Does lemming winter grazing impact vegetation in the Canadian Arctic?

In low-productivity environments such as the tundra, it has been proposed that regular, multi-annual population cycles of lemmings could be driven by winter food depletion in years of peak abundance. If lemming population dynamics is controlled by food resources, we predict that (1) winter grazing should negatively impact the abundance of food plants, (2) this impact should be proportional to lemming density and (3) high lemming winter grazing pressure should result in reduced plant growth during the following summer. We tested these predictions on Bylot Island, Nunavut, Canada, where two species of lemmings are present: the brown (Lemmus trimucronatus) and collared lemming (Dicrostonyx groenlandicus). We installed 16 exclosures in their preferred wintering habitat (snowbeds) and annually sampled plant biomass inside and outside exclosures at snow melt and at peak growth during the summers of 2009–2012, covering a full population cycle. Winter grazing had no impact on total vascular plant or moss biomass at snow melt in all years. Among plant families, only Caryophyllaceae, which was uncommon, showed a decline. In moss taxa, a negative effect was found on Polytrichum in only 1 year out of three. Overall, plant regrowth during the subsequent summer showed annual variation and tended to be reduced in the 2 years of high lemming abundance. However, this could be a consequence of summer grazing. Overall, the impact of lemming winter grazing on plants was weak and short-lived, even in years of high lemming abundance. Therefore, our results are not consistent with the hypothesis that food depletion during winter was the cause of the lemming decline following peak abundance at our study site. Other factors may limit lemming populations and prevent them from reaching densities high enough to exhaust their food resources.     

The effect of snow cover on lemming population cycles in the Canadian High Arctic

Rising temperatures and changes in the precipitation regime will have a strong impact on the quality of the snow cover in the Arctic. A snow cover of good quality protecting lemmings from cold temperatures and predators is thought to be an important factor for maintaining the cyclic dynamic of their populations in the tundra. We examined if the characteristics of annual fluctuations (amplitude and shape of phases) in brown lemming (Lemmus trimucronatus) density could be determined by snow depth, snow density, sub-nivean temperature and persistence of snow. Using an 18-year time series of brown lemming abundance on Bylot Island in the Canadian Arctic, we tested if snow variables could explain the residual variation between the observed lemming density and the one predicted by models where cyclicity had been accounted for. Our analysis provides support for the hypothesis that snow cover can affect the amplitude and possibly also the periodicity of lemming population cycles in the High Arctic. Summer abundance of brown lemmings was higher following winters with a deep snow cover and a low-density snow pack near the ground but was unaffected by the date of establishment or melting and duration of the snow cover. Two snow variables showed a temporal trend; mean winter snow depth tended to increase and date of establishment of the hiemal threshold occurred earlier over time. These temporal trends, which should be favourable to lemmings, may explain why healthy population cycles have apparently been maintained at our study site contrary to other Arctic sites.     

Functional and numerical responses of four lemming predators in high arctic Greenland

The high‐arctic tundra ecosystem has the world's simplest vertebrate predator–prey community, with only four predators preying upon one rodent species, the collared lemming (Dicrostonyx groenlandicus). We document the functional and numerical responses of all the four predators in NE Greenland. Using these data, we assess the impact of predation on the dynamics of the collared lemming with a 4 yr cycle and >100‐fold difference between maximum and minimum densities. All predator species feed mostly (>90%) on lemmings when lemming density is >1 ha^?1, but the shapes of the predators’ responses vary greatly. The snowy owl (Nyctea scandiaca) is present and breeds only when lemming densities at snowmelt are >2 ha^?1, giving rise to a step‐like numerical response. The long‐tailed skua (Stercorarius longicaudus) has a type III functional response and shifts from alternate food (mainly berries and insects) to lemmings with increasing lemming density. The skua surpasses all the other predators in summer by its total response. The type III functional response of the Arctic fox (Alopex lagopus) starts to increase at much lower lemming densities than the responses of the avian predators, but it has only a weak numerical response. Finally, the stoat (Mustela erminea) is the most specialized predator and the only one with a clearly delayed numerical response. According to their specific functional and numerical responses, each predator plays a key role at some point of the lemming cycle, but only the stoat has the potential to drive the lemming cycle. Stoat predation is greatly reduced in the winter preceding the lemming peak, and it reaches a maximum in the winter preceding the lowest lemming summer density. Stoat predation appears to maintain low lemming densities for at least two successive years. Our study provides empirical support for the specialist predator hypothesis about small mammal population cycles.     

Habitat use by lemmings near Barrow, Alaska

Brown lemmings reach much higher densities than collared lemmings near Barrow, Alaska, and captures from 19 summers of snap trapping confirm previous reports that brown lemmings prefer lower, wetter habitats than do collared lemmings. Data also support the hypothesis that brown lemmings concentrate in higher habitats during early summer when melt water floods lower habitats, then shift to lower habitats where preferred food is more available as the waters recede. This pattern appears similar to seasonal shifts in habitat use reported for Norwegian lemmings. Two hypotheses were not supported by our data: (1) Unlike Norwegian lemmings, brown lemmings did not expand their use of suboptimal habitats at higher population densities. Rather, absolute densities changed in concert so that the relative densities among habitats remained unchanged. (2) Preferential use of polygon troughs during winter, as indicated by patterns of winter grazing, was not simply a function of snow depth. Instead it appeared to be related to shoot density of preferred foods. Nearly all patterns of habitat use seemed to be linked to food availability. Other factors, such as protection from predators by vegetative cover in summer and increased insulation from deeper snow in winter, did not appear to influence the distribution of lemmings as strongly.     

Habitat selection,reproduction and predation of wintering lemmings in the Arctic

Snow cover has dramatic effects on the structure and functioning of Arctic ecosystems in winter. In the tundra, the subnivean space is the primary habitat of wintering small mammals and may be critical for their survival and reproduction. We have investigated the effects of snow cover and habitat features on the distributions of collared lemming (Dicrostonyx groenlandicus) and brown lemming (Lemmus trimucronatus) winter nests, as well as on their probabilities of reproduction and predation by stoats (Mustela erminea) and arctic foxes (Vulpes lagopus). We sampled 193 lemming winter nests and measured habitat features at all of these nests and at random sites at two spatial scales. We also monitored overwinter ground temperature at a subsample of nest and random sites. Our results demonstrate that nests were primarily located in areas with high micro-topography heterogeneity, steep slopes, deep snow cover providing thermal protection (reduced daily temperature fluctuations) and a high abundance of mosses. The probability of reproduction increased in collared lemming nests at low elevation and in brown lemming nests with high availability of some graminoid species. The probability of predation by stoats was density dependent and was higher in nests used by collared lemmings. Snow cover did not affect the probability of predation of lemming nests by stoats, but deep snow cover limited predation attempts by arctic foxes. We conclude that snow cover plays a key role in the spatial structure of wintering lemming populations and potentially in their population dynamics in the Arctic.     

Comparing the genetic population structure of two species of arctic lemmings: more local differentiation in Lemmus trimucronatus than in Dicrostonyx groenlandicus

Collared and brown lemmings ( Dicrostonyx groenlandicus and Lemmus trimucronatus ) are two largely sympatric and ecologically comparable species of arctic microtine rodents, differing however in some respects which allow us to hypothesise differences in the genetic structure of their populations. Collared lemmings are particularly well adapted to life at high latitude, they occasionally emerge to the surface of the snow and may disperse over larger distances than brown lemmings – possibly even over snow and ice. This should result in more local differentiation among populations of brown lemmings than among populations of collared lemmings. We compared the genetic population structure between the two lemming species in a fragmented landscape with small islands in the central Canadian Arctic using four microsatellite loci and partial mitochondrial control region sequences. Both types of genetic markers showed higher differentiation ( F _ST values) among local populations for brown lemmings than for collared lemmings. We discuss to what extent the observed genetic differences may be explained by differences in dispersal rates in addition to differences in average effective population size.     

Lemming winter habitat choice: a snow-fencing experiment

The insulative value of early and deep winter snow is thought to enhance winter reproduction and survival by arctic lemmings (Lemmus and Dicrostonyx spp). This leads to the general hypothesis that landscapes with persistently low lemming population densities, or low amplitude population fluctuations, have a low proportion of the land base with deep snow. We experimentally tested a component of this hypothesis, that snow depth influences habitat choice, at three Canadian Arctic sites: Bylot Island, Nunavut; Herschel Island, Yukon; Komakuk Beach, Yukon. We used snow fencing to enhance snow depth on 9-ha tundra habitats, and measured the intensity of winter use of these and control areas by counting rodent winter nests in spring. At all three sites, the density of winter nests increased in treated areas compared to control areas after the treatment, and remained higher on treated areas during the treatment. The treatment was relaxed at one site, and winter nest density returned to pre-treatment levels. The rodents’ proportional use of treated areas compared to adjacent control areas increased and remained higher during the treatment. At two of three sites, lemmings and voles showed significant attraction to the areas of deepest snow accumulation closest to the fences. The strength of the treatment effect appeared to depend on how quickly the ground level temperature regime became stable in autumn, coincident with snow depths near the hiemal threshold. Our results provide strong support for the hypothesis that snow depth is a primary determinant of winter habitat choice by tundra lemmings and voles.     

Long-term microtine dynamics in north Fennoscandian tundra: the vole cycle and the lemming chaos

Densities of microtine rodents in two habitat complexes in the tundra of Finnmarks-vidda, Norwegian Lapland, were studied during 1977-89 by means of snap trapping (Small Quadrat Method) Predator populations were studied by mapping breeding raptors and by snow-tracking small mustelids During 1977-85, snow-trackmg was conducted only during peak and decline years, whereas during 1986-89, snow-tracking was conducted every winter (November-December) and live-trapping (in August) was used as an additional method
Lowland vole populations had regular density fluctuations with peaks in 1978-79. 1982-84 and 1987-88 Highland vole populations fluctuated less regularly and at lower over-all densities Highland lemming populations had two outbreaks, in 1978 and 1988, ending in abrupt winter crashes In the lowland, outbreak levels were reached only in 1978 All microtine declines in relatively productive lowland habitats were accompanied by intense activity of small mustelids. whereas avian predators were common only in 1983 Lowland declines also showed clear between-habitat asynchrony they started in areas with an exceptional abundance of productive habitats and then spread to more barren areas These lowland data are consistent with the hypothesis of a mustelid-microtine limit cycle, although also several other hypotheses remain unrefuted The highland lemming data suggest a simple exploiter-victim interaction between lemmings and the vegetation     

Reproductive behaviour of female Siberian lemmings during the increase and peak phase of the lemming cycle

The reproduction of female Siberian lemmings in the increase and peak phases of the lemming cycle was investigated in connection with a ship-borne expedition along the Siberian arctic tundra. The cycle phase of each studied lemming population was determined using demographic analyses, i.e. current density indices (captured lemmings per 100 traps per 24 h), information on previous density (frequency of old lemming faeces and runways), and information from dendrochronological analyses revealing the most recent winters with a high intensity of willow-stem scarring caused by lemmings. The cycle phase determination was corroborated with data on the age profiles of the populations. The reproductive behaviour of female lemmings differed markedly in relation to cycle phase. In increase-phase populations, all captured females (including young and winter born) were reproducing (had embryos or were lactating), and females started to reproduce early in life, i.e. when <2 months old. By contrast, in peak-phase populations, only 6% of the young females and 63% of the winter-born ones were reproducing, and females did not start to reproduce until they were 5–6 months old. The average number of embryos per reproducing female was significantly higher in increase-phase populations than in peak-phase ones. It is concluded that the rapid population growth in lemmings during the increase phase can largely be explained by the early (young age) reproductive start and, consequently, the shorter generation time, the high proportion of females taking part in reproduction, and the large litters produced. Similarly, a delay in the start of reproduction, a lower proportion of reproducing females, and smaller litter sizes produced by peak-phase lemming populations can contribute substantially to the deceleration in the population increase and possibly lead to a decline. Received: 14 June 1999 / Accepted: 15 November 1999     

Asynchronous population dynamics of Siberian lemmings across the Palaearctic tundra

The synchrony of Siberian lemming (Lemmus sibiricus L.) population dynamics was investigated during a ship-borne expedition along the Palaearctic tundra coast in the summer of 1994. On 12 sites along the coast from the Kola Peninsula to Wrangel Island, relative densities of lemmings were recorded using a standardised snap-trapping programme. The phase position of the lemming cycle in each of the studied populations was determined based on current density estimates, signs of previous density and the age profile of each population (ageing based on eye lens mass). In addition, dendrochronological methods were used to determine when the last peak in the density of microtine populations occurred at each site. The examined lemming populations were in different phases of the lemming cycle. Some populations were in the peak phase, as indicated by high current densities, an age profile in which older individuals were well represented, and signs of high previous density (abundant old lemming faeces). Other populations were in the decline phase, as reflected in a moderate current density, a predominance of older individuals and signs of high previous density. Populations in the low phase had an extremely low current density and showed signs of high previous density, while populations in the increase phase had a moderate current density, a predominance of younger individuals and showed signs of low previous density. The results of phase determinations based on dendrochronological methods support the findings based on lemming demography. Recent Russian studies carried out on some of the sites also agreed with our phase determination results. Thus, on a regional scale (across the whole Palaearctic tundra), the population dynamics of Siberian lemmings can be considered asynchronous. However, sites situated adjacent to each other were often phase synchronous, suggesting a more fine-grained pattern of dynamics with synchrony over distances as long as 1000 km or so, e.g. the Yamal and Taymyr Peninsulas. Received: 19 August 1998 / Accepted: 1 March 1999     

Population dynamics of the collared lemming and the tundra vole at Pearce Point,Northwest Territories,Canada

From 1987 to 1989 we monitored population changes during summer of the collared lemming (Dicrostonyx groenlandicus) and the tundra vole (Microtus oeconomus) at Pearce Point, Northwest Territories, Canada (69° 48 N, 122° 40 W). Populations on four study areas did not cycle but remained at low density (<3/ha) each year and continued at low numbers for the following 3 years (Reid et al. 1995). Lemming numbers often declined throghout the summer in spite of continous reproduction, and population recovery occurred overwinter. Heavy predation losses of radio-collared lemmings occurred each summer and this lemming population may be trapped in a predator-pit. Collared lemmings breed in winter and only because of winter population growth do these populations persist. Tundra vole numbers increased rapidly in most summers but usually declined overwinter. Tundra voles do not seem able to sustain winter reproduction in this extreme environment and this prevents them from reaching high density because of the short summer. Population growth in both these rodents could be prevented by poor food or by predation losses, and landscape patchiness may also help to prevent population growth. For lemmings we do not think that a shortage of shelter or intrinsic limitations could be restricting population increase at Pearce Point. This is the first detailed study of a non-cyclic collared lemming population.     

Can Reindeer Overabundance Cause a Trophic Cascade?

Abstract The region Finnmark, in northernmost Europe, harbors dense populations of semi-domestic reindeer of which some exhibit characteristics of overabundance. Whereas overabundance is evident in terms of density-dependent reductions in reindeer body mass, population growth and abundance of forage plants, claims have been made that this reindeer overabundance also has caused a trophic cascade. These claims are based on the main premise that reindeer overgrazing negatively impacts small-sized, keystone tundra herbivores. We tested this premise by a large-scale study in which the abundance of small rodents, hares and ptarmigans was indexed across reindeer management districts with strong differences in stocking densities. We examined the scale-dependent relations between reindeer, vegetation and these small-sized herbivores by employing a spatially hierarchical sampling design within the management districts. A negative impact of reindeer on ptarmigan, probably as a result of browsing reducing tall Salix, was indicated. However, small rodents (voles and lemmings), which are usually the keystone herbivores in the plant-based tundra food web, were not negatively impacted. On the contrary, there was a strong positive relationship between small rodents and reindeer, both at the scale of landscape areas and local patches, with characteristics of snow-bed vegetation, suggesting facilitation between Norwegian lemmings and reindeer. We conclude that the recent dampening of the vole and lemming population cycle with concurrent declines of rodent predators in northernmost Europe were not caused by large herbivore overgrazing.     

Highly Overlapping Winter Diet in Two Sympatric Lemming Species Revealed by DNA Metabarcoding

Sympatric species are expected to minimize competition by partitioning resources, especially when these are limited. Herbivores inhabiting the High Arctic in winter are a prime example of a situation where food availability is anticipated to be low, and thus reduced diet overlap is expected. We present here the first assessment of diet overlap of high arctic lemmings during winter based on DNA metabarcoding of feces. In contrast to previous analyses based on microhistology, we found that the diets of both collared (Dicrostonyx groenlandicus) and brown lemmings (Lemmus trimucronatus) on Bylot Island were dominated by Salix while mosses, which were significantly consumed only by the brown lemming, were a relatively minor food item. The most abundant plant taxon, Cassiope tetragona, which alone composes more than 50% of the available plant biomass, was not detected in feces and can thus be considered to be non-food. Most plant taxa that were identified as food items were consumed in proportion to their availability and none were clearly selected for. The resulting high diet overlap, together with a lack of habitat segregation, indicates a high potential for resource competition between the two lemming species. However, Salix is abundant in the winter habitats of lemmings on Bylot Island and the non-Salix portion of the diets differed between the two species. Also, lemming grazing impact on vegetation during winter in the study area is negligible. Hence, it seems likely that the high potential for resource competition predicted between these two species did not translate into actual competition. This illustrates that even in environments with low primary productivity food resources do not necessarily generate strong competition among herbivores.     

Temporal variability in arctic fox diet as reflected in stable-carbon isotopes; the importance of sea ice

Consumption of marine foods by terrestrial predators can lead to increased predator densities, potentially impacting their terrestrial resources. For arctic foxes (Alopex lagopus), access to such marine foods in winter depends on sea ice, which is threatened by global climate change. To quantify the importance of marine foods (seal carrion and seal pups) and document temporal variation in arctic fox diet I measured the ratios of the stable isotopes of carbon (¹³C/¹²C) in hair of arctic foxes near Cape Churchill, Manitoba, from 1994 to 1997. These hair samples were compared to the stable carbon isotope ratios of several prey species. Isotopic differences between seasonally dimorphic pelage types indicated a diet with a greater marine content in winter when sea ice provided access to seal carrion. Annual variation in arctic fox diet in both summer and winter was correlated with lemming abundance. Marine food sources became much more important in winters with low lemming populations, accounting for nearly half of the winter protein intake following a lemming decline. Potential alternative summer foods with isotopic signatures differing from lemmings included goose eggs and caribou, but these were unavailable in winter. Reliance on marine food sources in winter during periods of low lemming density demonstrates the importance of the sea ice as a potential habitat for this arctic fox population and suggests that a continued decline in sea ice extent will disrupt an important link between the marine and terrestrial ecosystems.     

Population fluctuations of the wood lemming<Emphasis Type="Italic">Myopus schisticolor</Emphasis> in eastern and western Finland

Olavi Eskelinen, Pertti Sulkava and Risto Sulkava From 1982 to 2003 we studied fluctuations in populations of the wood lemmingMyopus schisticolor (Liljeborg, 1844) in the Heinävesi (eastern Finland) and Keuruu regions (western Finland) by counting field signs and dead animals in standardized field surveys. We compared the population fluctuations of lemmings to those of other voles, owls and small mustelids in these regions. The lemming population in Heinävesi fluctuated regularly in 3-year cycles and in synchrony with the field vole population. Populations of owls also fluctuated in synchrony with wood lemming and field vole populations. In the Keuruu region, oscillations in the wood lemming population were irregular, and neither lemming and vole populations nor lemming and owl populations were correlated. Although direct mechanistic evidence is lacking, specialist predators such as weasels and owls probably cause the cyclicity in the field vole and wood lemming populations in the Heinävesi area. On the other hand, scarcity of high-quality habitats, unfavourable winter weather conditions and generalist predators may prevent the development of cyclicity in the wood lemming population of Keuruu.     

Demography of two lemming species on Bylot Island, Nunavut, Canada

Lemmings play a key role in the tundra food web and their widely reported cyclic oscillations in abundance may have a strong effect on other components of the ecosystem. We documented seasonal and annual variations in population density, reproductive activity, survival, and body mass of two sympatric species, the brown (Lemmus trimucronatus) and collared lemmings (Dicrostonyx groenlandicus), over a 2-year period on Bylot Island, Nunavut, Canada. We live trapped and marked lemmings on two grids throughout the summer and we estimated demographic parameters using three different capture–recapture methods. All three methods are based on robust estimators and they yielded similar population density estimates. The density of brown lemmings declined markedly between the 2 years whereas that of collared lemmings was relatively constant. For brown lemmings, 2004 was a peak year in their cycle and 2005 a decline phase. Density of brown lemmings also decreased during the summer, but not that of collared lemmings. The recruitment of juvenile brown lemmings in the population increased during the summer and was higher in the peak year than in the year after, but no change was detected in collared lemmings. Survival rates of both species tended to be lower during the peak year than in the following year and body mass of brown lemmings was higher in the peak year than in the following year. We conclude that both changes in adult survival and juvenile recruitment occur during the population decline of brown lemmings.     

Landscapes of fear or competition? Predation did not alter habitat choice by Arctic rodents

In systems where predation plays a key role in the dynamics of prey populations, such as in Arctic rodents, it is reasonable to assume that differential patterns of habitat use by prey species represent adaptive responses to spatial variation in predation. However, habitat selection by collared (Dicrostonyx groenlandicus) and brown (Lemmus trimucronatus) lemmings depends on intra- and inter-specific densities, and there has been little agreement on the respective influences of food abundance, predators, and competition for habitat on lemming dynamics. Thus, we investigated whether predation affected selection of sedge-meadow versus upland tundra by collared lemmings in the central Canadian Arctic. We first controlled for the effects of competition on lemming habitat selection. We then searched for an additional signal of predation by comparing habitat selection patterns between 12 control plots and one large grid where lemmings were protected from predators by fencing in 1996 and 1997, but not during 5 subsequent years when we monitored habitat use in the grid as well as in the control plots. Dicrostonyx used upland preferentially over meadows and was more numerous in 1996 and 2011 than in other sample years. Lemmus was also more abundant in 1996 than in subsequent years, but its abundance was too low in the exclosure to assess whether exclusion of predators influenced its habitat selection. Contrary to the effects of competition, predation had a negligible impact on the spatial dynamics of Dicrostonyx, at least during summer. These results suggest that any differences in predation risk between the two habitats have little direct influence on the temporal dynamics of Dicrostonyx even if induced through predator–prey cycles.     

Phase dependence in winter physiological condition of cyclic voles

Lack of food resources has been suggested as a factor which limits the growth of cyclic vole populations. During peak phases of the cycle, vole population growth typically ceases during late autumn or early winter, and is followed by a decrease in density over the winter. To investigate whether this decrease is due to increased mortality induced by a depletion of food resources, we studied overwinter food consumption and physiological condition of field voles ( Microtus agrestis ) in western Finland in both an increase and a decrease phase of a three-year population cycle. The growth rate of vole populations was negatively related both to prevailing vole densities and to densities six months earlier. The condition index of voles, as well as their blood levels of haematocrit, proteins, free fatty acids and immunoglobulin G, were positively related to population growth rate when populations were declining. When populations were increasing, these parameters tended to be negatively related to population growth rate. The overall physiological condition of voles was lower in the winter of the decrease phase as compared to the increase phase. The return rate of voles, a proxy of survival, was also lower in the decrease than in the increase phase of the cycle and positively related to haematocrit levels. Almost 90% of all green vegetation shoots were consumed by voles during the winter of the decrease phase while only two thirds were eaten in the increase phase. Our results suggest that the winter decrease phase of cyclic vole populations is associated with both a deterioration in the physiological condition of voles and a significant depletion of winter food resources. This implies that malnutrition induces poor physiological condition in voles, which in turn may increase mortality either directly through starvation or indirectly through increased susceptibility to predators and pathogens.     

6. NESTING OF NYASA VIOLET-BACKED SUNBIRD ON ELVINGTON FARM,BEATRICE DISTRICT,SOUTHERN RHODESIA

Summary

Summers, R. W., Underhill, L. G, Waltner, M. &; Whitelaw, D. A. 1987. Population, biometrics and movements of the Sanderling Calidris alba in southern Africa. Ostrich 58:24-39.

The Sanderling in southern Africa is restricted to coastal habitats. The population during the austral summer was estimated to be 78000. Highest densities occurred along the west coast where the Benguela Up-welling System gives rise to enriched inter-tidal invertebrate communities, partly through the stranding of large kelps. Immigration from the breeding areas took place during September and November. Numbers were highest during mid summer and emigration took place in April. The summer population at Lange-baan Lagoon was biased towards males (72%). The percentage of first-year birds in the summer population fluctuated on a three-year cycle, coinciding with the lemming cycle in the Taimyr Peninsula in Siberia. It is suggested that the cyclic fluctuation is caused by Arctic Foxes, and perhaps other predators, feeding on lemmings in years of lemming abundance but subsisting on the eggs and chicks of birds when lemmings are scarce. First-year birds are similar in size (similar bill lengths) to older birds, but are lighter in mass during the summer. First-year birds do not accumulate the large fat reserves which older birds deposit between March and early May for northward migration. However, not all first-year birds remain in southern Africa. Trie 13000 km northward migration is completed within seven weeks and probably involves three long flights and two periods of replenishing fat reserves. Birds cross the Sahara to the Mediterranean, and also pass through the Caspian Sea to Siberia or through the North Sea coasts to either Siberia or Greenland. Return routes also involve the North, Caspian and Black Seas but there is little information about routes through Africa.     

The nature of lemming cycles on Wrangel: an island without small mustelids

Lemming cycles are a key process in the functioning of tundra ecosystems. Although it is agreed that trophic interactions are important in causing the cycles, the actual mechanism is disputed. Some researchers attribute a major role to predation by small mustelids such as stoats and least weasels. Here we present a 40-year time series of lemming dynamics from Wrangel Island and show statistically that lemmings do exhibit population cycles in the absence of small mustelids. The observed density fluctuations differed, however, from those observed elsewhere, with long cycles and possibly higher densities of lemmings during the low phase. These differences in the shape of the population cycles may be related to the unique species assemblage of Wrangel Island, where arctic foxes are the only year-round resident lemming predator, and to the high diversity of landscapes, microclimatic conditions, and plants on the island. Both spectral analysis and wavelet analysis show a change in period length from five?years in the 1970s to nearly eight?years in the 1990s and 2000s. This change in dynamics coincides with reports of dampening or fading out of lemming cycles that have been observed in several regions of the Arctic in recent decades. As in the other cases, the changed lemming dynamics on Wrangel Island may be related to ground icing in winter, which could delay peak years.