Population dynamics of a generalist rodent in relation to variability in pulsed food resources in a fragmented landscape

1. Pulsed food resources are often considered equivalent in their potential impact on the reproduction and population dynamics of consumers, but differences in the attributes of food pulses and their distribution in the landscape may cause differences in their effects. 2. We tested whether a perishable pulsed resource (periodical cicadas, Magicicada spp.) had similar effects on the population dynamics of a generalist forest rodent, Peromyscus leucopus, as have been reported for a cacheable pulsed resource (acorn mast). 3. Because the availability of periodical cicadas may vary between edge and interior habitat, we also tested whether habitat type (edge vs. interior) and fragment size affected the abundance of cicadas and P. leucopus. 4. Nearly 90% of the variation in the relative population densities of P. leucopus was explained by the variation in the relative densities of periodical cicadas, and fragments with more cicadas tended to have more reproductive female mice and litters. 5. We found more cicadas and more P. leucopus in edge than interior habitat, but no differences in the relative densities of either in relation to fragment size. 6. Data from a non-emergence year revealed no differences other than the presence of periodical cicadas that could explain the 50% higher relative densities of P. leucopus in the emergence year. 7. At the beginning of the emergence of periodical cicadas, the three fragments with the highest numbers of emergence holes had three times more mice than the fragments with the lowest numbers of emergence holes, suggesting that P. leucopus is able to anticipate the emergence of periodical cicadas and increase reproduction prior to the pulse. 8. Hence, despite differences in perishability, seasonal timing and nutritional quality of pulsed food resources in a fragmented landscape, they appear to have similar positive effects on the population dynamics of the generalist rodent, P. leucopus and, in fact, P. leucopus may be able to anticipate resource pulses.     

Microhabitat selection by small mammals in a southern Appalachian fen in the USA

Little ecological information is available on small mammals inhabiting wetlands in the southern Appalachian mountains of the USA. These wetland systems are becoming rare features in southern landscapes due to human activities. We investigated the small mammal fauna and examined the microhabitat associations of the two most abundant species in a southern Appalachian fen. Four species of small mammals were captured: the meadow jumping mouse (Zapus hudsonius), short-tailed shrew ( Blarina brevicauda), white-footed mouse ( Peromyscus leucopus), and golden mouse ( Ochrotomys nuttalli). Peromyscus and Ochrotomys, which were caught in the largest numbers, preferred sites characterized by moderate herbaceous cover and substantial canopy closure. Peromyscus, however, selected areas with greater canopy closure and higher tree densities, suggesting that they are greater habitat specialists than Ochrotomys in this wetland community.     

Cyclic voles and shrews and non-cyclic mice in a marginal grassland within European temperate forest

Cyclic population dynamics of small mammals are not restricted to the boreal and arctic zones of Eurasia and North America, but long-term data series from lower latitudes are still less common. We demonstrated here the presence of periodic oscillations in small mammal populations in eastern Poland using 22-year (1986–2007) trapping data from marginal meadow and river valley grasslands located in the extensive temperate woodland of Białowieża Primeval Forest. The two most common species inhabiting meadows and river valleys, root vole Microtus oeconomus and common shrew Sorex araneus, exhibited synchronous periodic changes, characterised by a 3-year time lag as indicated by an autocorrelation function. Moreover, the cycles of these two species were synchronous within both habitats. Population dynamics of the striped field mouse Apodemus agrarius was not cyclic. However, this species regularly reached maximum density 1 year before the synchronized peak of root voles and common shrews, which may suggest the existence of interspecific competition. Dynamics of all three species was dominated by direct density-dependent process, whereas delayed density dependent feedback was significant only in the root vole and common shrew. Climatic factors acting in winter and spring (affecting mainly survival and initial reproduction rates) were more important than those acting in summer and autumn and affected significantly only the common shrew. High temperatures in winter and spring had positive effects on autumn-to-autumn changes in abundance of this species, whereas deep snow in combination with high rainfall in spring negatively affected population increase rates in common shrew.     

Breeding timed to maximize reproductive success for a migratory songbird: the importance of phenological asynchrony

Phenological advances and trophic mismatches are frequently reported ecological consequences of climate warming. Trophic mismatches occur when phenological responses to environmental conditions differ among trophic levels such that the timing of resource demand by consumers becomes decoupled from supply. We used 25 years of demographic measurements of a migratory songbird (the black‐throated blue warbler Setophaga caerulescens) to compare its breeding phenology to the phenology of both its caterpillar prey and the foliage on which caterpillars feed. Caterpillar biomass in this forest did not show a predictable seasonal pulse. Nest initiation by warblers in this northern hardwood forest was therefore not timed to coincide with a peak in food availability for nestlings. Nonetheless, timing of first clutches was strongly associated with spring leaf expansion (slope ± SE = 0.56 ± 0.08 days per day of change in leaf phenology, R² = 0.66). Warblers adjusted the timing of breeding to early springs mainly by shortening the interval between arrival and clutch initiation, but this likely has limits because recent early springs are approaching the relatively inflexible dates when birds arrive on the breeding grounds. Although the timing of first nests did not match 1:1 with leaf‐out phenology, the adjustments in breeding time maximized mean annual reproductive success. Nest predation had the greatest effect on annual reproductive success, but the ability of nesting warblers to appropriately track leaf phenology accounted for effects on annual reproductive success comparable to the influence of variation in caterpillar abundance and conspecific density. Nesting phenology in black‐throated blue warblers was generally well matched to the timing of leaf‐out, even though the match was not 1:1. Without measurements of reproductive success, these unequal phenological shifts might otherwise have been interpreted as having negative ecological consequences.     

Bioenergetics of cross-ice movements by Microtus pennsylvanicus, Peromyscus leucopus and Blarina brevicauda

Laboratory and field experiments were conducted to investigate bioenergetics of winter dispersal and to compare winter (cross-ice) dispersal abilities of three small mammals: Microtus pennsylvanicus, Peromyscus leucopus and Blarina brevicauda . Total metabolic rates increased with running activity and decreased as ambient temperatures increased for all species. Thermal conductance was significantly higher for running than for resting Microtus and Peromyscus , but decreased significantly with activity for Blarina .
Winter dispersal abilities, calculated from treadmill experiments, increased with ambient temperature and with body size of the species. The superior dispersal ability of Microtus in comparison with Peromyscus results from the former's ability to utilize more energy reserves during running. The comparatively low winter dispersal ability of Blarina , which was less than a third of the two rodent species, resulted from its high weight specific cost of transport at winter temperatures and its relatively low energy stores and/or energy utilization during running.     

Warming climate advances breeding and improves synchrony of food demand and food availability in a boreal passerine

Global climate change affects ecosystems via several trophic levels. We investigated changes in the timing of breeding in the willow tit (Poecile montanus) and timing of its caterpillar food resource in relation to warming springs in a boreal forest. We used generalized linear mixed effect models to study the importance of synchrony between the timing of breeding in willow tits and the caterpillar food availability on the breeding success, measured as nestling survival rate and mean nestling weight. Both the timing of breeding and the timing of the caterpillar peak advanced during the last decades, and were well explained by spring temperatures. Unlike in most passerine populations studied, synchrony has improved with rising spring temperatures. However, it had only a modest although statistically significant positive influence on breeding success. Spring temperatures do not seem to be used as cues for the timing of caterpillar food availability by willow tits. We conclude that responses to climatic warming seem to be population, species and habitat specific, necessitating research in a wide range of taxa in different climatic zones.     

川西农区小型兽类分布研究

经3个监测点三年定位布夹监测数据分析发现,川西农区田间小兽的密度分布有明显的院落依托性,即离院落越近的农田中密度越高。分布模式呈Logistic曲线。家栖鼠(如褐家鼠、小家鼠)比野栖鼠(如大足鼠和黑线姬鼠)更集中于院落周围。不同季节的密度分布有显著差异,春、夏季多集中在离院落50—70米范围。秋、冬季别离院落越近,其密度越高。     

Female striped mice (Rhabdomys pumilio) change their home ranges in response to seasonal variation in food availability

Animals may respond to seasonally changing environments withphysiological and behavioral strategies. Whereas migration isa behavioral strategy used by many taxa, it may not be an optionfor small mammals. However, small mammals can seasonally varythe area of habitat in which they are active. The striped mouseRhabdomys pumilio in the semiarid Succulent Karoo of South Africalives in a seasonal environment, characterized by hot, dry summerswith low food abundance and cold, wet winters, followed by highfood abundance in spring. We radio tracked a total of 28 femalesduring the 2004 dry season, the following breeding season inspring, and the following dry season in 2005 and tested theprediction that females shift their home ranges in relationto food availability. Females shifted their home ranges froman area characterized by evergreen succulent shrubs in the vicinityof a dry riverbed in the dry season to sandy areas that werecharacterized by new plant growth of annuals in spring. Homeranges during the breeding season in spring had a higher percentageof annuals than dry season home ranges measured in spring. Femalehome range size increased during the breeding season. We suggestthat female striped mice shift their home ranges seasonallyto gain access to protein-rich young plant material, which isimportant for breeding.     

Snow evens fragmentation effects and food determines overwintering success in ground-dwelling voles

Climate instability strongly affects overwintering conditions in organisms living in a strongly seasonal environment and consequently their survival and population dynamics. Food, predation and density effects are also strong during winter, but the effect of fragmentation of ground vegetation on ground-dwelling small mammals is unknown. Here, we report the results of a winter experiment on the effects of habitat fragmentation and food on experimental overwintering populations of bank voles Myodes glareolus. The study was conducted in large outdoor enclosures containing one large, two medium-sized or four small habitat patches or the total enclosure area covered with protective tall-grass habitat. During the stable snow cover of midwinter, only food affected the overwintering success, body condition, trappability and earlier onset of breeding in bank voles. However, after the snow thaw in spring, habitat fragmentation gained importance again, and breeding activities increased the movements of voles in the most fragmented habitat. The use of an open, risky matrix area increased along the habitat fragmentation. Our experiment showed that long-lasting stable snow cover protects overwintering individuals in otherwise exposed and risky ground habitats. We conclude that a stable winter climate and snow cover should even out habitat fragmentation effects on small mammals. However, along prolonged snow-free early winter and in an earlier spring thaw, this means loss of protection by snow cover both in terms of thermoregulation and predation. Thus, habitat cover is important for the survival of small ground-dwelling boreal mammals also during the non-breeding season.     

Marked reduction in demographic rates and reduced fitness advantage for early breeding is not linked to reduced thermal matching of breeding time

Warmer springs may cause animals to become mistimed if advances of spring timing, including available resources and of timing of breeding occur at different speed. We used thermal sums (cumulative sum of degree days) during spring to describe the thermal progression (timing) of spring and investigate its relationship to breeding phenology and demography of a long‐distant migrant bird, the northern wheatear (Oenanthe oenanthe L.). We first compare 20‐year trends in spring timing, breeding time, selection for breeding time, and annual demographic rates. We then explicitly test whether annual variation in selection for breeding time and demographic rates associates with the degree of phenological matching between breeding time and thermal progression of spring. Both thermal progression of spring and breeding time of wheatears advanced in time during the study period. But despite breeding on average 7 days earlier with respect to date, wheatears bred about 4 days later with respect to thermal spring progression. Over the same time period, selection for breeding time changed from distinct within‐season advantage of breeding early to no or very weak advantage. Furthermore, demographic rates (nest success, fledgling production, recruitment, adult survival) and nestling weight declined markedly by 16%–79%. Those temporal trends suggest that a reduced degree of phenological matching may affect within‐season fitness advantage of early breeding and population demographic rates. In contrast, when we investigate links based on annual variation, we find no significant relationship between either demographic rates or fitness advantage of early breeding with annual variation in the degree of phenological matching. Our results show that corresponding temporal trends in phenological matching, selection for breeding time and demographic rates are inconclusive evidence for demographic effects of changed phenological matching. Instead, we suggest that the trends in selection for breeding time and demographic rates are due to a general deterioration of the breeding environment.     

Effects of predator removal on vertebrate prey populations: birds of prey and small mammals

We studied the effects of removal of breeding nomadic avian predators (the kestrel, Falco tinnunculus and Tengmalm's owl, Aegolius funereus) on small mammals (voles of the genera Microtus and Clethrionomys and the common shrew, Sorex araneus) during 1989–1992 in western Finland to find out if these predators have a regulating or limiting impact on their prey populations. We removed potential breeding sites of raptors from five manipulation areas (c. 3 km² each), whereas control areas had nest-boxes in addition to natural cavities and stick-nests. Densities of small mammals were monitored by snap-trapping in April, June, and August, and densities of mammalian predators (the least weasel, Mustela nivalis nivalis, the stoat, M. erminea and the red fox, Vulpes vulpes) by snow tracking in early spring and late autumn. The yearly mean number of raptor breeding territories was 0.2–1.0 in reduction areas and 3.0–8.2 in control areas. Breeding raptors alone did not regulate prey populations in the long term, but probably caused short-term changes in the population dynamics of both the main prey, the sibling vole (Microtus rossiaemeridionalis) and an alternative prey (the common shrew). The densities of an alternative prey, the bank vole (Clethrionomys glareolus) decreased in raptor reduction areas, most likely due to increased least weasel predation pressure in the absence of breeding avian predators.     

The influence of carotid arterial sounds on hearing sensitivity in mammals

Pulsations of the internal carotid and stapedial arteries produce unwanted sounds (noise) in the middle ear cavity which influence hearing in some mammals. 'Noise' pressure levels calculated from pulse rates, volume pulsations of the arteries, and Fourier analysis of arterial waveforms, correlate well with low frequency thresholds of hearing in the Long-eared hedgehog, Tree shrew and Kangaroo rat. In mammals adapted to hear low frequency sounds, such as the Kangaroo rat and fossorial insectivores, the arteries are enclosed in noise attenuating bony tubes. However, most small mammals possess extended high frequency hearing with little sensitivity at the low frequencies of the arterial sounds. In lower tetrapods such as anurans and most lizards, the broad connection between the middle ear cavity and the pharynx creates a leakage pathway which greatly reduces the noise from the stapedial artery. It is probably for these reasons that the large intratympanic arteries did not disturb hearing in early mammals or submammalian forms.     

Olfactory mucosa ultrastructure in the short-tailed shrews,Blarina brevicauda and Blarina carolinensis

The olfactory mucosae of the northern short-tailed shrew, Blarina brevicauda, and the southern short-tailed shrew, Blarina carolinensis, were examined by light and electron microscopy. A well-developed olfactory epithelium was observed that included all of the cells necessary to provide for a sensitive olfactory system, suggesting that olfaction plays a major role in the behavior of these animals. There were no significant differences between the olfactory mucosae of these two species. The general features of the olfactory epithelium in these shrews were similar to those reported for several other macrosmatic mammals. A new type of supporting cell, called the light supporting cell, was observed in these shrews. The light supporting cell cytoplasm exhibited very little staining by light microscopy and had low electron density by transmission electron microscopy compared to that of the more common dark supporting cell. The light supporting cell had a convex apical surface with microvilli and lacked the large amounts of smooth endoplasmic reticulum (SER) typical of the apical cytoplasm of the dark supporting cell. In the lamina propria of the mucosa, the Bowman's glands consisted of two cell types, one with electron-lucent, alcian blue-positive granules, and the other with electron-dense PAS-positive granules. The cell with electron-lucent granules contained large amounts of SER and small clumps of rough ER. The cells with electron-dense granules had large amounts of RER and little SER.     

Seasonal regulation in fluctuating small mammal populations: feedback structure and climate

We studied fluctuating populations of six small mammal species in the Appalachian Plateau of Pennsylvania, USA for 20 yr. We analyzed the feedback structure of these species using statistical time series models for spring and autumn abundances. All species showed a seasonal density-dependent structure, and in five of them first-order feedbacks were dominant in winter and summer. Instead, southern red-backed voles ( Clethrionomys gapperi ) showed a different feedback structure during winter and summer. In three species ( C. gapperi , Napaeozapus insignis and Peromyscus maniculatus ), environmental factors were more important during summer, while the opposite pattern was found in Blarina brevicauda and Tamias striatus . Snowfall influenced positively the winter population growth rates of southern red-backed voles, white-footed mice, woodland jumping mice and eastern chipmunks. We found seasonal differences in the effects of the small mammals assemblage on population growth rates of the two Peromyscus species. The common feedback structure between seasons observed in most of the species, particularly among voles and mice, points to a different feedback structure from northern cyclic small mammals. We conclude that a seasonal feedback structure dominated by intra- and inter-specific competitive interactions may be at the basis of the population dynamics of these species.     

The interacting effects of food,spring temperature,and global climate cycles on population dynamics of a migratory songbird

Although long‐distance migratory songbirds are widely believed to be at risk from warming temperature trends, species capable of attempting more than one brood in a breeding season could benefit from extended breeding seasons in warmer springs. To evaluate local and global factors affecting population dynamics of the black‐throated blue warbler (Setophaga caerulescens), a double‐brooded long‐distance migrant, we used Pradel models to analyze 25 years of mark–recapture data collected in New Hampshire, USA. We assessed the effects of spring temperature (local weather) and the El Niño Southern Oscillation index (a global climate cycle), as well as predator abundance, insect biomass, and local conspecific density on population growth in the subsequent year. Local and global climatic conditions affected warbler populations in different ways. We found that warbler population growth was lower following El Niño years (which have been linked to poor survival in the wintering grounds and low fledging weights in the breeding grounds) than La Niña years. At a local scale, populations increased following years with warm springs and abundant late‐season food, but were unaffected by spring temperature following years when food was scarce. These results indicate that the warming temperature trends might have a positive effect on recruitment and population growth of black‐throated blue warblers if food abundance is sustained in breeding areas. In contrast, potential intensification of future El Niño events could negatively impact vital rates and populations of this species.     

Muscle capillary supply in hind limb and diaphragm of the common shrew (Sorex araneus)

Shrew species of the subfamily Soricinae have unusually high metabolic rates when compared to Crocidurinae shrews and other similar-sized mammals. The aim of this study was to clarify whether the high basal metabolic rate of Soricinae shrews is reflected in a high capillary density in their muscles. To this end, the capillary supply of four limb muscles and diaphragm of the common shrew (Sorex araneus) was quantified from cross-sectioned muscles. The capillary densities of the limb muscles were 2575 ± 329, 3111 ± 299, 2812 ± 197 and 2752 ± 173 capillaries mm⁻² fibre area in gastrocnemius lateralis, g. medialis, plantaris and soleus, respectively. Capillary density of the shrew diaphragm (6691 ± 1057) was double that of the limb muscles. This value is among the highest ever measured in mammals. In general, the capillary supply in the hind limb of the common shrew is about 3–4 times higher than commonly found in the leg muscles of the laboratory rat or other bigger mammals, but similar to those in Crocidurinae shrews and some small rodents. Thus the high resting metabolism of the common shrew is not associated with an extraordinarily high capillary density. The apparent disparity between basal metabolic rate and muscle capillary supply in S. araneus is probably due to the small aerobic scope of shrews in the subfamily Soricinae. Accepted: 22 January 1998     

Differences in food abundance cause inter-annual variation in the breeding phenology of High Arctic waders

Previous work has shown that High Arctic waders in Greenland are “income breeders”, i.e. the resources used for egg formation are based almost entirely on biomass obtained on the breeding grounds. Thus, their breeding phenology is expected to be highly sensitive to inter-annual variation in food abundance during the pre-laying period. Early spring snow-cover may also influence timing of egg-laying either directly or mediated through food resources. Here, we report on the inter-annual variation in clutch initiation of three wader species breeding in High Arctic Greenland, Sanderling (Calidris alba), Dunlin (Calidris alpina) and Ruddy Turnstone (Arenaria interpres), in relation to spring snow-cover and spring arthropod abundance over ten breeding seasons at Zackenberg Research Station 1995–2005. Food abundance had the strongest effect on timing of clutch initiation, while the proportion of snow-free land had a weaker but still significant effect, i.e. more food and more snow-free land both result in earlier egg-laying. We hypothesize that food is most important when there is sufficient snow-free land to nest on, while snow-cover is of increasing importance in years with late snowmelt.     

Seasonal changes in density, demography and body composition of small mammals in a southern temperate forest

Eleven small mammal species (nine rodents and two shrews) were recorded in 16 months of trapping in the Karkloof Forest, Natal. Total densities varied seasonally, ranging from 60.7 animals/ha in the dry autumn and winter to 29.1 animals/ha at the onset of the rains in spring and early summer. Peaks in breeding coincided with high body fat and water content in the five common species during summer: in winter, when body fat and water levels were low, there was no breeding. This may be related to poor food quality at this time. Annual changes in density, breeding season and juvenile recruitment of Rhabdomys pumilio, Mastomys natalensis and Myosorex varius are the same as recorded in conspecific populations from other habitat types and do not seem modified by the habitat stability of the forest environment. Of the five common species, only Grammomys dolichurus , one of the few forest-associated small mammals in southern Africa, and Graphiurus murinus demonstrated stable, K-selectéd life-history styles.     

Demographic response of tundra small mammals to a snow fencing experiment

Snow cover is a key environmental component for tundra wildlife that will be affected by climate change. Change to the snow cover may affect the population dynamics of high‐latitude small mammals, which are active throughout the winter and reproduce under the snow. We experimentally tested the hypotheses that a deeper snow cover would enhance the densities and winter reproductive rates of small mammals, but that predation by mustelids could be higher in areas of increased small mammal density. We enhanced snow cover by setting out snow fences at three sites in the Canadian Arctic (Bylot Island, Nunavut, and Herschel Island and Komakuk Beach, Yukon) over periods ranging from one to four years. Densities of winter nests were higher where snow depth was increased but spring lemming densities did not increase on the experimental areas. Lemmings probably moved from areas of deep snow, their preferred winter habitat, to summer habitat during snow melt once the advantages associated with deep snow were gone. Our treatment had no effect on signs of reproduction in winter nests, proportion of lactating females in spring, or the proportion of juveniles caught in spring, which suggests that deep snow did not enhance reproduction. Results on predation were inconsistent across sites as predation by weasels was higher on the experimental area at one site but lower at two others and was not higher in areas of winter nest aggregations. Although this experiment provided us with several new insights about the impact of snow cover on the population dynamics of tundra small mammals, it also illustrates the challenges and difficulties associated with large‐scale experiments aimed at manipulating a critical climatic factor.