Effects of conspecific and heterospecific residents on patterns of immigration in two species of voles

Studies with birds have shown that presence and density of resident conspecifics and heterospecifics can influence patterns of habitat selection. There have been few studies on the effects of social cues on rates of immigration in mammals. We report results from a long-term live trapping study of immigration in two species of voles, Microtus ochrogaster Wagner, 1842 and Microtus pennsylvanicus Ord, 1815, in bluegrass habitat in east-central Illinois, USA. We compare immigration into control sites from which no individuals of either species were removed with immigration into experimental sites from which either conspecifics or heterospecifics were removed. We focus on characteristics of immigrants and rates of immigration in relation to density in destination habitats. Within each species, immigrants into control and removal sites were similar with respect to body mass and reproductive condition, indicating no major differences in the physical condition of immigrants into sites with established populations and sites without established populations. For both species, density of conspecifics at a site positively influenced rate of immigration at that site. Density of heterospecifics at destination sites did not significantly influence rate of immigration for either species. These results suggest that site selection by dispersing M. ochrogaster and M. pennsylvanicus is characterized by conspecific attraction.     

Demography of fluctuating prairie vole populations: comparison of demographic variables among phases of fluctuations

We tested for differences in the proportion of reproductively active males and females, proportion of the population composed of young and immigrants, and monthly survival (total, adult, young) among phases (trough, increase, and decline) and among habitats (alfalfa, bluegrass, and tallgrass) of 30 population fluctuations ofMicrotus ochrogaster Wagner, 1842 over 25 years in east-central Illinois USA. Total population survival and survival of adults and young were greatest during the increase phase, among fluctuations, irrespective of habitat. The proportion of reproductively active adult males and females was lowest during the decline phase, an effect of lower reproduction during the winter. These results suggest that phase-specific changes in survival were the primary demographic factor driving population fluctuations ofM. ochrogaster in our study sites. We conclude that small-scale spatially different population fluctuations may be explained by the same mechanisms that explain fluctuations within a population.     

Vole population dynamics: factors affecting peak densities and amplitudes of annual population fluctuations of<Emphasis Type="Italic">Microtus pennsylvanicus</Emphasis>

We studied factors affecting peak densities and amplitudes of fluctuation during 20 annual population fluctuations ofMicrotus pennsylvanicus Ord, 1815 in alfalfa and bluegrass habitats over a 25-year period. Survival was correlated with population density over the 25 years and was the most consistent variable associated with stoppage of population growth. Although not correlated with population density over the 25 years, a decline in the proportion of reproductively active adult females contributed to cessation of growth of population fluctuations that peaked in late autumn-winter, and to cessation of growth of eight of eleven population fluctuations that peaked during summer-early autumn. We conclude variation in survival to be the primary factor affecting peak densities and amplitudes of population fluctuation ofM. pennsylvanicus.     

Demography of fluctuating vole populations: Phase homogeneity of demographic variables

Most research on fluctuations of arvicoline rodents has focused on changes in demographic variables across phases. Here, we examined whether demographic variables varied within a phase (trough, increase, and decline) across 30 fluctuations of Microtus ochrogaster and 14 fluctuations of Microtus pennsylvanicus over 25 years. For each species, we tested for phase homogeneity (i.e., values of demographic variables were invariant across all fluctuations within a given phase) for the following demographic variables: monthly survival (total, adult, young), proportion of reproductively active adult males and females, proportion of young, and proportion of immigrants. We found phase homogeneity for 96.7% of the 582 phase/fluctutation data sets for the seven variables, in the three phases of across all fluctuations in the three habitats for M. ochrogaster and 98.0% of 254 of those for M. pennsylvanicus in two habitats. We conclude that for each of these two species demographic variables are relatively invariant both within a phase and across fluctuations.     

The development of social behaviour between siblings and non-siblings of the voles Microtus ochrogaster and Microtus pennsylvanicus

It has been suggested that the strength and duration of sibling relations during the juvenile period may reflect the social structure of mammal species. Two Microtus species with contrasting social systems, M. ochrogaster and M. pennsylvanicus, were chosen for a comparative study of juvenile social behaviour. Young were observed in sibling and non-sibling dyadic encounters from the second until the seventh week of age. Social interactions consisted mainly of body nosing of the partner, and included behaviours interpreted as investigation, socialization and settling. Only minor sex differences in body nosing scores occurred, but some ontogenetic trends were noted. When the species were compared, M. ochrogaster received higher scores for body nosing and sitting in contact, and displayed more reciprocity in their nosing, than did M. pennsylvanicus, whether siblings or non-siblings were considered. When the sibling types were compared, non-sibling dyads received higher nosing scores than did siblings in both species, but the disparity was greater in M. ochrogaster than in M. pennsylvanicus. These results are interpreted with reference to known behavioural and developmental correlates of the extended family system in naturally occurring M. ochrogaster and the less communal social system in M. pennsylvanicus.     

Demography in relation to population density in two herbivorous marsupials: testing for source–sink dynamics versus independent regulation of population size

We compared demography of populations along gradients of population density in two medium-sized herbivorous marsupials, the common brushtail possum Trichosurus vulpecula and the rufous bettong Aepyprymnus rufescens, to test for net dispersal from high density populations (acting as sources) to low density populations (sinks). In both species, population density was positively related to soil fertility, and variation in soil fertility produced large differences in population density of contiguous populations. We predicted that if source–sink dynamics were operating over this density gradient, we should find higher immigration rates in low-density populations, and positive relationships of measures of individual fitness—body condition, reproductive output, juvenile growth rates and survivorship—to population density. This was predicted because under source–sink dynamics, immigration from high-density sites would hold population density above carrying capacity in low-density sites. The study included 13 populations of these two species, representing a more than 50-fold range of density for each species, but we found that individual fitness, immigration rates and population turnover were similar in all populations. We conclude that net dispersal from high to low density populations had little influence on population dynamics in these species; rather, all populations appeared to be independently regulated at carrying capacity, with a balanced exchange of dispersers among populations. These two species have suffered recent reductions in range, and they are ecologically similar to other species that have declined to extinction in inland Australia. It has been argued that part of the cause of the vulnerability of species like these is that they exhibit source–sink dynamics, and disturbance to source habitats can therefore cause large-scale population collapses. The results of our study argue against this interpretation.     

Settlement in empty versus occupied habitats: an experimental study on bank voles

Despite the recognised importance of colonisation (settlement in empty habitats) and immigration (settlement in an established population) to species persistence and evolution few have investigated these processes in territorial mammals and how they affect species’ traits. We translocated female bank voles (Myodes glareolus) onto an island (2.58 ha) in a two-stage experiment (stage 1: colonisation of empty population space and stage 2: immigration into an established population) to test (1) if colonisers and immigrants differ in probability of settlement and pregnancy, and (2) if settlement is affected by cues of conspecifics, i.e., simulated deserted home ranges (SDHR) and resident presence. Density was kept well below saturation in 8 temporally distinct population replicates over 3 years. SDHR and resident presence neither attracted nor repelled colonisers and immigrants, respectively, and settlement was not different from a random model. Probability of settlement tended to be higher in colonisers than immigrants and the probability of pregnancy was significantly higher in colonisers; immigrants settling within the home range of residents had nearly zero probability of pregnancy. Colonisation of empty habitat patches selected based on physical or resource based habitat features is apparently the optimal settlement strategy of dispersing voles, because cues from conspecifics may provide ambiguous information and social factors may inhibit settlement or delay reproduction in immigrants even at low population density.     

Agonistic behaviour in males of five species of voles Microtus

Agonistic behaviour of Microtus pennsylvanicus, M. montanus, M. californicus, M. longicaudus, and M. ochrogaster was observed in 160 staged encounters; a natural observation of sympatric species is included. No difference in the character and movements of agonistic behaviour was apparent when arena size was reduced; only a difference in the frequency of postures and movements was observed. In the smaller arena, M. montanus and M. ochrogaster exhibited a significant increase in frequency of agonistic components. The other three species did not show a significant increase. In interspecific pairings, M. montanus won over M. pennsylvanicus, M. californicus and M. longicaudus; M. ochrogaster won over M. montanus and M. pennsylvanicus. Three species used mutual avoidance in maintaining individual distance.     

Selection against immigrants in wild seabird populations

Immigration is a major demographic parameter shaping population dynamics and is an important driver of eco‐evolutionary patterns, but the fitness consequences for individuals following their settlement to a new population (immigrants) remain poorly tested in wild animal populations, particularly among long‐lived species. Here we show that immigrants have a lower fitness than residents in three wild seabird populations (wandering albatross Diomedea exulans, southern fulmar Fulmarus glacialoides, snow petrel Pagodroma nivea). Across all species and during a 32‐year period, immigrants made on average ?9 to 29% fewer breeding attempts, had 5–31% fewer fledglings, had 2–16% lower breeding success and produced 6–46% fewer recruits. Female immigration and male residency were also favored through differences in breeding performance. We provide evidence for selection against immigrants in wild populations of long‐lived species and our results are consistent with female‐biased dispersal in birds being driven by asymmetric limiting resources and the competitive ability of dispersers vs. non‐dispersers.     

Quantifying the contribution of immigration to population dynamics: a review of methods,evidence and perspectives in birds and mammals

The demography of a population is often reduced to the apparent (or local) survival of individuals and their realised fecundity within a study area defined according to logistical constraints rather than landscape features. Such demographics are then used to infer whether a local population contributes positively to population dynamics across a wider landscape context. Such a simplistic approach ignores a fundamental process underpinning population dynamics: dispersal. Indeed, it has long been accepted that immigration contributed by dispersers that emigrated from neighbouring populations may strongly influence the net growth of a local population. To date however, we lack a clear picture of how widely immigration rate varies both among and within populations, in relation to extrinsic and intrinsic ecological conditions, even for the best‐studied avian and mammalian populations. This empirical knowledge gap precludes the emergence of a sound conceptual framework that ought to inform conservation and population ecology. This review, conducted on both birds and mammals, has thus three complementary objectives. First, we describe and evaluate the relative merits of methods used to quantify immigration and how they relate to widely applicable metrics. We identify two simple and unifying metrics to measure immigration: the immigration rate i_t defined as the ratio of the number of immigrants present in the population at time t + 1 and the total breeding population in year t, and π_t , the proportion of immigrants among new recruits (i.e. new breeders). Two recently developed methods are likely to provide the most valuable data on immigration in the near future: individual parentage (rather than population) assignments based on genetic sampling, and spatially explicit integrated population models combining multiple sources of demographic data (survival, fecundity and population counts). Second, we report on a systematic literature review of studies providing a quantitative measure of immigration. Although the diversity of methods employed precludes detailed analyses, it appears that the number of immigrants exceeds locally born individuals in recruitment for most avian populations (median π_t  = 0.57, N = 45 estimates from 37 studies), a figure twofold higher than estimated for mammalian populations (median π_t  = 0.26, N = 33 estimates from 11 studies). Third, recent quantitative studies reveal that immigration can be the main driver of temporal variation in population growth rates, across a wide array of demographic and spatial contexts. To what extent immigration acts as a regulatory process has however been considered only rarely to date and deserves more attention. Overall, it is likely that most populations benefit from immigrants without necessarily being sink populations. Furthermore, we suggest that quantitative estimates of immigration should be core to future demographic studies and plead for more empirical evidence about the ways in which immigration interacts with local demographic processes to shape population dynamics. Finally, we discuss how to tackle spatial population dynamics by exploring, beyond the classical source–sink framework, the extent to which populations exchange individuals according to spatial scale and type of population distribution throughout the landscape.     

Influential factors for natal dispersal in an avian island metapopulation

Dispersal is a key parameter in evolutionary, demographic and conservation theory, but the factors influencing dispersal between populations are rarely known, and the contribution of immigrants to population stability remains uncertain. Using dispersal data from nine island populations of song sparrows, we show that female and male immigrants responded differently to population structure: in females, immigration varied with adult sex ratio; whereas immigration by males was more influenced by population density. These patterns are consistent with the hypothesis that intra-sexual competition for breeding resources influenced recruitment patterns. Immigrants often constituted a substantial fraction of local population size, and in six cases immigration by females prevented the extirpation of that sex from the island. Breeding vacancies and extirpations may have been more likely in females because their apparent survival was lower than in males. Local recruitment and immigration varied markedly among islands, perhaps as consequence of island size and isolation. Overall, our results suggest that immigration varied with local demography in a sex-specific way, stabilized population numbers and reduced extinction rates in the smallest populations.     

Factors affecting immigration of adults: experimental and theoretical observations with rodents

We examined immigration in populations of Peromyscus leucopus (white-footed mice) by 1) monitoring natural immigration at 10 sites, 2) introducing experimental immigrants into eight populations, and 3) constructing qualitative models of immigration. Density of natural immigrants covaried positively with resident density, and successful assimilation was lower at low resident and immigrant densities. Females and males did not differ in their chance of achieving residency. Year, sex, and number introduced were significant predictors of assimilation by individuals. Experimentally introduced individuals had no effect on densities of those mice resident before the introductions. Results obtained from studying natural immigration differed from those obtained from studying experimental immigration. Signed digraphs and time averaging were used to model the consequences of different resident-immigrant relationships and to suggest how different sources of variation may have affected assimilation and led to differences in natural and experimental immigration. Resident and immigrant densities could have been positively correlated even if residents actively inhibited immigration. Variation in immigrant density and survival rather than resident territorial activity apparently determined patterns of assimilation.     

The dynamics of the yeast community of the Tagus river estuary: testing the hypothesis of the multiple origins of estuarine yeasts

Yeasts are common inhabitants of different types of aquatic habitats, including marine and estuarine waters and rivers. Although numerous studies have surveyed yeast occurrence in these habitats, the identification of autochthonous populations has been problematic because several yeast species seem to be very versatile and therefore mere presence is not sufficient to establish an ecological association. In the present study we investigated the dynamics of the yeast community in the Tagus river estuary (Portugal) by combining a microbiological study involving isolation, quantification, and molecular identification of dominant yeast populations with the analysis of hydrological and hydrographical data. We set out to test the hypothesis of the multiple origins of estuarine yeast populations in a transect of the Tagus estuary and we postulate four possible sources: open sea, terrestrial, gastrointestinal and the estuary itself in the case of populations that have become resident. Candida parapsilosis and Pichia guilliermondii were correlated with Escherichia coli, which indicated an intestinal origin. Other cream-colored yeasts like Debaryomyces hansenii and Candida zeylanoides had similar dynamics, but no association with E. coli and quite distinct ecological preferences. They might represent a group of resident estuarine populations whose primary origin is diverse and can include marine, terrestrial, and gastrointestinal habitats. Another major yeast population was represented by Rhodotorula mucilaginosa. The cosmopolitan nature of that species and its moderate association with E. coli point to terrestrial sources as primary habitats.     

Emigration to new habitats by voles: the cost of dispersal paradox

We tested the hypothesis that dispersal and philopatry are components of a mixed evolutionarily stable strategy (ESS). The hypothesis predicts that fitness of dispersers should be equal to that of philopatric individuals. Alternatively, fitness of dispersers could be lower (the resident fitness hypothesis) or greater (the cost of dispersal hypothesis) than that of philopatric individuals. We compared fitness of individuals that moved to new habitats (emigrants) and those that remained within habitat boundaries (residents) in populations of the prairie vole, Microtus ochrogaster, and the meadow vole, M. pennsylvanicus. We established vole populations in four enclosures (). Within each enclosure, voles were free to move between four types of habitats that varied in the availability of supplemental food and the amount of vegetative cover. We analysed two fitness components: the survival rates of all individuals, and pregnancy rates of females. Our study showed that emigrants generally had greater fitness than residents and that the difference in fitness was habitat dependent (i.e. was greater when individuals were emigrating from low-quality habitats than from high-quality habitats). High-food, high-cover habitats were the only habitat types for which fitness of emigrants was lower than that of residents. Similar patterns occurred in both prairie voles and meadow voles. Our results support the cost of dispersal hypothesis.     

The correlation between demography and metabolic rate: a test using the beach vole (Microtus breweri) and the meadow vole (Microtus pennsylvanicus)

Summary This study examines the hypothesis that mammalian species with wide fluctuations in population size will have greater metabolic rates than species with smaller population fluctuations. We tested this hypothesis using two microtine rodents — the beach vole (Microtus breweri) and the meadow vole (M. pennsylvanicus). Although these species experience similar climatic regimes, eat similar foods, and have a very close phylogenetic relationship, they show marked differences in demography. Microtus pennsylvanicus is prone to large supraannual fluctuations in population size, while M. breweri is essentially acyclical. Metabolic rate (oxygen consumption) of each species was measured using open-flow respirometry at ambient temperatures ranging from 2 to 34° C. Basal metabolic rate of M. pennsylvanicus (1.81 ml O₂ g^–1 h^–1) was significantly greater than that of M. breweri (1.39 ml O₂ g^–1 h^–1). The lower critical temperature, estimated by continuous two-phase regression, was 28.9° C for M. pennsylvanicus and 29.8° C for M. breweri. Regression lines below thermoneutrality did not differ in slope, but the elevation for M. pennsylvanicus was significantly higher. Thus, M. pennsylvanicus has a higher metabolic rate at all temperatures examined. These results support the hypothesis that metabolic rate is positively correlated with the extent of population fluctuation. We suggest that further evidence for, or against, this hypothesis should be found by comparing closely matched species pairs, rather than resorting to confounded allometric comparisons of ecologically and phylogenetically diverse taxa.     

The influence of stage-dependent dispersal on the population dynamics of three amphipod species

In metapopulations, the maintenance of local populations can depend on source–sink dynamics, where populations with positive growth rate seed populations with negative growth rate. The pattern and probability of successful dispersal among habitats can therefore be crucial in determining whether local populations will become rare or increase in abundance. We present here data on the dispersal strategy and population dynamics of three marine amphipods living in pen shells (Atrina rigida) in the Gulf of Mexico. The three amphipod species in this study disperse at different life stages. Neomegamphopus hiatus and Melita nitida disperse as adults, while Bemlos unicornis disperses as juveniles. The two species that disperse as adults have the highest initial population sizes when a new shell becomes available, likely caused by the arriving females releasing their brood into these recently occupied shells. This dispersal pattern results in initially higher population growth, but fewer occupied shells, as noted by their clumped distribution. In contrast, the species that disperses as juveniles accumulates more slowly and more evenly across habitats, eventually dominating the other two in terms of numerical abundance. The metapopulation dynamics of the three species seems to be highly dependent on the life history stage involved in dispersal. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Peripheral genetic structure of Helicoverpa zea indicates asymmetrical panmixia

Seasonal climatic shifts create peripheral habitats that alternate between habitable and uninhabitable for migratory species. Such dynamic peripheral habitats are potential sites where migratory species could evolve high genetic diversity resulting from convergence of immigrants from multiple regionally distant areas. Migrant populations of Helicoverpa zea (Boddie) captured during two different seasons were assessed for genetic structure using microsatellite markers and for host plant type using stable carbon isotope analysis. Individuals (N = 568) were genotyped and divided into 13 putative populations based on collection site and time. Fixation indices (F‐statistics), analysis of molecular variance (AMOVA), and discriminant analysis of principal components (DAPC) were used to examine within and among population genetic variation. Mean number of alleles per locus was 10.25 (± 3.2 SD), and allelic richness ranged from 2.38 to 5.13 (± 3.2 SD). The observed and expected heterozygosity ranged from 0.07 to 0.48 and 0.08 to 0.62, respectively. Low F_ST (0.01 to 0.02) and high F_IS (0.08 to 0.33) values suggest captured migrants originated from breeding populations with different allele frequencies. We postulate that high genetic diversity within migrant populations and low genetic differentiation among migrant populations of H. zea are the result of asymmetrical immigration due to the high dispersal and reproductive behavior of H. zea, which may hinder the adaptation and establishment of H. zea to peripheral habitat. These findings highlight the importance of assessing peripheral population structure in relation to ecological and evolutionary dynamics of this and other highly reproductive and dispersive species.     

The demographic drivers of local population dynamics in two rare migratory birds

The exchange of individuals among populations can have strong effects on the dynamics and persistence of a given population. Yet, estimation of immigration rates remains one of the greatest challenges for animal demographers. Little empirical knowledge exists about the effects of immigration on population dynamics. New integrated population models fitted using Bayesian methods enable simultaneous estimation of fecundity, survival and immigration, as well as the growth rate of a population of interest. We applied this novel analytical framework to the demography of two populations of long-distance migratory birds, hoopoe Upupa epops and wryneck Jynx torquilla, in a study area in south-western Switzerland. During 2002–2010, the hoopoe population increased annually by 11%, while the wryneck population remained fairly stable. Apparent juvenile and adult survival probability was nearly identical in both species, but fecundity and immigration were slightly higher in the hoopoe. Hoopoe population growth rate was strongly correlated with juvenile survival, fecundity and immigration, while that of wrynecks strongly correlated only with immigration. This indicates that demographic components impacting the arrival of new individuals into the populations were more important for their dynamics than demographic components affecting the loss of individuals. The finding that immigration plays a crucial role in the population growth rates of these two rare species emphasizes the need for a broad rather than local perspective for population studies, and the development of wide-scale conservation actions.     

Density-dependent immigration promotes population stability in a long-distance migratory bird

The spatial structure of populations determines the relative importance of reproduction, survival and movement on population dynamics. However, the mechanisms by which local individuals and immigrants interact and the subsequent effects of immigrants on productivity are poorly known. We developed an integrated population model (IPM) to study the extent and consequences of immigration on the dynamics of a neotropical migrant (American redstart, Setophaga ruticilla) over an 11-year period in Ontario, Canada. New immigrants represented the majority of the study population each year with higher immigration rates for males than females and for first-year breeders than breeders in their second year or older. Immigration was negatively density dependent, with immigrants replacing previously established breeders in a compensatory manner following their death or emigration. Because of the tradeoff between immigration and apparent survival, neither had a strong influence on population growth and reproductive output was most strongly correlated with a change in abundance between years. However, if immigration ceased, the study population would become locally extinct within 7 years and thus immigrants were essential for local population persistence. We found no evidence for reduced breeding success when immigrants represented a higher proportion of the study population. Our research highlights the importance of movement in the stability of open populations and the strong correlation between the fates of local breeders and the number of immigrants entering the population. We recommend the use of IPMs to address the spatial scale over which immigration occurs and how different scales influence its contribution to population dynamics.     

Genetic diversities of four little-known species of Malesherbia (Malesherbiaceae) endemic to the arid inter-Andean valleys of Peru

Few studies of genetic variation have been conducted on plants of the Pacific coastal desert and neighboring Andes of Peru, although the region has many endemic taxa. Enzyme electrophoresis was employed to examine allozyme diversities of four species of the family Malesher-biaceae, an endemic to the arid Andes and coastal desert. One population ofMalesherbia splendens and two ofM. tubulosa, both endemics of the department of Lima, one population ofM. weberbaueri var.weberbaueri, an endemic of the Andean department Huancavélica, and the two Lima populations ofM. scarlatiflora were studied. Fifteen loci were examined for all populations and an additional seven loci were resolved forM. weberbaueri andM. splendens. Malesherbia splendens, which is known from three populations, has a low mean number of alleles per locus (A), proportion of polymorphic loci (P), and expected heterozygosity (H_s) (A=1.214, P=0.214, H_s=0.057).Malesherbia weberbaueri (A=1.231, P=0.154, H_s=0.079) andM. scarlatiflora (A=1.364, P=0.273, H_t=0.083) both have average expected heterozygosities and relatively low mean numbers of alleles per locus and proportions of polymorphic loci. InM. tubulosa, all measures of genetic diversity are high in comparison with other endemics (A=1.818, P=0.364, H_t=0.206).Malesherbia tubulosa has high interpopulation differentiation, whereasM. scarlatiflora has low among-population diversity. The relatively low allozyme diversities, restricted habitats, narrow ranges ofM. splendens andM. weberbaueri, and the morphological similarities of all four species suggest that they evolved recently by founder events. Greater allozyme diversities inM. tubulosa could be attributable to its maintenance of larger populations in a greater variety of habitats.