Taxonomie, écologie et lutte biologique

Summary It is impossible to study modern taxonomy without taking into consideration the population genetic and ecology. Thé Linnean concept of monotypic species should be replaced by the biometric study of the frequency of the variability of the biological characters (particularly morphological) used in taxonomy. The study of populations offers a larger scientific interest than the one of isolated types. It gives the possibility of caracterizing the evolution stability of the species. So, Taxonomy can help considerably in biological control. By revealing large variability species, it gives a criterion of their adjusting plasticity. But, by definition, the biological control is based on the adjusting capacity of the entomophagous insects either to a new host or to a new environment. Information given by the taxonomist on the species stability has consequently a great value in the choice and use of the Entomophagous insects. — Ecologists using biological control, expect from the taxonomist to lie more than a label, which implies the use of biometric analysis methods in Entomophagous taxonomy.      

The What,Why and How of Primate Taxonomy

Taxonomy has a well-defined role, which is much more than simply stamp-collecting and pigeon-holing. Species are the units of classification, biogeography and conservation; as such they must be defined as objectively as possible. The biological species concept, still widely used in biology, though predominantly by non-taxonomists and all too often misunderstood, is a process-based concept, which offers no criterion for the classification of allopatric populations beyond inference and hypothesis. The phylogenetic species concept—a pattern-based concept—is as nearly objective as we are likely to get. Amount of difference is not a criterion for recognizing species. It is not possible to insist on monophyly at the specific level, but it is mandatory for the higher categories (genus, family, etc.). The rank we assign to a given supraspecific category should be determined by its time depth.     

Interbreeding, monophyly and the genetic yardstick: species concepts in parasites

The biological species concept defines species on the criterion of interbreeding. This may not be applicable to many parasites that are capable of self-fertilization and asexual reproduction. In this review, Alan Lymbery explores alternative concepts that may be applied to recognize species in such groups, using the cestode genus Echinococcus as an example. Two conclusions can be drawn. First, that the applicability of the biological species concept should not be dismissed without some knowledge o f the frequency of interbreeding in natural populations. Second, that where interbreeding is absent or rare, species should be delimited on the basis o f both monophyletic origin and genetic distinctness.     

Cryptic speciation in Fusarium subglutinans

Fusarium isolates that form part of the Gibberella fujikuroi species complex have been classified using either a morphological, biological, or phylogenetic species concept. Problems with the taxonomy of Fusarium species in this complex are mostly experienced when the morphological and biological species concepts are applied. The most consistent identifications are obtained with the phylogenetic species concept. Results from recent studies have presented an example of discordance between the biological and phylogenetic species concepts, where a group of F. subglutinans sensu stricto isolates, i.e., isolates belonging to mating population E of the G. fujikuroi complex, could be sub-divided into more than one phylogenetic lineage. The aim of this study was to determine whether this sub-division represented species divergence or intraspecific diversity in F. subglutinans. For this purpose, we included 29 F. subglutinans isolates belonging to the E-mating population that were collected from either maize or teosinte, from a wide geographic range. DNA sequence data for six nuclear regions in each of these isolates were obtained and used in phylogenetic concordance analyses. These analyses revealed the presence of two major groups representing cryptic species in F. subglutinans. These cryptic species were further sub-divided into a number of smaller groups that appear to be reproductively isolated in nature. This suggests not only that the existing F. subglutinans populations are in the process of divergence, but also that each of the resulting lineages are undergoing separation into distinct taxa. These divergences did not appear to be linked to geographic origin, host, or phenotypic characters such as morphology.     

The spatial analysis of biological interactions: morphological variation responding to the co‐occurrence of competitors and resources

By sharing geographic space, species are forced to interact with one another and the contribution of this process to evolutionary and ecological patterns of individual species is not fully understood. At the same time, species turnover makes that species composition varies from one area to another, so the analysis of biological interaction cannot be uncoupled from the spatial context. This is particularly important for clades that show high degree of specialization such as hummingbirds, where any variation in biotic pressures might lead to changes in morphology. Here, we describe the influence of biological interactions on the morphology of Hylocharis leucotis by simultaneously considering potential competition and diet resources. We characterized the extent of local potential competition and local available floral resources by correlating two measurements of hummingbird diversity, floral resources and the size of morphological space of H. leucotis along its geographic distribution. We found that H. leucotis shows an important morphological variability across its range and two groups can be recognized. Surprisingly, morphological variation is not always linked to local hummingbird richness or the phylogenetic similarity of. Only in the southern part of its distribution, H. leucotis is morphologically more variable in those communities where it coexist with closely related hummingbird species. We also found that morphological variation in H. leucotis is independent from the availability of floral resources. Our results suggest that abiotic factors might be responsible for morphological differences across populations in Hylocharis leucotis being biological interactions of minor importance.     

Comparison of Biological,Molecular, and Morphological Methods of Species Identification in a Set of Cultured Panagrolaimus Isolates

We have developed a molecular barcode system that uses the small subunit ribosomal RNA (SSU) sequence to define molecular operational taxonomic units (MOTU) of soil nematodes. Here we attempt to differentiate five cultured isolates of a taxonomically difficult genus, Panagrolaimus, using morphological, molecular, and biological (breeding) criteria. The results indicated that the five culture populations belonged to two reproductively isolated species. The available morphological criteria, including scanning electron microscopy (SEM), were insufficient to differentiate among them, and all five could be classified as one morphospecies. Within-culture variation of the morphometrical data did not discern between the two biological species. Sequence data clearly separated the populations into two groups that supported the breeding results. Given this study represented only five populations of one genus, we suggest a congruence of MOTU analysis with the biological species concept. This multifaceted approach is promising for future identification of nematodes as it is simple, comparable, and transferable.     

Systematics,reproductive isolation and species boundaries in monogonont rotifers

The typological concept of rotifer species and the morphological basis of rotifer systematics is reviewed and alternatives proposed. Occasional sexuality in the cyclical parthenogenetic life cycle of monogononts permits application of the biological species concept to this group. Data from cross-mating experiments with Asplanchna, Brachionus and Epiphanes illustrate the usefulness of reproductive isolation as a criterion for species boundaries. Populations from different geographic regions are often interfertile indicating that rotifer species are genetically integrated over wide areas. The main types of isolating mechanisms operating in monogononts are reviewed. The role of behavioral reproductive isolation in maintaining species boundaries is examined. The use of a mate recognition bioassay which estimates the probability of copulation and quantifies the degree of isolation is described. Recent work of the mechanism of mate recognition is reviewed. It is concluded that the biological species concept is applicable to rotifers and that a more experimental approach to determining species boundaries is both feasible and desirable.     

Speciation in the fossil record

It is easy to claim that the fossil record says nothing about speciation because the biological species concept (which relies on interbreeding) cannot be applied to it and genetic studies cannot be carried out on it. However, fossilized organisms are often preserved in sufficient abundance for populations of intergrading morphs to be recognized, which, by analogy with modern populations, are probably biological species. Moreover, the fossil record is our only reliable documentation of the sequence of past events over long time intervals: the processes of speciation are generally too slow to be observed directly, and permanent reproductive isolation can only be verified with hindsight. Recent work has shown that some parts of the fossil record are astonishingly complete and well documented, and patterns of lineage splitting can be examined in detail. Marine plankton appear to show gradual speciation, with subsequent morphological differentiation of lineages taking up to 500000 years to occur. Marine invertebrates and vertebrates more commonly show punctuated patterns, with periods of rapid speciation followed by long-term stasis of species lineages.     

Variation in infection parameters among populations within parasite species: intrinsic properties versus local factors

Within any parasite species, variation among populations in standard infection parameters (prevalence, intensity and abundance) is an accepted fact. The proportion of hosts infected and the mean number of parasites per host are not fixed values across the entire geographic range of any parasite species. The question is whether this inter-population variation occurs within a narrow, species-specific range and is thus driven mainly by the biological features of the parasite, or whether it is substantial and unpredictable, leaving population parameters at the mercy of local conditions. Here, the repeatability of estimates of prevalence, intensity and abundance of infection was assessed across populations of the same parasite species, for 77 metazoan parasite species of Canadian freshwater fishes. Overall, parameter values from different populations of the same parasite species were more similar to each other and more different from those of other species, than expected by chance alone. Much of the variation in parameter values in the dataset was associated with differences between parasite species, rather than with differences among populations within species. This was particularly true for intensity and abundance of infection; in contrast, prevalence values, while somewhat repeatable among populations of the same species, still showed considerable variation. Among the higher taxa investigated (monogeneans, trematodes, cestodes, nematodes, acanthocephalans, copepods), there was no evidence that species of one taxon display intrinsically greater variation in population parameters than species of other taxa. Overall, the results suggest that intensity and abundance of infection are real species characters, though somewhat variable. This conclusion supports the view that the biological features of parasite species can potentially override local environmental conditions in driving parasite population dynamics.     

Putting the Biological Species Concept to the Test: Using Mating Networks to Delimit Species

Although interfertility is the key criterion upon which Mayr’s biological species concept is based, it has never been applied directly to delimit species under natural conditions. Our study fills this gap. We used the interfertility criterion to delimit two closely related oak species in a forest stand by analyzing the network of natural mating events between individuals. The results reveal two groups of interfertile individuals connected by only few mating events. These two groups were largely congruent with those determined using other criteria (morphological similarity, genotypic similarity and individual relatedness). Our study, therefore, shows that the analysis of mating networks is an effective method to delimit species based on the interfertility criterion, provided that adequate network data can be assembled. Our study also shows that although species boundaries are highly congruent across methods of species delimitation, they are not exactly the same. Most of the differences stem from assignment of individuals to an intermediate category. The discrepancies between methods may reflect a biological reality. Indeed, the interfertility criterion is an environment-dependant criterion as species abundances typically affect rates of hybridization under natural conditions. Thus, the methods of species delimitation based on the interfertility criterion are expected to give results slightly different from those based on environment-independent criteria (such as the genotypic similarity criteria). However, whatever the criterion chosen, the challenge we face when delimiting species is to summarize continuous but non-uniform variations in biological diversity. The grade of membership model that we use in this study appears as an appropriate tool.     

Limited gene flow may enhance adaptation to local optima in isolated populations of the Roesel's bush cricket (Metrioptera roeselii)

Variation in morphological traits along latitudinal gradients often manifests as size clines. In insects, both positive and negative correlations are seen, and the mechanism behind the response is unclear. We studied variation in seven morphological traits of Roesel's bush cricket, Metrioptera roeselii, sampled from seven latitude-matched-pair populations that were either geographically isolated from or connected to the species continuous distribution range. The aim was to examine whether morphological traits differed between isolated and continuous populations, and whether latitudinal variation was apparent. The data were used to indicate whether variation in trait means originates from plastic responses to the environment or genetic adaptation to local conditions. To evaluate the influence of gene flow on trait means, we analysed the genetic variation in seven microsatellites. Data showed that individuals from isolated populations display a positive relationship between latitude and body size, whereas individuals from continuous populations show little or no such relationship. The combined morphological and genetic data suggest that the isolated populations have adapted to local optima, while gene flow between continuous populations appears to counteract this process.     

The debate about the biological species concept - a review

The importance of the species concept in biology has led to a continuing debate about the definition of species. This paper summarizes the recent literature in relation to the ‘biological species concept’ (MAYR 1942). Among the general attributes demanded, possible limitations of the universality and applicability of a species definition are discussed. Three different areas of criticism of the biological species concept are considered: 1. The impracticability of the criterion of reproductive isolation. The demand for more practical criteria is rejected, because reproductive isolation is seen as the factor that produces and maintains species as discrete entities in nature. 2. The inapplicability to non-bisexual organisms. A brief survey of modes of uniparental reproduction and their relative importance suggests that obligatory apomicts are of little evolutionary significance. 3. The inapplicability to multidimensional situations. Despite practical difficulties, the biological species concept is held to apply to organisms separated in space. The impossibility to delimit species in time by reproductive isolation is recognized. Out of two ways to divide continuous evolutionary lineages in time, the phylogenetic approach, which considers only speciation events (cladogenesis), is preferred as it is more objective. A list of recently published alternative definitions of species, none of which is found acceptable, is given. It is concluded that the biological species concept needs not be changed or dismissed on the basis of the discussed criticisms.     

Studies on sexual compatibility betweenEctocarpus siliculosus (Phaeophyceae) from Chile and the Mediterranean Sea

Clonal isolates of the brown algaEctocarpus siliculosus (Ectocarpales) originating from Naples (Mediterranean Sea) and southern Chile were compared in laboratory culture studies. The two isolates showed distinctly different morphological characters, but very similar details of life history and sexual reproduction. Gametes are sexually compatible; hybrid zygotes are formed and sporophytes develop, which are fertile on the basis of mitotic spores. However, unilocular sporangia were abortive, indicating segregational sterility caused by chromosomal mismatch during meiosis. Although the biological species concept in a strict sense does not apply, and appreciable morphological variability exists in this cosmopolitan taxon, local populations are considered as representatives of the same species. Dedicated to Dr. Dr. h. c. Peter Kornmann on the occasion of his eightieth birthday.     

Gen‐morph species concept—A new and integrative species concept for outbreeding organisms

The present article briefly reviewed the prevailing species concepts, especially biological, genetic, evolutionary, phylogenetic, ecological, and several taxonomic species concepts. The former five reflect the properties of species from diverse aspects and in different degrees, while taxonomic species concepts all contain more or less subjective elements, except for Hedberg's taxonomic method (not species concept). So far, there is no species concept that is both theoretically rational and practically operable. The present article outlined recent studies on the genus Paeonia L. (Paeoniaceae) in biology, particularly in morphology, biogeography, molecular phylogeny, and reproductive behavior, which provided insight into the relationship between variation of morphological characteristics and phylogeny. Taking the study on Paeonia L. as a case, referring to studies on some other plant groups, and incorporating the merits of the prevailing species concepts into our consideration, “gen‐morph species concept” is proposed here formally as new for outbreeding organisms. The new species concept has three special features: (i) a bridge linking morphological aspect with genetic and other aspects of species; (ii) proposal of a concrete morphological criterion for species definition, and (iii) considering quantitative and qualitative characteristics as equally valuable for species definition and introducing statistics into the concept to handle such characteristics. Therefore, the gen‐morph species concept is an integrative species concept, both theoretically objective and practically operable.     

Geographic variation and plasticity to water and nutrients in Pelargonium australe

Here, patterns of phenotypic plasticity and trait integration of leaf characteristics in six geographically discrete populations of the perennial herb Pelargonium australe were compared. It was hypothesized that populations would show local adaptation in trait means, but similar patterns of plasticity and trait integration. Further, it was questioned whether phenotypic plasticity was positively correlated with environmental heterogeneity and whether plasticity for water-use traits in particular was adaptive. Seedlings were grown in a glasshouse at six combinations of water and nutrient availability. Leaf anatomical, morphological and gas exchange traits were measured. High amounts of plasticity in leaf traits were found in response to changes in growth conditions and there was evidence of local adaptation among the populations. While there were significant correlations between plasticity and environmental heterogeneity, not all were positive. Notably, patterns of plasticity and trait integration varied significantly among populations. Despite that variation, some of the observed plasticity was adaptive: fitness was correlated with conservative water use when water was limiting. Pelargonium arrived in Australia approximately 5 million yr ago. It is concluded here that high amounts of plasticity, in some cases adaptive, and weak integration among traits may be key to the spread and success of this species.     

Should I stay or should I go? The Ectodysplasin locus is associated with behavioural differences in threespine stickleback

Adaptive divergence may be facilitated if morphological and behavioural traits associated with local adaptation share the same genetic basis. It is therefore important to determine whether genes underlying adaptive morphological traits are associated with variation in behaviour in natural populations. Positive selection on low-armour alleles at the Ectodysplasin (Eda) locus in threespine stickleback has led to the repeated evolution of reduced armour, following freshwater colonization by fully armoured marine sticklebacks. This adaptive divergence in armour between marine and freshwater populations would be facilitated if the low allele conferred a behavioural preference for freshwater environments. We experimentally tested whether the low allele is associated with preference for freshwater by measuring the preference of each Eda genotype for freshwater versus saltwater after acclimation to either salinity. We found no association between the Eda low allele and preference for freshwater. Instead, the low allele was significantly associated with a reduced preference for the acclimation environment. This behaviour may facilitate the colonization of freshwater habitats from the sea, but could also hinder local adaptation by promoting migration of low alleles between marine and freshwater environments.     

Strict ESS for n-species systems

A system of n asexual populations is considered where both intra- and interspecific frequency-dependent game conflicts with lack of information take place. The concept of a strict n-species ESS is introduced which implies local asymptotic stability of the replicator dynamics of pure phenotypes. The dynamical concept of strict stability is also introduced which turns out to be equivalent to the strict n-species ESS concept. The above notions are also related to similar concepts considered in the literature for the same biological situation.     

Co-occurrence patterns and morphological similarity of semiaquatic insects (Hemiptera: Gerromorpha) in streams of Eastern Amazonia

1. To understand how the semiaquatic bug communities are shaped, it was first verified whether there was a pattern of co-occurrence between species. It was subsequently tested whether the pattern found was related to environmental variables. Lastly, it was verified whether morphological divergence between species was present in the morphological and functional traits selected. 2. A pattern of species co-occurrence was found in the studied assemblage, related to both environmental variables and interspecific biological interactions. Pairs of species with negative, positive, and random co-occurrence were found. Traits related to predation competition, sexual competition, and interaction with the environment showed significant morphological divergence. 3. Therefore, more than one process defines species co-occurrence patterns in semiaquatic bug communities. It is suggested that environmental influence is related to species microhabitat preference. On the other hand, the morphological divergence found is related to competition and sharing of food resources.     

Morphological Consequences of Developmental Plasticity in <Emphasis Type="Italic">Rana temporaria</Emphasis> are not Accommodated into Among-Population or Among-Species Variation

Environmental induced developmental plasticity occurs in many organisms and it has been suggested to facilitate biological diversification. Here we use ranid frogs to examine whether morphological changes derived from adaptive developmental acceleration in response to pool drying within a species are mirrored by differences among populations and across species. Accelerated development in larval anurans under pool drying conditions is adaptive and often results in allometric changes in limb length and head shape. We examine the association between developmental rate and morphology within population, among populations in divergent environments, and among species inside the Ranidae frog family, combining experimental approaches with phylogenetic comparative analyses. We found that frogs reared under decreasing water conditions that simulated fast pool drying had a faster development rate compared to tadpoles reared on constant water conditions. This faster developmental rate resulted in different juvenile morphologies between the two pool drying conditions. The association between developmental rate and morphology found as a result of plasticity was not mirrored by differences among populations that differed in development, neither was it mirrored among species that differed in development rate. We conclude that morphological differences among populations and species were not driven by variation in developmental time per se. Instead, selective factors, presumably operating on locomotion and prey choice, seem to have had a stronger evolutionary effect on frog morphology than evolutionary divergences in developmental rate in the ranid populations and species studied.     

Climate change effects on behavioral and physiological ecology of predator–prey interactions: Implications for conservation biological control

Habitat management under the auspices of conservation biological control is a widely used approach to foster conditions that ensure a diversity of predator species can persist spatially and temporally within agricultural landscapes in order to control their prey (pest) species. However, an emerging new factor, global climate change, has the potential to disrupt existing conservation biological control programs. Climate change may alter abiotic conditions such as temperature, precipitation, humidity and wind that in turn could alter the life-cycle timing of predator and prey species and the behavioral nature and strength of their interactions. Anticipating how climate change will affect predator and prey communities represents an important research challenge. We present a conceptual framework—the habitat domain concept—that is useful for understanding contingencies in the nature of predator diversity effects on prey based on predator and prey spatial movement in their habitat. We illustrate how this framework can be used to forecast whether biological control by predators will become more effective or become disrupted due to changing climate. We discuss how changes in predator–prey interactions are contingent on the tolerances of predators and prey species to changing abiotic conditions as determined by the degree of local adaptation and phenotypic plasticity exhibited by species populations. We conclude by discussing research approaches that are needed to help adjust conservation biological control management to deal with a climate future.