. Location and Methods: We characterized wind dispersal potential of > 1100 Central European species using measured V_term To quantify the habitat effect on wind dispersal, we measured meteorological key‐parameters and simulated dispersal distance spectra of nine selected species under typical alpine conditions (foreland of the Scaletta‐glacier, Switzerland) and typical lowland conditions (grassland in Bad Lippspringe, Germany). Results: Lowland species had higher V_term compared to alpine species. However, this difference is absent when only species of species of open habitats are concerned. The meteorological measurements showed that the alpine habitat was mainly characterized by higher frequency and strength of updrafts. The simulations showed that under alpine conditions long distance dispersal occurred much more frequent. Conclusions: More than 50 % of the alpine species have a fair chance to be dispersed by wind over long distances, while this proportion is less than 25 % for species from open habitats in the lowland. The more prominent role of wind dispersal in alpine habitats is mainly a result of differences in environmental conditions, namely more intense vertical turbulence in the alpine habitat, and does not result from prominent differences in plant traits, namely V_term, between alpine and lowland species.     

Primate species separation in relation to secondary plant chemicals

Food selection of different primate species is reviewed in relation to secondary plant chemicals such as alkaloids and tannins. Different species discriminate differently against these chemicals. The tolerance of potentially toxic substances (e.g. alkaloids) is not restricted to species with enlarged foreguts but is also found among lemurs with rather unspecialized digestive systems. On the community level different reactions towards food chemicals seem to be an important mechanism contributing to species separation and thus allowing coexistence.     

Biologic interactions determining geographic range size: a one species response to phylogenetic community structure

Range size variation in closely related species suggests different responses to biotic and abiotic heterogeneity across large geographic regions. Species turnover generates a wide spectrum of species assemblages, resulting in different competition intensities among taxa, creating restrictions as important as environmental constraints. We chose to adopt the widely used phylogenetic relatedness (NRI) measurement to define a metric that depicts competition strength (via phylogenetic similarity), which one focal species confronts in its environment. This new approach (NRI_focal) measures the potential of the community structure effect over performance of a single species. We chose two ecologically similar Peucaea sparrows, which co‐occur and have highly dissimilar range size to test whether the population response to competition intensity is different between species. We analyzed the correlation between both Peucaea species population sizes and NRI_focal using data from point counts. Results indicated that the widespread species population size was not associated with NRI_focal, whereas the population of restricted‐sized species exhibited a negative relationship with competition intensity. Consequently, a species' sensitivity to competition might be a limiting factor to range expansion, which provides new insights into geographic range analysis and community ecology.     

Distribution of plant species and vegetation types in relation to climate

The distribution of plant species and vegetation types in the Prairie provinces of Canada can be related to climatic phenomena. The ratio of summer/spring precipitation appears to be especially important, with temperatures of less importance. Climatic areas can be outlined; within these areas differences in soils are important.