Modelling the mating system of polar bears: a mechanistic approach to the Allee effect

Allee effects may render exploited animal populations extinction prone, but empirical data are often lacking to describe the circumstances leading to an Allee effect. Arbitrary assumptions regarding Allee effects could lead to erroneous management decisions so that predictive modelling approaches are needed that identify the circumstances leading to an Allee effect before such a scenario occurs. We present a predictive approach of Allee effects for polar bears where low population densities, an unpredictable habitat and harvest-depleted male populations result in infrequent mating encounters. We develop a mechanistic model for the polar bear mating system that predicts the proportion of fertilized females at the end of the mating season given population density and operational sex ratio. The model is parametrized using pairing data from Lancaster Sound, Canada, and describes the observed pairing dynamics well. Female mating success is shown to be a nonlinear function of the operational sex ratio, so that a sudden and rapid reproductive collapse could occur if males are severely depleted. The operational sex ratio where an Allee effect is expected is dependent on population density. We focus on the prediction of Allee effects in polar bears but our approach is also applicable to other species.     

Variable fortunes in a patchy landscape —The habitat templet of an insect migrant

Migration by flight is essential for insects living in patchy landscapes and knowledge about variability in habitat patch structure and quality is important for the development of theory concerning insect dispersal polymorphisms. However, few studies provide more than anecdotal evidence about habitat change in time and space and its effects on insect survival and reproduction. Here I show how habitats and resource density of a lygaied bug,Lygaeus equestris, change in a patchy landscape over a seventeen year period. Although habitat patches per se are very stable, there are large temporal fluctuations in per capita food resources. Food seems to be limiting only in some years, and there may be periods of several years when populations change due to weather disturbance. Between-patch variation in reproductive success is large. Furthermore, the relative favourability of patches changes between years. There are also distance effects; in some years distances to suitable hibernation sites affect reproductive success. Long fliers experience more variable success, and sometimes extreme success, when compared with short fliers. The importance of movements from outlying areas also changes with time. The insect is thus faced with a habitat templet which varies strongly on many temporal and spatial scales.     

Comparative spatial ecology of fished spiny lobsters <Emphasis Type="Italic">Panulirus argus</Emphasis> and an unfished congener <Emphasis Type="Italic">P. guttatus</Emphasis> in an isolated marine reserve at Glover's Reef atoll,Belize

Palinurid lobsters are being exploited with increasing intensity in coral reef ecosystems, but marine protected areas may play a key role in preventing overfishing and local extinctions. In order to define the spatial requirements for protection, we compared the spatial and temporal patterns in distribution, density, biomass, size structure, and reproductive seasonality of Caribbean spiny lobsters Panulirus argus and the congeneric spotted lobsters P. guttatus on coral patch reef, forereef, and deep reef habitat at Glover's Reef, Belize. The relative impact of fishing on P. argus was also examined in an isolated marine reserve and adjacent fished habitats, in comparison with the relatively unfished distribution of P. guttatus. Over a 5-year period, both species co-occurred in all major reef habitats, but aspects of their population dynamics differed markedly due to both habitat and fishing effects. All size classes of spiny lobsters P. argus occupied shallow patch reefs, but large adults were predominant on the deep wall reef. Panulirus guttatus also occupied patch reefs in the lagoon, but spur-and-groove forereef appeared to be the primary habitat of this species. Density and exploitable (adult) biomass of P. argus increased significantly over time in the protected patch reef habitat of the lagoon but remained stable on deep reef habitat. The biomass of spotted lobsters P. guttatus in all habitats was at least an order of magnitude less than that of exploitable P. argus. Reproductive activity by both species was evident most of the year in all habitats, but breeding P. argus females were concentrated on the deep reef. Commercial fisheries for spotted lobsters P. guttatus are currently being considered for development, but data from this and other studies suggest that such a fishery may be relatively unproductive and may lead to rapid localized extinctions. Spiny lobsters P. argus used a variety of coral reef habitats, but spotted lobsters P. guttatus were habitat specialists restricted to shallow reef habitat. The protection needs of both species are similar in one aspect: large protected areas. However, P. argus required large areas with heterogeneous habitats including coral reefs and seagrass beds, whereas P. guttatus required large areas of coral reef habitat.     

Seed production in a threatened Aloe is not affected by bird exclusion or population size

Small populations of many plant species have been shown to exhibit ecological Allee effects. These effects are expected to be pronounced in plants which are obligate outcrossers and rely on pollinators which forage preferentially in larger populations with greater nectar availability. We examined the breeding and pollination systems, level of pollen limitation and seed production in populations of a threatened “ornithophilous” species, Aloe pruinosa. Experimental hand-pollinations showed that A. pruinosa is genetically self-incompatible and thus an obligate outcrosser. Experimental exclusion of birds from inflorescences did not affect seed production, suggesting that insects are effective pollinators. Supplemental hand-pollinations in several populations showed that seed production in A. pruinosa is not pollen limited. Further, there were no significant relationships between population size and any measure of reproductive success in this Aloe species. Small populations of A. pruinosa are thus viable in terms of pollination processes and should be protected from more direct threats, such as habitat alteration.     

Habitat selection reduces extinction of populations subject to Allee effects

Theoretical studies indicate that a single population under an Allee effect will decline to extinction if reduced below a particular threshold, but the existence of multiple local populations connected by random dispersal improves persistence of the global population. An additional process that can facilitate persistence is the existence of habitat selection by dispersers. Using analytic and simulation models of population change, I found that when habitat patches exhibiting Allee effects are connected by dispersing individuals, habitat selection by these dispersers increases the likelihood that patches persist at high densities, relative to results expected by random settlement. Populations exhibiting habitat selection also attain equilibrium more quickly than randomly dispersing populations. These effects are particularly important when Allee effects are large and more than two patches exist. Integrating habitat selection into population dynamics may help address why some studies have failed to find extinction thresholds in populations, despite well-known Allee effects in many species.     

破碎化生境中粗柄独尾草种群大小对繁殖特性的影响

景观破碎化是目前存在的一种普遍现象,导致物种生存的原有生境总面积减小,产生许多大小不一的斑块种群。不同斑块生境中植物种群的大小直接反映植物对环境的响应特征和适应对策,与植物繁殖有关的开花结实特性、种子形态及萌发特性的研究将有助于深入了解植物在不同生境中的生存策略。以粗柄独尾草(Eremurus inderiensis)为研究对象,以野外调查为基础,对比研究不同斑块粗柄独尾草种群的种群大小和生殖成功,揭示不同生境中粗柄独尾草的生态适应性。结果表明:(1)不同斑块生境中,种群大小差异较大,且较小的种群幼苗比例也较小,种群大小与种群幼苗比例呈显著相关;(2)较小的种群自然座果率、结籽率较低,种群间差异显著;(3)随着生境破碎化程度增加,种子千粒重,种子长、宽等形态特征呈减小趋势,但种翅占比增大;(4)在粗柄独尾草最适萌发温度下,随着生境破碎化程度增加,种子萌发率有下降趋势。针对人类干扰程度强的衰退型粗柄独尾草种群,亟需减少人为干扰,依据不同生境中的干扰因素及种群生存现状,制定科学与切实可行的保护和恢复策略。     

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.     

Temporal constraints on the potential role of fry odors as cues of past reproductive success for spawning lake trout

Deciding where to reproduce is a major challenge for most animals. Many select habitats based upon cues of successful reproduction by conspecifics, such as the presence of offspring from past reproductive events. For example, some fishes select spawning habitat following odors released by juveniles whose rearing habitat overlaps with spawning habitat. However, juveniles may emigrate before adults begin to search for spawning habitat; hence, the efficacy of juvenile cues could be constrained by degradation or dissipation rates. In lake trout (Salvelinus namaycush), odors deposited by the previous year's offspring have been hypothesized to guide adults to spawning reefs. However, in most extant populations, lake trout fry emigrate from spawning reefs during the spring and adults spawn during the fall. Therefore, we postulated that the role of fry odors in guiding habitat selection might be constrained by the time between fry emigration and adult spawning. Time course chemical, physiological, and behavioral assays indicated that the odors deposited by fry likely degrade or dissipate before adults select spawning habitats. Furthermore, fry feces did not attract wild lake trout to constructed spawning reefs in Lake Huron. Taken together, our results indicate fry odors are unlikely to act as cues for lake trout searching for spawning reefs in populations whose juveniles emigrate before the spawning season, and underscore the importance of environmental constraints on social cues.     

To produce many small or few large eggs: a size-independent reproductive tactic of fish

Summary We demonstrate here the existence of a range of size-independent reproductive tactics in teleostean fish involving the allocation of a size-dependent reproductive effort between fecundity and egg size. Despite considerable evidence that larger eggs and the larvae hatching from them are more likely to survive than smaller ones, we found no evidence of evolutionary trends towards greater egg sizes. Fish with pelagic eggs tend to spawn many, and therefore small, eggs, whereas demersal spawners tend to produce large, and therefore few, eggs. Maximizing egg number should increase the number of eggs hatching in suitable locations in the patchy pelagic environment and, hence, increase the reproductive success of pelagic spawners. On the other hand, the reproductive success of demersal spawners, which reduce the variance in growing conditions experienced by the off spring, should be more dependent on the survival of the individual larvae, which increases as egg size increases.     

Assortative mating,sexual selection,and their consequences for gene flow in Littorina

When divergent populations are connected by gene flow, the establishment of complete reproductive isolation usually requires the joint action of multiple barrier effects. One example where multiple barrier effects are coupled consists of a single trait that is under divergent natural selection and also mediates assortative mating. Such multiple-effect traits can strongly reduce gene flow. However, there are few cases where patterns of assortative mating have been described quantitatively and their impact on gene flow has been determined. Two ecotypes of the coastal marine snail, Littorina saxatilis, occur in North Atlantic rocky-shore habitats dominated by either crab predation or wave action. There is evidence for divergent natural selection acting on size, and size-assortative mating has previously been documented. Here, we analyze the mating pattern in L. saxatilis with respect to size in intensively sampled transects across boundaries between the habitats. We show that the mating pattern is mostly conserved between ecotypes and that it generates both assortment and directional sexual selection for small male size. Using simulations, we show that the mating pattern can contribute to reproductive isolation between ecotypes but the barrier to gene flow is likely strengthened more by sexual selection than by assortment.