Regulation of local populations of a coral reef fish via joint effects of density- and number-dependent mortality

Density-dependent mortality can regulate local populations - effectively minimizing the likelihood of local extinctions and unchecked population growth. It is considered particularly important for many marine reef organisms with demographically open populations that lack potential regulatory mechanisms tied to local reproduction. While density-dependent mortality has been documented frequently for reef fishes, few studies have explored how the strength of density-dependence varies with density, or how density-dependence may be modified by numerical effects (i.e., number-dependent mortality). Both issues can have profound effects on spatial patterns of abundance and the regulation of local populations. I address these issues through empirical studies in Moorea, French Polynesia, of the six bar wrasse (Thalassoma hardwicke), a reef fish that settles to isolated patch reefs. Per capita mortality rates of newly settled wrasse increased as a function of density and were well approximated by the Beverton-Holt function for both naturally formed and experimentally generated juvenile cohorts. Average instantaneous mortality rates were a decelerating function of initial densities, indicating the per capita strength of density-dependence decreased with density. Results of a factorial manipulation of density and group size indicate that per capita mortality rates were simultaneously density- and number-dependent; fish at higher densities and/or in groups had higher probabilities of disappearing from patch reefs compared with fish that were solitary and/or at lower densities. Mortality rates were ~30% higher for fish at densities of 0.5 fish/m² than at 0.25 fish/m². Similarly, mortality rates increased by ~45% when group size was increased from 1 to 2 individuals per patch, even when density was kept constant. These observations suggest that the number of interacting individuals, independent of patch size (i.e., density-independent effects) can contribute to regulation of local populations. Overall, this work highlights a greater need to consider numerical effects in addition to density effects when exploring sources of population regulation.     

Temporal changes in the strength of density-dependent mortality and growth in intertidal barnacles

1. In demographically open marine systems, the extent to which density-dependent processes in the benthic adult phase are required for population persistence is unclear. At one extreme, represented by the recruitment limitation hypothesis, larval supply may be insufficient for the total population size to reach a carrying capacity and density-independent mortality predominates. At the opposite extreme, populations are saturated and density-dependent mortality is sufficiently strong to reshape patterns established at settlement. 2. We examined temporal variation in the way density-independent and density-dependent mortality interact in a typical sessile marine benthic invertebrate, the acorn barnacle Semibalanus balanoides (L.), over a 2-year period. 3. Recruitment was manipulated at two high recruitment sites in north Wales, UK to produce recruit densities covering the range naturally found in this species. Following manipulation, fixed quadrats were monitored using digital photography and temporal changes in mortality and growth rate were examined. 4. Over a 2-year period there was a clear, spatially consistent, over-compensatory relationship between the density of recruits and adult abundance indicating strong density-dependent mortality. The strength of density dependence intensified with increasing recruitment. 5. Density-dependent mortality did not operate consistently over the study period. It only operated in the early part of the benthic phase, but the pattern of adult abundance generated was maintained throughout the whole 2-year period. Thus, early life-history processes dictated adult population abundance and dynamics. 6. Examination of the natural recruitment regime in the area of study indicated that both positive and negative effects of recruitment will occur over scales varying from kilometres to metres.     

What makes a species common? No evidence of density-dependent recruitment or mortality of the sea urchin Diadema antillarum after the 1983–1984 mass mortality

A potential consequence of individuals compensating for density-dependent processes is that rare or infrequent events can produce profound and long-term shifts in species abundance. In 1983–1984 a mass mortality event reduced the numbers of the abundant sea urchin Diadema antillarum by 95–99 % throughout the Caribbean and western Atlantic. Following this event, the abundance of macroalgae increased and the few surviving D. antillarum responded by increasing in body size and fecundity. These initial observations suggested that populations of D. antillarum could recover rapidly following release from food limitation. In contrast, published studies of field manipulations indicate that this species had traits making it resistant to density-dependent effects on offspring production and adult mortality; this evidence raises the possibility that density-independent processes might keep populations at a diminished level. Decadal-scale (1983–2011) monitoring of recruitment, mortality, population density and size structure of D. antillarum from St John, US Virgin Islands, indicates that population density has remained relatively stable and more than an order of magnitude lower than that before the mortality event of 1983–1984. We detected no evidence of density-dependent mortality or recruitment since this mortality event. In this location, model estimates of equilibrium population density, assuming density-independent processes and based on parameters generated over the first decade following the mortality event, accurately predict the low population density 20 years later (2011). We find no evidence to support the notion that this historically dominant species will rebound from this temporally brief, but spatially widespread, perturbation.     

Density-dependent reproductive output in relation to a drastically varying food supply: getting the density measure right

When a limiting resource (e.g. food) varies drastically between years, and population density is measured in the conventional way as individuals per area, demographic processes such as productivity and survival may erroneously be considered density-independent. We tested the hypothesis that if the variation in a limiting resource is not taken into account in the density measure, this may lead to erroneous conclusions about the density-dependence of demographic variables. We studied the food-related variation in productivity of bramblings Fringilla montifringilla , an insectivorous passerine bird, using 19 years of standardised insect censusing, bird censusing and mist-netting of birds in subalpine birch forest in Swedish Lapland. The yearly variation in our measure of brambling per capita productivity (numbers of juveniles per adult trapped) was explained to 30–40% by the larvae abundance of the moth Epirrita autumnata . Taking larvae density into account, no other environmental variable (inferred predation pressure, breeding phenology, and summer temperature) was significantly related to variation in reproductive output. There was no effect of brambling population density on per capita productivity, that is, when density was measured the conventional way, productivity seemed density-independent. However, per capita productivity was significantly and negatively correlated to the food-related population density (population density divided by larval density), supporting the hypothesis that not including a limiting resource into the density measure may indeed lead to erroneous conclusions about the density-dependence of demographic variables.     

Biology ofhyphantria cunea drury (lepidoptera : Arctiidae) in Japan. IX. population dynamics

Fluctuations in population density of Hyphantria cunea in Japan are characterized by a gradation-like pattern. Analysis of the life table data taken from two stations during eight successive generations showed that (1) mortality during egg and early larval stages was density-independent, (2) mortality during later larval stages was inversely density-dependent, and (3) mortality during prepupal and pupal stages was density-independent. Thus the overall mortality process from egg to adults eclosion was inversely density-dependent. The inverse density-dependence in mortality process during later larval instars was mainly attributed to the ‘escape’ (VOÛTE, 1946) of H. cunea populations from the predation pressure of polyphagous predators such as birds and Politses wasps. This inverse density-dependence was considered to be a cause of gradation-like fluctuation. Field collection of egg-masses showed that the mean number of eggs per egg-mass, which was believed to be a good representation of mean fecundity, varied from 425 to 1050 during 4 years. Density-dependent reduction in the mean number of eggs per egg-mass was demonstrated, and this reduction was a factor regulating the population density. Assuming fixed sex ratio and survival rate of adults, a preliminary population model was constructed. The number of eggs laid in the survey station could be predicted well by the model based on the number of eggs laid in the previous generation, in 9 out of 13 cases. An attempt to apply a model of the same type to mimic the fluctuation of abundance (peak number of larval colonies per tree) on road-side trees suggested another density-dependent process, that is, insecticide application by man. Discussion was also presented on the causes responsible for the turn of population trend from decreasing to increasing in the 1st generation of 1968.     

Post-settlement emigration affects mortality estimates for two Bahamian wrasses

Understanding the dynamics of open marine populations is inherently complex, and this complexity has led to decades of debate regarding the relative importance of pre- versus post-settlement processes in structuring these populations. Movement between patches may be an important modifier of patterns established at settlement, yet local immigration and emigration have received less attention than other demographic rates. I examined loss rates from tagged populations of juvenile wrasses (yellowhead wrasse Halichoeres garnoti and bluehead wrasse Thalassoma bifasciatum) at two sites in the Bahamas. Assuming that all losses were due solely to mortality would have significantly underestimated survivorship of yellowhead wrasse by 29% and bluehead wrasse by 14%. On average, per capita mortality and emigration rates were higher for yellowhead than bluehead wrasse, but neither demographic rate differed between sites for either species. With respect to within-species density, bluehead wrasse mortality was density-dependent at the patch reef site, but mortality rates of yellowhead wrasse were consistently density-independent. Evaluating the effects of between-species density, yellowhead wrasse mortality increased with a decrease in bluehead wrasse density, but this effect was limited to the patch reef site. Emigration rates were not a function of either within-species or between-species density, but instead varied inversely with isolation distance. Numerous previous studies of coral-reef fish, conducted on patch reefs separated by only a few meters of sand and often using untagged fish, may have confounded losses due to emigration with those due to mortality. A better understanding of the factors affecting emigration in marine fishes is important to their effective management using spatial tools such as marine protected areas.     

The significance of variable and density-independent post-recruitment mortality in local populations of reef fishes

Abstract In this paper I focus on how post-settlement mortality may modify initial patterns of settlement in reef fish. Infrequent recruitment surveys may underestimate the role of early post-settlement mortality as most mortality in reef fishes occurs shortly after settlement. Consequently, results from infrequent recruitment surveys shed little light on the mechanisms producing patterns of abundance because these surveys ignore early post-settlement mortality. Variation in density-independent mortality may be a common mechanism that can prevent a positive relationship between larval settlement and subsequent population abundance. Although density-dependent mortality is the most commonly recognized mechanism that can disrupt the correlation between settlement and adult abundance, density-independent mortality’ can also destroy this correlation if the variance associated with post-settlement mortality is greater than variance in settlement. This point is illustrated with a simulation model in which I modelled two populations: a piscivorous fish population that was recruitment-limited with constant mortality, and a prey population that had variable recruitment and mortality that was a function of the size of the predator population. The results of this model indicate that even when mortality of prey is density-independent, predation can determine prey abundance when variation in piscivore recruitment is high relative to prey recruitment. Thus, initial patterns of prey settlement can be modified by a recruitment-limited predator population.     

Plants as reef fish: fitting the functional form of seedling recruitment

The life histories of many species depend first on dispersal to local sites and then on establishment. After dispersal, density-independent and density-dependent mortalities modify propagule supply, determining the number of individuals that establish. Because multiple factors influence recruitment, the dichotomy of propagule versus establishment limitation is best viewed as a continuum along which the strength of propagule or establishment limitation changes with propagule input. To evaluate the relative importance of seed and establishment limitation for plants, we (1) describe the shape of the recruitment function and (2) use limitation and elasticity analyses to quantify the sensitivity of recruitment to perturbations in seed limitation and density-independent and density-dependent mortality. Using 36 seed augmentation studies for 18 species, we tested four recruitment functions against one another. Although the linear model (accounting for seed limitation and density-independent mortality) fitted the largest number of studies, the nonlinear Beverton-Holt model (accounting for density-dependent mortality) performed better at high densities of seed augmentation. For the 18 species, seed limitation constrained population size more than other sources of limitation at ambient conditions. Seedling density reached saturation with increasing seed density in many studies, but at such high densities that seedling density was primarily limited by seed availability rather than microsite availability or density dependence.     

Density-dependent parameters and demographic equilibrium in open populations

Populations near their equilibrium are expected to show density-dependence through a negative feedback on at least one demographic parameter, e.g. survival and/or productivity. Nevertheless, it is not always clear which vital rate is affected the most, and even less whether this dependence holds in open populations in which immigration and emigration are also important. We assessed the relative importance of population density in the variation of local survival, recruitment, proportion of transients (emigrants) and productivity through the analysis of detailed life-histories of 4286  seabirds from a colony that reached an apparent demographic equilibrium after a period of exponential increase. We provide evidence that the role of population density and resource availability changes according to the demographic parameter considered. Estimates indicated that transients increased from 5% to 20% over the study period, suggesting an average turnover of about 1400 individuals per year. The parameters most influenced by population density alone were local survival and probability of transience. Recruitment was negatively associated with population density during the increasing phase but unexpected high values were also recorded at high population levels. These high values were explained by a combination of population size and food availability. Mean productivity varied with food availability, independently from population variations. The population density alone explained up to a third of the yearly variation of the vital rates considered, suggesting that open populations are equally influenced by stochastic and density-independent events (such as environmental perturbations) than by intrinsic (i.e. density-dependent) factors.     

Population regulation by post-settlement mortality in two temperate reef fishes

 Input of individuals dispersing into open populations can be highly variable, yet the consequences of such variation for subsequent population densities are not well understood. I explored the influence of variable input (”supply”) on subsequent densities of juveniles and adults in open local populations of two temperate reef fishes, the bluebanded goby (Lythrypnus dalli) and the blackeye goby (Coryphopterus nicholsii). Variable recruitment was simulated by stocking a natural range of densities of young fishes on replicate patch reefs. Density and mortality of the stocked cohorts were followed over time, until the fishes reached maturity. Over the first day of the experiments, mortality of both species was significantly density-dependent; however, there was still a very strong relationship between density on day 1 and density on day 0 (i.e., simulated recruitment was still an excellent predictor of population density). At this point in the study, the main effects of density-dependent mortality were to reduce mean densities and variation about the mean. Over the period from the start of the experiments until the time when maturity was reached by each species (about 1 and 3 months for Lythrypnus and Coryphopterus, respectively), mortality was strongly density-dependent. Such strong density-dependent mortality virtually eliminated any linear relationship between adult density and ”recruit” density. However, for both species, the relationship between these two variables was well fit by an asymptotic curve, with the asymptotic density of adults equal to c. 3/m² for Coryphopterus, and c. 10/m² for Lythrypnus. Natural recruitment (via settlement of larvae) to the reefs over the period of the study (9 months) was above the asymptotic densities of adults for the two species, even though the study did not encompass the periods of peak annual recruitment of either species. This suggests that adult populations of these two gobies may often be limited, and regulated, by post-settlement processes, rather than by input of settlers. Other studies have shown that mortality of the two species is density-independent, or only weakly density-dependent, on reefs from which predators have been excluded. Hence, it appears that predators cause density-dependent mortality in these fishes. Received: 26 November 1996 / Accepted: 5 April 1997     

The survival, growth and diet of pike fry, Esox lucius L., stocked at different densities in experimental ponds

Yolk-sac pike fry were stocked at densities of 0.74 – 81.4 m⁻² in two ponds, each divided into eight sectors (mean area 155.8 m²). Growth and survival were recorded from May to December 1985. The growth rates were variable within each sector. The size-range of sampled fish increased throughout the year, but showed no significant correlation with density. Fry survival was initially density-independent but switched by late June/July to density-dependence, ranging from 0.5 to 43.6% of initial stocking numbers. The highest mean daily mortality rates occurred during May-July. The final survival in December ranged between sectors from 0.059 to 11.25% of the starting stock densities. The final biomass per unit area of pike surviving in December was not related to initial stocking density. In Pond 1 the mean biomass produced was 2.21 gm⁻² and in Pond 2 was 3.49gm⁻².
Pike fry < 30 mm fed only on invertebrates; those 30–100 mm took a wide range of invertebrates, cyprinids. sticklebacks and other pike. Cannibalism occurred at most densities between 5.45 and 81.4 fish m⁻².
Where attempts are made to increase pike production in managed populations by releasing small fry, an upper stock density of 5 fry m⁻² is advised if large, density-dependent mortalities are to be avoided.     

The dynamics of abundance and incidence of annual plant species during colonization in a desert

We studied colonization of annual plants in small-scale disturbances and undisturbed soil for four years in the northern Negev desert. The experiment consisted of 24 patches of I m²: eight undisturbed patches, eight 30 cm deep pits and eight 20 cm high mounds. Disturbance removed the seed bank from the pits and mounds. Rainfall was average during 1992 and 1993 (163.5 and 157.0 mm), very low in 1994 (97.5 mm) and very high in 1995 (283.0 mm). For all ca 100 species in the assemblage, we measured abundance, as the average local density in patches occupied by the species, and incidence, the proportion of patches occupied. Abundance and incidence were positively correlated in the four years of the study (R^2-= 0.71, 0.56. 0. 56 and 0.64). The relationship became steeper during the course of colonization. A minority of species colonized all patches rapidly and increased exponentially in abundance. These high incidence and abundance species responded little or not at all to variations in rainfall. Most of the species experienced frequent local extinctions and colonizations in few patches. These low abundance and low to medium incidence species responded strongly to rainfall by fluctuations in abundance. Differences per species between abundance and/or incidence in pits, mounds and matrix were evident in many species along the entire relationship. These were mostly in favor of pits. We propose that the positions and trajectories of the individual species in the abundance-incidence phase plane signify constraints on population growth along the abundance axis, and on colonization and population persistence along the incidence axis. Since species with lower incidence and abundance respond strongly to rainfall variation, we conclude that their local populations and spatial distribution are to a large extent site-limited. High-abundance and high-incidence species have few or weak constraints and experience density-independent population growth, which is only seed-limited. The causes of site limitation and its effect on population growth and on eotonization and extinction processes differ for different species. In a number of species, abundance and incidence may be limited by dispersal, in conjunction with dispersal mode and patch-specific seed capture. Based on our case study we suggest that the trajectories of the species in the abundance-incidence phase plane are a useful tool for investigating assemblage dynamics.     

Quantifying density-independent mortality of temperate tree species

Forest resilience to climate change is a topic of national concern as our standing assets and future forests are important to our livelihood. Many tree species are predicted to decline or disappear while others may be able to adapt or migrate. Efforts to quantify and disseminate the current condition of forests are urgently needed to guide management and policy. Here, we develop a new indicator to summarize raw density-independent mortality of forested stands by species from the last decade of the 20th century to the first decade of the 21st century using forest inventory data. We define density-independent mortality to be stand mortality by species due to processes unrelated to natural mortality from succession or stand maturation, which is marked by overall increase in tree girth at the expense of density of individuals. We assess trends for 22 species on national forests in two U.S. states, Washington and Oregon. Populations of some species including timber species have no or low overall levels of density-independent mortality (Juniperus occidentalis, Abies procera, Thuja plicata, Tsuga heterophylla, Pinus ponderosa, and Pseudotsuga menziesii). In contrast, other species demonstrate unsustainable levels of density-independent mortality (Pinus monticola, Arbutus menziesii, Pinus albicaulis, Abies lasiocarpa, Taxus brevifolia, Pinus contorta, Abies grandis, Picea englemanii, and Larix occidentalis). Additionally, the net potential for unsustainable levels of density-independent mortality in standing populations does not necessarily warrant concern when examined across species for our study area and time period; however, when examined by species, the number of species in decline exceeds the number of species where mortality can be generally attributed to endogenous self-thinning. We argue that this work can aid management and policy decisions and our understanding of complex vegetation dynamics in a changing climate. Application of the indicator at larger spatial scales and in conjunction with data on migration and establishment may be used to address questions such as, how can we make cost-effective management decisions to ensure long-term sustainability of tree species and forests? Tree species distributions across the landscape are complex systems, and raw characterization of current trends occurring in forest inventories is important especially given the uncertainty associated with the modeling and prediction of complex systems such as tree species.     

Population regulation in trichostrongylids of ruminants

Since regulation of population size requires the existence of one or more density-dependent processes affecting parasite numbers, the literature was examined for evidence of density-dependence in establishment, reproduction and mortality of trichostrongylids of ruminants. Because of differences between the two environments required for completion of the trichostrongylid life-cycle, this evidence was sought in processes occurring within the host, rather than in dung-pats or on pasture. Evidence was found of at least one density-dependent population process in several economically important species of the genera Haemonchus, Ostertagia and Trichostrongylus, although further observations on populations derived from continuous infection at several rates of larval intake are required to substantiate much of this evidence. It is suggested that populations of trichostrongylids of ruminants are regulated at the level of the suprapopulation by density-dependent constraints on egg production of constituent infrepopulations; rates of larval intake may play a central role in coordinating regulation within each ecosystem.     

Population variability among three small mammal species in the semiarid Neotropics: the role of density-dependent and density-independent factors

We addressed the role of density-dependent (direct and delayed) and density-independent (precipitation) factors in shaping the dynamics of fluctuating populations of three small mammal species. Using a stepwise regression procedure, we tested the effects of nonlagged population density (log₁₀ N_t-1), lagged population density (log₁₀ N_t-2), and annual precipitation on the per capita rate of population change of Phyllotis darwini, Akodon olivaceus , and Thylamys elegans in two habitat types of a semiarid region of Chile. The most irruptive species ( P. darwini ) showed direct and delayed density-dependent effects in equatorial subpopulation, and only direct density-dependence in polar subpopulation. The per capita rates of population change of A. olivaceus showed direct density-dependent and precipitation effects in both habitats types, while T. elegans showed direct density-dependence and precipitation effects in the equatorial subpopulation but only a marginal effect of direct density-dependence in the polar subpopulation. The presence of delayed density-dependent strongly suggests the importance of biological interactions in shaping the dramatic irruptions exhibited by P. darwini.     

The role of density-dependent dispersal in source-sink dynamics

I investigate two aspects of source-sink theory that have hitherto received little attention: density-dependent dispersal and the cost of dispersal to sources. The cost arises because emigration reduces the per capita growth rate of sources, thus predisposing them to extinction. I show that source-sink persistence depends critically on the interplay between these two factors. When the emigration rate increases with abundance at an accelerating rate, dispersal costs to sources is the lowest and risk of source-sink extinction the least. When the emigration rate increases with abundance at a decelerating rate, dispersal costs to sources is the highest and the risk of source-sink extinction the greatest. Density-independent emigration has an intermediate effect. Thus, density-dependent dispersal per se does not increase or decrease source-sink persistence relative to density-independent dispersal. The exact mode of dispersal is crucial. A key point to appreciate is that these effects of dispersal on source-sink extinction arise from the temporal density-dependence that dispersal induces in the per capita growth rates of source and sink populations. Temporal density-dependence due to dispersal is beneficial at low abundances because it rescues sinks from extinction, and detrimental at high abundances because it drives otherwise viable sources to extinction. These results are robust to the nature of population dynamics in the sink, whether exponential or logistic. They provide a means of assessing the relative costs and benefits of preserving sink habitats given three biological parameters.     

Alternate attractors in the population dynamics of a tree-killing bark beetle

Among the most striking changes in ecosystems are those that happen abruptly and resist return to the original condition (i.e., regime shifts). This frequently involves conspicuous changes in the abundance of one species (e.g., an oubreaking pest or keystone species). Alternate attractors in population dynamics could explain switches between low and high levels of abundance, and could underlie some cases of regime shifts in ecosystems; this longstanding theoretical possibility has been difficult to test in nature. We compared the ability of an alternate attractors model versus two competing models to explain population fluctuations in the tree-killing bark beetle, Dendroctonus frontalis. Frequency distributions of abundance were distinctly bimodal, a prediction of the alternate attractors model, strongly indicating the lack of a single, noisy equilibrium. Time series abundance data refuted the existence of strong delayed density-dependence or nonlinearities, as required by the endogenous cycles model. The model of alternate attractors was further supported by the existence of positive density-dependence at intermediate beetle abundances. Experimental manipulations show that interactions with competitors and shared enemies could create a locally stable equilibrium in small populations of D. frontalis. High variation among regions and years in the abundance of predators and competitors could permit switches between alternate states. Dendroctonus frontalis now provides the strongest case that we know of for alternate attractors in natural population dynamics. The accompanying demographic instability appears to underlie spatially extensive outbreaks that have lasting impacts on forest ecosystems. Understanding feedbacks in populations with alternate attractors can help to identify thresholds underlying regime shifts, and potentially manage them to avoid undesirable impacts.     

Role of predators in the early post-settlement demography of coral-reef fishes

Populations with dispersive larvae are often demographically open such that local reproduction and subsequent larval settlement are not linked. Thus, understanding whether and how settlement patterns are established and subsequently modified is central to understanding local demography. Settlement is typically not measured directly, but rather it is estimated by recruitment, which is the observation of new individuals sometime after settlement. At Lizard Island on the Great Barrier Reef, I examined how patterns of recruitment of coral-reef fishes were modified across a range of natural recruit densities in the presence and absence of resident predators. Resident predators decreased recruitment and increased mortality for all species, but these effects varied considerably among species. The effects of predators on recruitment were at least partly due to mortality within 2 days after settlement. At their most extreme, predators caused recruitment failure of several species of butterflyfish. For one species of damselfish (Pomacentrus amboinensis), predators both induced weakly density-dependent mortality and obscured any relationship between recruitment and subsequent abundance, while for another damselfish (Neopomacentrus cyanomos), mortality was density-independent and subsequent abundance was a function of recruitment. These contrasting results may reflect differences in prey behavior. P. amboinensis tended to feed near or within the branches of coral inhabited by resident predators, while N. cyanomos tended to feed higher in the water column above the reefs, and thus farther away from resident predators. These results highlight the speed and extent to which patterns of settlement are modified, indicating that caution should be exercised when attributing patterns of recruitment to patterns of settlement. Tremendous between-species variation in how patterns of recruitment, and presumably settlement, were modified by predation indicates that generalizations or between-species extrapolations about the magnitude of these effects may be unwarranted.     

The population dynamics of the voleClethrionomys rufocanus in Hokkaido, Japan

Population dynamics of the gray sided-vole,Clethrionomys rufocanus, in Hokkaido, Japan were described on the basis of 225 time series (being from 12 to 31 years long); 194 of the time series have a length of 23 years or longer. The time series were classified into 11 groups according to geographic proximity and topographic characteristics of the island of Hokkaido. Mean abundance varied among populations from 1.07 to 21.07 individuals per 150 trap-nights. The index of variability for population fluctuation (s-index) ranged from 0.204 to 0.629. Another index for population variability (amplitude on log-10 scale) ranged from 0.811 to 2.743. Mean abundance and variability of populations were higher in the more northern and eastern regions of the island. Most populations, except for the southernmost populations, exhibited significant direct density-dependence in population growth. Detection rate for delayed density-dependence varied among groups from 0% to 22.6%. Both direct and delayed density-dependence tended to be stronger in the more northern and eastern populations. The proportion of cyclic populations was higher in the northern-eastern areas than that in the southern-western areas. There was a clear gradient from the asynchronous populations in southwest, to the highly synchronized populations in the northeast.