Performance of wild animals with “broken” traits: Movement patterns in nature of moose with leg injuries

Organismal traits are presumed to be well suited for performance in the tasks required for survival, growth, and reproduction. Major injuries to such traits should therefore compromise performance and prevent success in the natural world; yet some injured animals can survive for long periods of time and contribute to future generations. We here examine 3 years of camera trap observations along a remote trail through old‐growth forest in northern British Columbia, Canada. The most common observations were of moose (2966), wolves (476), and brown bears (224). The moose overwhelmingly moved in one direction along the trail in the late fall and early winter and in the other direction in the spring. This movement was clustered/contagious, with days on which many moose traveled often being interspersed with days on which few moose traveled. On the video recordings, we identified 12 injured moose, representing 1.4% of all moose observations. Seven injuries were to the carpus, three were to the antebrachium, and two were to the tarsus—and they are hypothesized to reflect damage to ligaments, tendons, and perhaps bones. The injured moose were limping in all cases, sometimes severely; and yet they did not differ noticeably from uninjured moose in the direction, date, contagiousness, or speed of movement along the trail. We discuss the potential relevance of these findings for the action of natural selection in the evolution of organismal traits important for performance.     

COVID‐19 lockdown measures impacted citizen science hedgehog observation numbers in Bavaria,Germany

The COVID‐19 pandemic has led to temporary changes in human–animal interactions due to changes in human activities. Here, we report on a surge in hedgehog observations during the first COVID‐19 lockdown in Germany in 2020, on the citizen science Web portal “Igel in Bayern” (Hedgehogs in Bavaria) in Germany. This increase in comparison with previous years was attributed to an increase in the number of people reporting hedgehog observations, rather than an increase in the number of hedgehog observations made by each observer. Additionally, in contrast to other studies on the effects of a COVID‐19 lockdown on observations recorded by citizen science projects, the share of observations made in more urbanized areas during the lockdown time was not higher than the change observed in less urbanized areas. This is possibly a result of the differences in COVID‐19 measures between Germany and other countries where preceding studies were carried out, in particular the lack of measures limiting traveling outdoor activities for citizens.     

Drastic increase in the prevalence of Echinococcus multilocularis in foxes (Vulpes vulpes) in southern Bavaria, Germany

A survey on the prevalence of Echinococcus multilocularis in foxes was conducted in the administrative district of Starnberg (federal state of Bavaria, Germany) and some adjacent municipalities from October 2002 to March 2003. The background to the study was the scarcity of recent data for Bavaria, where, in contrast to neighbouring regions, a general increase in the prevalence of the parasite has not yet been demonstrated. To estimate the current infection rate, a total of 268 shot foxes were examined using the intestinal scraping technique, resulting in an overall prevalence of 51%. This was compared with retrospective data collected during the period from 1989 to 2001. For the corresponding area and season, the retrospective prevalence was estimated at 32%, based on 222 shot foxes. The prevalence and its temporal development differ considerably on small spatial scales. The most conspicuous change has taken place in the western part of the study area, where a previous prevalence of 35% has increased to 80% in 2002/2003.     

Mobility of moose—comparing the effects of wolf predation risk,reproductive status,and seasonality

In a predator–prey system, prey species may adapt to the presence of predators with behavioral changes such as increased vigilance, shifting habitats, or changes in their mobility. In North America, moose (Alces alces) have shown behavioral adaptations to presence of predators, but such antipredator behavioral responses have not yet been found in Scandinavian moose in response to the recolonization of wolves (Canis lupus). We studied travel speed and direction of movement of GPS‐collared female moose (n = 26) in relation to spatiotemporal differences in wolf predation risk, reproductive status, and time of year. Travel speed was highest during the calving (May–July) and postcalving (August–October) seasons and was lower for females with calves than females without calves. Similarly, time of year and reproductive status affected the direction of movement, as more concentrated movement was observed for females with calves at heel, during the calving season. We did not find support for that wolf predation risk was an important factor affecting moose travel speed or direction of movement. Likely causal factors for the weak effect of wolf predation risk on mobility of moose include high moose‐to‐wolf ratio and intensive hunter harvest of the moose population during the past century.     

The declining occurrence of moose (Alces alces) at the southernmost edge of its range raise conservation concerns

The border region between Austria, the Czech Republic, and Germany harbors the most south‐western occurrence of moose in continental Europe. The population originated in Poland, where moose survived, immigrated from former Soviet Union or were reintroduced after the Second World War expanded west‐ and southwards. In recent years, the distribution of the nonetheless small Central European population seems to have declined, necessitating an evaluation of its current status. In this study, existing datasets of moose observations from 1958 to 2019 collected in the three countries were combined to create a database totaling 771 records (observations and deaths). The database was then used to analyze the following: (a) changes in moose distribution, (b) the most important mortality factors, and (c) the availability of suitable habitat as determined using a maximum entropy approach. The results showed a progressive increase in the number of moose observations after 1958, with peaks in the 1990s and around 2010, followed by a relatively steep drop after 2013. Mortality within the moose population was mostly due to human interactions, including 13 deadly wildlife‐vehicle collisions, particularly on minor roads, and four animals that were either legally culled or poached. Our habitat model suggested that higher altitudes (ca. 700–1,000 m a.s.l.), especially those offering wetlands, broad‐leaved forests and natural grasslands, are the preferred habitats of moose whereas steep slopes and areas of human activity are avoided. The habitat model also revealed the availability of large core areas of suitable habitat beyond the current distribution, suggesting that habitat was not the limiting factor explaining the moose distribution in the study area. Our findings call for immediate transboundary conservation measures to sustain the moose population, such as those aimed at preventing wildlife‐vehicle collisions and illegal killings. Infrastructure planning and development activities must take into account the habitat requirements of moose.     

Morphological and ecological characteristics of moose, <Emphasis Type="Italic">Alces</Emphasis> cf. <Emphasis Type="Italic">alces</Emphasis> (Artiodactyla), from the southeast of Western Siberia in the Late Neopleistocene and Holocene

Fossil, subfossil, and recent materials of moose from the southeast of Western Siberia, including the remains of the skull, antlers, mandibles, and postcranial skeleton, has been analyzed. Adaptive features revealed in the structure of the mandible (a high pars dentalis) suggest that forests were a less typical habitat for ancient moose, compared to recent A. a. alces. The high pars dentalis of the mandible and the shape of antlers, with a long basal part and bifurcated spatula, are indicative of their close relation to A. americanus from Eastern Siberia. Thus, moose from the Late Neopleistocene and Holocene appear to represent the same species as A. americanus. It cannot be excluded, however, that A. cf. alces is a special type of European A. a. alces moose in which the characters convergent to Eastern Siberian A. americanus developed independently, due to habitation in similar open (forest-steppe and steppe) landscapes.     

Challenges and opportunities for estimating abundance of a low-density moose population

Monitoring large herbivores across their core range has been readily accomplished using aerial surveys and traditional distance sampling. But for peripheral populations, where individuals may occur in patchy, low-density populations, precise estimation of population size and trend remains logistically and statistically challenging. For moose (Alces alces) along their southern range margin in northern New York, USA, we sought robust estimates of moose distribution, abundance, and population trend (2016–2019) using a combination of aerial surveys (line transect distance-sampling), repeated surveys in areas where moose were known to occur to boost the number of detections, and density surface modeling (DSM) with spatial covariates. We achieved a precise estimate of density (95% CI = 0.00–0.29 moose/km²) for this small population (656 moose, 95% CI = 501–859), which was patchily distributed across a large and heavily forested region (the 24,280-km² Adirondack Park). Local moose abundance was positively related to active timber management, elevation, and snow cover, and negatively related to large bodies of water. As expected, moose abundance in this peripheral population was low relative to its core range in other northern forest states. Yet, in areas where abundance was greatest, moose densities in New York approached those where epizootics of winter tick (Dermacentor albipictus) have been reported, underscoring the need for effective and efficient monitoring. By incorporating autocorrelation in observations and landscape covariates, DSM provided spatially explicit estimates of moose density with greater precision and no additional field effort over traditional distance sampling. Combined with repeated surveys of areas with known moose occurrence to achieve viable sample sizes, DSM is a useful tool for effectively monitoring low density and patchy populations.     

Range expansion by moose into coastal temperate rainforests of British Columbia, Canada

Ranges of species are dynamic and respond to long‐term climate change and contemporary effects such as habitat modification. We report here that moose (Alces alces) have recently colonized coastal temperate rainforests of British Columbia, Canada. Contrary to recent publications, field observations of moose and their sign, combined with their occurrence in wolf (Canis lupus) faeces, suggest that moose are now widespread on the coastal mainland and occur on least three islands. Traditional ecological knowledge (information accumulated by aboriginal peoples about their environment) suggests that colonization occurred during the mid 1900s, concomitant with logging of major watersheds that bisect the Coast Mountain Range. Range expansion by moose may have ecological consequences such as alteration of predator–prey dynamics and transmission of disease to native deer (Odocoileus hemionus).     

Age and sex-specific variation in detectability of moose (<Emphasis Type="Italic">Alces alces</Emphasis>) during the hunting season: implications for population monitoring

The sustainability of wild ungulate harvests can be greatly enhanced if monitoring data are available to permit an adaptive management approach. Utilising data provided by hunters is potentially the most cost-effective approach. In Scandinavia, observations recorded by moose (Alces alces) hunters provide a range of indices of population density, composition and reproductive performance. These are routinely used in practical management, but there are still many questions about their accuracy and precision. In this study, we availed of the fact that virtually all individual moose on the island of Vega in central Norway were marked during the period 1992–2005. Thus, we were able to compare the observation indices provided by hunters to the known size and composition of the population. The results indicate that the hunter observation indices provided accurate estimates of variation in moose cow recruitment and twinning rates. The estimates of sex ratio closely followed the pattern of annual variation but showed a consistent pattern of over-estimating the proportion of males. Thus, males seem to visually expose themselves more often for hunters than do females. The density index, measured as number of moose seen per hunter day did not to the same extent follow the variation in population density, possibly because moose were more naive in the first years after hunting was introduced or because of reduced detection conditions due to increasing scrub and tree encroachment during the study period. The results are discussed in light of their application to management and the evolutionary pressures in moose anti-predator behaviour.     

Perinatal mortality in Bavaria, Germany, after the Chernobyl reactor accident

As has been shown by the authors of a paper recently published in this journal, a deviation from a long-term trend in perinatal mortality within the former Federal Republic of Germany occurred in 1987, i.e. 1 year following the Chernobyl disaster. It is the aim of this study to make a comparison between the areas of the state Bavaria, Germany, with different fallout levels as well as between the observed and expected numbers of perinatal deaths relating to these areas. The expected numbers of perinatal deaths, defined as external standard, were derived from the remainder of the former FRG. Testing an a priori formulated hypothesis revealed no differences in the temporal development of perinatal mortality between the areas with different fallout levels and subsequent exposures. Including May 1986 into the analysis revealed a significant increase during the first 3 months after the accident, which is due to an excess in May alone. Since no elevated radiation risks for the last days in utero are known, the additional Chernobyl radiation exposure is not plausible as a causative agent. Further analyses on stillbirths showed an increase in Southern Bavaria during the first 2 years following the accident. Later on, the rates were comparable to the expected values again. Received: 23 January 1997 / Accepted in revised form: 4 April 1997     

<Emphasis Type="Italic">Rosselia socialis</Emphasis> in the Upper Marine Molasse of southwestern Germany

Sediments of the Miocene Upper Marine Molasse in southern Germany (southwestern Bavaria) represent a shallow marine, siliciclastic, and tide-influenced depositional environment. Three sections cutting through these sediments contain a distinct horizon of trace fossils. Preserved in a fine-grained quartz-sandstone are numerous more or less vertical burrows that are usually funnel-shaped, closely spaced, and locally branching. The burrows belong to the ichnospecies Rosselia socialis. The sequence-stratigraphic relevance of the Rosselia-horizons is discussed, and a hypothesis is put forward on the correlation of selected sections of the Upper Marine Molasse based on these layers.     

Effects of willow nutrition and morphology on calving success of moose

Across much of North America, populations of moose (Alces alces) are declining because of disease, predation, climate change, and anthropogenic-driven habitat loss. Contrary to this trend, populations of moose in Colorado, USA, have continued to grow. Studying successful (i.e., persistent or growing) populations of moose can facilitate continued conservation by identifying habitat features critical to persistence of moose. We hypothesized that moose using habitat with higher quality willow (Salix spp.) would have a higher probability of having a calf-at-heel (i.e., calving success). We evaluated moose calving success using repeated ground observations of collared individuals with calves in an occupancy model framework to account for detection probability. We then evaluated the impact of willow habitat quality and nutrition on moose calving success by studying 2 spatially segregated populations of moose in Colorado. Last, we evaluated correlations between willow characteristics (browse intensity, height, cover, leaf length, and species) and willow nutrition (dry matter digestibility [DMD]) to assess the utility of using those characteristics to assess willow nutrition. We found willow height and cover had a high probability of being positively associated with higher individual-level calving success. Willow DMD, browse intensity, and leaf length were not predictive of individual moose calving success; however, the site with higher mean DMD consistently had higher mean estimates of calving success for the same year. Our results suggest surveying DMD is likely not a useful metric for assessing differences in calving success of individual moose but may be of use at population levels. Further, the assessment of willow morphology and density may be used to identify areas that support higher levels of moose calving success.     

Early Miocene (Ottnangian) Mollusca of the Western Paratethys—ontogenetic strategies and palaeo-environments

Early ontogenetic shells of 25 species of brackish water and freshwater molluscs from the Ottnangian (Lower Miocene) Oncophora Beds (Lower Bavaria, South Germany) and Kirchberg Formation (Upper Bavaria, South Germany) are described for the first time. Taxonomic implications are discussed. The investigated bivalves (Cardiidae and Dreissenidae) were characterised by an indirect development with inclusion of a planktonic veliger stage. Among the gastropods only three species of the genus CtyrokiaSchlickum, 1965 were characterised by veliger larvae, all the other gastropod species were direct developers, which hatched as crawling young. The species Agapilia schlickumi nov. sp. (Neritidae), Nematurella pseudozilchi nov. sp. and Nematurella strauchi nov. sp. (Hydrobiidae) are introduced. Our study revealed the co-occurrence of 10 molluscan species in the Oncophora Basin of Lower Bavaria and the Kirchberg Basin of Upper Bavaria and thus indicates similar oligohaline to mesohaline coastal swamp milieus for both depositional environments. The presence of planktonic larval development in gastropods indicates a connection to the open sea.     

Early jurassic (Upper Pliensbachian) gastropods from the Herforder Liasmulde (Bielefeld, Northwest Germany)

An Early Jurassic (Upper Pliensbachian) gastropod fauna from the Herforder Liasmulde (Herford Lias Syncline) near Bielefeld, Northwest Germany is described. This is one of the first detailed and illustrated reports of Pliensbachian gastropods from this area. Sixteen species are reported. Surface collecting yielded relatively large vetigastropods (pleurotomarioids amd trochomorphs). Bulk samples yielded abundant small caenogastropods (especiallyLevipleura blainvillei andKalchreuthia frankei) and cylindrobullinid opisthobranchs (e.g.Cylindrobullina dornend). These small species are also abundant in the Pliensbachian of South Germany (N Bavaria, Franconia). Therefore, this characteristic Pliensbachian microgastropod assemblage was widespread in Germany and probably also in other comparable marine deposits of Central Europe (e.g. the Paris Basin).      

Managing for Elevated Yield of Moose in Interior Alaska

ABSTRACT Given recent actions to increase sustained yield of moose (Alces alces) in Alaska, USA, we examined factors affecting yield and moose demographics and discussed related management. Prior studies concluded that yield and density of moose remain low in much of Interior Alaska and Yukon, Canada, despite high moose reproductive rates, because of predation from lightly harvested grizzly (Ursus arctos) and black bear (U. americanus) and wolf (Canis lupus) populations. Our study area, Game Management Unit (GMU) 20A, was also in Interior Alaska, but we describe elevated yield and density of moose. Prior to our study, a wolf control program (1976–1982) helped reverse a decline in the moose population. Subsequent to 1975, moose numbers continued a 28-year, 7-fold increase through the initial 8 years of our study (λ_B1 = 1.05 during 1996–2004, peak density = 1,299 moose/1,000 km²). During these initial 8 hunting seasons, reported harvest was composed primarily of males ( = 88%). Total harvest averaged 5% of the prehunt population and 57 moose/1,000 km², the highest sustained harvest-density recorded in Interior Alaska for similar-sized areas. In contrast, sustained total harvests of <10 moose/1,000 km² existed among low-density, predator-limited moose populations in Interior Alaska (≤417 moose/1,000 km²). During the final 3 years of our study (2004–2006), moose numbers declined (λ_B2 = 0.96) as intended using liberal harvests of female and male moose ( = 47%) that averaged 7% of the prehunt population and 97 moose/1,000 km². We intentionally reduced high densities in the central half of GMU 20A (up to 1,741 moose/1,000 km² in Nov) because moose were reproducing at the lowest rate measured among wild, noninsular North American populations. Calf survival was uniquely high in GMU 20A compared with 7 similar radiocollaring studies in Alaska and Yukon. Low predation was the proximate factor that allowed moose in GMU 20A to increase in density and sustain elevated yields. Bears killed only 9% of the modeled postcalving moose population annually in GMU 20A during 1996–2004, in contrast to 18–27% in 3 studies of low-density moose populations. Thus, outside GMU 20A, higher bear predation rates can create challenges for those desiring rapid increases in sustained yield of moose. Wolves killed 8–15% of the 4 postcalving moose populations annually (10% in GMU 20A), hunters killed 2–6%, and other factors killed 1–6%. Annually during the increase phase in GMU 20A, calf moose constituted 75% of the predator-killed moose and predators killed 4 times more moose than hunters killed. Wolf predation on calves remained largely additive at the high moose densities studied in GMU 20A. Sustainable harvest-densities of moose can be increased several-fold in most areas of Interior Alaska where moose density and moose: predator ratios are lower than in GMU 20A and nutritional status is higher. Steps include 1) reducing predation sufficient to allow the moose population to grow, and 2) initiating harvest of female moose to halt population growth and maximize harvest after density-dependent moose nutritional indices reach or approach the thresholds we previously published.     

Reintroduction of the endangered and endemic plant species Cochlearia bavarica—Implications from conservation genetics

Population reintroduction is a common practice in conservation, but often fails, also due to the effects of inbreeding or outbreeding depression. Cochlearia bavarica is a strongly endangered plant species endemic to Bavaria in Germany, constantly declining since the late 1980s. Therefore, population reintroduction is intended. In this study, we analyzed genetic diversity within and genetic differentiation between all 32 remnant populations of the species in Swabia and Upper Bavaria using amplified fragment length polymorphisms. Our aim was to increase reintroduction success by providing data to avoid negative effects of inbreeding and outbreeding and to preserve the natural genetic pattern of the species. Genetic diversity within populations was low but similar to other rare and endemic species and varied strongly between populations but did not depend on population size. Our analysis revealed a strong geographic pattern of genetic variation. Genetic differentiation was strongest between Swabia and Upper Bavaria and at the population level, whereas differentiation between subpopulations was comparatively low. Isolation by distance and genetic differentiation was stronger among populations from Upper Bavaria than from Swabia. From the results of our study, we derived recommendations for a successful reintroduction of the species. We suggest using rather genetically variable than large populations as reintroduction sources. Moreover, the exchange of plant material between Swabia and Upper Bavaria should be completely avoided. Within these regions, plant material from genetically similar populations should preferably be used for reintroduction, whereas the exchange among subpopulations seems to be possible without a negative impact on genetic variation due to natural gene flow.     

Simulated responses of moose populations to browsing‐induced changes in plant architecture and forage production

We and others have previously shown that browsing by large mammalian herbivores can alter the fractal dimension and other architectural properties of plant crowns. Using data from an experiment in northern Sweden which manipulated moose population densities from 0 to 50 moose per 1000 ha and measured architectural responses of birch, Betula pendula and B. pubescens, and Scots pine, Pinus sylvestris, we constructed a simulation model to examine how these changes in plant crown architecture affect moose population dynamics. Architectural changes in birch tree crowns caused forage availability to increase from low to moderate moose population densities, then decline at higher densities. In contrast, forage availability of pine decreased monotonically with increasing moose population density. The model, which incorporated equations fitted to these experimental results, predicted realistic moose population densities for northern Sweden. More importantly, the model also predicted that the quadratic responses of birch crowns to moose browsing results in moose population oscillations on high productivity sites because the moose population density overshoots the maximum birch forage availability then declines. Changes in the geometry of plant canopies caused by mammalian browsers and soil fertility may feed back on the population dynamics of the browsers themselves and even result in complex dynamics such as population oscillations.     

Genetic and morphological divergence among Gravel Bank Grasshoppers, <Emphasis Type="BoldItalic">Chorthippus pullus</Emphasis> (Acrididae), from contrasting environments

Gravel Bank Grasshopper (Chorthippus pullus) populations inhabit two contrasting environments, pebbly gravel banks with scarce vegetation cover in mountainous areas along the Alps and lowland grasslands dominated by Common Heather (Calluna vulgaris). Heath populations of C. pullus have been rediscovered only recently, and show a distribution scattered across Central Europe. The wings are reduced in this species; thus, it has low potential for long-distance dispersal. We used sequence data on a newly developed non-coding nuclear marker from three gravel-bank and four heath populations to test whether grasshoppers from the two environments represent distinct lineages. Gravel-bank populations were studied in southern Germany (Bavaria), heath populations in eastern Germany (Brandenburg and Saxony) and Ukraine. We compared those genetic data with an analysis of variation in a suite of morphometric traits. Finally, we combined genetic and morphometric data to reconstruct a plausible scenario for the ecological shift observed in C. pullus. Our newly developed marker did not sort populations from contrasting environments in two monophyletic lineages. Nevertheless, we found a general lack of gene flow between the gravel-bank and heath populations. There was pronounced variation among populations in morphometric traits. That variation was partially partitioned by habitat type, and populations from the same habitat tended to be more similar than those from different habitats. Our data suggest that heath populations originated through northward expansion from multiple southern European refugia, and that the gravel-bank populations represent one of these sources. Patterns of genetic and morphometric divergence suggest that gravel-bank and heath populations may be in the process of incipient speciation.     

Cascading effects of moose (<Emphasis Type="Italic">Alces alces</Emphasis>) management on birds

Large herbivores often have key functions in their ecosystems, and may affect ecosystem processes with cascading effects on other animals. The mechanisms often involve relocations of resources of various kinds, including reduction in resource availability following foraging and increase in resources from animal excreta. As large herbivore populations in Europe generally are intensely managed, management activities may interact with the activities of the herbivores themselves in the effect on other ecosystem components. We investigated the effects of moose (Alces alces) winter browsing, together with the effect of net nutrient input via supplementary winter feeding of moose on functional composition and species richness of birds in a boreal forest. Supplementary feeding stations for moose had a net zero effect on bird species richness and abundance, because negative effects of moose browsing were balanced by positive effects of nutrient input. Sites with a similar browsing intensity as at feeding stations but without nutrient input had lower abundance and species richness than feeding stations. Functional groups of bird species showed differing responses: birds nesting at or below browsing height were negatively affected by moose browsing, whereas species nesting above the browsing zone were positively affected by moose browsing. Insect-eating species responded negatively to moose browsing on birch but positively to nutrient input at feeding stations, whereas seed-eating species responded positively to birch browsing and negatively to feeding stations. This study showed that both high levels of cervid activity and human management interventions influence bird communities.     

Moose (Alces alces) reacts to high summer temperatures by utilizing thermal shelters in boreal forests – an analysis based on airborne laser scanning of the canopy structure at moose locations

The adaptation of different species to warming temperatures has been increasingly studied. Moose (Alces alces) is the largest of the ungulate species occupying the northern latitudes across the globe, and in Finland it is the most important game species. It is very well adapted to severe cold temperatures, but has a relatively low tolerance to warm temperatures. Previous studies have documented changes in habitat use by moose due to high temperatures. In many of these studies, the used areas have been classified according to how much thermal cover they were assumed to offer based on satellite/aerial imagery data. Here, we identified the vegetation structure in the areas used by moose under different thermal conditions. For this purpose, we used airborne laser scanning (ALS) data extracted from the locations of GPS‐collared moose. This provided us with detailed information about the relationships between moose and the structure of forests it uses in different thermal conditions and we were therefore able to determine and differentiate between the canopy structures at locations occupied by moose during different thermal conditions. We also discovered a threshold beyond which moose behaviour began to change significantly: as day temperatures began to reach 20 °C and higher, the search for areas with higher and denser canopies during daytime became evident. The difference was clear when compared to habitat use at lower temperatures, and was so strong that it provides supporting evidence to previous studies, suggesting that moose are able to modify their behaviour to cope with high temperatures, but also that the species is likely to be affected by warming climate.