The time-energy budget of a moose

As the first step in developing an optimal foraging model for moose (Belovsky 1978), a time-energy budget for a moose was developed. For the summer period (late May through early September) at Isle Royale National Park, Michigan, we made various behavioral observations on feeding moose and inventoried their food resources. Employing these various sets of data, we were able to determine the quantity of each food consumed by moose, the time required to acquire the foods, preferences for different food plants, and differences demonstrated by the sexes. In addition to this information, we collected data on the moose's ability to utilize each of the forage plants and the population densities of moose in different years (1972–1974) in two forest types. This enabled us to examine energy flow within each of the populations and determine in what manner energy limits the moose population.     

Increased osteoarthritis in moose from Isle Royale

Over the past 30 yr, moose (Alces alces) in Isle Royale National Park (Michigan, USA) exhibited a several-fold increase in the prevalence of osteoarthritis, or degenerative joint disease (DJD). Available evidence points to an environmental explanation for this change. Greater physical activity among afflicted moose is not a likely contributing factor, nor is genetic change in the population. The possible introduction of an unspecified disease agent cannot be dismissed at this time. Moose exhibiting the highest prevalence of DJD were those born during a period of severe undernutrition, and it is hypothesized that nutritional stress early in life was responsible for increased joint disease during senescence. Such an etiology for osteoarthritis has not been suggested previously for any species.     

Prevalence of neutralizing antibody to Jamestown Canyon virus (California group) in populations of elk and moose in northern Michigan and Ontario, Canada

Blood samples were collected from free-ranging elk (Cervus elaphus) harvested in Michigan's northern Lower Peninsula, from moose (Alces alces) relocated from Ontario's Algonquin Provincial Park to Michigan's Upper Peninsula, and from moose from Michigan's Isle Royale National Park. Sera were tested by serum dilution neutralization tests in Vero cell culture for neutralizing antibody to California serogroup viruses, in particular Jamestown Canyon (JC), La Crosse/snowshoe hare (LAC/SSH), and trivittatus (TVT) viruses. Specific neutralizing antibody to JC virus was detected in 71% of 31 and 65% of 20 moose from Algonquin and Isle Royale, respectively. An additional six moose from Algonquin and five from Isle Royale showed evidence of multiple infection. One juvenile moose from Isle Royale had specific neutralizing antibody to TVT virus. Specific neutralizing antibody to JC virus was detected also in 54% of 50 elk from Michigan; 20 of the 50 elk showed evidence of multiple infection. While no single serum sample showed specific neutralizing antibody only to LAC/SSH virus, its presence in sera from some animals may have been masked by the high prevalence of antibody to JC virus.     

Carfentanil and xylazine for immobilization of moose (Alces alces) on Isle Royale

Twenty-one moose were captured on Isle Royale between 28 May and 2 June 1984 at licks on the western end of the island. The animals were darted, at ranges of 10 to 35 m, with 3-cc dart syringes containing 3 or 4 mg of carfentanil and 100 or 175 mg of xylazine. Three animals were drugged with carfentanil alone. Immobilization time ranged from 2.5 to 6 min. There was no excitement phase evident in 18 of the animals and they rarely moved more than 30 m after darting. Reversal was begun at 30 to 90 min after darting using naloxone and diprenorphine given intramuscularly (i.m.) and intravenously (i.v.). Recovery time varied from 10 min to 3 hr with the longer times occurring at the higher doses of immobilizing drugs. Two animals died within 30 hr, one as the result of aspiration of rumen contents and the other was unable to get on its feet and was euthanized. Analysis of blood samples from the 18 moose immobilized with the drug combination yielded hemoglobin values of 13.1 +/- 0.3 g/dl, hematocrit: 37.6 +/- 0.7%, red blood cells: 5.46 +/- 0.1 million/microliter, leucocytes: 6.1 +/- 0.4 thousand/microliter, and serum urea nitrogen: 29.3 +/- 1.6 mg/dl. Our experience indicated that quiet, undisturbed moose can be immobilized with 3 mg carfentanil and 100 mg or less of xylazine.     

Patterns of mineral lick visitation by spider monkeys and howler monkeys in Amazonia: are licks perceived as risky areas?

Mineral licks--also known as "salados," "saladeros," or "collpas"--are specific sites in tropical and temperate ecosystems where a large diversity of mammals and birds come regularly to feed on soil. Although the reasons for vertebrate geophagy are not completely understood, animals are argued to obtain a variety of nutritional and health benefits from the ingestion of soil at mineral licks. We studied the temporal patterns of mineral lick use by white-bellied spider monkey (Ateles belzebuth) and red howler monkey (Alouatta seniculus) in a lowland rain forest in Amazonian Ecuador. Using camera and video traps at four different mineral licks, combined with behavioral follows of one group of spider monkeys, we documented rates of mineral lick visitation by both primate species and the relative frequency and intensity of mineral lick use by spider monkeys. On the basis of 1,612 days and 888 nights of mineral lick monitoring, we found that A. belzebuth and A. seniculus both visit mineral licks frequently throughout the year (on average ～14% of days for both species), and mineral lick visitation was influenced by short-term environmental conditions (e.g. sunny and dry weather). For spider monkeys, the area surrounding the lick was also the most frequently and most intensively used region within the group's home range. The fact that spider monkeys spent long periods at the lick area before coming to the ground to obtain soil, and the fact that both species visited the lick preferentially during dry sunny conditions (when predator detectability is presumed to be relatively high) and visited simultaneously more often than expected by chance, together suggest that licks are indeed perceived as risky areas by these primates. We suggest that howler and spider monkeys employ behavioral strategies aimed at minimizing the probability of predation while visiting the forest floor at risky mineral lick sites.     

Moose teeth as monitors of environmental isotopic parameters

Summary The radiocarbon activities of amino acids isolated from crown first molar dentin of moose born between 1948 and 1984 on Isle Royale National Park, USA closely follows the bomb radiocarbon signal generated from atmospheric nuclear weapons testing. This demonstrates that these amino acids are metabolically inert and have recorded the isotopic parameters of the environment during the first year or two of the animal's life. The moose teeth amino acids provide both carbon and nitrogen isotopic chronologies for Isle Royale for a period of nearly four decades. The carbon isotopic record reflects both anthropogenic alterations of the global atmospheric carbon cycle (the Suess effect), and changes in forest ecology and moose feeding habitat. The nitrogen isotopic chronology is more variable than the carbon record and is the result of environmental and biological factors which are poorly understood.     

Elemental analysis of mineral lick soils from the Serengeti National Park, the Konza Prairie and Yellowstone National Park

We analyzed mineral lick soils and non-mineral lick soils from the Serengeti National Park, the Konza Praine, and Yellowstone National Park The concentrations of 14 different elements considered important for ungulate nutrition were analyzed to determine the relative importance of Na vs other elements in attracting animals to lick areas Only Na was consistently higher in the mineral lick soils of the three ecosystems Mean Na concentrations were c 20x greater in Serengeti licks, 10x greater in Konza licks and 3x greater in Yellowstone licks compared to the respective non-lick soils Despite the numerous elements analyzed for this survey, our results suggest that Na is the primary element initiates mineral lick use in these widely separated ecosystems The widespread distribution of Na-deficient forage likely explains the importance of Na in most mineral licks     

Geophagy in brown spider monkeys (Ateles hybridus) in a lowland tropical rainforest in Colombia

Spider monkeys and howler monkeys are the only Neotropical primates that eat soil from mineral licks. Not all species within these genera visit mineral licks, and geophagy has been restricted to populations of Ateles belzebuth belzebuth,Ateles belzebuth chamek and Alouatta seniculus in western Amazonian rainforests. With the aid of a camera trap we studied the visitation patterns of a group of brown spider monkeys (Ateles hybridus) to a mineral lick at Serrania de Las Quinchas, in Colombia. Spider monkeys visited the lick frequently throughout the year, with a monthly average of 21.7 ± 7.2 visits per 100 days of camera trapping (n = 14 months). Spider monkeys visited the mineral lick almost always on days with no rain, or very little (<3 mm) rain, suggesting that proximate environmental variables might determine spider monkeys' decisions to come to the ground at the licks. This study expands the geographical occurrence of mineral lick use by spider monkeys providing additional data for future assessments on the biogeographical correlates of mineral lick use by platyrrhines.     

Effects of predation risk on the grouping patterns of white‐bellied spider monkeys (Ateles belzebuth belzebuth) in Western Amazonia

Predation is proposed to be one of the most important factors influencing the evolution of mammalian societies. Although predation risk is thought to influence both the behavior and grouping patterns of most diurnal primates, evidence supporting this hypothesis is still limited. The spatial and temporal patterns of mineral lick use by one group of white‐bellied spider monkeys (Ateles belzebuth) were evaluated, based on the growing evidence that mineral licks are perceived as areas of relative high predation risk by Neotropical primates. The area immediately surrounding the mineral lick was the most intensively used area within the home range of the study group, particularly by large subgroups of monkeys, and there were differences in mean subgroup size on days of mineral lick visitation versus days without lick visits. Additionally, on days of mineral lick visitation, subgroup size reached its maximum specifically during the period of lick visitation. Finally, on visit days subgroups showed a greater increase in size and higher fusion rates in the 2 hr before arriving at the lick in comparison with matched time windows on non‐visit days. Together, these results provide an example of how primates employ behavioral strategies that might reduce the effects of predation. This study also demonstrates how taxa characterized by a high degree of fission‐fusion dynamics can allow us to test hypotheses regarding the effects of socioecological variables on primate grouping patterns. Am J Phys Anthropol 150:579–590, 2013. © 2013 Wiley Periodicals, Inc.     

Some aspects of geophagia in Wyoming bighorn sheep (<Emphasis Type="Italic">Ovis canadensis</Emphasis>)

Geophagia has been commonly reported for bighorn sheep (Ovis canadensis) and other ungulates worldwide. The phenomenon is often attributed to the need to supplement animal diets with minerals available in the soil at mineral lick locations. Sodium is the mineral most frequently cited as being the specific component sought, although this has not been found universally. In this study area, bighorn sheep left normal summer-range to make bimonthly 26-km, 2,000-m-elevation round-trip migrations, the apparent purpose of which was to visit mineral licks on normal winter-range. Lick soil and normal summer-range soil were sampled for their available mineral content and summer-range forage was sampled for total mineral content, and comparisons were made to determine the specific components sought at the lick by bighorn sheep consuming soil. It was concluded that bighorn sheep were attracted to the lick by a desire for sodium but that geophagia also supplemented a diet deficient in the trace element selenium. Where sheep are denied access to licks, populations may be limited by mineral deficiency.     

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.     

Daily rhythm and seasonal pattern of lick use in sika deer (Cervus nippon) in China

Many ungulates are reported to use natural or artificial licks with seasonal patterns around the world. From December 2016 to August 2017, we used infrared camera to record the use of artificial licks in wild sika deer (Cervus nippon) in Zhejiang Qingliangfeng National Nature Reserve, China. We explored the daily rhythm, seasonal pattern and sex difference in lick utilization. In total, 12,043 videos and 22,901 pictures were collected. Our results showed that: (1) the lick visiting frequency was higher at night than that during daytime; (2) the difference in lick visiting frequency between females and males disappeared after taking into account of sex ratio; (3) the lick duration peaked in April during a year. These findings suggested that there were clear daily rhythm and seasonal pattern of lick use in sika deer. Seasonal change in lick use intensity was consistent with our prediction. These variations in lick use might be driven by both the physiological needs of the mineral elements in different life stages and the seasonal changes in climate and food. The reserve management authority should pay more attention to the supplement of licks in spring and summer to fulfill animal’s physiological needs.     

Behavior of Moose Relative to a Road Network

Abstract Roads often negatively affect terrestrial wildlife, via habitat loss or fragmentation, noise, and direct mortality. We studied moose (Alces alces) behavior relative to a road network, in an area with a history of moose-vehicle accidents, to determine when moose were crossing roadways or using areas near roads and to investigate if environmental factors were involved in this behavior. We tracked 47 adult moose with Global Positioning System collars in a study area crossed by highways and forest roads. We hypothesized that moose would avoid crossing roads but would make occasional visits to roadsides to feed on sodium-rich vegetation and avoid biting insects. Further, we expected moose avoidance to be greater for highways than forest roads. We recorded 196,710 movement segments but only observed 328 highway and 1,172 forest-road crossings (16 and 10 times lower than expected by chance). Moose usually avoided road proximity up to ≥500 m on each side but 20% of collared moose made visits to areas within 50 m of highways, which might have resulted from moose searching for sodium in vegetation and roadside salt pools. In fact, vegetation along highways had higher sodium concentrations and was browsed in similar proportions to vegetation in adjacent forest, despite moose avoidance of these zones. Moose, however, did not use areas near roads more during periods of biting insect abundance. Our results supported the hypothesis of scale-dependent selection by moose; avoidance of highways at a coarse scale may confer long-term benefits, whereas selection of highway corridors at finer scales may be part of a strategy to overcome short-term limiting factors such as sodium deficiency. We found a positive relationship between home-range size and the proportion of road axes they contained, suggesting that moose either compensated for habitat loss or made specific movements along highways to gather sodium. The presence of sodium along highways likely increases moose-vehicle accident risks. Removal of salt pools or use of a de-icing salt other than sodium chloride should render highway surroundings less attractive to moose.     

Behavioral Adaptations of Moose to Roadside Salt Pools

Abstract: Sodium has many fundamental physiological functions in animals but is rare in boreal ecosystems where moose (Alces alces) thrive. In Québec (Canada), sodium is readily available in aquatic vegetation and in salt pools that form along highways. We do not know if moose are adopting specific behaviors to access sodium sources or if they simply use the sodium sources they encounter during their movements. We tested the hypothesis that moose modify both space and habitat use to gather sodium from salt pools. We expected moose to use salt pools mostly in spring and early summer, when needs are greatest and before aquatic vegetation has fully developed. We fitted 47 moose with Global Positioning System telemetry collars and collected data for 2 to 36 months between 2003 and 2006. We rarely located moose at salt pools (0.12% among the 95,007 locations collected). As we expected, use of salt pools was highest in late spring and in early summer, and we observed a time lag between peak use of salt pools compared to use of lakes and waterways, indicating moose fulfilled their sodium requirements in salt pools before aquatic vegetation was available. Moose selected salt pools over lakes and waterways when these 2 sodium sources were present in their home range and moved rapidly over large distances to reach them. Our results were consistent with moose using salt pools when they are likely to be sodium deficient. Salt pools were less accessible, required long-distance movements, and were located in habitually avoided areas along highways. Elimination of roadside salt pools should be considered among strategies to reduce cervid-vehicle collision risks in boreal environments.     

Moose and snowshoe hare competition and a mechanistic explanation from foraging theory

Summary Moose (Alces alces) and snowshoe hare (Lepus americanus) appear to compete with each other. This was determined using the natural experiments of populations found in sympatry and allopatry on islands at Isle Royale National Park, Michigan, and manipulated exclosures. The population densities from these areas are fit to a series of competition models based upon different competitive mechanisms (Schoener 1974a), using non-linear regression techniques. A model of competition for food where the food can be separated into exclusively used and shared categories is found to predict observed densities of moose and hare best. Finally, the competition model's parameters (fraction of food shared and competition coefficients) are shown to agree with values predicted independently from a foraging model.     

Predator disease out-break modulates top-down, bottom-up and climatic effects on herbivore population dynamics

Human-introduced disease and climatic change are increasingly perturbing natural ecosystems worldwide, but scientists know very little about how they interact to affect ecological dynamics. An outbreak of canine parvovirus (CPV) in the wolf population on Isle Royale allowed us to test the transient effects of an introduced pathogen and global climatic variation on the dynamics of a three-level food chain. Following the introduction of CPV, wolf numbers plummeted, precipitating a switch from top-down to bottom-up regulation of the moose population; consequently, the influence of climate on moose population growth rate doubled. This demonstrates that synergistic interactions between pathogens and climate can lead to shifts in trophic control, and suggests that predators in this system may play an important role in dampening the effects of climate change on the dynamics of their prey.     

Differences in urinary chemistry profiles of moose on Isle Royale during winter

During winters 1987-1988 (I) and 1988-1989 (II), we assessed the physiological status of moose (Alces alces) residing on the west and east ends of Isle Royale (Michigan, USA) by collecting and chemically analyzing urine deposited in snow (snow-urine) from January to early March. Samples were assayed for urea nitrogen (U), sodium (Na), potassium (K), calcium (Ca), phosphorus (P) and creatinine (C). Throughout both winters, elevated urinary U:C ratios in snow-urine samples collected from east-end moose compared to west-end moose indicated greater dietary energy deprivation and accelerated net catabolism of endogenous protein. Sodium: C ratios were low throughout the study and were similar between moose from both ends of the island, except during the middle of winter I. Greater K:C, P:C and Ca:C ratios in east-end moose compared to west-end moose throughout winter I, and increases in these ratios and U:C in east-end moose from middle to late winter during the second year provided additional evidence of a greater deterioration in condition in east-end moose. The superior nutrition provided to moose on the west end of the island was associated with more developed soils and diverse vegetation and a lower stem density of balsam fir compared to the east end.     

SOME ASPECTS OF BIG-GAME UTILIZATION OF MINERAL LICKS IN YANKARI GAME RESERVE, NIGERIA

At least ten major mineral licks occur along the Gaji River which bisects the Yankari Game Reserve. The licks show varying degrees of utilization and have craters of up to 1200m³ in volume. Three hundred and eighty five hours of observation were kept at one of the licks. Its use by different species was highly variable. Some species occurring in the reserve made no visits to the lick, whereas Western hartebeest (51 groups), waterbuck (53 groups) and warthog (276 groups) made frequent visits. Lick use was much greater in the daytime than at night, with the peak occurring around 1400 h. A limited number of soil samples (29) were analysed but no single element can be considered as the factor attracting wildlife. However, the lick areas were richer in most minerals than were samples of soil from drier habitat in the open woodland. It is possible that significantly increased lick use might in the future indicate degrading environmental conditions in the drier parts of the reserve. If hides are constructed, the commonly utilized licks will provide excellent opportunities for behaviour observatioii and photography by tourists.     

Salt licks do not increase local densities of the deer ked,Lipoptena cervi,an abundant ectoparasite of cervids

The deer ked, Lipoptena cervi (Diptera: Hippoboscidae), is a common ectoparasite of the moose, Alces alces (Artiodactyla: Cervidae). Salt licks are widely used to manipulate moose movements to prevent damage to saplings and traffic accidents. They may cause moose to gather in small areas, which could create aggregates of deer ked pupae as the parasite is a short‐distance flyer and its dispersion depends on its hosts. We investigated whether the population density of flying deer keds could be influenced by manipulating salt licks and how environmental variables affect parasite density. Densities were estimated in 40 experimental sites with four treatments (no salt licks, introduced salt licks, removed salt licks, permanent salt licks) in September during 2007–2010. Forest edges, mixed forests on mineral soil and coniferous forests on peat soil were the habitats with high numbers of parasites. The manipulation of salt licks seemed to be ineffective in reducing the density of deer keds as the only factor to show statistical significance with parasite numbers in the mixed‐model analysis was year of determination. Annual deer ked densities correlated with the abundance of moose in the region. Moreover, high spring and summer temperatures seemed to increase the numbers of flying imagos.     

Mineral diversity in wild plants: benefits and bane for moose

We studied the mineral nutrition ecology of moose, Alces alces , by a comprehensive examination of concentration levels of 18 elements in browsing plants, and in rumen and distal colon contents, respectively. The plants, in total 14 species, included deciduous and evergreen trees, shrubs, herbs, grasses and aquatic macrophytes. Our study was done in four sites in southern Norway and the plants were sampled in spring, summer, autumn and winter to document seasonal trends. The plants were generally characterized by low concentrations of major elements, very low phosphorus: calcium ratios, and the willow Salix aurita – a preferred browse – had high cadmium concentrations, up to 9 ppm (dry mass). Sodium concentrations in terrestrial plants were significantly under the dietary requirement, while levels were sufficient in aquatic plants, which, however, had high concentrations of toxic heavy metals. We suggest that the moose experience a nutritional dilemma as a strong physiological need for sodium results in an overintake of heavy metals. A selective intake of plants rich in heavy metals was corroborated by high heavy metal concentrations in rumen and distal colon. Over all, the plants were characterized by a marked mineral diversity due to element concentration levels that were highly variable between species and sites. Thus, there is a strong link between plant species diversity and mineral diversity, which has a pivotal role for herbivores. It is concluded that a diversity of plant species is a prerequisite for the mineral diversity that the moose needs to obtain essential minerals, not only in sufficient amounts, but also in physiologically balanced proportions.