Modelling habitat-suitability using museum collections: an example with three sympatric Apodemus species from the Alps

Aim, Location Although the alpine mouse Apodemus alpicola has been given species status since 1989, no distribution map has ever been constructed for this endemic alpine rodent in Switzerland. Based on redetermined museum material and using the Ecological-Niche Factor Analysis (ENFA), habitat-suitability maps were computed for A. alpicola , and also for the co-occurring A. flavicollis and A. sylvaticus .
Methods In the particular case of habitat suitability models, classical approaches (GLMs, GAMs, discriminant analysis, etc.) generally require presence and absence data. The presence records provided by museums can clearly give useful information about species distribution and ecology and have already been used for knowledge-based mapping. In this paper, we apply the ENFA which requires only presence data, to build a habitat-suitability map of three species of Apodemus on the basis of museum skull collections.
Results Interspecific niche comparisons showed that A. alpicola is very specialized concerning habitat selection, meaning that its habitat differs unequivocally from the average conditions in Switzerland, while both A. flavicollis and A. sylvaticus could be considered as 'generalists' in the study area.
Main conclusions Although an adequate sampling design is the best way to collect ecological data for predictive modelling, this is a time and money consuming process and there are cases where time is simply not available, as for instance with endangered species conservation. On the other hand, museums, herbariums and other similar institutions are treasuring huge presence data sets. By applying the ENFA to such data it is possible to rapidly construct a habitat suitability model. The ENFA method not only provides two key measurements regarding the niche of a species (i.e. marginality and specialization), but also has ecological meaning, and allows the scientist to compare directly the niches of different species.     

Space Use and Habitat Selection by Female Louisiana Black Bears in the Tensas River Basin of Louisiana

Abstract: Studies of space use and habitat selection of endangered species are useful for identifying factors that influence fitness of individuals and viability of populations. However, there is a lack of published information regarding these behaviors for the federally threatened Louisiana black bear (Ursus americanus luteolus). We documented space use and habitat selection for 28 female black bears in 2 subpopulations of the Tensas River Basin population in northeast Louisiana, USA. The Tensas subpopulation inhabits a relatively large (>300-km²) contiguous area of bottomland hardwood forest, whereas the Deltic subpopulation exists mainly in 2 small (<7-km²) forested patches surrounded by an agricultural matrix. Females on Deltic maintained smaller seasonal and annual home ranges than females on Tensas (all P < 0.04), except for females with cubs during spring. On Tensas, females with cubs maintained smaller home ranges than females without cubs during spring (P = 0.01), but we did not detect this difference on Deltic or in other seasons. Females on Tensas and Deltic exhibited differences in habitat selection when establishing home ranges and within home ranges (P < 0.001). Deltic females selected mature bottomland hardwood forests and avoided agricultural habitats at both spatial scales. Tensas females selected a mixture of swamps, mature and regenerating forests, and exhibited variation in selection across scale, season, and reproductive status. We suggest that differences in space use and habitat selection between Tensas and Deltic are at least partially due to habitat differences at the landscape (i.e., amount of forested habitat) and patch (i.e., food availability) scales. Our results contribute to the understanding of factors that influence space use and habitat selection by black bears and provide specific information on habitat types selected by Louisiana black bears to agencies involved in habitat protection and restoration for this threatened subspecies.     

First parturition of tigers in a semi-arid habitat,western India

Long-term data of large felids is important to understand their reproductive biology and behavior for effective conservation planning. We used camera trap data and direct sightings from 2005 to 2013 to estimate the age of the first parturition of Bengal tigers (Panthera tigris) in a semi-arid habitat in India. We monitored 11 females in the Ranthambhore Tiger Reserve (RTR) from when they were 2–6 months old. The mean age at first reproduction (impregnation leading to cubs) was 51.3?±?(SE) 4.5 months. The tiger population in RTR is an important source population and genetic pool in the western most distribution of tiger. Thus, continuous monitoring of tiger populations is important to develop an understanding of reproductive biology.     

Predicting the potential distribution of invasive ring-necked parakeets Psittacula krameri in northern Belgium using an ecological niche modelling approach

The threat to biodiversity due to invasive alien species is considered second only to that of habitat loss. Given the large number of species that are currently invading ecosystems all over the world, we need to distinguish invaders with minor effects from those with large effects in order to prioritize management efforts. Ecological niche models can be used to predict the potential distribution of an invasive species from occurrence records and environmental data layers. We used the Ecological Niche Factor Analysis (ENFA), a presence-only predictive modelling approach, to describe the invasive ring-necked parakeets’ realized niche and to identify areas suitable for the parakeet in northern Belgium. ENFA proved to be a robust and reliable modelling technique, able to gauge the ecological requirements of an invasive species without the need to include historical information on the starting point of the invasion. ENFA shows that the parakeets tend to occupy relatively rare habitats compared to the main environmental conditions in northern Belgium, although they show some tolerance for environmental conditions inside parks and forests. The general distribution of the ring-necked parakeet is governed primarily by the amount of older forest patches, parks and built-up area in the landscape—reflecting the parakeets’ need for suitable nesting cavities and its reliance upon urban areas to forage. Our resulting habitat suitability maps show that the parakeets have ample room to further increase their range in northern Belgium. Our results indicate some concern for increased competition between parakeets and the nuthatches, native cavity nesters known to suffer from competition with parakeets, as some regions known as nuthatch strongholds are highly likely to be invaded by the parakeets.     

Age-related patterns of reproductive success in house sparrows Passer domesticus

A common life history pattern in many organisms is that reproductive success increases with age. We report a similar pattern in house sparrows Passer domesticus , older individuals performed better than yearlings for most measures of reproductive success. Older males and females began breeding earlier in a given season and fledged more young than their yearling counterparts. Individual males also fledged more young in their second breeding season than they did in their first, but individual females did not show consistent improvement in reproductive success from year one to two. A path analysis indicated that age in both sexes acted primarily through the timing of breeding; earlier nesters laid more eggs and hence fledged more young but did not have more nesting attempts. We tested whether the increased reproductive success with age arose from high quality individuals surviving to be older (selection hypothesis). In contrast to the main prediction of this hypothesis that reproductive success and survival should be positively related, we found that survival from one year of age to two years of age was negatively related to reproductive success in the first year for males and females combined. Additionally, individuals that survived to breed as two-year-olds did not differ in total young fledged in their first year from those that did not survive to their second season of breeding. Our results indicate that fledgling production increases with age due to improvements in timing of breeding, particularly in females, and not because of the loss of poor breeders or increased output. Mechanisms producing age-related differences in timing of breeding warrant further study.     

Hierarchical habitat selection by barren-ground grizzly bears in the central Canadian Arctic

Using resource selection functions, we examined habitat selection patterns of barren-ground grizzly bears (Ursus arctos) in the central Canadian Arctic among and within home ranges. There was no difference between the sexes with regard to habitat selection patterns at the home range level (Wilks' 5, approx. F_11,11=1.27, P=0.37). Bear home ranges contain more esker habitat, tussock/hummock successional tundra, lichen veneer, birch seep, and tall shrub riparian areas relative to the proportional availability of habitats in the study area. We observed differences in habitat selection within home ranges among levels of sex/reproductive status (Wilks' 5, approx. F_20,412=3.32, P<0.001) and by season (Wilks' 5, approx. F_30,605=2.71, P<0.001). Eskers and tall shrub riparian zones were the habitats most preferred by bears throughout the year. Tussock/hummock successional tundra was also favored by males at varying times during the year and lichen veneers were favored in spring and autumn by most bears. Females with cubs tended to avoid the highest ranked habitat for males throughout the year. This pattern of habitat selection was not observed for females without accompanying young. Results of this study underline the importance of scale dependence in habitat selection. Failure to view habitat selection as a hierarchical process may result in a narrow and possibly misleading notion of habitat selection patterns.     

Density and population structure of the jaguar (<Emphasis Type="Italic">Panthera onca</Emphasis>) in a protected area of Los Llanos,Venezuela, from 1 year of camera trap monitoring

Density is crucial for understanding large carnivore ecology and conservation, but estimating it has proven methodologically difficult. We conducted 1 year of camera trapping to estimate jaguar (Panthera onca) density and population structure in the Los Llanos region of Venezuela on the Hato Piñero ranch, where hunting is prohibited and livestock are excluded from half of ranch lands. We identified 42 different jaguars and determined their sex, age class, and reproductive status. We estimated adult jaguar densities with spatial capture-recapture models, using sex/reproductive state and session as covariates. Models without temporal variation received more support than models that allowed variation between sessions. Males, reproductive females, and nonreproductive females differed in their density, baseline detectability, and movement. The best estimate of total adult jaguar population density was 4.44 individuals/100 km². Based on reproductive female density and mean number of offspring per female, we estimated cub density at 3.23 individuals/100 km² and an overall density of 7.67 jaguars/100 km². Estimated jaguar population structure was 21% males, 11% nonreproductive females, 26% reproductive females, and 42% cubs. We conclude that extending the sampling period to 1 year increases the detectability of females and cubs and makes density estimates more robust as compared to the more common short studies. Our results demonstrate that the Venezuelan Llanos represent important jaguar habitat, and further, they emphasize the importance of protected areas and hunting restrictions for carnivore conservation.     

Assessing <Emphasis Type="Italic">Octopus vulgaris</Emphasis> distribution using presence-only model methods

The distribution of the common octopus (Octopus vulgaris) in the Mediterranean and Eastern Atlantic waters is evaluated using two presence-only analyses: The maximum entropy model (Maxent) and the ecological niche factor analysis (ENFA). Maxent predicts those geographical areas that satisfy the environmental or abiotic requirements of a species while ENFA explores the niche and habitat preferences of O. vulgaris. The analyses were implemented recovering the spatial information from 213 octopus presence data collected from surveys and bibliographical records. Together, these analyses provided reasonable estimates of the species distribution and the octopus habitat. Among the gathered set of explanatory environmental variables, sea bottom temperature, sea bottom salinity, surface dissolved oxygen and sea surface chlorophyll-α appear as the main variables involved in O. vulgaris distribution. These results were confronted with the available literature.     

The Influence of Age on Male Mate‐Searching Behaviour in Thornbug Treehoppers

One prediction from life‐history theory is that males should increase investment in reproductive effort as they age because the opportunity for future reproductive events declines. However, older males may not be able to increase their reproductive effort if condition declines with age. The effect of age‐related changes in condition may be especially important for energetically costly activities such as moving within and between habitat patches while searching for mates. Although such searching is a component of many mating systems, the relationship between age and active mate searching has not been investigated. We investigated whether mate‐searching effort increased with age in the thornbug treehopper, Umbonia crassicornis (Hemiptera: Membracidae). In this species, males search for females using a ‘fly‐call‐walk’ strategy consisting of three phases: (1) flying from one plant to another; (2) walking and signalling while on a plant; and (3) close‐range courtship of encountered females. We measured several aspects of mate‐searching behaviour over the month‐long period of a male’s reproductive lifetime. Over the relevant period of male sexual activity (19–33 d), male condition remained stable. However, older males (25–33 d) did not search more actively than younger males as predicted; instead, younger males (19 d) had greater plant‐to‐plant flight activity and found females faster. Within‐plant walking rates and courtship duration did not differ among age classes. These results suggest that thornbug males may be investing so heavily in mate searching at younger ages that they are unable to increase investment in searching effort when they get older. As a result, older males are likely to be at a competitive disadvantage when active searching is required to locate sparsely distributed females.     

The ontogeny of postmaturation resource allocation in turtles

Resource-allocation decisions vary with life-history strategy, and growing evidence suggests that long-lived endothermic vertebrates direct resources toward growth and self-maintenance when young, increasing allocation toward reproductive effort over time. Few studies have tracked the ontogeny of resource allocation (energy, steroid hormones, etc.) in long-lived ectothermic vertebrates, limiting our understanding of the generality of life-history strategies among vertebrates. We investigated how reproductively mature female painted turtles (Chrysemys picta) from two distinct age classes allocated resources over a 4-yr period and whether resource-allocation patterns varied with nesting experience. We examined age-related variation in body size, egg mass, reproductive frequency, and yolk steroids and report that younger females were smaller and allocated fewer resources to reproduction than did older females. Testosterone levels were higher in eggs from younger females, whereas eggs from second (seasonal) clutches contained higher concentrations of progesterone and estradiol. These allocation patterns resulted in older, larger females laying larger eggs and producing second clutches more frequently than their younger counterparts. We conclude that resource-allocation patterns do vary with age in a long-lived ectotherm.     

Role-Reversed Nepotism Among Cubs and Sires in the Spotted Hyena (Crocuta crocuta)

Nepotism is widespread among parents and offspring, and is typically associated with fundamental asymmetries; parents can usually do more to help their offspring than vice versa, and the cost of a given nepotistic behavior is usually less for parents than for offspring. This may not be so among spotted hyenas (Crocuta crocuta); sires never achieve social rank equal to that of their cubs, to whom they provide no obvious paternal care. Nepotism by sires towards their cubs is thus socially constrained. However, the higher social status of cubs might offset cubs’ developmental disadvantages and permit nepotistic treatment of sires by offspring. We therefore sought to determine whether the interactions among spotted hyena cubs and adult males were influenced by kinship, using long‐term observations from one clan of spotted hyenas. Sires exhibited no overt nepotism towards their cubs in most behaviors, but they did associate more closely with their daughters than with unrelated control females. Sires also associated more closely with their daughters than with their sons, with whom they associated no more closely than they did with unrelated control males. Cubs favored their sires by directing less intense aggression towards them than towards unrelated control males. Cubs of both sexes also associated more closely with their sires than with control males after the cubs were independent of the clan's communal den. Although sires evidently recognize their offspring, paternal nepotism by sires towards their cubs is weaker than filial nepotism by cubs towards their sires. The small size and immaturity of cubs thus appear to be weaker constraints on nepotism than is the low social status of their sires.     

Spatial pattern in a troop of yellow baboons (Papio cynocephalus) in Tanzania

Observations of adult and subadult yellow baboons (Papio cynocephalus) revealed class differences in spatial pattern both during movement and when at rest. males tended to spend slightly more time on the periphery than did females, and travelled more at the front. Positions of males reflected their association with females: younger and older adults were more peripheral, while prime adults were focal. Positions of adult females reflected their reproductive state: mothers of young infants were gregarious, but pregnant females were peripheral and travelled at the sides. Lower-ranking females were also more peripheral. These patterns are discussed in relation to male and female reproductive differences as well as to predation.     

Behavioural Differences between Single Scandinavian Brown Bears (Ursus arctos) and Females with Dependent Young When Experimentally Approached by Humans

Carnivore-human encounters that result in human injury present a conservation and management challenge and it is therefore important to understand under what conditions such incidents occur. Females with cubs are often involved when humans are injured by brown bears Ursus arctos. In Scandinavia, this is particularly true for unarmed recreational forest users. Our aim was to document behavioural differences between single bears and females with cubs in order to develop recommendations to minimize the risk of injuries to recreational forest users. We documented the reactions of GPS-collared females with cubs and single brown bears to experimental approaches by humans to 50 m from the bear on 42 and 108 occasions, respectively. The majority of females with cubs (95%) and single bears (89%) left when approached. Bears that left were passed at shorter distances and were in more open areas than those that stayed. Both groups had similar flight initiation distances, which were longer for bears that were active at the time of the disturbance. Females with cubs selected more open habitat than single bears, also for the new site they selected following disturbance. Females with cubs, particularly active females with cubs of the year, moved greater distances and spent more time active following the approach. Females with cubs and single bears were seen or heard in 26% and 14% of the approaches, respectively. None of the bears displayed any aggressive behaviour during the approaches. Females with cubs selected more open habitat, perhaps predisposing them to encountering people that are not involved in hunting activities, which might be the primary explanation why females with cubs are most frequently involved when unarmed people are injured by bears in Scandinavia. To mitigate injury risks, one must consider factors that bring bears closer to human activity in the first place.     

Serengeti real estate: density vs. fitness-based indicators of lion habitat quality

Habitat quality is typically inferred by assuming a direct relationship between consumer density and resource abundance, although it has been suggested that consumer fitness may be a more accurate measure of habitat quality. We examined density vs. fitness-based measures of habitat quality for lions in the Serengeti National Park, Tanzania. A 40-year average of female reproductive success (yearling cubs per female) was best explained by proximity to river confluences, whereas patterns of productivity (yearling cubs per km²) and adult female density (individuals per km²) were associated with more general measures of habitat quality and areas of shelter in poor habitat. This suggests that density may not accurately distinguish between high-quality 'source' areas and low-quality sites that merely provide refuges for effectively non-reproductive individuals. Our results indicate that density may be a misleading indicator of real estate value, particularly for populations that do not conform to an ideal free distribution.     

Reproductive biology and ecology of female polar bears (Ursus maritimus)

Data on age-specific natality rates, litter size, interbirth interval, age of first reproduction, reproductive senescence, age of weaning and cub survival were determined for a free-ranging population of polar bears inhabiting Hudson Bay, Canada, near the southern limit of the species range. Serum progesterone levels were also determined for females at different stages of their reproductive cycle to provide corroborative support for the reproductive parameters described. Animals were live captured using immobilizing drugs and each animal uniquely marked for future identification. First parturition occurred at four or five years of age and the age-specific natality rate increased with age until approximately 20 years, after which it dropped markedly. At least 40% of adult females displayed two-year interbirth intervals and 55% of cubs in their second year were independent of their mother. Mean size of cub litters in spring was 1.9 and 13% of litters had three or more cubs. The natality rate for 5–20-year-old females was estimated as 0.9, higher than that reported for any more northerly polar bear populations where two-year interbirth intervals are rare, fewer than 5% of yearling cubs are weaned and triplet litters occur with less than 1% frequency. Cub mortality was initially high and declined with age. Although cubs in western Hudson Bay were weaned at a younger age and a lighter weight than their counterparts in more northern populations, cub mortality rates were similar. The reason for the marked differences in reproductive parameters in the western Hudson Bay population is not known. We speculate that sea-ice conditions may be sufficiently different to allow weaned bears at a lighter body weight to hunt seals more successfully there than further north.     

Adaptive female choice for middle-aged mates in a lekking sandfly

Most theoretical models of age-related mate choice predict that females should prefer older males because they have proven survival ability. An alternative view is that older males represent inferior mates because of negative genetic correlations between early and late fitness components, or because older males have traded off longevity against other fitness components, have accumulated deleterious germ-line mutations, or are less well adapted to current conditions than more recently born individuals. While numerous studies have reported female choice for older males, few have explicitly examined the fitness consequences of such a preference. We present evidence from a lekking sandfly, Lutzomyia longipalpis, showing that choosy females discriminate against older males and gain a fitness benefit from their choice. When permitted free choice from an aggregation consisting of males aged zero to two days (young), four to six days (middle-aged) and eight to ten days (old), females preferentially mated with middle-aged males, but all measures of female reproductive success were independent of male age. In contrast, when a second set of females was randomly assigned single virgin males of known age, the eggs of those paired to old mates exhibited lower hatching success than the eggs of females mated to young or middle-aged males. These results suggest that females avoid mating with older males because they represent poorer quality mates. Age-related differences in male quality may have a genetic basis, but could equally well arise through a phenotypic decline in sperm quality or sperm transfer ability with male age. The lack of evidence of female discrimination against older males from other studies may be because these did not explore the reproductive success of the full age range of males.     

Landscape-scale evaluation of habitat uses by sympatric mammals foraging for bark and buds in a heavy snowfall area of northern Japan

This study focused on influences of three different growing populations of mammals foraging for bark and buds—Japanese serows (Capricornis crispus), Tohoku hares (Lepus brachyurus angustidens), and Japanese macaques (Macaca fuscata)—on a cool-temperate forest in northern Japan. To acquire the initial knowledge to facilitate ecological monitoring of the influences, we evaluated habitat uses by each species on a landscape scale by using ecological niche factor analysis (ENFA), and identified commonalities and differences among the uses by using discriminant analysis (DA). Within a 50-km² area, we selected five 5-km transects and recorded tracks of each mammal on the snow surface along transects during the month of March in 2008 and 2009. The track data were used as the proof of species presence for entry into ENFA and DA. Marginality and specialization of species habitat use, computed by ENFA, indicated that the macaque habitat represented the most heterogeneous distribution and the lowest environmental tolerance, which was strictly limited by altitude and terrain conditions to prevent exposure to severe cold climate conditions. This refuge selection by macaques resulted in habitat segregation from serows, although it appears that both mammals prefer the same young broadleaf forests. Meanwhile, the serow was observed to be distinctly inclined to use mountainous forests and to avoid artificial environments. As a result, its habitat could be differentiated from that of the hare, as confirmed by coefficients of a discriminant factor. These findings indicate that the impact of three different mammals foraging on the native vegetation could be quite restrictive.     

Using presence‐only modelling to predict Asian elephant habitat use in a tropical forest landscape: implications for conservation

Aim Asian elephants, Elephas maximus, are threatened throughout their range by a combination of logging, large scale forest conversion and conflict with humans. We investigate which environmental factors, both biotic and abiotic, constrain the current distribution of elephants. A spatially explicit habitat model is constructed to find core areas for conservation and to assess current threats. Location Ulu Masen Ecosystem in the province of Nanggroe Aceh Darussalam on the island of Sumatra, Indonesia. Methods A stratified survey was conducted at 12 sites (300 transects) to establish the presence of elephants. Presence records formed the basis to model potential habitat use. Ecological niche factor analysis (ENFA) is used to describe their niche and to identify key factors shaping elephant distribution. An initial niche model was constructed to describe elephant niche structure, and a second model focused on identifying core areas only. To assess the threat of habitat encroachment, overlap between the elephants’ optimal niche and the occurrence of forest encroachment is computed. Results Elephants were recorded throughout the study area from sea level to 1600 m a.s.l. The results show that the elephant niche and consequently habitat use markedly deviates from the available environment. Elephant presence was positively related to forest cover and vegetation productivity, and elephants were largely confined to valleys. A spatially explicit model showed that elephants mainly utilize forest edges. Forest encroachment occurs throughout the elephants range and was found within 80% of the elephants’ ecological niche. Main conclusions In contrast to general opinion, elephant distribution proved to be weakly constrained by altitude, possibly because of movement routes running through mountainous areas. Elephants were often found to occupy habitat patches in and near human‐dominated areas. This pattern is believed to reflect the displacement of elephants from their former habitat.     

Reproductive characteristics of female Bengal tigers,in Ranthambhore Tiger Reserve,India

Reproductive characteristics of tigers (Panthera tigris) are important to understand population viability. We studied the reproductive parameters of female Bengal tigers (P. t. tigris) in a dry, tropical, deciduous habitat in Ranthambhore Tiger Reserve (RTR), western India, from April 2005 to March 2010. We monitored tigers by direct observation and with cameras placed throughout their habitat. The potential breeding population included 13 adult females. The average age at first reproduction was 3.3 years; 34 cubs were born during the study period (6.2?±?0.82 per year). Sixty-six percent of the births occurred between October and December. Mean litter size was 2.26?±?0.52 (n?=?13, range?=?1–3). The sex ratio of 32 cubs was 1.29 M:1.00 F. The survival rate of cubs (<12 months) was 85 % (95 % CI?=?0.68–0.94), whereas that of juveniles (12–24 months), and subadults (24–36 months) was 79 % (95 % CI?=?0.61–0.91). All breeding females were >3 years old. Only 2 of the 13 females reproduced twice during the 5 years of the study. The birth interval was 33.4?±?3.7 months (range 24–65 months). The mean reproductive rate was 0.59?±?0.23 cubs/female/year. Our study indicates that tiger populations can grow rapidly if the habitat provides adequate protection, an adequate population of prey, and minimal to no poaching.