Light requirements for regeneration in tropical forest plants: Taxon-level and ecological attribute effects

Abstract Understanding co-adapted traits and strategies in organisms has been increasingly shown to require a broader view of structure, function, taxonomic affiliation (phylogeny) and their possible interactions. In order to understand mechanisms of recruitment dynamics in tropical forest, nine juvenile (fruit, seed and seedling) and seven adult ecological and morphological traits were examined in 89 woody plant species, from 75 genera, 35 families and 21 orders. The species were from tropical rainforest in northern Queensland and varied widely in light requirements and hence regeneration niches. The importance of taxonomic (phylogenetic) constraints was considered using various techniques. Data were collected mostly from herbarium specimens, augmented by observations in the field. The main source of variation in most traits was at the species level, although fruit size and light requirement varied most at the genus level, and flower size at the level of order. Covariation among attributes increased in strength with increasing taxonomic level. Cluster analyses of the species arising either from five or three groups reflected juvenile rather than adult attributes. However, these groups were not morphologically distinct. Sets of juvenile traits poorly predict adult functions (and vice-versa) at species, genus and family levels, but do so more strongly at the order level. Analyses of variance and multivariate analyses, with and without the effects of phylogeny, indicated that fruit size, seed size, hardness of seed coat, seedling cotyledon function, fruit maturation period and, to a lesser extent, seed number per fruit and wood density could predict light requirement, and hence regeneration niche. Flower size showed a strong phylogenetic effect, and thus is conserved along taxonomic lines. It is concluded that as different traits may vary at different taxonomic levels, the influence of phylogeny on interspecific variation needs to be routinely recognized by ecologists involved in comparative analyses of plant life-history functions.     

Seed Size,Fecundity and Postfire Regeneration Strategy Are Interdependent in Hakea

Seed size is a key functional trait that affects plant fitness at the seedling stage and may vary greatly with species fruit size, growth form and fecundity. Using structural equation modelling (SEM) and correlated trait evolution analysis, we investigated the interaction network between seed size and fecundity, postfire regeneration strategy, fruit size, plant height and serotiny (on-plant seed storage) among 82 species of the woody shrub genus, Hakea, with a wide spectrum of seed sizes (2–500 mg). Seed size is negatively correlated with fecundity, while fire-killed species (nonsprouters) produce more seeds than resprouters though they are of similar size. Seed size is unrelated to plant height and level of serotiny while it scales allometrically with fruit size. A strong phylogenetic signal in seed size revealed phylogenetic constraints on seed size variation in Hakea. Our analyses suggest a causal relationship between seed size, fecundity and postfire regeneration strategy in Hakea. These results demonstrate that fruit size, fecundity and evolutionary history have had most control over seed size variation among Hakea species.     

Plant dispersal strategies and the colonization of Araucaria forest patches in a grassland‐forest mosaic

Questions: In a natural grassland‐forest mosaic: What is the influence of phylogeny and diaspore traits related to disperser attraction (DAT) on (1) seed size/number trade‐off (SSNT) in woody species colonizing forest patches; (2) on the frequency of the species? 3. What is the influence of forest patch area on mean seed size and number. 4. Do phylogeny and DAT expressed at the species level affect this relationship? Location: Campos grassland and Araucaria forest in São Francisco de Paula, RS, Brazil, at ca. 29°28’ S; 50° 13’ W. Methods: Forest patches of different sizes in a grassland site recovering for ten years since human disturbances were surveyed by the relative abundance of vertebrate‐dispersed woody saplings. We described colonizer species according to taxonomic phylogenetic relationships and diaspore type, size and color. We analyzed with a variation partitioning method their influence on SSNT and on species frequency in the patches. At the community level we regressed mean seed size and number on forest patch area and evaluated how these relationships were affected by phylogeny and DAT at the species level. Results: 1. Phylogeny and DAT mostly explained seed size and seed number per diaspore variation. 2. By controlling phylogeny and DAT influence the frequency of species in forest patches was positively associated with their seed number in the diaspores, and negatively associated with their seed size. 3. Mean seed size and seed number at the community level were positively associated with patch area. 4. When phylogeny and DAT influences on seed size were removed this relationship was stronger for seed size and weaker for seed number. Conclusions: 1. Energy allocation to dispersal in detriment of offspring survival increased the successful establishment of colonizer species in forest patches, despite phylogenetic relationships and DAT variation in their diaspores. 2. Although patch area exerted a selective pressure on seed size, habitat preferences of dispersers may also influence patch colonization.     

Shared nucleotide composition biases among species and their impact on phylogenetic reconstructions of the Drosophilidae

Compositional changes are a major feature of genome evolution. Overlooking nucleotide composition differences among sequences can seriously mislead phylogenetic reconstructions. Large compositional variation exists among the members of the family Drosophilidae. Until now, however, base composition differences have been largely neglected in the formulations of the nucleotide substitution process used to reconstruct the phylogeny of this important group of species. The present study adopts a maximum-likelihood framework of phylogenetic inference in order to analyze five nuclear gene regions and shows that (1) the pattern of compositional variation in the Drosophilidae does not match the phylogeny of the species; (2) accounting for the heterogeneous GC content with Galtier and Gouy's nucleotide substitution model leads to a tree that differs in significant aspects from the tree inferred when the nucleotide composition differences are ignored, even though both phylogenetic hypotheses attain strong nodal support in the bootstrap analyses; and (3) the LogDet distance correction cannot completely overcome the distorting effects of the compositional variation that exists among the species of the Drosophilidae. Our analyses confidently place the Chymomyza genus as an outgroup closer than the genus Scaptodrosophila to the Drosophila genus and conclusively support the monophyly of the Sophophora subgenus.     

Seed mass of Indiana Dune genera and families: Taxonomic and ecological correlates

Summary Several surveys have documented an association among species between habitat type and seed mass, suggesting that habitat attributes impose a direct selective force on seed mass. Previous comparative surveys, however, have not controlled for the statistical effects of shared phylogenetic history (at the genus or family level) and life form when evaluating the relationship between habitat and seed mass. This study of the Indiana Dunes angiosperm flora provides statistical control of genus and family membership by: (i) partitioning out the statistical effect of genus membership prior to measuring the effect of habitat on seed mass, and (ii) seeking an association between habitat and seed mass within eight genera (206 species) and ten families (366 species). To measure the associations between ecological factors, taxonomic membership and seed mass, I examined life form, phenological schedules and seed mass among species in 8 genera distributed among 13 habitat types (assigned to 1 of 4 categories of inferred water and light availability). One-way ANOVAS indicated that genus, life form, habitat, water/light category, the onset of flowering and the duration of flowering accounted for 71%, 51%, 10%, 4%, 14% and 14% of the variance in seed mass, respectively. However, multi-factor ANOVAS measured the variance in seed mass accounted for by each variable independently of the others: only genus explained a significant proportion (11%). Genus membership is strongly associated with the other ecological factors, accounting for the difference between one-way and multi-factor ANOVAS. Within the ecologically widespread genera and families of this study, there was no significant association between water/light category and seed mass, even though this association can be detected across taxa. Among congeners and confamilials, interspecific variation in seed mass (measured as the coefficient of variation) was as high within habitat types as among them, suggesting that habitats do not provide upper limits to the range of seed mass exhibited by the species within them. A previous study of 648 Indiana Dune species showed that species segregate among habitats on the basis of seed size; large-seeded species tend to occupy closed habitats and small-seeded species tend to inhabit open habitats. This segregation creates the ecologically meaningful observation that low-light habitats support larger-seeded species than high-light habitats, even though this pattern cannot be detected independently of taxonomic membership. Generalist taxa may occupy a wide range of habitats for reasons other than their seed size. If this is a common feature of ecological generalists, it may not be possible to detect an association between habitat and seed size independently of taxonomic membership.     

Cracking the case: Seed traits and phylogeny predict time to germination in prairie restoration species

Traits are important for understanding how plant communities assemble and function, providing a common currency for studying ecological processes across species, locations, and habitat types. However, the majority of studies relating species traits to community assembly rely upon vegetative traits of mature plants. Seed traits, which are understudied relative to whole‐plant traits, are key to understanding assembly of plant communities. This is particularly true for restored communities, which are typically started de novo from seed, making seed germination a critical first step in community assembly and an early filter for plant establishment. We experimentally tested the effects of seed traits (mass, shape, and embryo to seed size ratio) and phylogeny on germination response in 32 species commonly used in prairie grassland restoration in the Midwestern USA, analyzing data using time‐to‐event (survival) analysis. As germination is also influenced by seed dormancy, and dormancy break treatments are commonly employed in restoration, we also tested the effects of two pretreatments (cold stratification and gibberellic acid application) on time to germination. Seed traits, phylogeny, and seed pretreatments all affected time to germination. Of all traits tested, variables related to seed shape (height and shape variance) best predicted germination response, with high‐variance (i.e., pointier and narrower) seeds germinating faster. Phylogenetic position (the location of species on the phylogenetic tree relative to other tested species) was also an important predictor of germination response, that is, closely related species showed similar patterns in time to germination. This was true despite the fact that all measured seed traits showed phylogenetic signal, therefore phylogeny provided residual information that was not already captured by measured seed traits. Seed traits, phylogenetic position, and germination pretreatments were important predictors of germination response for a suite of species commonly used in grassland restoration. Shape traits were especially important, while mass, often the only seed trait used in studies of community assembly, was not a strong predictor of germination timing. These findings illustrate the ecological importance of seed traits that are rarely incorporated into functional studies of plant communities. This information can also be used to advance restoration practice by guiding restoration planning and seed mix design.     

An AFLP marker approach to lower-level systematics in Eucalyptus (Myrtaceae)

Genus Eucalyptus, with over 700 species, presents a number of systematic difficulties including taxa that hybridize or intergrade across environmental gradients. To date, no DNA marker has been found capable of resolving phylogeny below the sectional level in the major subgenera. Molecular markers are needed to support taxonomic revision, assess the extent of genetic divergence at lower taxonomic levels, and inform conservation efforts. We examined the utility of 930 amplified fragment length polymorphisms (AFLPs) for analyzing relationships among Tasmanian taxa of subgenus Symphyomyrtus section Maidenaria. Phenetic and cladistic analyses resolved species into clusters demonstrating significant genetic partitioning, largely concordant with series defined in the most recent taxonomic revision of Eucalyptus. Some departures from current taxonomy were noted, indicating possible cases of morphological convergence and character reversion. Although the resolution obtained using AFLP was greatly superior to that of single sequence markers, the data demonstrated high homoplasy and incomplete resolution of closely related species. The results of this study and others are consistent with recent speciation and reticulate evolution in Maidenaria. We conclude that a combination of phylogenetic and population genetic approaches using multiple molecular markers offers the best prospects for understanding taxonomic relationships below the sectional level in Eucalyptus.     

Predicting plant responses to mycorrhizae: integrating evolutionary history and plant traits

We assessed whether (1) arbuscular mycorrhizal colonization of roots (RC) and/or plant responses to arbuscular mycorrhizae (MR) vary with plant phylogeny and (2) MR and RC can be more accurately predicted with a phylogenetic predictor relative to a null model and models with plant trait and taxonomic predictors. In a previous study, MR and RC of 95 grassland species were measured. We constructed a phylogeny for these species and found it explained variation in MR and RC. Next, we used multiple regressions to identify the models that most accurately predicted plant MR. Models including either phylogenetic or phenotypic and taxonomic information similarly improved our ability to predict MR relative to a null model. Our study illustrates the complex evolutionary associations among species and constraints of using phylogenetic information, relative to plant traits, to predict how a plant species will interact with AMF.     

Relationships among ecologically important dimensions of plant trait variation in seven neotropical forests

BACKGROUND AND AIMS: When ecologically important plant traits are correlated they may be said to constitute an ecological 'strategy' dimension. Through identifying these dimensions and understanding their inter-relationships we gain insight into why particular trait combinations are favoured over others and into the implications of trait differences among species. Here we investigated relationships among several traits, and thus the strategy dimensions they represented, across 2134 woody species from seven Neotropical forests. METHODS: Six traits were studied: specific leaf area (SLA), the average size of leaves, seed and fruit, typical maximum plant height, and wood density (WD). Trait relationships were quantified across species at each individual forest as well as across the dataset as a whole. 'Phylogenetic' analyses were used to test for correlations among evolutionary trait-divergences and to ascertain whether interspecific relationships were biased by strong taxonomic patterning in the traits. KEY RESULTS: The interspecific and phylogenetic analyses yielded congruent results. Seed and fruit size were expected, and confirmed, to be tightly related. As expected, plant height was correlated with each of seed and fruit size, albeit weakly. Weak support was found for an expected positive relationship between leaf and fruit size. The prediction that SLA and WD would be negatively correlated was not supported. Otherwise the traits were predicted to be largely unrelated, being representatives of putatively independent strategy dimensions. This was indeed the case, although WD was consistently, negatively related to leaf size. CONCLUSIONS: The dimensions represented by SLA, seed/fruit size and leaf size were essentially independent and thus conveyed largely independent information about plant strategies. To a lesser extent the same was true for plant height and WD. Our tentative explanation for negative WD-leaf size relationships, now also known from other habitats, is that the traits are indirectly linked via plant hydraulics.     

Rapid divergence of microsatellite abundance among species of Drosophila

Among major taxonomic groups, microsatellites exhibit considerable variation in composition and allele length, but they also show considerable conservation within many major groups. This variation may be explained by slow microsatellite evolution so that all species within a group have similar patterns of variation, or by taxon-specific mutational or selective constraints. Unfortunately, comparing microsatellites across species and studies can be problematic because of biases that may exist among different isolation and analysis protocols. We present microsatellite data from five Drosophila species in the Drosophila subgenus: D. arizonae, D. mojavensis, and D. pachea (three cactophilic species), and D. neotestacea and D. recens (two mycophagous species), all isolated at the same time using identical protocols. For each species, we compared the relative abundance of motifs, the distribution of repeat size, and the average number of repeats. Dimers were the most abundant microsatellites for each species. However, we found considerable variation in the relative abundance of motif size classes among species, even between sister taxa. Frequency differences among motifs within size classes for the three cactophilic species, but not the two mycophagous species, are consistent with other studied Drosophila. Frequency distributions of repeat number, as well as mean size, show significant differences among motif size classes but not across species. Sizes of microsatellites in these five species are consistent with D. virilis, another species in the subgenus Drosophila, but they have consistently higher means than in D. melanogaster, in the subgenus Sophophora. These results confirm that many aspects of microsatellite variation evolve quickly but also are subject to taxon-specific constraints. In addition, the nature of microsatellite evolution is dependent on temporal and taxonomic scales, and some variation is conserved across broad taxonomic levels despite relatively high rates of mutation for these loci.     

Phylogeny of the genus Aphis Linnaeus, 1758 (Homoptera: Aphididae) inferred from mitochondrial DNA sequences

Aphis is the largest aphid genus in the world and contains several of the most injurious aphid pests. It is also the most reluctant aphid genus to any comprehensive taxonomic treatment: while most species are easily classified into "species groups" that form well defined entities, numerous species within these groups are difficult to tell apart morphologically and identification keys remain ambiguous and mostly rely on host plant affiliation. In this paper, we used partial sequences of COI/COII and CytB genes to reconstruct the first phylogeny of Aphis and discuss the present systematics. The monophyly of the subgenus Bursaphis and of the tree major species groups, Black aphid, Black backed aphid and frangulae-like species was recovered by all phylogenetic analyses. However our data suggested that the nominal subgenus was not monophyletic. Relationships between major species groups were often ambiguous but "Black" and "Black backed" species groups appeared as sister clades. The most striking result of this study was that our molecular data met the same limits as the morphological characters used in classifications: mitochondrial DNA did not allow the differentiation of species that are difficult to identify. Further, interspecies relationships within groups of species for which taxonomic treatment is difficult stayed unresolved. This suggests that species delineation in the genus Aphis is often ambiguous and that diversification might have been a rapid process.     

Geographic variation in seed traits within and among forty‐two species of Rhododendron (Ericaceae) on the Tibetan plateau: relationships with altitude,habitat, plant height,and phylogeny

Seed mass and morphology are plant life history traits that influence seed dispersal ability, seeding establishment success, and population distribution pattern. Southeastern Tibet is a diversity center for Rhododendron species, which are distributed from a few hundred meters to 5500 m above sea level. We examined intra‐ and interspecific variation in seed mass and morphology in relation to altitude, habitat, plant height, and phylogeny. Seed mass decreased significantly with the increasing altitude and increased significantly with increasing plant height among populations of the same species. Seed mass differed significantly among species and subsections, but not among sections and subgenera. Seed length, width, surface area, and wing length were significantly negative correlated with altitude and significantly positive correlated with plant height. Further, these traits differed significantly among habitats and varied among species and subsection, but not among sections and subgenera. Species at low elevation had larger seeds with larger wings, and seeds became smaller and the wings of seeds tended to be smaller with the increasing altitude. Morphology of the seed varied from flat round to long cylindrical with increasing altitude. We suggest that seed mass and morphology have evolved as a result of both long‐term adaptation and constraints of the taxonomic group over their long evolutionary history.     

染色体数据的挖掘及其在植物多样性进化研究中的利用

多倍化（或全基因组加倍）是植物物种形成的重要途径,现存的被子植物可能都发生过一次甚至多次多倍化事件。多倍化传统的定义是染色体数目相对于祖先类群呈整倍性增加。其中最常用的研究方法是核型分析,核型能够提供物种的基本细胞学参数,包括染色体数目、倍性水平、核型不对称性、核型变异系数等。目前核型研究的趋势表现出从物种基本核型参数分析逐渐演化到多类群、多学科交叉融合的特点：一方面植物核型分析从种群、物种、科属的类群到生命之树,探讨染色体核型在各支系的进化特征、趋势以及驱动植物系统进化的细胞学机制;另一方面探讨和分析区域或生态系统植物区系的染色体谱或倍性等细胞学特征,可以探究区域地质环境变化或生态环境对染色体倍性等的影响,或通过区域染色体谱的构建,分析区域植物区系的形成和进化历史。因而,植物核型研究为系统发育、分子系统进化、生命之树以及植物区系地理的起源和演化研究提供了新思路。越来越多的新方法、新手段在植物核型分析与多倍化研究中得到运用,从而揭示了植物类群或植物区系的染色体进化以及细胞地理特征。今后植物细胞学研究趋势会向多学科交叉融合,整合各研究领域证据,从不同水平角度综合分析植物核型多样性形成的原因及意义,从而更加全面地认识和理解植物物种多样化与物种形成原因。     

Hidden responses to environmental variation: maternal effects reveal species niche dimensions

Species responses to fluctuating environments structure population and community dynamics in variable ecosystems. Although offspring number is commonly used to measure these responses, maternal effects on offspring quality may be an important but largely unrecognised determinant of long‐term population growth. We selected 29 species across a Mediterranean annual plant phylogeny, and grew populations of each species in wet and dry conditions to determine responses in seed number and maternal effects (seed size, seed dormancy, and seedling growth). Maternal effects were evident in over 40% of species, but only 24% responded through seed number. Despite a strong trade‐off between seed size and seed number among species, there was no consistent trade‐off within species; we observed correlations that ranged from positive to negative. Overall, species in this plant guild show a complex range of responses to environmental variation that may be underestimated when only seed number responses are considered.     

Evolution of genome size across some cultivated Allium species.

Allium L. (Alliaceae), a genus of major economic importance, exhibits a great diversity in various morphological characters and particularly in life form, with bulbs and rhizomes. Allium species show variation in several cytogenetic characters such as basic chromosome number, ploidy level, and genome size. The purpose of the present investigation was to study the evolution of nuclear DNA amount, GC content, and life form. A phylogenetic approach was used on a sample of 30 Allium species, including major vegetable crops and their wild allies, belonging to the 3 major subgenera Allium, Amerallium, and Rhizirideum and 14 sections. A phylogeny was constructed using internal transcribed spacer (ITS) sequences of 43 accessions representing 30 species, and the nuclear DNA amount and the GC content of 24 Allium species were investigated by flow cytometry. For the first time, the nuclear DNA content of Allium cyaneum and Allium vavilovii was measured, and the GC content of 16 species was measured. We addressed the following questions: (i) Is the variation in nuclear DNA amount and GC content linked to the evolutionary history of these edible Allium species and their wild relatives? (ii) How did life form (rhizome or bulb) evolve in edible Allium? Our results revealed significant interspecific variation in the nuclear DNA amount as well as in the GC content. No correlation was found between the GC content and the nuclear DNA amount. The reconstruction of nuclear DNA amount on the phylogeny showed a tendency towards a decrease in genome size within the genus. The reconstruction of life form history showed that rhizomes evolved in the subgenus Rhizirideum from an ancestral bulbous life form and were subsequently lost at least twice independently in this subgenus.     

Papilio phylogeny based on mitochondrial cytochrome oxidase I and II genes

Butterflies of the genus Papilio have served as the basis for numerous studies in insect physiology, genetics, and ecology. However, phylogenetic work on relationships among major lineages in the genus has been limited and inconclusive. We have sequenced 2.3 kb of DNA from the mitochondrial cytochrome oxidase I and II genes (COI and COII) for 23 Papilio taxa and two outgroups, Pachliopta neptunus and Eurytides marcellus, in order to assess the potential of these genes for use in Papilio phylogenetics and to examine patterns of gene evolution across a broad taxonomic range. Nucleotide and amino acid variation is distributed heterogeneously, both within and between genes. Structural features of the proteins are not always reliable predictors of variation. In a combined analysis, these sequences support a nearly fully resolved topology within subgenera and species groups, though higher level relationships among species groups require additional study. The most noteworthy findings are that neither Papilio alexanor nor P. xuthus belongs in the machaon group and that the subgenus Pterourus is paraphyletic with respect to the subgenus Pyrrhosticta. We leave relationships among members of the phorcas species group as a trichotomy. These two protein coding genes, particularly COI, show excellent performance in resolving relationships at the level of species and species groups among Papilionidae. We strongly endorse a similar approach for future studies aimed at these levels.     

Karyomorphology and GC-rich heterochromatin pattern in Passiflora (Passifloraceae) wild species from Decaloba and Passiflora subgenera

Thirteen wild species of Passiflora were analyzed using conventional and CMA/DA/DAPI staining to evaluate the karyotype diversity between and within the subgenus Decaloba and Passiflora. The karyotypic features indicate that both subgenera have a conserved chromosome number, as reported before for several species. Submetacentric (sm) chromosomes were found in species from both subgenera, suggesting that sm chromosomes are not restricted to a particular subgenus. The analysis of the karyotypic heterogeneity enabled to distribute the species in three groups, but with no support to phylogenetic and taxonomic levels. The application of fluorochromes allowed for the visualization of CMA⁺/DAPI⁻ blocks, which in our studies always correlated with the occurrence of satellites, showing that occurrence of two chromosome pairs with satellites per cell is a characteristic shared by some species from both subgenera. This feature does not always have relationship with the basic chromosome number. The data found in this study will help to understand the phylogeny, cytotaxonomy, and evolution of the genus Passiflora showing that karyotypic variation can be seen between and within the subgenus Decaloba and Passiflora.     

Partitioning variance in a physiological trait: desiccation resistance in keratin beetles (Coleoptera, Trogidae)

1. Although variation in physiological traits forms the substance of evolutionary physiology, the way that this variation is partitioned among different hierarchical levels (e.g. population, species, genus) is not well known.
2. In this study variance partitioning is examined in body size, water content, lipid content, rate of water loss, maximum tolerable water loss and survival time at four levels (individual, population, species, genus) in southern African keratin beetles.
3. It is found that most variance in body size, and the physiological traits that are strongly influenced by body size (water and lipid content, maximum tolerable water loss, rate of water loss), is partitioned at the generic level (50–70%), then at the species level (20–50%) and finally at the population (1–9%) and individual (2–18%) levels.
4. On the other hand, variance in survival time, and variance in rate of maximum water loss once the effects of body size have been taken into account, are partitioned mostly at the species level (40–70%), whereas maximum tolerable water loss and lipid and water content show greatest variance at the individual level (63–75%). This is largely a consequence of differences in the extent of scaling of these traits.
5. The results suggest that where possible the effects of phylogeny should be controlled for when examining desiccation resistance in insects using comparative methods, but that confidence in conclusions from previous studies, which have used mass-specific data, is not unwarranted.     

Nuclear markers confirm taxonomic status and relationships among highly endangered and closely related right whale species

Right whales (genus: Eubalaena) are among the most endangered mammals, yet their taxonomy and phylogeny have been questioned. A phylogenetic hypothesis based on mitochondrial DNA (mtDNA) variation recently prompted a taxonomic revision, increasing the number of right whale species to three. We critically evaluated this hypothesis using sequence data from 13 nuclear DNA (nuDNA) loci as well as the mtDNA control region. Fixed diagnostic characters among the nuclear markers strongly support the hypothesis of three genetically distinct species, despite lack of any diagnostic morphological characters. A phylogenetics analysis of all data produced a strict consensus cladogram with strong support at nodes that define each right whale species as well as relationships among species. Results showed very little conflict among the individual partitions as well as congruence between the mtDNA and nuDNA datasets. These data clearly demonstrate the strength of using numerous independent genetic markers during a phylogenetics analysis of closely related species. In evaluating phylogenetic support contributed by individual loci, 11 of the 14 loci provided support for at least one of the nodes of interest to this study. Only a single marker (mtDNA control region) provided support at all four nodes. A study using any single nuclear marker would have failed to support the proposed phylogeny, and a strong phylogenetic hypothesis was only revealed by the simultaneous analysis of many nuclear loci. In addition, nu DNA and mtDNA data provided complementary levels of support at nodes of different evolutionary depth indicating that the combined use of mtDNA and nuDNA data is both practical and desirable.     

Segregating the Effects of Seed Traits and Common Ancestry of Hardwood Trees on Eastern Gray Squirrel Foraging Decisions

The evolution of specific seed traits in scatter-hoarded tree species often has been attributed to granivore foraging behavior. However, the degree to which foraging investments and seed traits correlate with phylogenetic relationships among trees remains unexplored. We presented seeds of 23 different hardwood tree species (families Betulaceae, Fagaceae, Juglandaceae) to eastern gray squirrels (Sciurus carolinensis), and measured the time and distance travelled by squirrels that consumed or cached each seed. We estimated 11 physical and chemical seed traits for each species, and the phylogenetic relationships between the 23 hardwood trees. Variance partitioning revealed that considerable variation in foraging investment was attributable to seed traits alone (27–73%), and combined effects of seed traits and phylogeny of hardwood trees (5–55%). A phylogenetic PCA (pPCA) on seed traits and tree phylogeny resulted in 2 “global” axes of traits that were phylogenetically autocorrelated at the family and genus level and a third “local” axis in which traits were not phylogenetically autocorrelated. Collectively, these axes explained 30–76% of the variation in squirrel foraging investments. The first global pPCA axis, which produced large scores for seed species with thin shells, low lipid and high carbohydrate content, was negatively related to time to consume and cache seeds and travel distance to cache. The second global pPCA axis, which produced large scores for seeds with high protein, low tannin and low dormancy levels, was an important predictor of consumption time only. The local pPCA axis primarily reflected kernel mass. Although it explained only 12% of the variation in trait space and was not autocorrelated among phylogenetic clades, the local axis was related to all four squirrel foraging investments. Squirrel foraging behaviors are influenced by a combination of phylogenetically conserved and more evolutionarily labile seed traits that is consistent with a weak or more diffuse coevolutionary relationship between rodents and hardwood trees rather than a direct coevolutionary relationship.