Life history patterns in female moose (Alces alces): the relationship between age,body size,fecundity and environmental conditions

I examined the relationship between age, body size and fecundity in 833 female moose (Alces alces) from 14 populations in Sweden sampled during 1989–1992. Data on population density, food availability and climatic conditions were also collected for each population. Age and body mass were both significantly positively related to fecundity, measured as ovulation rate, among female moose. The relationship between the probability of ovulation and body mass was dependent on age with (1) a higher body mass needed in younger females for attaining a given fecundity, and (2) body mass having a stronger effect on fecundity in yearling (1.5 year) than in older (2.5 year) females. Thus, a 40 kg increase in yearling body mass resulted in a 42% increase in the probability of ovulation as compared to a 6% increase in older females. The lower reproductive effort per unit body mass, and the relatively stronger association between fecundity and body mass in young female moose compared to older ones, is likely to primarily represent a mechanism that trades off early maturation against further growth, indicating a higher cost of reproduction in young animals. In addition to age and body mass, population identity explained a significant amount of the individual variation in fecundity, showing that the relationship between body mass and fecundity was variable among populations. This variation was in turn related to the environment, in terms of climatic conditions forcing female moose living in relatively harsh/more seasonal climatic conditions to attain a 22% higher body mass to achive the same probability of multiple ovulation (twinning) as females living in climatically milder/less seasonal environments. The results suggests that the lower fecundity per unit body mass in female moose living in climatically harsh/more seasonal environments may be an adaptive response to lower rates of juvenile survival, compared to females experiencing relatively milder/less seasonal climatic conditions.     

Lack of compensatory body growth in a high performance moose Alces alces population

Considerable work has been done on disentangling important factors determining early development in body size, yet our knowledge of the extent to which animals living under varying conditions can achieve catch-up growth for a bad start in life is limited. Here, we investigated how body mass at the age of 8 months influenced adult body mass in a moose Alces alces population living under excellent environmental conditions on the island of Vega in northern Norway. We also investigated if mother age and birth date effects on calf body mass persisted until adulthood. We show that neither males nor females were able to show compensatory growth before they reached adulthood, and that part of the variation in adult body mass may have been due to variation in mother age and date of birth. The pattern observed in males can be related to their growth strategy in relation to reproduction, while such results were not expected in females where size-dependent start of reproduction is likely to interact with body growth. We suggest that the good environmental conditions experienced on Vega led to females having small somatic costs of an early start of reproduction or that larger females were able to acquire more resources for growth than their smaller conspecifics. In both cases, females that postpone their first reproduction may not be able to achieve catch-up growth for their lower early body mass compared to females that start reproduction at an early age. Our results concur with previous studies indicating that compensatory growth is weak in moose, increasing the likelihood that variation in life history characters are also transferred between generations.     

Aspects of the life cycle and pathogenesis of Elaphostrongylus alces in moose (Alces alces).

Aspects of the migratory life cycle and pathogenesis of Elaphostrongylus alces were studied in 7 randomly selected moose calves and 7 yearlings killed during August to November. One calf and 1 yearling were uninfected. The 6 infected calves had recent infections, whereas the 6 yearlings showed older infections from the summer of the previous year. The 2 calves killed in September had a total of 26 adult E. alces in the epidural space of the caudal vertebral canal and none in the skeletal muscles, whereas the remaining calves killed 1 to 2 mo later had 25 nematodes in the caudal and cranial vertebral canal and 7 in the skeletal muscles. The yearlings had a total of 101 adult E. alces in the skeletal muscles and 2 in the vertebral canal. There were no findings indicating involvement of the central nervous system in the life cycle of E. alces. Our findings suggest that E. alces migrates directly from the gut to the epidural space of the caudal vertebral canal where development to the adult stage takes place. During development, the nematode produces inflammation of the epidural tissue and spinal nerves. Development in the caudal vertebral canal is followed by some anterior dispersion of nematodes along the canal, and migration into skeletal muscles. Here the nematodes seem to live in reproductive pairs and groups. The predilection site for E. alces in moose is the muscles of the thigh.     

Aquatic feeding by moose Alces alces in a Canadian lake

The aquatic feeding behaviour of moose, and the abundance, species, and chemical composition of aquatic plants, were studied in a small Canadian lake which attracted many animals. Feeding was much more common in June than later in the summer, and somewhat more common during the morning and evening than in (he afternoon. Individual adult moose appeared to use the lake intensively during 1- to 4-day visits. Compared to deciduous browse, the aquatic plants had high levels of sodium and iron, less fat, and similar levels of crude protein, crude fibre, sulphate and other minerals. In the preferred feeding areas, compared to other parts of the lake, plants were more abundant, had a different species composition, and were richer in iron and calcium. Recent flooding, a flow of water through the feeding areas, and a bicarbonate-rich tributary may all have contributed to the lake's attraction for moose.     

Present genetic structure revealed by microsatellites reflects recent history of the Finnish moose (Alces alces)

Genetic structures of Holarctic species are largely formed by Pleistocene colonisation history, dispersal capacity and interactions between biotic and abiotic factors, even though the human impact can also be significant. The Holarctic moose (Alces alces) arrived in Fennoscandia around 9,000–8,000 years ago, and it has been exploited by humans ever since. During the last 400 years, the Finnish moose population has suffered from several population declines, and even local and regional extirpations have occurred. The purpose of the present study is to describe the genetic variation and population structure of the Finnish moose in order to clarify how historical events and human exploitation have influenced the present-day genetic patterns. Altogether 130 moose individuals from seven sampling sites in Finland were analysed at ten microsatellite loci. A variety of population genetic and coalescent-based methods was applied. The Finnish moose population was found to be divided into southern and northern subpopulations with additional lower hierarchical genetic structure. The estimated time of divergence between these two subpopulations was about 96–238 years ago. In addition, an isolation-by-distance pattern was discovered.     

Genetic diversity of moose (Alces alces L.) in Eurasia

Polymorphism of nucleotide sequence of D-loop fragment of the mitochondrial DNA was studied in 20 moose from several local populations on the territory of Eurasia. Three main haplotype variants of D-loop were detected by molecular phylogenetic method, which formed three clusters named European, Asian and American. Intraspecies variation in the length of HVSI of D-loop of the mitochondrial DNA of moose was revealed. In the Far Eastern and Yakutian moose, haplotypes with a 75-bp deletion were found, which were most similar with haplotypes (also with the deletion), earlier observed in North American moose [1]. The highest diversity of the haplotypes of mitochondrial DNA is characteristic of Yakutia and the Far East (where three haplotype variants were found), which demonstrates the probable role of the region as the center of the species or as the region of ancient population mixture. The geographic region might be considered as a probable source of ancient moose migrations from Asia to America, basing on the data of distribution of mitochondrial haplotypes of D-loop and alleles of MhcAlal-DRB1. Divergence of nucleotide sequences of haplotypes with the 75-bp deletion (forming the American cluster on the phylogenetic tree) was the lowest (0.4%), which evidences respectively recent origin of the group of haplotypes. In Europe, only haplotypes of mitochondrial DNA referred to European variant were observed. Basing on analysis of variation of nucleotide sequences of D-loop, exon 4 of kappa-Casein and exon 2 of MhcALal-DRB1, we demonstrated that Eurasian moose studied belong to the unique species, which has probably passed through a bottle neck. The time of the origin of modern diversity of D-loop haplotypes of the species was estimated as 0.075-0.15 Myr ago.     

Dermacentor albipictus on moose (Alces alces) in Ontario.

Fifty-five moose (Alces alces) collected from 1963 to 1965 in the Chapleau Crown Game Preserve of northern Ontario were examined for ectoparasites. Dermacentor albipictus was the only parasite recovered. D. albipictus was absent from 15 moose examined during June, July and August; first observed on 1 or 2 moose examined in September; and present on all 38 moose collected from October to May. Ticks were not evenly distributed on the body. Infestations varied from 32 to 13,490 ticks/moose for 14 individuals. Levels of infestation observed did not have an obvious adverse effect on the health of the moose.     

Elaeophora schneideri in moose (Alces alces) from Colorado.

Two adult moose (Alces alces) from Colorado (USA) were naturally infected with Elaeophora schneideri. Both animals had patent infections indicating that moose may serve as definitive hosts. Gross and histological lesions were characterized by fibromuscular intimal proliferation within the carotid arteries and rete mirabile cerebri. This is the first report of Elaeophora schneideri in moose from Colorado.     

Geographical and latitudinal variation in growth patterns and adult body size of Swedish moose (Alces alces)

We examined the geographical pattern in growth and adult body size among 14 populations of Swedish moose (Alces alces) using data from 4,294 moose (1.5 years old) killed during the hunting season in 1989–1992. In both sexes, adult body mass was significantly positively correlated with latitude. Moose in northern populations had a 15–20% larger adult body mass than moose in the south. Juvenile body mass was correlated with neither latitude nor adult body mass. Thus, variation in time (years) and rate of body growth after the juvenile stage were responsible for most of the variation in adult body mass among populations. Moose in northern populations grew for approximately 2 more years of life than southern moose. In contrast to adult body mass, skeletal size (measured as jawbone length) was not correlated with latitude, suggesting that variation in adult body mass was primarily due to differences in fat reserves. Discrimination between population characteristics, such as moose density, climate, and the amount of browse available to moose, showed climatic harshness to be the most important variable explaining geographical variation in body mass among populations. The results support the notion that in mammals body size increases with latitude in accordance with Bergmann's rule. We conclude that (1) variation in patterns of growth after the juvenile stage is the main cause of the latitudinal trend in adult body size in moose, and (2) climatic conditions are a more important factor than population density and availability of food in explaining geographical variation in growth patterns and adult body mass between populations of Swedish moose.     

The effects of productivity and seasonality on life history: comparing age at maturity among moose (Alces alces) populations

The productivity hypothesis in respect of an animal species’ geographical range predicts that whereas higher productivity at the equatorial periphery of a species’ range favours superior competitors, lower productivity at the centre of a species’ range favours high reproduction and reduced competitive traits. I test whether life‐history patterns follow this hypothesis, using demographic data from 15 Canadian moose (Alces alces) populations. Two models are contrasted; the first assumes that intraspecific variation in age at maturity is explained proximately by density and juvenile mortality. Age at maturity was found to increase with decreasing juvenile mortality (P = 0.01) and increasing density (P = 0.006). To test the productivity hypothesis, the second model additionally included primary productivity and seasonality as geographical explanatory variables that would ultimately influence age at maturity via juvenile mortality and density. Path analysis indicated that including productivity and seasonality improved the model predictions of variation in age at maturity (R_a² 0.56 and 0.85). In bivariate comparisons, seasonality was negatively associated (P = 0.01) with age at maturity. In the best model, however, primary productivity was the environmental variable that explained 25% of the variance in age at maturity, and forest cover replaced seasonality as an explanatory variable. The positive association between primary productivity and age at maturity is consistent with the productivity hypothesis. Relative to populations that lived at the centre of the species’ range (51°N), moose populations living in relatively high productivity and low seasonality environments (equatorial periphery of species’ range; 48°N) experienced less juvenile mortality, more variable year‐to‐year density, higher relative density and slower life history (slower growth rate, later age at maturity, lower fecundity).     

Patterns of hunting mortality in Norwegian moose (Alces alces) populations

We used a simple life table approach to examine the age-specific patterns of harvest mortality in eight Norwegian moose populations during the last 15 years and tried to determine if the observed patterns were caused by hunter selectivity. The general opinion among local managers is that hunters prefer to shoot female moose not in company with calves to keep a high number of reproductive females in the population (and because of the emotional stress involved in leaving the calf/calves without a mother), and relatively large males because of the higher return with respect to meat and trophy. In support of the former view, we found the harvest mortality of adult females to be higher among pre-prime (1–3 years old) than prime-aged age classes (4–7 years old). This is probably because prime-aged females are more fecund and, therefore, more likely to be in company with one or two calves during the hunting season. As the season progressed, however, the selection pressure on barren females decreased, probably due to more productive females becoming ‘legal’ prey as their calf/calves were harvested. In males, we did not find any evidence of strong age-specific hunter selectivity, despite strong age-dependent variation in body mass and antler size. We suggest that this was due to the current strongly female-biased sex ratio in most Norwegian moose populations, which leaves the hunters with few opportunities to be selective within a relatively short and intensive hunting season. The management implications of these findings and to what extent the results are likely to affect the future evolution of life histories in Norwegian moose populations are discussed.Electronic Supplementary Material Supplementary material is available for this article at and accessible for authorised users.     

Redefining physiological responses of moose (Alces alces) to warm environmental conditions

We tested the concept that moose (Alces alces) begin to show signs of thermal stress at ambient air temperatures as low as 14 °C. We determined the response of Alaskan female moose to environmental conditions from May through September by measuring core body temperature, heart rate, respiration rate, rate of heat loss from exhaled air, skin temperature, and fecal and salivary glucocorticoids. Seasonal and daily patterns in moose body temperature did not passively follow the same patterns as environmental variables. We used large changes in body temperature (≥1.25 °C in 24hr) to indicate days of physiological tolerance to thermal stressors. Thermal tolerance correlated with high ambient air temperatures from the prior day and with seasonal peaks in solar radiation (June), ambient air temperature and vapor pressure (July). At midday (12:00hr), moose exhibited daily minima of body temperature, heart rate and skin temperature (difference between the ear artery and pinna) that coincided with daily maxima in respiration rate and the rate of heat lost through respiration. Salivary cortisol measured in moose during the morning was positively related to the change in air temperature during the hour prior to sample collection, while fecal glucocorticoid levels increased with increasing solar radiation during the prior day. Our results suggest that free-ranging moose do not have a static threshold of ambient air temperature at which they become heat stressed during the warm season. In early summer, body temperature of moose is influenced by the interaction of ambient temperature during the prior day with the seasonal peak of solar radiation. In late summer, moose body temperature is influenced by the interaction between ambient temperature and vapor pressure. Thermal tolerance of moose depends on the intensity and duration of daily weather parameters and the ability of the animal to use physiological and behavioral responses to dissipate heat loads.     

Geographical variation of antler morphology of moose (Alces alces) in Russia

We attempt to identify geographical subdivisions within the range of moose according to antler size for a more correct trophy evaluation in the СIC system (The International Council for Game and Wildlife Conservation). Traditionally, this division coincides with subspecific division of trophy animal species. That is why we partially touch upon the problem of the geography of moose subspecies in Eurasia. Catalogs of hunters’ trophies that were available to us served as the research material (1,047 trophies in total). We were choosing the same measurements for moose antlers evaluated by different methods. They are overall spread, circumference of beam, width of palm, and number of tines. We applied multivariate statistic methods to the findings, enlarging the geographical division groups step by step. As a result, we suggest four geographical groups: European, Siberian, the north of the Far East, and the south of the Far East. In principle, this division corresponds to the subspecies: European (Alces alces alces L., 1758), east Siberian (A. a. pfizenmayeri Zukowsky, 1910), Chukotka or Kolyma (A. a. buturlini Chernyavski et. Zhelesnov, 1982), and Ussuri (А. а. cameloides Milne-Edwards, 1867). We consider it incorrect to draw the line between the European and Siberian groups along the river Yenisei. It is more correct to move it to the Urals.     

The role of the urals in the genetic diversity of the European moose subspecies (Alces alces alces)

The genetic diversity of the Uralian moose population and the role of the Ural region in the phylogeographic structure of the European moose were evaluated based on sequence polymorphisms of the mtDNA control region. The nucleotide diversity of the Ural moose was low, whereas haplotype diversity was rather high. It was found that the haplotype pool of the Ural moose reflects both the unique features of their mitochondrial lineages and their connection with Alces alces alces populations of Europe and West Siberia. The structure of median networks and the territorial haplotype distribution support the hypothesis that the mitochondrial lineages typical for this part of the European moose area originate from a late Pleistocene refugium that was located in the Urals.     

Mitochondrial phylogeography of moose (Alces alces): late pleistocene divergence and population expansion

We examined phylogeographic relationships of moose (Alces alces) worldwide to test the proposed existence of two geographic races and to infer the timing and extent of demographic processes underpinning the expansion of this species across the Northern Hemisphere in the late Pleistocene. Sequence variation within the left hypervariable domain of the control region occurred at low or moderate levels worldwide and was structured geographically. Partitioning of genetic variance among regions indicated that isolation by distance was the primary agent for differentiation of moose populations but does not support the existence of distinct eastern and western races. Levels of genetic variation and structure of phylogenetic trees identify Asia as the origin of all extant mitochondrial lineages. A recent coalescence is indicated, with the most recent common ancestor dating to the last ice age. Moose have undergone two episodes of population expansion, likely corresponding to the final interstade of the most recent ice age and the onset of the current interglacial. Timing of expansion for the population in the Yakutia--Manchuria region of eastern Asia indicates that it is one of the oldest populations of moose and may represent the source of founders of extant populations in North America, which were colonized within the last 15,000 years. Our data suggest an extended period of low population size or a severe bottleneck prior to the divergence and expansion of extant lineages and a recent, less-severe bottleneck among European lineages. Climate change during the last ice age, acting through contraction and expansion of moose habitat and the flooding of the Bering land bridge, undoubtedly was a key factor influencing the divergence and expansion of moose populations.     

Environmental variation and moose Alces alces density as determinants of spatio-temporal heterogeneity in browsing

Understanding temporal variation in habitat selection and browsing intensity by large herbivores is fundamental because of their large impact on the ecosystems. I studied the annual variation in winter browsing pressure on young trees and habitat selection by moose Alces alces over a ten year period. Specifically, the relationships between browsing pressure on Scots pine Pinus sylvestris and two birch species ( Betula ssp.) and three explanatory variables – 1) availability of forage, 2) moose density (estimated by pellet group counts) and 3) snow cover was studied. At a larger spatial scale (forest stand level) the relationship between moose habitat selection between three different habitat types (forest <30 yr, forest>30 yr and mire) and two explanatory variables, 1) snow condition and 2) moose density, were studied. Browsing pressure on Scots pine, the dominating food plant, was related to forage availability, moose density and snow condition. No significant relationships between any of the three explanatory variables and browsing pressure on the two birch species were found. Moose selection for certain habitats varied between years and was affected by number of days with >0.10 m of snow.
Habitat selection was not significantly related to moose density and the relationship between overall moose density and habitat specific moose densities was proportional within the studied density range. These findings have implications for understanding varying browsing patterns – and will affect both the ability to predict herbivores' effect on the forest ecosystem. A snow dependent browsing pattern also indicates that one can expect a long term decrease in browsing pressure on the tree and shrub layer as a consequence of the ongoing large-scale climate change.     

Vegetation diversity influences endozoochoric seed dispersal by moose (Alces alces L.)

Research on moose-mediated seed dispersal is limited. However, its potential role in transferring seeds in patchy landscapes may be of great importance. In this work we examined how seasons and vegetation diversity influence the species richness and abundance of seeds dispersed endozoochorically by moose. Samples of moose faeces were collected year-round, fortnightly, from contrasting vegetation types, dominated by diverse, species-rich wetland or poor, dry pine forest. The viable seed content of dung was studied by the seedling emergence method. The mean number of emerged seedlings per 0.8 L sample and the mean number of plant species per 0.8 L sample were several times higher in the diverse wetland vegetation than in the poor pine forest vegetation. Maximum species richness and seed abundance was observed during the fructification period, and the minimum during spring. The species richness of samples did not differ between winter and the growing season, although the composition of plant species was different. The results of this study suggest that moose are efficient seed vectors, especially of grasses typical for grasslands and wetlands. The species richness and abundance of dispersed seeds coincides with the diversity of the vegetation of the animal’s habitat.