Can environmental fluctuation prevent competitive exclusion in sympatric flycatchers?

Ecology has been characterized by a central controversy for decades: namely, whether the distribution and abundance of organisms are determined by species interactions, such as competitive exclusion, or by environmental conditions. In part, this is because competitive exclusion has not been convincingly demonstrated in open, natural systems. In addition, traditional theoretical models cannot predict the outcome of competitive interactions in the presence of environmental variability. In this paper we document the limiting influence of strong interspecific competition on population dynamics and nestling mortality in a mixed population of pied flycatchers (Ficedula hypoleuca) and collared flycatchers (F. albicollis in a narrow zone of sympatry. Whereas the former species was limited mainly by interspecific competition, the latter species was limited by the concerted influences of intraspecific competition and climate. The analysis suggests a march towards competitive exclusion of the pied flycatcher during warm periods. However, competitive exclusion is apparently prohibited on a local scale because intraspecific competition among individual collared flycatchers intensifies when they are forced to cope with severe environmental conditions, promoting the temporary and local presence of pied flycatchers.     

How do natural and sexual selection contribute to sympatric speciation?

I use explicit genetic models to investigate the importance of natural and sexual selection during sympatric speciation and to sort out how genetic architecture influences these processes. Assortative mating alone can lead to speciation, but rare phenotypes' disadvantage in finding mates and intermediate phenotypes' advantage due to stabilizing selection strongly impede speciation. Any increase in the number of loci also decreases the likelihood of speciation. Sympatric speciation is then harder to achieve than previously demonstrated by many theoretical studies which assume no mating disadvantage for rare phenotypes and consider a small number of loci. However, when a high level of assortative mating evolves, sexual selection might allow populations to split into dimorphic distributions with peaks corresponding to nearly extreme phenotypes. Competition then works against speciation by favouring intermediate phenotypes and preventing further divergence. The interplay between natural and sexual selection during speciation is then more complex than previously explained.     

When do parasites fail to speciate in response to host speciation?

Cospeciation generally increases the similarity between host and parasite phylogenies. Incongruence between host and parasite phylogenies has previously been explained in terms of host switching, sorting, and duplication events. Here, we describe an additional process, failure of the parasite to speciate in response to host speciation, that may be important in some host-parasite systems. Failure to speciate is likely to occur when gene flow among parasite populations is much higher than that of their hosts. We reconstructed trees from mitochondrial and nuclear DNA sequences for pigeons and doves (Aves: Columbiformes) and their feather lice in the genus Columbicola (Insecta: Phthiraptera). Although comparisons of the trees from each group revealed a significant amount of cospeciation, there was also a significant degree of incongruence. Cophylogenetic analyses generally indicated that host switching may be an important process in the history of this host-parasite association. Using terminal sister taxon comparisons, we also identified three apparent cases where the host has speciated but the associated parasite has not. In two of these cases of failure to speciate, these comparisons involve allopatric sister taxa of hosts whose lice also occur on hosts sympatric with both of the allopatric sisters. These additional hosts for generalist lice may promote gene flow with lice on the allopatric sister species. Relative rate comparisons for the mitochondrial cytochrome oxidase I gene indicate that molecular substitution occurs about 11 times faster in lice than in their avian hosts.     

What,if anything,is sympatric speciation?

Sympatric speciation has always fascinated evolutionary biologists, and for good reason; it pits diversifying selection directly against the tendency of sexual reproduction to homogenize populations. However, different investigators have used different definitions of sympatric speciation and different criteria for diagnosing cases of sympatric speciation. Here, we explore some of the definitions that have been used in empirical and theoretical studies. Definitions based on biogeography do not always produce the same conclusions as definitions based on population genetics. The most precise definitions make sympatric speciation an infinitesimal end point of a continuum. Because it is virtually impossible to demonstrate the occurrence of such a theoretical extreme, we argue that testing whether a case fits a particular definition is less informative than evaluating the biological processes affecting divergence. We do not deny the importance of geographical context for understanding divergence. Rather, we believe this context can be better understood by modelling and measuring quantities, such as gene flow and selection, rather than assigning cases to discrete categories like sympatric and allopatric speciation.     

Year class coexistence or competitive exclusion for strict biennials?

 We consider a discrete time model of semelparous biennial population dynamics. Interactions between individuals are modelled with the aid of an ``environmental' variable I. The impact on and the sensitivity to the environmental condition is age specific. The main result is that competitive exclusion between the year classes is possible as is their coexistence. For moderate values of the basic reproduction ratio R ₀ there is a strict dichotomy: depending on the other parameters we either find competitive exclusion or coexistence. We characterize rather precisely the patterns of age specific impact and sensitivity that lead to either of these outcomes. Received: 13 July 2001 / Revised version: 26 June 2002 / Published online: 19 November 2002 Key words or phrases: Competitive exclusion – Semelparous species – Periodical insects     

From elaborate to compact seasonal plant epidemic models and back: is competitive exclusion in the details?

Seasonality, or periodic host absence, is a central feature in plant epidemiology. In this respect, seasonal plant epidemic models take into account the way the parasite overwinters and generates new infections. These are termed primary infections. In the literature, one finds two classes of models: high-dimensional elaborate models and low-dimensional compact models, where primary infection dynamics are explicit and implicit, respectively. Investigating a compact model allowed previous authors to show the existence of a competitive exclusion principle. However, the way compact models derive from elaborate models has not been made explicit yet. This makes it unclear whether results such as competitive exclusion extend to elaborate models as well. Here, we show that assuming primary infection dynamics are fast in a standard elaborate model translates into a compact form. Yet, it is not that usually found in the literature. Moreover, we numerically show that coexistence is possible in this original compact form. Reversing the question, we show that the usual compact form approximates an alternate elaborate model, which differs from the earlier one in that primary infection dynamics are density dependent. We discuss to which extent these results shed light on coexistence within soil- and air-borne plant parasites, such as within the take-all disease of wheat and the grapevine powdery mildew cryptic species complexes, respectively.     

Incipient speciation driven by phenotypic plasticity? Evidence from sympatric populations of Arctic charr

Recent models suggest that the existence of environmentally induced polymorphisms within a single population (especially those related to foraging) facilitates the process of evolutionary divergence within a single gene pool by generating distinct phenotypic modes that are exposed to differential selection. In order to test a prediction of the phenotypic plasticity model of divergence, we used a well-documented polymorphism to disentangle the relative effects of morph and rearing environment in generating phenotypic variance. We reared first-generation offspring of two sympatric morphs of Arctic charr Salvelinus alpinus in the laboratory and compared their head morphology with that of their wild parents. Morphological characters with a known functional role in foraging were highly plastic. Rearing environment accounted for the largest component of the variation in expressed phenotype, but this environmental effect overlaid a clear (but small) genetic effect. We conclude that phenotypic plasticity has played a significant role in the evolution of this trophic polymorphism, but that the evolutionary process has progressed to the point that the gene pool is now segregated.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 611–618.     

The Midas cichlid species complex: incipient sympatric speciation in Nicaraguan cichlid fishes?

Abstract Sympatric speciation is a contentious concept, although theoretical models as well as empirical evidence support its relevance in evolutionary biology. The Midas cichlid species complex (Amphilophus citrinellus, labiatus, zaliosus) from several crater lakes in Nicaragua fits several of the key characteristics of a sympatric speciation model. In particular, in A. citrinellus (i) strong assortative mating on the basis of colour polymorphism and (ii) ecological differentiation based on morphological polymorphisms involving the feeding apparatus and body shape might both be mechanisms of incipient speciation. Seven microsatellite markers and mtDNA control region sequences [836 base pairs (bp)] were used to study the population genetic structure of 519 specimens of Midas cichlid populations from the two Great Lakes Managua and Nicaragua, and three crater lakes in Nicaragua, Central America. The three named species of the species complex occupy different ecological niches, are morphologically distinct and can be distinguished genetically. We uncovered allopatric genetic differentiation of populations of A. citrinellus from different lakes and distant locations within Lake Managua and, more interestingly, incipient genetic differentiation of several sympatric populations based on colouration (in A. citrinellus and A. labiatus) but not on the morphology of the pharyngeal jaws (in A. citrinellus). Sexual selection and assortative mating might be the driven forces of diversification within named species. The Midas cichlid species complex in Nicaragua is an excellent model system for the study of the incipient stages of adaptation, speciation and the formation of species flocks.     

Is resource overlap disadvantageous to three sympatric legumes?

Abstract The sympatric legumes Dillwynia hispida, Dillwynia uncinata and Pultenaea densifolia overlap in flowering phenology, floral characters and native bee pollinators. Tests for pollen-limitation were conducted to determine whether or not the overlap in pollinator use was detrimental to individuals of these species. Pollen limitation was only detected in D. hispida. Pollen loads on bees were examined and it was found that pollen loads from Dillwynia and P. densifolia were spatially segregated on the pollinator's body reducing the likelihood of pollen transfer between these species. The pollen of D. hispida and D. uncinata overlapped on the heads of bees suggesting that interspecific pollinations may be responsible for lowered fruit-set in D. hispida. However, hand pollinations involving heterospecific pollen and controlled floral visits by bees to the two Dillwynia species revealed that heterospecific pollen loads did not significantly depress fruit-set in either D. hispida or D. uncinata. Pollinator preferences were examined in detail and it was found that D. uncinata flowers received twice as many bees in a day compared with D. hispida and P. densifolia. Competition through pollinator preference is implicated here as a mechanism of competition. Intraspecific competition is also implicated as D. hispida was pollen limited in the absence of D. uncinata and P. densifolia.     

Batteries: what causes them to fail?

Rechargeable batteries do not die suddenly but gradually get weaker over time. The ability to hold charge diminishes through aging, usage (cycling), handling and environmental conditions. Batteries seem to have their own inherited life span. A fixed replacement plan, in which all batteries are replaced at a certain age, does not work well. Rather, each battery should periodically be analyzed and only those replaced whose capacity drops below an acceptable performance threshold. Implementing such a maintenance program reduces unexpected downtime caused by weak batteries and saves money. Unexpected downtime due to failing batteries can cause tense moments for the equipment operator. Some people may even seek legal compensation for personal damages suffered by a failing battery.     

Parapatric divergence of sympatric morphs in a salamander: incipient speciation on Long Island?

Speciation is often categorized based on geographic modes (allopatric, parapatric or sympatric). Although it is widely accepted that species can arise in allopatry and then later become sympatrically or parapatrically distributed, patterns in the opposite direction are also theoretically possible (e.g. sympatric lineages or ecotypes becoming parapatric), but such patterns have not been shown at a macrogeographic scale. Here, we analyse genetic, climatic, ecological and morphological data and show that two typically sympatric colour morphs of the salamander Plethodon cinereus (redback and leadback) appear to have become parapatrically distributed on Long Island, New York, with pure‐redback populations in the west and pure‐leadback populations in the east (and polymorphic populations in between and on the mainland). In addition, the pure‐leadback populations in eastern Long Island are genetically, ecologically and morphologically divergent from both mainland and other Long Island populations, suggesting the possibility of incipient speciation. This parapatric separation seems to be related to the different ecological preferences of the two morphs, preferences which are present on the mainland and across Long Island. These results potentially support the idea that spatial segregation of sympatric ecotypes may sometimes play an important part in parapatric speciation.     

Incipient sympatric speciation in Midas cichlid fish from the youngest and one of the smallest crater lakes in Nicaragua due to differential use of the benthic and limnetic habitats?

Understanding how speciation can occur without geographic isolation remains a central objective in evolutionary biology. Generally, some form of disruptive selection and assortative mating are necessary for sympatric speciation to occur. Disruptive selection can arise from intraspecific competition for resources. If this competition leads to the differential use of habitats and variation in relevant traits is genetically determined, then assortative mating can be an automatic consequence (i.e., habitat isolation). In this study, we caught Midas cichlid fish from the limnetic (middle of the lake) and benthic (shore) habitats of Crater Lake Asososca Managua to test whether some of the necessary conditions for sympatric speciation due to intraspecific competition and habitat isolation are given. Lake As. Managua is very small (<900 m in diameter), extremely young (maximally 1245 years of age), and completely isolated. It is inhabited by, probably, only a single endemic species of Midas cichlids, Amphilophus tolteca. We found that fish from the limnetic habitat were more elongated than fish collected from the benthic habitat, as would be predicted from ecomorphological considerations. Stable isotope analyses confirmed that the former also exhibit a more limnetic lifestyle than the latter. Furthermore, split‐brood design experiments in the laboratory suggest that phenotypic plasticity is unlikely to explain much of the observed differences in body elongation that we observed in the field. Yet, neutral markers (microsatellites) did not reveal any genetic clustering in the population. Interestingly, demographic inferences based on RAD‐seq data suggest that the apparent lack of genetic differentiation at neutral markers could simply be due to a lack of time, as intraspecific competition may only have begun a few hundred generations ago.     

Why do cladocerans fail to control algal blooms?

Field studies show that even at high nutrient loads phytoplankton may be kept at low levels by filter-feeding zooplankton for a period of weeks (spring clear water phase in lakes) or months (low-stocked fish-ponds). In the absence of planktivorous fish, large-bodied cladocerans effectively control the abundance of algae of a broad size spectrum. Laboratory experiments show that, although difficult to handle and of poor nutritional value, filamentous algae can also be utilized by large-bodiedDaphnia and prevented from population increase, exactly as the principles of the biomanipulation approach would predict. This is not always the case, however. Even when released from predation, large cladocerans often cannot grow and reproduce fast enough to prevent bloom formation. Sometimes, they disappear when the bloom becomes dense, and the biomanipulation approach is not applicable any more. Recent experimental data on four differently-sizedDaphnia species are used in an attempt to (1) explain why cladocerans fail to control filamentous cyanobacteria when filament density is high, and (2) determine the critical filament density at whichDaphnia becomes ineffective. At this critical concentration,Daphnia growth and reproduction is halted, and no positive numerical response to growing phytoplankton standing crop should be expected fromDaphnia population. Bloom formation thus becomes irreversible. The question of what can be done to overcome this bottleneck of the biomanipulation approach may become one of the most challenging questions in plankton ecology in the nearest future.