Clarifying Baker's Law

Background

Baker''s Law states that colonization by self-compatible organisms is more likely to be successful than colonization by self-incompatible organisms because of the ability for self-compatible organisms to produce offspring without pollination agents. This simple model has proved very successful in plant ecology and has been applied to various contexts, including colonizing or ruderal species, islands colonizers, invasive species or mating system variation across distribution ranges. Moreover, it is one of the only models in population biology linking two traits of major importance in ecology, namely dispersal and mating system. Although Baker''s Law has stimulated a large number of empirical studies reporting the association of self-fertilization and colonizing ability in various contexts, the data have not established a general pattern for the association of traits.

Scope

In this paper, a critical position is adopted to discuss and clarify Baker''s Law. From the literature referring to Baker''s Law, an analysis made regarding how mating success is considered in such studies and discrepancies with population genetics theory of mating systems are highlighted. The data reporting the association of self-fertilization and colonizing ability are also briefly reviewed and the potential bias in interpretation is discussed. Lastly, a recent theoretical model analysing the link between colonizing ability and self-fertilization is considered.

Conclusions

Evolutionary predictions are actually more complex than Baker''s intuitive arguments. It appears that Baker''s Law encompasses a variety of ecological scenarios, which cannot be considered a priori as equivalent. Questioning what has been considered as self-evident for more than 50 years seems a reasonable objective to analyse in-depth dispersal and mating system traits.     

Effects of population size and metapopulation dynamics on a mating-system polymorphism

The evolutionary dynamics of neutral alleles under the Wright-Fisher model are well understood. Similarly, the effect of population turnover on neutral genetic diversity in a metapopulation has attracted recent attention in theoretical studies. Here we present the results of computer simulations of a simple model that considers the effects of finite population size and metapopulation dynamics on a mating-system polymorphism involving selfing and outcrossing morphs. The details of the model are based on empirical data from dimorphic populations of the annual plant Eichhornia paniculata, but the results are also of relevance to species with density-dependent selfing rates in general. In our model, the prior selfing rate is determined by two alleles segregating at a single diploid locus. After prior selfing occurs, some remaining ovules are selfed through competing self-fertilisation in finite populations as a result of random mating among gametes. Fitness differences between the mating-system morphs were determined by inbreeding depression and pollen discounting in a context-dependent manner. Simulation results showed evidence of frequency dependence in the action of pollen discounting and inbreeding depression in finite populations. In particular, as a result of selfing in outcrossers through random mating among gametes, selfers experienced a "fixation bias" through drift, even when the mating-system locus was selectively neutral. In a metapopulation, high colony turnover generally favoured the fixation of the outcrossing morph, because inbreeding depression reduced opportunities for colony establishment by selfers through seed dispersal. Our results thus demonstrate that population size and metapopulation processes can lead to evolutionary dynamics involving pollen and seed dispersal that are not predicted for large populations with stable demography.     

POLLINATOR-MEDIATED COMPETITION,REPRODUCTIVE CHARACTER DISPLACEMENT,AND THE EVOLUTION OF SELFING IN ARENARIA UNIFLORA (CARYOPHYLLACEAE)

Ecological factors that reduce the effectiveness of cross-pollination are likely to play a role in the frequent evolution of routine self-fertilization in flowering plants. However, we lack empirical evidence linking the reproductive assurance value of selfing in poor pollination environments to evolutionary shifts in mating system. Here, we investigated the adaptive significance of prior selfing in the polymorphic annual plant Arenaria uniflora (Caryophyllaceae), in which selfer populations occur only in areas of range overlap with congener A. glabra. To examine the hypothesis that secondary contact between the two species contributed to the evolution and maintenance of selfing, we used field competition experiments and controlled hand-pollinations to measure the female fitness consequences of pollinator-mediated interspecific interactions. Uniformly high fruit set by selfers in the naturally pollinated field arrays confirmed the reproductive assurance value of selfing, whereas substantial reductions in outcrosser fruit set (15%) and total seed production (20–35%) in the presence of A. glabra demonstrated that pollinator-mediated interactions can provide strong selection for self-pollination. Heterospecific pollen transfer, rather than competition for pollinator service, appears to be the primary mechanism of pollinator-mediated competition in Arenaria. Premating barriers to hybridization between outcrossers and A. glabra are extremely weak. The production of a few inviable hybrid seeds after heterospecific pollination and intermediate seed set after mixed pollinations indicates that A. glabra pollen can usurp A. uniflora ovules. Thus, any visit to A. uniflora by shared pollinators carries a potential female fitness cost. Moreover, patterns of fruit set and seed set in the competition arrays relative to controls were consistent with the receipt of mixed pollen loads, rather than a lack of pollinator visits. Competition through pollen transfer favors preemptive self-pollination and may be responsible for the evolution of a highly reduced floral morphology in A. uniflora selfers as well as their current geographical distribution.     

Heterogeneity in local density allows a positive evolutionary relationship between self‐fertilisation and dispersal

Despite empirical evidence for a positive relationship between dispersal and self‐fertilization (selfing), theoretical work predicts that these traits should always be negatively correlated, and the Good Coloniser Syndrome of high dispersal and selfing (Cf. Baker's Law) should not evolve. Critically, previous work assumes that adult density is spatiotemporally homogeneous, so selfing results in identical offspring production for all patches, eliminating the benefit of dispersal for escaping from local resource competition. We investigate the joint evolution of dispersal and selfing in a demographically structured metapopulation model where local density is spatiotemporally heterogeneous due to extinction‐recolonization dynamics. Selfing alleviates outcrossing failure due to low local density (an Allee effect) while dispersal alleviates competition through dispersal of propagules from high‐ to low‐density patches. Because local density is spatiotemporally heterogeneous in our model, selfing does not eliminate heterogeneity in competition, so dispersal remains beneficial even under full selfing. Hence the Good Coloniser Syndrome is evolutionarily stable under a broad range of conditions, and both negative and positive relationships between dispersal and selfing are possible, depending on the environment. Our model thus accommodates positive empirical relationships between dispersal and selfing not predicted by previous theoretical work and provides additional explanations for negative relationships.     

Inferring demographic processes from the genetic structure of a metapopulation of <Emphasis Type="Italic">Boltonia decurrens</Emphasis>(Asteraceae)

Boltonia decurrens(Asteraceae), a federally listed, threatened floodplain species, requires regular flooding for suitable habitat and seed dispersal. Flood suppression and habitat destruction have resulted in fewer than 25 populations remaining throughout its 400 km range. Because individual populations are widely interspaced (>10 km) and subject to frequent extinction and colonization, seed dispersal along the river, not pollen flow, is likely the primary determinant of population genetic structure. We used neutral genetic markers (isozymes) assayed for fourteen populations to determine which demographic processes contribute to the genetic structure of B. decurrens. Significant genetic differentiation was detected among populations (F _ST=0.098, P< 0.05) but not among regions (F _RT=0.013, P> 0.05), suggesting that long-distance dispersal events occur and involve seed from a small number of populations. Correspondingly, we found no evidence of isolation by distance, and admixture analyses indicate that colonization events involve seed from 3 to 5 source populations. Individual populations exhibited high levels of fixation (mean F _IS=0.192, P< 0.05), yet mean population outcrossing rates were high (t _m=0.87–0.95) and spatial autocorrelation analyses revealed no fine-scale within population structure, indicating that inbreeding alone cannot explain the observed fixation. Rather, genetic bottlenecks, detected for 12 of 14 populations, and admixture at population founding may be important sources of fixation. These observations are consistent with a metapopulation model and confirm the importance of regular flooding events, capable of producing suitable habitat and dispersing seed long distances, to the long-term persistence of B. decurrens.     

Pollination in vitro: effects on the growth of pollen tubes,seed set and gametophytic self-incompatibility in Trifolium pratense L. and T. repens L.

Summary Growth of pollen tubes and seed set were compared after hand pollination in situ and in vitro in two self-incompatible species, Trifolium pratense and Trifolium repens. Adhesion of pollen grains to the stigma was greater in vitro for both species. After cross-pollination, in vitro culture gave a significant increase in the cumulative growth of pollen tubes in pistils of T. pratense compared to in situ conditions. After selfing in T. repens, pollen tube growth was significantly increased by in vitro culture of florets. Seed set after crossing in situ and in vitro was similar for both species. Seed set after selfing in vitro was not increased in T. pratense. Several genotypes of T. repens were classified as very good, good and poor selfers based on their capacity for seed set following selfing in situ. In vitro pollination increased self seed formation by 1.7-, 18.0- and 31.0-fold for each class, respectively. Ovules located nearest to the style were fertilized more often after selfing than after crossing.     

Allee effect and self-fertilization in hermaphrodites: reproductive assurance in a structured metapopulation

Reproductive assurance through selfing during colonization events or when population densities are low has often been put forward as a mechanism selecting for the evolution of self-fertilization. Such arguments emphasize on the role of both local demography and metapopulation processes. We developed a model for the evolution of self-fertilization in a structured metapopulation in which local densities are not steady because of population growth. Reproduction by selfing is density-independent (reproductive assurance) but selfed seeds endure inbreeding depression, whereas reproduction by outcrossing is density-dependent (Allee effect). First, we derived an analytical criterion for metapopulation viability as a function of the selfing rate and metapopulation parameters. We show that outcrossers can develop a viable metapopulation when they produce a high amount of dispersal seeds that counterbalances their incapacity to found new populations from low densities. Second, the model shows there is a positive feedback between demography and outcrossing rates, leading to either complete outcrossing or selfing. Specifically, we illustrate that inbreeding depression can paradoxically favor the evolution of selfing because of its negative effect on density. Also, complete outcrossing can be selected despite pollen limitation, although it does not provide a full seed set. This model underlines the influence of the mating system both on demography and gene dynamics in a metapopulation context.     

Dances with fire: Tracking metapopulation dynamics of<Emphasis Type="Italic">polygonella basiramia</Emphasis> in Florida scrub (USA)

The regional persistence of species subject to local population colonization and extinction necessarily depends on how landscape features and disturbance affect metapopulation dynamics. Here, we characterize the metapopulation structure and short-term dynamics ofPolygonella basiramia. This rare, short-lived perennial herb is endemic to Florida scrublands and lacks a seed bank. Fires create the open sand gaps within a shrub matrix that support this species but also kill established plants. Thus, persistence depends on frequent colonization of unoccupied gaps. We are monitoring population dynamics within and among 1204 gaps distributed among 19 shrub patches. Considerable subpopulation turnover is evident at the gap level with rates of gap extinction exceeding rates of colonization in the first year. Whether declines in overall abundance continue is likely to depend on patterns of disturbance and regional stochasticity in this dynamic landscape.Polygonella is more likely to occupy larger and less isolated gaps, demonstrating that landscape features and disturbance strongly affect metapopulation dynamics. BecausePolygonella basiramia displays characteristics, occupancy patterns, and turnover dynamics consistent with metapopulation theory, it represents a model system for studying plant metapopulations.     

Metapopulation dynamics in an aphid-parasitoid system

Metapopulation theory makes a number of predictions concerning the effects of dispersal on the persistence of predator-prey or host-parasitoid systems. While the stabilising effects of dispersal have been shown in a number of laboratory studies, evidence from field studies remains scarce due to a lack of suitable model systems. I describe a host-parasitoid system that shows a classical metapopulation structure with frequent extinctions and colonisations consisting of the aphidiid Lysiphlebus hirticornisand the aphid Metopeurum fuscoviride. Both the parasitoid and the aphid are specialists on their respective hosts. I followed the dynamics of host and parasitoid on individually marked tansy (Tanacetum vulgare) plants, the host of M. fuscoviride. Dynamics of host and parasitoid populations were characterized by frequent extinctions and colonisations. Mean longevity of aphid colonies was only 3.1 weeks. Parasitism by L. hirticorniswas a main cause of extinction for the aphid as rates of parasitism often reached 100%, in particular towards the end of the field season. Patchiness in this system occurs at two spatial scales. Aphid colonies form on single tansy ramets = shoots but movements of aphid individuals among ramets within a particular tansy genet are frequent. Because aphids can persist on a genet for a large numer of generations, it is argued that local populations form on genets rather than ramets. The number of host and parasitoid extinctions described in this study exceeds the number of extinctions usually observed in field studies of host-parasitoid metapopulations. It is suggested that aphid-parasitoid systems such as the one studied in this paper may be good models to test the predictions of metapopulation theory.     

HYDROTHERMAL-VENT ALVINELLID POLYCHAETE DISPERSAL IN THE EASTERN PACIFIC. 2. A METAPOPULATION MODEL BASED ON HABITAT SHIFTS

Marine organisms typically fall into two main categories: those with a high level of population structuring and those with a low one. The first are often found to be poor dispersers, following isolation by distance or stepping-stone theoretical predictions. The second are commonly associated with high-dispersal taxa and are best described by the island model. Deep-sea hydrothermal vent systems represent a good model for studying one-dimensional metapopulations. Whereas isolation by distance might be expected to be the rule in such a system for species with limited dispersal capabilities, a biological paradox can be observed: an apparent genetic homogeneity in some vent species with short-scale dispersal potential, in a one-dimensional fragmented habitat. This can be explained if one key assumption of the existing models is not met: gene flow between populations and genetic drift may not have the time to equilibrate. Geophysical models revealed that hydrothermal convection is intrinsically unstable, inducing processes of coalescence or splitting of venting areas in a chaotic manner. This is likely to generate frequent extinctions and recolonizations. Theoretical genetic predictions derived from extinctions/recolonizations cannot satisfactorily model a situation where habitat shifts are frequent and constantly affect the metapopulation equilibrium. Because neither the island and the stepping-stone models nor the classical metapopulation models resemble the hydrothermal vent reality, we present here a realistic model developed to provide a compromise between existing conceptual models and what is currently known of the biology and ecology of one of the most peculiar and best-studied vent species, the polychaete Alvinella pompejana. This model allows us to define the boundaries between which the metapopulation is evolutionary stable in an unstable context. Simulations show different patterns in which metapopulation size and recolonization vary but reach an equilibrium despite chaotic vent extinctions. In contrast, the model also shows that displacing habitat continuously affects the equilibrium between gene flow and drift. As a consequence, the time required to balance these evolutionary forces can never be attained, leading to chaotic fluctuations in F-statistics. Those fluctuations are mainly due to stochastic changes of the interpatch distance which affect migration rates. The shifting of active zones of venting can episodically counterbalance differentiation and allow a long-term genetic homogenization at the ridge scale. These findings lead to a new concept in which the exchanges between populations would mainly depend on the habitat's movements along the ridge axis rather than the organim's dispersal. We therefore propose a new model based on patch-network displacements in which transient contact zones allow low levels of gene flow throughout the metapopulation.     

Optimum reproduction and dispersal strategies of a clonal plant in a metapopulation: a simulation study with Hieracium pilosella

Clonal spread is favoured in many plants at the expense of seed production in order to expand rapidly into open habitats or to occupy space by forming dense patches. However, for the dynamics of a population in a patchy landscape seed dispersal remains important even for clonal plants. We used a spatially explicit individual-based metapopulation model to examine the consequences of two trade-offs in Hieracium pilosella L: first, between vegetative and sexual reproduction, and second, between short and far-distance dispersal of seeds. Our main question was, what are the environmental conditions that cause a mixed strategy of vegetative and sexual reproduction to be optimal. The model was parameterised with field data on local population dynamics of H. pilosella. Patch dynamics were given firstly by disturbance events that opened patches in a matrix of a clonal grass that were colonisable for H. pilosella, and secondly by the gradual disappearance of H. pilosella patches due to the expanding grass. Simulations revealed opposing selection pressures on traits determined by the two trade-offs. Vegetative reproduction is favoured by local dynamics, i.e. the need for maintenance and expansion of established populations, whereas seed production is favoured by the necessity to colonise empty habitats. Similar pressures act on the proportion of seeds dispersed over short and far distances. Optimum reproductive and dispersal strategies depended on habitat quality (determined by seedling establishment probability), the fraction of dispersed seeds, and the fraction of seeds lost on unsuitable ground. Under habitat conditions supporting moderate to low seedling establishment, between 20% and 40% of reproductive effort in H. pilosella should be devoted to sexual reproduction with at least 10% of the seeds dispersed over distances suitable to attain empty patches. We conclude that in a spatially heterogeneous landscape sexual seed production in a clonal plant is advantageous even at the expense of local vegetative growth.     

Metapopulation viability of an endangered holoparasitic plant in a dynamic landscape

By creating transient patch mosaics, disturbance can influence the dynamics of interacting populations in many ecosystems. In European heathland, traditional land use created such dynamic systems favourable for both early and later successional species. Little empirical evidence is, however, available on the impact of current management on metapopulations occurring in such landscapes. This paper looks at the metapopulation viability of the endangered holoparasite Cuscuta epithymum, a species that typically occurs in early successional stages of recently managed heathlands. We used both observational and experimental data from a 4‐yr study to parameterise a spatially explicit metapopulation model. This model explores the impact of demographic characteristics and spatiotemporal landscape patterns created by management events on metapopulation viability. Both occasional long‐distance dispersal and dormant seeds are shown to be critical for the long‐term survival of C. epithymum in a dynamic heathland landscape subjected to a fixed rotational mowing of 15 yr. A relatively high management frequency (<15 yr between two consecutive mowing events) appeared to be necessary to sustain a viable C. epithymum metapopulation. When there is a longer interval between management events, grazing can counterbalance the negative effects of vegetation succession. Our results indicate that small‐scale cyclical management events combined with extensive grazing are the most appropriate management strategy to maintain viable populations of C. epithymum instead of the current large‐scale management events. Our results further emphasise the importance of incorporating both spatiotemporal patch availability and key demographic characteristics, especially seed banks, for a realistic view of metapopulation dynamics in disturbed landscapes. This study clearly demonstrates the usefulness of metapopulation models to understand the impact of management events and to provide new ecological insights into processes acting at a landscape scale.     

Long-distance seed dispersal in plant populations

Long-distance seed dispersal influences many key aspects of the biology of plants, including spread of invasive species, metapopulation dynamics, and diversity and dynamics in plant communities. However, because long-distance seed dispersal is inherently hard to measure, there are few data sets that characterize the tails of seed dispersal curves. This paper is structured around two lines of argument. First, we argue that long-distance seed dispersal is of critical importance and, hence, that we must collect better data from the tails of seed dispersal curves. To make the case for the importance of long-distance seed dispersal, we review existing data and models of long-distance seed dispersal, focusing on situations in which seeds that travel long distances have a critical impact (colonization of islands, Holocene migrations, response to global change, metapopulation biology). Second, we argue that genetic methods provide a broadly applicable way to monitor long-distance seed dispersal; to place this argument in context, we review genetic estimates of plant migration rates. At present, several promising genetic approaches for estimating long-distance seed dispersal are under active development, including assignment methods, likelihood methods, genealogical methods, and genealogical/demographic methods. We close the paper by discussing important but as yet largely unexplored areas for future research.     

The effects of seed abundance on seed predation and dispersal by rodents in <Emphasis Type="Italic">Castanopsis fargesii</Emphasis> (Fagaceae)

Based on the animal dispersal hypothesis and the predator satiation hypothesis, we examined the effects of seed abundance at both population (i.e., mast seeding) and community levels on seed predation and dispersal of Castanopsis fargesii (Fagaceae), a rodent-dispersed mast species in Eastern Asia. In a subtropical evergreen broadleaved forest in the Dujiangyan region of Sichuan Province, China, individual seeds with coded tin tags were tracked in two contrasting stands (seed-poor and seed-rich) over two years (2000, a low-seed year; 2001, a high-seed year). Our results showed that: (1) small rodents did not harvest the tagged seeds of C. fargesii more rapid in the high-seed year than in the low-seed year in either stand. But, seed harvest was significantly faster in the seed-rich stand than in the seed-poor stand. (2) The removal proportion was significantly lower in the high-seed year than in the low-seed year for either stand, but the removal proportion was slightly higher in the seed-poor stand than in the seed-poor stand. This indicates that high seed abundance decreases seed removal (predator satiation hypothesis). (3) There were only small differences about seed caching, seed survival and seedling establishment of C. fargesii between years and stands. During the survey, no cached seeds survived to geminate in the spring for both stands and years. (4) Mean dispersal distances of the cached seeds are much shorter in the high-seed year (3.1 m) than in the low-seed year (8.1 m) in the seed-rich stand, though similar trend is not examined in the seed-poor stand. Our results indicate that seed predation and dispersal of C. fargesii are influenced by both mast seeding and community-level seed abundance, which is not completely consistent with either the animal dispersal hypothesis or the predator satiation hypothesis, but seems more related to the predator satiation hypothesis.     

Human‐facilitated metapopulation dynamics in an emerging pest species,Cimex lectularius

The number and demographic history of colonists can have dramatic consequences for the way in which genetic diversity is distributed and maintained in a metapopulation. The bed bug (Cimex lectularius) is a re‐emerging pest species whose close association with humans has led to frequent local extinction and colonization, that is, to metapopulation dynamics. Pest control limits the lifespan of subpopulations, causing frequent local extinctions, and human‐facilitated dispersal allows the colonization of empty patches. Founder events often result in drastic reductions in diversity and an increased influence of genetic drift. Coupled with restricted migration, this can lead to rapid population differentiation. We therefore predicted strong population structuring. Here, using 21 newly characterized microsatellite markers and approximate Bayesian computation (ABC), we investigate simplified versions of two classical models of metapopulation dynamics, in a coalescent framework, to estimate the number and genetic composition of founders in the common bed bug. We found very limited diversity within infestations but high degrees of structuring across the city of London, with extreme levels of genetic differentiation between infestations (F_ST = 0.59). ABC results suggest a common origin of all founders of a given subpopulation and that the numbers of colonists were low, implying that even a single mated female is enough to found a new infestation successfully. These patterns of colonization are close to the predictions of the propagule pool model, where all founders originate from the same parental infestation. These results show that aspects of metapopulation dynamics can be captured in simple models and provide insights that are valuable for the future targeted control of bed bug infestations.     

Levels of allozyme diversity in closely related toadflaxes (<Emphasis Type="Italic">Linaria</Emphasis>, Plantaginaceae) and their correspondence with the breeding systems of the species

Toadflaxes (Linaria Miller) represent the largest genus of the tribe Antirrhineae (formerly included in the Scrophulariaceae) of the Plantaginaceae family. These taxa vary according to their ranges (narrow-regional–widespread), biological traits (duration, flower morphology, breeding system) and ecological preferences. No comparative genetic studies have been performed up to the moment and the levels and distribution of genetic diversity within this genus remain undocumented. We have conducted an allozyme study in 8 taxa of Section Supinae. Five of them are narrow endemic, self-incompatible, insect-pollinated outcrossers that inhabit relatively unaltered habitats. The other three taxa are widespread, small-flowered selfers that inhabit crops subjected to high levels of recurrent disturbance. Linaria taxa showed moderate to high levels of genetic diversity. Parameters of genetic diversity were directly addressed to differences in breeding systems. Mean number of alleles per locus, total genetic diversity and genetic diversity within populations were higher among the outcrossers than among the selfers, (A _e = 2.28 vs. 1.23; H _T = 0.24 vs. 0.13; H _S = 93.99% vs. 54.2%) while genetic diversity among populations and population differentiation were lower for the outcrossers than for the selfers (D _ST = 0.02% vs. 45.98%; G _ST = 0.03 vs. 0.52). Gene flow within and among populations produced a higher number of different genotypes (mean N _G = 16 for outcrossers vs. 5.4 for selfers) and migrants (mean N _m = 4.58 for outcrossers vs. 0.84 for selfers) that were attributed to an effective pollinator service in outcrossing populations. Differences in the levels and distribution of the genetic diversity are discussed in relation to biological and autoecological constraints of each group of taxa.     

The founding of Mauritian endemic coffee trees by a synchronous long‐distance dispersal event

The stochastic process of long‐distance dispersal is the exclusive means by which plants colonize oceanic islands. Baker's rule posits that self‐incompatible plant lineages are unlikely to successfully colonize oceanic islands because they must achieve a coordinated long‐distance dispersal of sufficiently numerous individuals to establish an outcrossing founder population. Here, we show for the first time that Mauritian Coffea species are self‐incompatible and thus represent an exception to Baker's rule. The genus Coffea (Rubiaceae) is composed of approximately 124 species with a paleotropical distribution. Phylogenetic evidence strongly supports a single colonization of the oceanic island of Mauritius from either Madagascar or Africa. We employ Bayesian divergence time analyses to show that the colonization of Mauritius was not a recent event. We genotype S‐RNase alleles from Mauritian endemic Coffea, and using S‐allele gene genealogies, we show that the Mauritian allelic diversity is confined to just seven deeply divergent Coffea S‐RNase allelic lineages. Based on these data, we developed an individual‐based model and performed a simulation study to estimate the most likely number of founding individuals involved in the colonization of Mauritius. Our simulations show that to explain the observed S‐RNase allelic diversity, the founding population was likely composed of fewer than 31 seeds that were likely synchronously dispersed from an ancestral mainland species.     

Population dynamics of epiphytic orchids in a metapopulation context

Background and Aims

Populations of many epiphytes show a patchy distribution where clusters of plants growing on individual trees are spatially separated and may thus function as metapopulations. Seed dispersal is necessary to (re)colonize unoccupied habitats, and to transfer seeds from high- to low-competition patches. Increasing dispersal distances, however, reduces local fecundity and the probability that seeds will find a safe site outside the original patch. Thus, there is a conflict between seed survival and colonization.

Methods

Populations of three epiphytic orchids were monitored over three years in a Mexican humid montane forest and analysed with spatially averaged and with spatially explicit matrix metapopulation models. In the latter, population dynamics at the scale of the subpopulations (epiphytes on individual host trees) are based on detailed stage-structured observations of transition probabilities and trees are connected by a dispersal function.

Key Results

Population growth rates differed among trees and years. While ignoring these differences, and averaging the population matrices over trees, yields negative population growth, metapopulation models predict stable or growing populations because the trees that support growing subpopulations determine the growth of the metapopulation. Stochastic models which account for the differences among years differed only marginally from deterministic models. Population growth rates were significantly lower, and extinctions of local patches more frequent in models where higher dispersal results in reduced local fecundity compared with hypothetical models where this is not the case. The difference between the two models increased with increasing mean dispersal distance. Though recolonization events increased with dispersal distance, this could not compensate the losses due to reduced local fecundity.

Conclusions

For epiphytes, metapopulation models are useful to capture processes beyond the level of the single host tree, but local processes are equally important to understand epiphyte population dynamics.     

Size and genetic composition of the colonizing propagules in a butterfly metapopulation

The amount and spatial distribution of genetic variation that is maintained in a metapopulation depends critically on the colonization process. Here, we use molecular markers to determine the number and genetic relatedness of individuals establishing new local populations in a large metapopulation of the Glanville fritillary butterfly Melitaea cinxia. The empirical results are compared with the predictions of a dispersal model based on a diffusion approximation of correlated random walk, which serves as a base‐line hypothesis about the rate and pattern of colonization. The results show that half of the new local populations consisted of a single larval group of full sibs and hence necessarily of the offspring of a single female. If the colonization involved two or more larval groups, these were usually oviposited by two different females that were unrelated to each other. The pattern of colonizations is thus intermediate between the propagule pool and the migrant pool models. These results elucidate the generation of genetic stochasticity, which may influence the dynamics of small populations. The dispersal model predicted well the pattern of habitat occupancy and the pattern of colonizations in relation to landscape structure, though which particular habitat patches became colonized was influenced also by measures of habitat quality not included in the model.     

Ant-mediated expansion of an obligate seeder species during the first years after fire

Most obligate seeder species build up a soil seed bank that is associated with massive seed germination in the year immediately after a fire. These species are also shade‐intolerant and disappear when vegetation cover closes, creating unsuitable conditions for seedling recruitment. The only way for these plants to expand their populations is when habitats suitable for seedling recruitment arise (i.e. in years immediately after a fire). However, short primary seed dispersal of obligate seeders does not allow these plants to colonise the suitable habitats, and these habitats can only be colonised by secondary seed dispersion. We hypothesised that Fumana ericoides, an obligate‐seeding small shrub, not only establishes abundantly in the first year after fire, but also expands its local range in the following years due to secondary dispersal by ants while suitable habitats are still available. We tested this hypothesis using experimental studies and a simulation model of potential population expansion in a recently burned area. Results showed that F. ericoides not only established prolifically in the year immediately after fire, but was also able to recruit new individuals and expand its population in the years following the fire, despite a low germination rate and short primary seed dispersal. Ant‐mediated seed dispersal and availability of suitable habitats were key factors in this phenomenon: ants redistributed seeds in suitable habitats while they were available, which accelerated the expansion of F. ericoides because new plants established far away from the core population.