Phylogenetic Correction Research Articles

IMPORTANCE OF SPECIES TRAITS FOR SPECIES DISTRIBUTION IN FRAGMENTED LANDSCAPES

Knowledge of the relationship between species traits and species distribution in fragmented landscapes is important for understanding current distribution patterns and as background information for predictive models of the effect of future landscape changes. The existing studies on the topic suffer from several drawbacks. First, they usually consider only traits related to dispersal ability and not growth. Furthermore, they do not apply phylogenetic corrections, and we thus do not know how considerations of phylogenetic relationships can alter the conclusions. Finally, they usually apply only one technique to calculate habitat isolation, and we do not know how other isolation measures would change the results. We studied the issues using 30 species forming congeneric pairs occurring in fragmented dry grasslands. We measured traits related to dispersal, survival, and growth in the species and recorded distribution of the species in 215 grassland fragments. We show many strong relationships between species traits related to both dispersal and growth and species distribution in the landscape, such as the positive relationship between habitat occupancy and anemochory and negative relationships between habitat occupancy and seed dormancy. The directions of these relationships, however, often change after application of phylogenetic correction. For example, more isolated habitats host species with smaller seeds. After phylogenetic correction, however, they turn out to host species with larger seeds. The conclusions also partly change depending on how we calculate habitat isolation. Specifically, habitat isolation calculated from occupied habitats only has the highest predictive power. This indicates slow dynamics of the species. All the results support the expectation that species traits have a high potential to explain patterns of species distribution in the landscape and that they can be used to build predictive models of species distribution. The specific conclusions are, however, dependent on the technique used, and we should carefully consider this when comparing among different studies. Since different techniques answer slightly different questions, we should attempt to use analyses both with and without phylogenetic correction and explore different isolation measures whenever possible and compare the results.

What youngsters say about adults: seedling roots reflect clonal traits of adult plants

1 Grime's theory on plant strategies predicts that the set of traits present in established plants is not correlated with that found in the regenerative stage of the life cycle. We tested this prediction and further investigated whether clonal growth traits, which are found in adult plants but also affect regeneration, are correlated with seedling traits. 2 We characterized seedling root systems by their total length, number of root tips and several architectural parameters (length of exterior and interior root links and two topological indices). These below-ground traits were supplemented by the ratio of leaf area to root length, representing relative investment into photosynthesizing surface. We compared seedling traits with clonal growth traits, adult plant heights, and species positions on gradients of nitrogen and water availability. 3 Plant species with limited horizontal spread by clonal growth exhibited a larger relative investment in photosynthetic area and also developed larger root systems as seedlings. 4 Seedlings of species with taller shoots and those which occur naturally at nutrient-rich sites developed both larger roots and more dichotomously branched root systems (with higher total length and more branches). 5 Taking phylogenetic inertia into account showed that this explained large parts of the variation in seedling traits. Relationships between clonal spread and seedling traits were strengthened by phylogenetic correction. 6 Our study shows that some of the traits of clonal growth affect both the established and the regenerative stages of the life cycle, suggesting that an evolutionary trade-off exists between the ability to spread clonally and performance at the seedling stage. Species not able to escape from less favourable conditions by extensive clonal spread seem to be more able to exploit the location in which they germinate.

Scaling of insect metabolic rate is inconsistent with the nutrient supply network model

Summary The nutrient supply network model of the metabolic theory of ecology predicts that metabolic rate scales as mass0·75 at all hierarchical levels. An alternative, cell size, model suggests that the scaling of metabolic rate is a by‐product of the way in which body size changes, by cell size or number, or some combination thereof. It predicts a scaling exponent of mass0·75 at the widest interspecific level, but values of mass0·67−1·0 for lower taxonomic groups or within species. Here these predictions are tested in insects using 391 species for the interspecific analysis, and the size‐polymorphic workers of eight ant species at the intraspecific level. In the latter, the contribution of ommatidium size and number to variation in body length, which is closely related to eye size, is used to assess the relative contributions of changes in cell size and number to variation in body size. Before controlling for phylogeny, metabolic rate scaled interspecifically as mass0·82. Following phylogenetic correction, metabolic rate scaled as mass0·75. By contrast, the intraspecific scaling exponents varied from 0·67 to 1·0. Moreover, in the species where metabolic rate scaled as mass1·0, cell size did not contribute significantly to models of body size variation, only cell number was significant. Where the scaling exponent was < 1·0, cell size played an increasingly important role in accounting for size variation. Data for one of the largest groups of organisms on earth are therefore inconsistent with the nutrient supply network model, but provide support for the cell size alternative.

Selective extinction of late Neogene bivalves on the temperate Pacific coast of South America

AbstractWe assessed selective extinction patterns in bivalves during a late Neogene mass extinction event observed along the temperate Pacific coast of South America. The analysis of 99 late Neogene and Quaternary fossil sites (recorded from 7°S to 55°S), yielding ∼2800 occurrences and 118 species, revealed an abrupt decline in Lyellian percentages during the late Neogene–Pleistocene, suggesting the existence of a mass extinction that decimated ∼66% of the original assemblage. Using the late Neogene data set (n= 59 species, 1346 occurrences), we tested whether the extinction was nonrandom according to taxonomic structure, life habit, geographic range, and body size. Our results showed that the number of higher taxa that went extinct was not different than expected by random. At first sight, extinction was selective only according to life habit and geographic range. Nevertheless, when phylogenetic effects were accounted for, body size also showed significant selectivity. In general, epifaunal, small-sized (after phylogenetic correction), and short-ranged species tended to have increased probability of extinction. This is verified by strong interactions between the variables herein analyzed, suggesting the existence of nonlinear effects on extinction chances. In the heavily decimated epifaunal forms, survival was not enhanced by widespread ranges or larger body sizes. Conversely, the widespread and large-sized infaunal forms tended to have lower probability of extinction. Overall, the ultimate extinction of late Neogene bivalve species along the Pacific coast of South America seems to have been determined by a complex interplay of ecological and historical (phylogenetic) effects.

Selective extinction of late Neogene bivalves on the temperate Pacific coast of South America

AbstractWe assessed selective extinction patterns in bivalves during a late Neogene mass extinction event observed along the temperate Pacific coast of South America. The analysis of 99 late Neogene and Quaternary fossil sites (recorded from 7°S to 55°S), yielding ∼2800 occurrences and 118 species, revealed an abrupt decline in Lyellian percentages during the late Neogene–Pleistocene, suggesting the existence of a mass extinction that decimated ∼66% of the original assemblage. Using the late Neogene data set (n = 59 species, 1346 occurrences), we tested whether the extinction was nonrandom according to taxonomic structure, life habit, geographic range, and body size. Our results showed that the number of higher taxa that went extinct was not different than expected by random. At first sight, extinction was selective only according to life habit and geographic range. Nevertheless, when phylogenetic effects were accounted for, body size also showed significant selectivity. In general, epifaunal, small-sized (after phylogenetic correction), and short-ranged species tended to have increased probability of extinction. This is verified by strong interactions between the variables herein analyzed, suggesting the existence of nonlinear effects on extinction chances. In the heavily decimated epifaunal forms, survival was not enhanced by widespread ranges or larger body sizes. Conversely, the widespread and large-sized infaunal forms tended to have lower probability of extinction. Overall, the ultimate extinction of late Neogene bivalve species along the Pacific coast of South America seems to have been determined by a complex interplay of ecological and historical (phylogenetic) effects.

Patterns of plant traits in annual vegetation of man-made habitats in central Europe

Man-made habitats in central Europe can be broadly divided into arable land with weed vegetation, and settlements and their surroundings, harbouring ruderal vegetation. The former is a predictable environment with frequent, regular and large-scale disturbances, while the latter is an unpredictable environment with irregular disturbances of varying spatial extent producing heterogeneous mosaics of different successional stages. We hypothesize that these differences in disturbance regimes select for different sets of biological and ecological plant traits in these two habitats. A data set of 2715 vegetation plots sampled in man-made habitats dominated by annual plants in the Czech Republic was combined with data on biological and ecological traits of vascular plants, mostly taken from the BiolFlor database. Differences due to temporal variation and location of plots in different climatic zones were partialled out using partial canonical correspondence analysis. Then the differences in traits of the plants growing on arable fields and in settlements were analysed using logistic and least-square regression models, both with and without phylogenetic correction. Plants growing on arable land were more often annuals, R-strategists, with overwintering green leaves, insect or self-pollinated, reproducing by seeds, with persistent seed banks and archaeophytes (i.e. those aliens that arrived prior to 1500). Plants growing in human settlements were more often biennials or perennials, C-strategists, wind-pollinated, flowering in mid summer, reproducing both by seeds and vegetatively, dispersed by wind or humans, neophytes (i.e. those aliens that arrived after 1500), species with high demands for light and nutrients and with more continental distribution ranges. Most associations between plant traits and habitats did not change after taking phylogenetic relationships into account. Traits strongly linked to phylogeny were especially modes of pollination and dispersal. By contrast, traits weakly linked to phylogeny included life strategy and alien status.

A COMMENT ON PHYLOGENETIC CORRECTION

Abstract Statistical methods are now commonly used to take into account the expected lack of independence of observations across different species (due to their phylogenetic relatedness) when computing correlations or regressions among traits. The methods are often interpreted as removing that part of the regression or correlation that is an artifact due to phylogeny and there is an expectation that the corrected regression or correlation coefficients will usually be closer to zero. It is shown here that this is not an accurate way to interpret these methods. The effect of taking phylogeny into account is to reduce the variance of the estimated regression or correlation coefficients. Their means are not because since estimates of regression coefficients are unbiased whether or not the correct phylogeny is taken into account. Estimates of correlations are only slightly biased (and in the opposite direction that many expect).

A COMMENT ON PHYLOGENETIC CORRECTION

Statistical methods are now commonly used to take into account the expected lack of independence of observations across different species (due to their phylogenetic relatedness) when computing correlations or regressions among traits. The methods are often interpreted as removing that part of the regression or correlation that is an artifact due to phylogeny and there is an expectation that the corrected regression or correlation coefficients will usually be closer to zero. It is shown here that this is not an accurate way to interpret these methods. The effect of taking phylogeny into account is to reduce the variance of the estimated regression or correlation coefficients. Their means are not because since estimates of regression coefficients are unbiased whether or not the correct phylogeny is taken into account. Estimates of correlations are only slightly biased (and in the opposite direction that many expect).

Functional coordination between leaf gas exchange and vulnerability to xylem cavitation in temperate forest trees

We examined functional coordination among stem and root vulnerability to xylem cavitation, plant water transport characteristics and leaf traits in 14 co-occurring temperate tree species. Relationships were evaluated using both traditional cross-species correlations and phylogenetically independent contrast (PIC) correlations. For stems, the xylem tension at which 50% of hydraulic conductivity was lost (psi50) was positively associated (P < 0.001) with specific conductivity (K(S)) and with mean hydraulically weighted xylem conduit diameter (D(h-w)), but was only marginally (P = 0.06) associated with leaf specific conductivity (K(L)). The PIC correlation for each of these relationships, however, was not statistically significant. There was also no relationship between root psi50 and root K(S) in either cross-species or PIC analysis. Photosynthetic rate (A) and stomatal conductance (g(s)) were strongly and positively correlated with root psi50 in the cross-species analysis (P < 0.001), a relationship that was robust to phylogenetic correction (P < 0.01). A and g(s) were also positively correlated with stem psi50 in the cross-species analysis (P = 0.02 and 0.10, respectively). However, only A was associated with stem psi50 in the PIC analysis (P = 0.04). Although the relationship between vulnerability to cavitation and xylem conductivity traits within specific organs (i.e. stems and roots) was weak, the strong correlation between g(s) and root psi50 across species suggests that there is a trade-off between vulnerability to cavitation and water transport capacity at the whole-plant level. Our results were therefore consistent with the expectation of coordination between vulnerability to xylem cavitation and the regulation of stomatal conductance, and highlight the potential physiological and evolutionary significance of root hydraulic properties in controlling interspecific variation in leaf function.

Evolution of prey specialization in aphidophagous syrphids of the genera Melanostoma and Platycheirus (Diptera: Syrphidae) 1. Body size, development and prey traits

Interactions between syrphid predators and their prey are poorly known. The adaptations of syrphids to aphid defences and the consequences for the evolution of life history traits in these predators especially are mostly unstudied. This is the first of two papers investigating the evolution of prey specialization in aphidophagous hoverflies. The study focuses on two questions: (1) Are differences in the body size of syrphid predators reflected in differences in the size of their prey? (2) Are differences in body size, body mass and development time of the syrphid predators correlated with the defence strategies of their aphid prey (e.g. mobility, toxicity)? Platycheirus clypeatus (Meigen, 1822), Platycheirus fulviventris (Macquart, 1829), Melanostoma mellinum (Linnaeus, 1758), and Melanostoma scalare (Fabricius, 1794), which differ considerably in their prey specialization, but are closely related, were chosen as model species. Life history data for these syrphids came from a laboratory study, and that for the aphids from a literature survey. These syrphid species can be arranged on a gradient of increasing prey specialization, from 32 prey species for the generalist M. mellinum and only 3 for the specialist P. fulviventris. Differences in prey specialization were even more evident when the defence ability of the various species of aphid prey was considered. For instance, the specialization on ant-attended aphids in M. scalare. Larvae exhibited a one or two weeks diapause which made the determination of developmental time imprecise. Body size of the predators was not reflected in that of their aphid prey. The postulated relationship between body size of the predator and the defence strategies of their prey was not supported by our data. A comparison of a wider range of syrphid species from different taxonomic groupings together with a phylogenetic correction procedure might reveal clearer trends. The second part of this paper (Dziock, in prep.) will investigate the correlation between prey specialization and reproductive strategies (i.e. clutch size, egg size and number) and will put the results into a broader framework.

Profiling invasive fish species: the importance of phylogeny and human use

ABSTRACTUnderstanding the ecological differences between native and invasive species is of considerable scientific and practical interest. We examined such differences between native and invasive inland fish species from the Iberian Peninsula in order to analyse the importance of phylogenetic correction and variability (in addition to central tendency). We collected 26 quantitative and qualitative variables on the ecology, life‐history traits and human use of the 69 inland fish species of the Iberian Peninsula, including native, invasive and migratory species. The taxonomic distribution of invasive fish species deviated significantly from world freshwater richness and in contrast to native species, invasive fish belongs to only five taxonomic orders but to a wide spectrum of families not native to the Iberian Peninsula. Because the life‐history traits were highly dependent on taxonomy, the results, with or without applying phylogenetic methods, differed and after accounting for phylogeny, invasive species displayed higher and wider latitude in general and a different reproductive season mainly among salmonids and cyprinids. Human use was also significantly different between native and invasive fish species and produced more variability in life‐history traits of invasive species and uneven taxonomic distribution because of the high diversity of species introduced. We show that accounting for taxonomy and studying variability in addition to central tendency is important in the comparison of life‐history traits between native and invasive species.

WHERE HAVE ALL THE CLADES GONE? A SYSTEMATIST'S TAKE ON INFERRING PHYLOGENIES1

Joseph Felsenstein published four papers between 1978 and 1985 that enabled phylogenetic biology to take giant steps forward. In chronological order, Felsenstein showed that parsimony could be misleading (1978), derived maximum likelihood methods for DNA phylogenies (1981), constructed a method for establishing significance of trees (bootstrapping: 1985a), and invented a phylogenetic correction to the comparative method (independent contrasts: 1985b). With the publication of Inferring Phylogenies 20 years later, he reviews a monumental portion of the increasingly difficult literature on phylogenetic methods that he helped foster. His book provides insights into extraordinarily difficult problems that few biologists are trained to understand and exposes the reader to the bleeding edge of a discipline undergoing rapid change. His emphasis on the very latest theoretical results is tempered by a consistently scholarly, if occasionally controversial, account of the history of the field. It is not much use as a beginner's guide to phylogenetic inference-one can search in vain through its nearly 600 pages for the most basic of definitions-nor does it adequately treat a large literature on parsimony-based phylogenetics (the reader is better off with Kitching et al.'s [1998] book), but no one with a serious interest in the quantitative aspects of the subject will be able to live without it. How quantitative the field has become will be apparent to the reader quickly. Something has happened to phylogenetic theory in the last 10 years. Previously, a few biologists with real expertise in math, like Felsenstein, together with a very small number of applied mathematicians-professional theorists, let's call them-were joined by a larger cadre of enthusiastic amateurs in moving the field forward. Recently, a New Wave of mathematicians and computer scientists has begun to play a much more dominant role. If you think Inferring Phylogenies is mathematically daunting, pick up a copy of Semple and Steel's Phylogenetics (2003), or the recently published edited volume by Olivier Gascuel on Mathematics of Evolution and Phylogeny (2005). Felsenstein's book builds a bridge between biologists and this New Wave. If you don't happen to recall Laplace or Fourier transforms, appreciate the utility of a uniformly most powerful statistical test, or enjoy the beauty of a good generating function, a few chapters will contain stumbling blocks. However, more often than not, he offers what may be the last best hope for those with persistence and at least a passing familiarity with basic calculus and probability theory. As arcane as the Had-

Correlations between adult mimicry and larval host plants in ithomiine butterflies

The apparent paradox of multiple coexisting wing pattern mimicry 'rings' in tropical butterflies has been explained as a result of microhabitat partitioning in adults. However, very few studies have tested this hypothesis. In neotropical forests, ithomiine butterflies dominate and display the richest diversity of mimicry rings. We show that co-mimetic species occupy the same larval host-plant species significantly more often than expected in two out of five communities that we surveyed; in one of these, the effect remains significant after phylogenetic correction. This relationship is most probably a result of a third correlated variable, such as microhabitat. Host-plant microhabitat may constrain adult movement, or host-plant choice may depend on butterfly microhabitat preferences and mimicry associations. This link between mimicry and host plant could help explain some host-plant and mimicry shifts, which have been important in the radiation of this speciose tropical group.

Reproductive traits of tropical rain-forest trees in New Caledonia

Reproductive traits of 123 species of rain-forest tree from ultramafic regions of New Caledonia were assessed, mainly from herbarium specimens. Most species had extremely small, simple, pale-coloured flowers that are probably mainly pollinated by small insects, but not including bees. The seeds of most species were considered to be bird dispersed. However, wind is also important for pollination and seed dispersal. The phenological trend was for an increase in the number of species flowering and fruiting around the end of the warm dry season/start of the hot wet season, followed by a decline at the end of the wet season, and lower proportions during the cooler season. Seed size was significantly correlated with fruit size. Other correlations, between flower size and fruit size, and between seed size and seed number, were significant using species as independent observations, but did not hold following phylogenetic correction. Compared with non-dioecious species, dioecious species had significantly larger seeds, and a greater proportion of species with biotic dispersal, abiotic pollination and solitary (female) flowers. The long-term persistence of at least the larger-seeded tree species in New Caledonia is precarious, since the endemic giant pigeon, Ducula goliath, is probably their principal effective disperser, and this species is in decline.

Phylogenetic Analysis and Comparative Data: A Test and Review of Evidence

The question is often raised whether it is statistically necessary to control for phylogenetic associations in comparative studies. To investigate this question, we explore the use of a measure of phylogenetic correlation, lambda, introduced by Pagel (1999), that normally varies between 0 (phylogenetic independence) and 1 (species' traits covary in direct proportion to their shared evolutionary history). Simulations show lambda to be a statistically powerful index for measuring whether data exhibit phylogenetic dependence or not and whether it has low rates of Type I error. Moreover, lambda is robust to incomplete phylogenetic information, which demonstrates that even partial information on phylogeny will improve the accuracy of phylogenetic analyses. To assess whether traits generally show phylogenetic associations, we present a quantitative review of 26 published phylogenetic comparative data sets. The data sets include 103 traits and were chosen from the ecological literature in which debate about the need for phylogenetic correction has been most acute. Eighty-eight percent of data sets contained at least one character that displayed significant phylogenetic dependence, and 60% of characters overall (pooled across studies) showed significant evidence of phylogenetic association. In 16% of tests, phylogenetic correlation could be neither supported nor rejected. However, most of these equivocal results were found in small phylogenies and probably reflect a lack of power. We suggest that the parameter lambda be routinely estimated when analyzing comparative data, since it can also be used simultaneously to adjust the phylogenetic correction in a manner that is optimal for the data set, and we present an example of how this may be done.

Metabolic and thermal physiology of pigeons and doves.

Pigeons and doves (Columbidae) are an interesting group to examine for physiological adaptations to climate and diet because this cosmopolitan family comprises more than 300 species that are mostly granivores, although some are specialized frugivores. We determined allometric and phylogenetic effects on body temperature (T(b)), basal metabolic rate (BMR; J h(-1)), and wet thermal conductance (C(wet); J h(-1) C(-1)), and we examined mass (M) and phylogenetically corrected residuals for further effects of climate, diet, and landmass size (mainland or island). Independent contrasts, correlograms, autoregression, and phylogenetic eigenvector regression (PVR) were used to examine phylogenetically related effects. We found a small but significant phylogenetic pattern for body mass of columbids. For T(b), there was no significant effect of mass or phylogeny. There was a significant effect of climate on T(b) and no significant effects of diet or landmass without mass or phylogenetic correction, but after mass and phylogenetic correction, there were no effects of climate, diet, or landmass. For BMR, there was a strong allometric effect, and residuals were significantly lower for arid and tropical species but not for temperate species, compared to predictions for nonpasserine birds. There was a nearly significant autoregressive phylogenetic relationship for BMR parl0;r=0.44), and the strong allometry of BMR remained for independent contrasts (slope=0.731), autoregressive residuals (0.698), and PVR (0.705). Residuals, from regression of autoregression and PVR residuals of M and BMR, were significantly associated with climate: arid pigeons had a lower BMR residual than tropical and temperate pigeons. PVR residuals were significantly affected by landmass (island columbids had a smaller residual than mainland columbids), but autoregression residuals were not. There was no association of autoregression or PVR residuals with diet. For C(wet), there was a strong allometric effect, and residuals for columbids were significantly higher compared to other birds. There was no significant relationship for C(wet) of columbids to climate, diet, or landmass. There was no significant autoregressive or PVR relationship for C(wet), and the strong allometry remained after phylogenetic analysis by independent contrasts (slope=0.501), autoregression (0.509), and PVR (0.514). Residuals from autoregression and PVR were not significantly correlated with climate, diet, or landmass (mainland/island).

Rarity, specialization and extinction in primates

AimTo determine and explain biological traits that distinguish rare from common primate taxa.LocationAfrica, Americas, Asia, Madagascar.MethodsWe compare the biology of rare primate taxa with the biology of common taxa. Rarity is defined by (1) small size of geographic range; (2) small geographic range plus low local population density; and (3) small geographic range plus low local density plus narrow habitat specificity. After a linear comparison of size of geographic range with various biological traits, globally and by realm, extremes of rarity and commonness per realm are identified, and then combined for a global analysis. Tests are done both with genera treated as independent data points (n=62), and also with phylogenetic control by use of an independent contrasts test. Extinction risk in vertebrates, including primates, often correlates with high resource requirements, slow population recovery rate, and specialization. The three indices of rarity are therefore compared with these three general traits. Measures of resource use are body mass, local density, annual range size, and group size; of recovery rate, interbirth interval, and maximum intrinsic rate of natural population increase; and of degree of specialization, variety of diet, of habitats, maximum latitude, and morphological variety. All data come from the literature. Because several measures are compared, probabilities are Bonferroni corrected.ResultsIf rarity in primates correlates with any biological attribute, it consistently correlates with only measures of specialization, and not with measures of high resource use, or slow population recovery rate. Without phylogenetic correction, the first two indices of rarity associate significantly with all four measures of specialization, and the third with maximum latitude. With phylogenetic correction, the first index still associates with all four, the second with two (maximum latitude, number of species per genus), and the third shows no significant associations. While the four measures of specialization are strongly interrelated, stepwise regressions on geographic range indicate that maximum latitude has the strongest effect, followed by dietary variety and number of species per genus and, finally, habitat variety.Main conclusionsThe most commonly demonstrated traits of susceptibility to extinction are those of high resource use, slow recovery rate, and specialization. Yet, while rarity (almost however, it is defined) is an inevitable precursor to extinction, specialization is the only trait found to correlate with rarity in this study. We cannot explain this apparent contradiction.

Phylogenetic analysis of stomach adaptation in digestive strategies in African ruminants

The stomach morphology of 28 species of artiodactyls that differ in feeding style (browser, mixed feeder, grazer) was analysed using a multivariate approach and phylogenetic correction in order to test whether stomach morphology was correlated with feeding style when body mass was controlled for. A total of 25 morphological traits of the stomach were used in the analysis. After the effects of body mass and phylogeny on stomach morphology were taken into account, there was no significant grouping of species according to feeding style. When information about the feeding style of each species was included in the analysis, the set of morphological traits separated the mixed feeders from the other two feeding styles, but grazers and browsers had similar morphological features. Most of the variance in stomach morphology was explained by body mass and a lesser proportion by phylogeny. The main morphological features that have previously been proposed as being adaptations in grazing species, namely, lengthening of the retention time of ingesta to achieve an increase in their fibre digestion capability by means of a larger relative stomach capacity, a greater subdivision of chambers and smaller openings, are not supported by the findings of this study. Thus, there is no consistent evidence to support a significant adaptive effect of stomach morphology to different diets in the Artiodactyla.

FOREBRAIN SIZE AND INNOVATION RATE IN EUROPEAN BIRDS: FEEDING, NESTING AND CONFOUNDING VARIABLES

Abstract Previous work has shown a positive correlation between relative forebrain size and feeding innovation frequency, corrected for species number, over different taxonomic groups of birds. Several confounding variables could account for this relationship: ornithologists could notice and report innovations more often in certain taxa because of biased expectations, greater research effort, editorial bias in journals or large population sizes of the taxa. The innovationforebrain correlation could also be spuriously caused by phylogeny or juvenile development mode. We examined these possibilities by entering species number per taxon, population size, number of full length papers, expectations (assessed by a questionnaire), journal source and development mode in multiple regressions that also included relative forebrain size. We did this with and without phylogenetic corrections and tested two behavioural categories, feeding and nesting, where flexibility and learning are clearly thought to differ, but confounds should have similar effects. Through an exhaustive survey covering 30 years in 11 journals, a total of 683 innovations was gathered for the northwestern part of Europe, 507 for feeding and 176 for nesting. Species number per taxon was the only significant confound for both feeding and nesting reports; as predicted, forebrain size was a second significant predictor for feeding innovations, but not for nesting. The frequency of feeding innovations in the short notes of ornithology journals thus appears to be a valid and reliable way to operationalise behavioural flexibility in birds.

Comparing the preferences of three herbivore species with resistance traits of 15 perennial dicots: the effects of phylogenetic constraints

Differences in palatability between 15 perennial dicotyledonous herbaceous plant species were investigated in a multiple choice test, involving three generalist herbivores. Palatability was compared with two categories of plant traits: defence characteristics and growth properties. On theoretical grounds, we expected a positive relationship between growth rate and palatability, and a negative relationship between palatability and defence traits. The possible effect of phylogeny on the outcome of the comparisons was investigated by analysing the data using phylogenetically independent contrasts. The three herbivores showed remarkably similar preferences, which indicates that the results of the choice experiment are a good representation of leaf palatability. The significant negative correlations found in a direct comparison between herbivore preference and relative growth rate, and between herbivore preference and lignin and dry matter content, did not hold after phylogenetic correction. On the other hand, one relationship emerged only when comparing phylogenetically independent contrasts: contrary to expectation, a negative relation was noted between juvenile growth rate and plant palatability. A possible explanation for the contradictory nature of this result is the uniformly ready availability of resources in our experiment.