The role of life history traits in mammalian invasion success

Why some organisms become invasive when introduced into novel regions while others fail to even establish is a fundamental question in ecology. Barriers to success are expected to filter species at each stage along the invasion pathway. No study to date, however, has investigated how species traits associate with success from introduction to spread at a large spatial scale in any group. Using the largest data set of mammalian introductions at the global scale and recently developed phylogenetic comparative methods, we show that human‐mediated introductions considerably bias which species have the opportunity to become invasive, as highly productive mammals with longer reproductive lifespans are far more likely to be introduced. Subsequently, greater reproductive output and higher introduction effort are associated with success at both the establishment and spread stages. High productivity thus supports population growth and invasion success, with barriers at each invasion stage filtering species with progressively greater fecundity.     

Life history traits of two dwarf cichlids species in the white waters of the Amazonian floodplain

The reproductive parameters are among the most important life history aspects of fishes influenced by environmental variation. During recent years, the main life history strategies of Amazonian fish species were defined mostly by a set of reproductive parameters. In the present work, we sought to describe important life history parameters, in particular on reproductive characteristics of Apistogramma agassizii and Apistogramma bitaeniata, found in floodplain lakes of the Brazilian Amazonia. The species presented a positive sexual dimorphism, and males were significantly bigger than females. For both sexes, four developmental phase of gonad maturation were detected, and based on those it was possible to identify mature, reproductive specimens throughout the entire period of the study. From the ovaries of mature females, fecundity and spawning type were determined. Low fecundity, short spawning periods, possibly separated only by few months, and total spawning are all good indications that A. agassizii and A. bitaeniata evolved an opportunistic strategy in their life history.     

Coexistence of three differentDrosophila species by rescheduling their life history traits in a natural population

We present evidence for coexistence of three differentDrosophila species by rescheduling their life history traits in a natural population using the same resource, at the same time and same place.D. ananassae has faster larval development time (DT) and faster DT(egg-fly) than other two species thus utilizing the resources at maximum at both larval and adult stages respectively. Therefore,D. ananassae skips the interspecific competition at preadult stage but suffers more from intraspecific competition. However,D. melanogaster andD. biarmipes have rescheduled their various life history traits to avoid interspecific competition. Differences of ranks tests for various life history traits suggest that except for DT(egg-pupa), the difference of ranks is highest for the combination ofD. melanogaster andD. ananassae for all other life history traits. This difference is maintained by tradeoffs between larval development time and pupal period and between pupal period and DT(egg-pupa) inD. ananassae.     

Theoretical habitat templets, species traits, and species richness: floodplain vegetation in the Upper Rhône River

• 1 The floodplain vegetation at approximately 100 sites located in nine different habitat types of the Upper Rhône River, France, was surveyed three times over the past 27 years. Information on species traits of the higher plants comprising the Rhône floodplain vegetation was based on studies conducted between Geneva, Switzerland, and Lyon, France.
• 2 These data were structured using a ‘fuzzy coding’ technique and then examined using ordination analyses to investigate: (i) relationships among species traits; (ii) habitat utilization; (iii) the relationship between species traits and habitat utilization; and (iv) trends of species traits and species richness in the framework of spatial–temporal habitat variability to test predictions of the river habitat templet and the patch dynamics concept.
• 3 Size, number of descendants per reproductive cycle, number of reproductive cycles per individual, and the regeneration potential of an individual were positively related with each other, whereas the degree of attachment to the soil decreased, and the reproductive period shifted from autumn/late summer towards early summer/spring, as size increased.
• 4 The habitat utilization by the higher plants of the floodplain revealed a double lateral gradient: the first was from the banks of the temporary waters to terrestrial flats; the second from aggrading pebble to aggrading silt habitats. These gradients were related to gradients in water saturation, oxygen conditions, nutrient loading, and nutrient retention of the soils.
• 5 A significant relationship between species traits and habitat utilization was observed for the floodplain vegetation, i.e. plant communities used particular habitat types with a particular set of species trait modalities (= categories).
• 6 Patterns of species trait modalities were significantly related to temporal and spatial habitat variability but only modalities of the trait ‘parental care’ conformed to trends predicted from theory.
• 7 No trends were observed when species richness of different habitat types was considered in the framework of spatial–temporal habitat variability.
• 8 Although the habitats of the Upper Rhône clearly act as a templet for the species traits of the floodplain vegetation, the lack of agreement between observations and predictions on trends in species traits and richness in terms of habitat variability suggest that important elements of theory should be rejected. However, human-induced changes in these habitats are too recent when compared with the longer time periods required for floodplain vegetation to respond to such changes.

     

Evolution of the G-matrix in life history traits in the common frog during a recent colonisation of an island system

Studies of genetic correlations between traits that ostensibly channel the path of evolution away from the direction of natural selection require information on key aspects such as ancestral phenotypes, the duration of adaptive evolution, the direction of natural selection, and genetic covariances. In this study we provide such information in a frog population system. We studied adaptation in life history traits to pool drying in frog populations on islands of known age, which have been colonized from a mainland population. The island populations show strong local adaptation in development time and size. We found that the first eigenvector of the variance–covariance matrix (g _max) had changed between ancestral mainland populations and newly established island populations. Interestingly, there was no divergence in g _max among island populations that differed in their local adaptation in development time and size. Thus, a major change in the genetic covariance of life-history traits occurred in the colonization of the island system, but subsequent local adaptation in development time took place despite the constraints imposed by the genetic covariance structure.     

Grasshopper response to reductions in habitat area as mediated by subfamily classification and life history traits

Although the loss of species is often attributed to reductions in habitat area, heterogeneity and connectivity, species specific traits and taxonomic relatedness can be important in explaining which species groups may be most impacted by the fragmentation process. In this study, using urban grassland fragments within the Front Range of northern Colorado, USA, we quantified the degree to which grasshopper species numbers declined with reductions in habitat area. We then examined the degree to which subfamily grouping (Gomphocerinae, Melanoplinae, Oedipodinae) and variation in life history characteristics (body size, dispersal ability, feeding preference and juvenile emergence time) explained which grasshopper species groups might be most impacted by reductions in habitat size. Our results showed a strong species-area relationship for grasshoppers across the urban fragments. Grasshoppers in different subfamily groupings were found to respond differentially, with the Melanoplinae being least affected by reductions in habitat area, while the other subfamilies responded similarly and lost species at a higher rate. Species with different feeding strategies were also found to respond differentially, with forbivorous species being least affected by the reductions in habitat area, while the graminivorous and herbivorous (mixed-feeders) responded similarly and lost species at a higher rate. As Melanoplinae tend to be forb feeders, this may partially explain why the subfamily was not as affected by the fragmentation process as the Gomphocerinae and Oedipodinae (who tend to be grass and mixed-feeders, respectively). Species that differed in body size and dispersal ability did not display differential responses to reductions in habitat area.     

Theoretical habitat templets, species traits, and species richness: fish in the Upper Rhône River and its floodplain

• 1 Recent developments in ecological theory concerned with habitat templets, species assemblages, and life history traits were examined for the riverine fish communities of the Upper Rhône River, France, in the context of spatial–temporal habitat variability. Relationships among species traits, habitat utilization of species, the relationship between species traits and habitat utilization, and trends of species traits and species richness in the spatial–temporal variability of the habitat types were analysed.
• 2 Relationships among twelve species traits, and utilization of eight habitats were examined for twenty-five fish species using correspondence analysis; the relationship between species traits and habitat utilization was investigated by co-inertia analysis.
• 3 Positive relationships among species traits were observed for size, fecundity, and the number of reproductive cycles per individual. However, species were not well differentiated according to the habitat utilization, except for habitats rarely connected with the main channel (i.e. two types of oxbow lakes).
• 4 No significant relationship was found between species traits and habitat utilization, nor for either species traits or species richness when examined in the framework of spatial–temporal habitat variability. Only two species traits corresponded (with slight trends) to predictions in a river habitat templet: (i) the number of descendants per reproductive cycle increased along with temporal variability; and (ii) the number of reproductive cycles per individual was either low or high at low temporal variability and intermediate at elevated temporal variability.
• 5 The discrepancy between the predictions of the river habitat templet as well as of the patch dynamics concept and the results observed for the fish in the Upper Rhône was explained in terms of scale problems, the evolutionary ecology of the European fish fauna, and the history of the Rhône River.

     

Theoretical habitat templets, species traits, and species richness: Trichoptera in the Upper Rhône River and its floodplain

• 1 For Trichoptera occurring in two sites of the Upper Rhône River (France) we examine: (i) relationships among species traits; (ii) habitat utilization of Trichoptera species; (iii) the relationship between species traits and habitat utilization; (iv) trends of species traits in the framework of spatial–temporal habitat variability to test predictions of the habitat templet concept; and (v) trends of species richness in the framework of spatial–temporal habitat variability to test predictions of the patch dynamics concept.
• 2 Of the sixteen species traits selected, twelve have high correlation ratios for the seventy-five species used in this analysis; these traits are related to behavioural, morphological, or physiological aspects. Traits related to reproduction or life cycle have lower correlation ratios.
• 3 An ordination by species traits separates the five main families into three groups: (i) Hydropsychidae and Polycentropodidae; (ii) Hydroptilidae; and (iii) Leptoceridae and Limnephilidae. An ordination of the habitat utilization of the species in ten habitats indicates that the Hydropsychidae occur preferentially in the main channel, Hydroptilidae, Polycentropodidae, and Limnephilidae occur in backwaters or oxbow lakes, and the Leptoceridae are ubiquitous.
• 4 The Hydropsychidae exhibit a relationship between species traits and habitat utilization, i.e. they use similar habitat types with similar species traits. The species traits of the other four families are similar but their habitat utilization is quite different.
• 5 The Hydropsychidae occur in lowest spatial–temporal variability habitats and Limnephilidae in the highest. Therefore, net spinners and filterers are characteristic of habitats with a low spatial–temporal variability, whereas shredders and case makers using plant material are characteristic of habitats with high spatial–temporal variability. The trends in species traits show little agreement with trends predicted from the river habitat templet.
• 6 Trends of species richness in the framework of spatial and temporal variability do not follow the predictions of the patch dynamics concept because richness is similar in all superficial habitats. This implies that each habitat, in spite of large differences in their spatial and temporal variability, offers Trichoptera a similar but limited number of ecological niches.

     

Theoretical habitat templets, species traits, and species richness: birds in the Upper Rhône River and its floodplain

• 1 To test predictions of the river habitat templet and the patch dynamics concept, twenty species traits and the species richness of eighty-one bird species were examined in the context of the spatial – temporal variability of ninety-one sites at Jons, a section of the Upper Rhône River, France.
• 2 The basic information was obtained either from the literature (for species traits) or by observation, then structured by a fuzzy coding technique, and examined using multivariate analyses.
• 3 The relationships between species traits separated three groups of birds: (i) large species that are long lived, have long breeding cycles, and use open sites to feed, breed, and sing (e.g. Podicipidae, Laridae); (ii) medium-sized species with intermediate longevities and breeding cycles that use less open sites (e.g. Columbidae, Corvidae); and (iii) small species that are short lived, have short breeding cycles, and use closed sites (e.g. Sylviidae).
• 4 The distribution of birds in the floodplain habitats of the river was closely related to the spatial–temporal variability of sites.
• 5 No trends in species traits were observed along a gradient of spatial–temporal variability of the sites.
• 6 Species richness of birds was relatively constant over the gradient of spatial–temporal variability of the ninety-one sites, but decreased at low spatial and high temporal variability because of human activities and vegetation changes that occurred at these sites.
• 7 These observations on species traits and richness were related to the high mobility of birds (which is a scale phenomenon) and the high intercorrelation between parameters used to determine the spatial and temporal variability of the sites (which makes separation of the spatial from the temporal variability impossible).

     

Theoretical habitat templets, species traits, and species richness: amphibians in the Upper Rhône River and its floodplain

• 1 The purpose of this study was to test predictions of the habitat templet and the patch dynamics concept by analysing the relationship between either the species traits or species richness of amphibians and the spatial–temporal variability of eight habitat types of the Upper Rhône River and its floodplain, France.
• 2 The information on species traits of the twelve amphibian species was accumulated primarily from the literature; that on habitat utilization was based on field surveys. The information was ‘fuzzy coded’, analysed by ordination techniques, and finally linked to the spatial–temporal variability of habitat types elaborated elsewhere in this issue.
• 3 After elimination of the variance caused by the differences between urodeles and anurans, correspondence analysis of the matrix of species traits demonstrated that the flexibility of the temporal pattern of reproduction was the major source of variation among amphibian species. A less significant source of variation within anurans was related to traits usually linked to the concept of r–K selection (potential number of descendants per reproductive cycle and ratio of egg size to female size).
• 4 A correspondence analysis of habitat utilization by amphibian species separated four groups of habitat types, corresponding to different degrees of influence that the active channel had on adjacent waters in the floodplain (i.e. from habitats that are most frequently disturbed by floods, to habitats of braided sections, old oxbow lakes, and the more terrestrialized habitats).
• 5 A co-inertia analysis of the relationship between species traits and habitat utilization separated two habitat types (terrestrialized former meanders and oxbow lakes) from the others. Species of terrestrialized habitats had a reproductive display, a low number of descendants per reproductive cycle, and a high ratio of egg size to female size. Species of oxbow lakes were early or synchronous breeders, and laid a single clutch.
• 6 Neither the predictions of the habitat templet concept nor those of the patch dynamics concept were validated. Species traits were not significantly correlated to the axes of the spatial–temporal variability templet. The highest species richness was observed in habitat types with the highest temporal variability; in addition, richness did not peak at highest spatial variability.
• 7 Deviation from the predictions of the river habitat templet could be explained by: (i) the over-simplification of this model with regard to the evolution of the complex life cycles of the amphibians; and (ii) an underestimation of the importance of phylogenetic constraints and the evolution of community interactions.

     

The role of seed dispersal and seedling traits in colonization and coexistence ofSalix species in a seasonally flooded habitat

Regeneration traits of six co-occurringSalix species were studied on a floodplain of the Sorachi River, central Hokkaido, Japan, and their colonization success and coexistence in a local habitat were discussed. MixedSalix communities contained sixSalix species; dominant:S. sachalinensis; four subordinates:S. rorida, S. pet-susu, S. miyabeana andS. subfragilis; rare species:S. jessoensis. Their phenology, falling velocity and longevity of seeds, and the effects of microtopography and soil texture on seedling establishment were studied. The sixSalix species had overlapped seed dispersal periods that coincided with the decrease of water level after a predictable spring flood. This coincidence was crucial for the colonization success because the seedlings were established on wet soils left by the decreasing water level. They showed two types of regeneration trait, specialization and generalization.S. rorida andS. subfragilis showed contrasting regeneration traits; early vs. late seed dispersal, large vs. small seeds, seedling distribution on coarse vs. fine soils, respectively. These two species rarely co-occurred. On the other hand, the dominantS. sachalinensis had an intermediate seed size and dispersal timing, and a wide range of seedling distribution from coarse to fine soils. These results revealed that flooding seasonality influenced the colonization success together with the regeneration traits ofSalix species, and that coexistence of theSalix species was facilitated primarily by regeneration niche separation related to flooding seasonality and soil heterogeneity.     

Theoretical habitat templets, species traits, and species richness: aquatic oligochaetes in the Upper Rhône River and its floodplain

• 1 This paper summarizes twenty years of ecological research on aquatic oligochaetes of the Upper Rhône River and its alluvial floodplain (France). Species traits of fifty species of the ninety taxa recorded from two areas Gons and Brégnier-Cordon) were used to examine the relationships among species traits, habitat utilization of these species, whether a relationship exists between species traits and habitat utilization, and the applicability of predictions from the river habitat templet and the patch dynamics concept in the framework of spatial and temporal habitat variability. We used fourteen habitat types and sixteen species traits in this analysis.
• 2 When examined by correspondence analysis, species traits separate the Naididae (with a higher potential for reproduction, small size, high mobility, and opportunistic diet) from all other families.
• 3 Habitat utilization by oligochaetes demonstrates two gradients: a vertical gradient that arranges species by their affinity for interstitial habitats (stygophily) and a transversal gradient that arranges them by their affinity for main channel habitats (rheophily).
• 4 No significant relationship was found between species traits and habitat utilization in a co-inertia analysis.
• 5 Trends observed for species traits within the framework of spatial-temporal habitat variability show only minor agreement with predictions of the river habitat templet.
• 6 Species richness is generally higher in superficial and interstitial habitats that are permanently connected with the main channel, and peaks in the superficial parapotamons (backwaters that are permanently connected with the main channel) characterized by intermediate levels of spatial as well as temporal variability; this pattern only partially fits with predictions of the patch dynamics concept.

     

Theoretical habitat templets, species traits, and species richness: aquatic Coleoptera in the Upper Rhône River and its floodplain

• 1 To test predictions of the river habitat templet and the patch dynamics concept, trends in species traits and species richness of aquatic beetles were related to the spatial-temporal variability of eighteen habitat types in the alluvial floodplain of the French Upper Rhône River. One hundred and twenty species of beetles were used in this analysis.
• 2 The basic information was obtained either from the literature (for most of the species traits) or from observations made at approximately 500 sampling sites in the Brégnier-Cordon and Jons sections over the past 19 years (for habitat utilization). This information was structured by a fuzzy coding technique and examined by ordination analyses.
• 3 Analyses of the relationships among nineteen species traits revealed a clear distinction according to traits such as body form (for adults), functional feeding type and food (adults and larvae), attachment to the substrate and dissemination potential (adults and larvae), and patterns of aquatic and/or terrestrial life of adults and larvae. Species traits such as number of descendants per reproductive cycle, and number of reproductive cycles per year or per individual showed less contrast, because these traits are rather homogeneous in aquatic beetles.
• 4 Analyses of the habitat utilization by the aquatic beetles revealed a vertical gradient that separates interstitial from superficial habitats, and a transverse gradient for the superficial habitats, which extends from the main channel towards permanent oxbow lakes and temporary waters.
• 5 The significant relationship betweeen species traits and habitat utilization demonstrates that most beetle species use a particular set of habitat types with a particular set of species trait modalities.
• 6 Species traits of aquatic beetles are homogeneous but evidently very successful and are adapted to many potential conditions of spatial–temporal variability. Because of this homogeneity, observations on aquatic beetles do not support trends of traits in the framework of spatial–temporal variability predicted from the river habitat templet.
• 7 The observed species richness of aquatic beetles is low in habitat types with a low spatial–temporal variability, increases as spatial variability increases, and tends to be highest at intermediate temporal variability. This pattern matches predictions of the patch dynamics concept.

     

Theoretical habitat templets, species traits, and species richness: aquatic insects in the Upper Rhône River and its floodplain

• 1 For five orders of Insecta (Plecoptera, Ephemeroptera, Odonata, Trichoptera, and Coleoptera) in two sites of the Upper Rhône River (France), the following are examined: (i) relationships among nineteen species traits; (ii) habitat utilization of species; (iii) the relationship between species traits and habitat utilization; and (iv) trends of species traits and species richness in a templet of spatial-temporal habitat variability.
• 2 The species traits having the highest correlations correspond to reproduction, life cycle, nutritional, and morphological features. Species trait characteristics of Coleoptera are distinctly contrasted with those of Plecoptera and Ephemeroptera; Odonata and Trichoptera are intermediate to these orders.
• 3 The distribution of species in fourteen habitat types of the Upper Rhône River floodplain demonstrates a transverse gradient from the main channel to the oxbow lakes and the temporary water habitats, and a vertical gradient from interstitial to superficial habitats.
• 4 Despite a significant relationship between species traits and habitat utilization, superposition between species traits and habitat utilization is limited. At the order level, species form usually one (Ephemeroptera, Trichoptera, Odonata) or several (Coleoptera) groups of relatively homogeneous species traits; however, the species of each of these groups utilize rather different habitat types.
• 5 Only for some life history traits, e.g. the minimum age of reproduction or the number of reproductive cycles per year, do the trends observed in the framework of spatial—temporal variability of habitat types agree with the predictions from the river habitat templet. This mismatch mainly results from the unique phylogenetic history of the Coleoptera compared with that of the other four orders.
• 6 Species richness peaks at an intermediate level of temporal variability; however, it does not gradually increase with increasing spatial variability, nor increase from low to intermediate temporal variability.

     

Theoretical habitat templets, species traits, and species richness: aquatic macrophytes in the Upper Rhône River and its floodplain

• 1 The objective of this study, which is based on forty-two species of hydrophytes and helophytes, is to investigate: (i) relationships among species traits; (ii) habitat utilization by species; (iii) the relationship between species traits and habitat utilization; (iv) trends in species traits in the framework of spatial–temporal habitat variability, and if trends match predictions from the river habitat templet; and (v) trends in species richness in the framework of spatial–temporal habitat variability, and if trends match predictions of the patch dynamics concept.
• 2 Two data sets were used for this analysis: species traits (mainly reproductive and morphological characteristics) were documented from the literature; and species distribution across eight habitat types was from field surveys conducted in the floodplain of the Upper Rhone River, France. This information was structured by a fuzzy coding technique and analysed by ordination methods.
• 3 Several species traits, which are related to disturbances and reflect resistance (e.g. attachment to soil or substrate) or resilience (e.g. potential for regeneration of an individual), are closely related for aquatic macrophytes.
• 4 Habitat utilization by aquatic macrophytes separates the habitat types along a gradient of connectivity with the main channel, which corresponds to a gradient in flood disturbance frequency and the permanence of the different water-bodies.
• 5 The relationship between species traits and habitat utilization is highly significant, indicating that a particular set of habitat types is used by taxa with a particular set of species trait modalities.
• 6 Observations in one habitat templet (in which scaling of the templet is primarily based on water level fluctuations for the temporal variability axis and on substrate characteristics for the spatial variability axis) generally do not support predictions on trends in species traits but do support predictions on trends in species richness.
• 7 Observations in an alternative habitat templet (in which scaling of the templet is based on frequency of flood scouring for the temporal variability axis and on heterogeneity of the substrate for the spatial variability axis) support theoretical predictions on trends for about half of the species traits for which predictions were available. However, trends in species richness in this alternative habitat templet are only partly in agreement with predictions.

     

Theoretical habitat templets, species traits, and species richness: 548 plant and animal species in the Upper Rhône River and its floodplain

• 1 For practical reasons, conceptual developments in community ecology are usually based on studies of a restricted systematic group. The cooperation of thirty or so specialists in the synthesis of long-term ecological research on the Upper Rhône River, France, provided a unique occasion to investigate relationships among species traits, the habitat utilization by species, the relationship between species traits and habitat utilization, and trends of species traits and species richness in the framework of spatial-temporal habitat variability for 548 species of plants (Hyphomycetes, aquatic macrophytes, floodplain vegetation) and animals (Tricladida, Oligochaeta, several groups of Crustacea, Insecta and Vertebrata).
• 2 Using correspondence analysis, 100 modalities of eighteen species traits were examined; the resulting typology demonstrates that systematic groups are the most important elements for separating species traits such as size, fecundity of individuals, parental care, mobility, body form, and food type. Small species have an intermediate number of descendants per reproductive cycle and few reproductive cycles both per year and per individual; in contrast, large species have a high number of descendants per reproductive cycle and few reproductive cycles per year but many potential reproductive cycles per individual.
• 3 The analysis of habitat utilization in the Upper Rhône River and its floodplain by the 548 species demonstrated a vertical gradient separating interstitial from superficial habitats; a transverse gradient for superficial habitats from the main channel towards more terrestrial ones is also evident.
• 4 Because of a significant (P < 0.01) relationship between species traits and habitat utilization, traits such as size, fecundity of individuals, parental care, tolerance to variation in humidity, and respiration are arranged along the vertical and transverse habitat gradient. Size, the number of reproductive cycles per individual, and the tolerance to variation of humidity increases from permanent waters to temporary waters, aggrading habitats, and terrestrial habitats.
• 5 Species traits showed significant (P < 0.01) trends in the framework of spatial-temporal habitat variability and were compared with predictions based on the river habitat templet. Although each habitat showed a mixture of species traits at low temporal and spatial variability, and at high variability sites, trends corresponded to predictions for three traits (number of descendants per reproductive cycle, number of reproductive cycles per individual, attachment to soil or substrate) along a gradient of increasing temporal habitat variability.
• 6 The species richness of each habitat within the Upper Rhône River and its floodplain significantly (P = 0.03) increased as the spatial variability of habitats increased but there is no statistical correlation between spedes richness and temporal variability. An altemative hypothesis predicting that fewer spedes per resource occur in temporally stable habitats is also not supported.

     

Theoretical habitat templets, species traits, and species richness: Plecoptera and Ephemeroptera in the Upper Rhône River and its floodplain

• 1 Ephemeroptera and Plecoptera in two sites of the Upper Rhône River (France) were examined using multivariate analyses to determine: (i) relationships among seventeen species traits; (ii) habitat utilization of the fifty-five species present; (iii) the relationship between species traits and habitat utilization; (iv) trends of species traits and species richness in a framework of spatial and temporal habitat variability.
• 2 The species traits having the highest correlation ratios correspond to reproduction or life cycle, behavioural, and morphological characteristics. According to their traits, species of Baetidae, Caenidae, and Leptophlebiidae (Ephemeroptera) are opposite species of Perlidae and Perlodidae (Plecoptera).
• 3 The distribution of species in thirteen habitat types of the Upper Rhône River floodplain demonstrates a transverse gradient from the main channel to the oxbow lakes. Plecoptera are restricted to the different main channel habitats; in contrast, Ephemeroptera families have a broader distribution with Baetidae and Leptophlebiidae occurring in most floodplain habitats.
• 4 Plecoptera exhibit a significant relationship between species traits and habitat utilization but no relationship is evident for Ephemeroptera. Baetidae use many habitat types and have diverse species traits; in contrast, Leptophlebiidae, Heptageniidae, and Caenidae use many habitat types but each family has a rather uniform set of traits.
• 5 Trends in species traits were significantly related to both the spatial and temporal variability of habitats. Considering only temporal variability, the distribution of species trait modalities (= categories) corresponded well to predictions on trends in the river habitat templet for ‘minimum age at reproduction’ and ‘potential longevity’, and in general for ‘descendants per reproductive cycle’, ‘reproductive cycles per year’, ‘potential size’, and ‘body flexibility’ trends in six other traits did not match predictions.
• 6 No trends in species richness were evident in spatial–temporal framework of habitat variability.

     

The ontogeny of bone growth in two species of dormice: Reconstructing life history traits

Though bone histology has become a powerful tool to reconstruct life history strategies and physiology in living and extinct reptiles and amphibians, it is of limited use in mammals. Dormice (Myoxidae) are good candidates for assessing the relation between bone microstructure and life history due to their long life span, marked physiological cycles and negligible bone remodelling. We carried out the most comprehensive study so far analyzing 16 wild individuals of unknown age belonging to two different species of dormice, Glis glis and Eliomys quercinus. Our study shows a high degree of consistency in the number of resting lines present in bones of the same individual, with femora providing the most accurate age estimations. Moreover, the presence of a single LAG in some juveniles allows discerning between offspring from different reproductive events (early or late litters).     

Interactions between alien plant species traits and habitat characteristics in agricultural landscapes in Finland

The survival and success of alien plant species is determined by species traits (i.e., invasiveness) and the characteristics of the habitats in the region of introduction (i.e., invasibility). However, little is known about species traits as related to habitat characteristics. We assessed the characteristics of successful invaders and the interaction of environmental factors and life-history traits for alien plant species. The vascular plants were recorded from 52 agricultural landscapes in Finland. We compared the traits of native and alien plant species with Fisher’s exact test and used a three table ordination analysis, RLQ analysis, to relate species traits to environmental conditions. Species were clustered according to their position on the RLQ axes, and the clusters were tested for phylogenetic independence. The successful alien plant species were associated with life form and preferences for moisture and nitrogen, but the trait composition varied according to the habitat type. Two RLQ axes explained 80.5% of the variation, and the species traits were significantly associated with environmental variables. The clustering showed that the occurrence of alien plant species in agricultural habitats was driven by invasion history, traits related to dispersal (dispersal type, seed mass) and habitat preferences, as well as environmental features, such as geographical location, temperature and the quality and disturbance regime of the habitats. All clusters were phylogenetically non-independent. Thus, the clusters of alien species comprised species of diverse taxonomic affinities, although, they shared the traits explaining their occurrence in particular habitats. This information is useful for understanding the link between species traits and the environmental conditions of the habitats, and complexity of the invasion process.