Covariates affecting spatial variability in bison travel behavior in Yellowstone National Park.

Understanding mechanisms influencing the movement paths of animals is essential for comprehending behavior and accurately predicting use of travel corridors. In Yellowstone National Park (USA), the effects of roads and winter road grooming on bison (Bison bison) travel routes and spatial dynamics have been debated for more than a decade. However, no rigorous studies have been conducted on bison spatial movement patterns. We collected 121 380 locations from 14 female bison with GPS collars in central Yellowstone to examine how topography, habitat type, roads, and elevation affected the probability of bison travel year-round. We also conducted daily winter bison road use surveys (2003-2005) to quantify how topography and habitat type influenced spatial variability in the amount of bison road travel. Using model comparison techniques, we found the probability of bison travel and spatial distribution of travel locations were affected by multiple topographic and habitat type attributes including slope, landscape roughness, habitat type, elevation, and distances to streams, foraging areas, forested habitats, and roads. Streams were the most influential natural landscape feature affecting bison travel, and results suggest the bison travel network throughout central Yellowstone is spatially defined largely by the presence of streams that connect foraging areas. Also, the probability of bison travel was higher in regions of variable topography that constrain movements, such as in canyons. Pronounced travel corridors existed both in close association with roads and distant from any roads, and results indicate that roads may facilitate bison travel in certain areas. However, our findings suggest that many road segments used as travel corridors are overlaid upon natural travel pathways because road segments receiving high amounts of bison travel had similar landscape features as natural travel corridors. We suggest that most spatial patterns in bison road travel are a manifestation of general spatial travel trends. Our research offers novel insights into bison spatial dynamics and provides conceptual and analytical frameworks for examining movement patterns of other species.     

Behavioral responses of bison and elk in Yellowstone to snowmobiles and snow coaches.

Managers of public lands are charged with protecting some of our most important natural resources and ecosystems, while providing for their use and enjoyment by visitors. Almost one million visitors entered Yellowstone National Park by motorized means on snowmobiles (87%) or snow coaches (13%) during 1992-2003. Most vehicles toured the central portion of the park where bison (Bison bison) and elk (Cervus elaphus) concentrate in geothermal areas. We sampled >6500 interactions between groups of these species and groups of snowmobiles and snow coaches (collectively, OSV, over-snow vehicles) during five winters (1999-2000, 2002-2004). Multinomial logits models were used to identify conditions leading to behavioral responses. Elk responded three times as often (52%) as bison (19%) during interactions with groups of snowmobiles and snow coaches due to increased vigilance responses (elk, 44%; bison, 10%). However, the frequency of higher-intensity movement responses by bison and elk were similar (6-7% travel, 1-2% flight, <1% defense) and relatively low compared to other studies of ungulates and snowmobile disturbance. The likelihood of active responses by bison and elk increased significantly if animals were on or near roads, groups were smaller, or humans approached. The likelihood of an active response by bison decreased within winters having the largest visitation, suggesting some habituation to snowmobiles and snow coaches. There was no evidence that snowmobile use during the past 35 years affected the population dynamics or demography of bison or elk. Thus, we suggest that regulations restricting levels and travel routes of over-snow vehicles (OSVs) were effective at reducing disturbances to bison and elk below a level that would cause measurable fitness effects. We recommend park managers consider maintaining OSV traffic levels at or below those observed during our study. Regardless, differing interpretations of the behavioral and physiological response data will continue to exist because of the diverse values and beliefs of the many constituencies of Yellowstone.     

Temporal variability of deep-sea zooplankton in the Arabian Sea

Mesozooplankton samples from two stations in the Arabian Sea (WAST, 4,050 m, 16°15′N, 60°20′E; CAST, 3,950 m, 14°30′N, 64°30′E) were collected from the surface down to 20 m above bottom during three monsoon periods: the autumn intermonsoon in October 1995, the spring intermonsoon in April 1997, and the NE monsoon in February 1998. The main goal of this study is to enhance our knowledge on the effect of spatial and temporal differences in primary production and particle flux rates on the abundance and distribution of mesozooplankton, with special attention to the deep sea. Literature data indicate episodically high rates of primary production and particle flux in the region during the SW monsoon and the autumn intermonsoon. Set in this context, the zooplankton showed an in-phase coupling in biomass and abundance with the primary production in the surface 150 m. In the mesopelagic realm (150–1,050 m), the seasonal coupling was less clear. In the bathypelagic zone, below 1,050 m, there was no evidence of in-phase coupling, though temporal differences in the distribution of zooplankton abundance and biomass with depth between seasons could be shown by an analysis of covariance and an a posteriori test. The results suggest that the bathypelagic community responds to increased particle flux rates, but with longer time gaps than in the epipelagic zone. This is probably due to longer development and response times of zooplankton in the cold, deep-water environment, independent of possible lateral advection processes.     

Temporal variability of forest fires in eastern Amazonia

Widespread occurrence of fires in Amazonian forests is known to be associated with extreme droughts, but historical data on the location and extent of forest fires are fundamental to determining the degree to which climate conditions and droughts have affected fire occurrence in the region. We used remote sensing to derive a 23-year time series of annual landscape-level burn scars in a fragmented forest of the eastern Amazon. Our burn scar data set is based on a new routine developed for the Carnegie Landsat Analysis System (CLAS), called CLAS-BURN, to calculate a physically based burn scar index (BSI) with an overall accuracy of 93% (Kappa coefficient 0.84). This index uses sub-pixel cover fractions of photosynthetic vegetation, non-photosynthetic vegetation, and shade/burn scar spectral end members. From 23 consecutive Landsat images processed with the CLAS-BURN algorithm, we quantified fire frequencies, the variation in fire return intervals, and rates of conversion of burned forest to other land uses in a 32 400 km2 area. From 1983 to 2007, 15% of the forest burned; 38% of these burned forests were subsequently deforested, representing 19% of the area cleared during the period of observation. While 72% of the fire-affected forest burned only once during the 23-year study period, 20% burned twice, 6% burned three times, and 2% burned four or more times, with the maximum of seven times. These frequencies suggest that the current fire return interval is 5-11 times more frequent than the estimated natural fire regime. Our results also quantify the substantial influence of climate and extreme droughts caused by a strong El Ni?o Southern Oscillation (ENSO) on the extent and likelihood of returning forest fires mainly in fragmented landscapes. These results are an important indication of the role of future warmer climate and deforestation in enhancing emissions from more frequently burned forests in the Amazon.     

Temporal patterns of shell-gape in Mytilus edulis

Hitherto published evidence of the presence or absence of endogenous activity rhythms in bivalve molluscs is equivocal. Mytilus edulis L. were collected from a North Wales (UK) estuary in 1985, and shell-gaping was investigated in individual mussels under constant conditions in the laboratory. Results suggest that there is no endogenous circatidal rhythm of shell-gaping in this species, This is consistent with the view that, unlike mobile species, sessile intertidal species are much more likely to exhibit exogenous rather than endogenous responses to tidal fluctuations. There is some evidence of weak circadian rhythmicity of shell-gaping in M. edulis, with greater duration of shell-closure during hours of expected daylight. Such behaviour could represent an adaptational defence against visually-feeding predators.     

Temporal patterns in the vibratory courtship signals of the wandering spider Cupiennius salei Keys.

Summary The vibratory courtship signal generated by abdominal and leg movements of male Cupiennius salei on plants consists of impulses, which in groups of about ten form syllables. A syllable together with the preceding silent pause is called a sequence. Up to 50 sequences chained together form a series. A series together with the preceding silent period is called an interseries. A complete courtship is built up from a row of successive interseries. An additional signal produced by palpal drumming only occurs between syllables. Females respond to ca. 40% of the male signal series with a single irregular signal produced by leg oscillations and lasting for 113–1790 ms. Their response reduces the duration of the male interseries significantly from an average of 27.9 s to 20.6 s. The female signal follows a male signal series within a narrow time frame of 0.89±0.53 s. Analysis over long periods of time and of complete courtships showed the male signalling to be highly structured in the time domain and to contain many cues potentially usable by the female for both such precise timing and conspecific recognition: (1) On average, the interval between the last two syllables is significantly longer than the preceding ones. (2) During the course of a series, syllable durations increase nearly linearly from 93 ms to 123 ms. (3) The twelfth sequence (T-12) prior to the final one (T) is outstanding since here, roughly 4 s before the end of a series, the duration of sequences and pauses, the signal amplitudes and the occurrence of palpal drumming abruptly start to increase. Also, the frequencies contained in the syllables shift to higher values. (4) Between sequences T-4 and T-6 pause duration and sequence duration reach their minimum whereas the values for signal acceleration and the occurrence of palpal drumming are highest.     

Temporal patterns of populations in a warming world: a modelling framework

In this paper, we present an approach for describing the environmentally induced temporal pattern of structured populations by partial integro-differential equations. Populations are structured according to size or stage. Growth, energy allocation and stage transitions are affected by environmental conditions of which temperature, photoperiod, water depth and food supply were taken into account. The resulting modelling framework was applied to describe, analyse and predict alterations in populations with continuous development, populations with distinct state structures and interacting populations. Our exemplary applications consider populations of freshwater Amphipoda, Isopoda and Odonata. The model was capable of simulating life cycle alterations in dependence on temperature in interaction with other environmental factors: (1) population dynamics, (2) seasonal regulation, (3) water depth-dependent dispersal, (4) intraguild predation and (5) consumer-resource dynamics.     

Determining patterns of variability in ecological communities: time lag analysis revisited

All ecological communities experience change over time. One method to quantify temporal variation in the patterns of relative abundance of communities is time lag analysis (TLA). It uses a distance-based approach to study temporal community dynamics by regressing community dissimilarity over increasing time lags (one-unit lags, two-unit lags, three-unit lags). Here, we suggest some modifications to the method and revaluate its potential for detecting patterns of community change. We apply Hellinger distance based TLA to artificial data simulating communities with different levels of directional and stochastic dynamics and analyse their effects on the slope and its statistical significance. We conclude that statistical significance of the TLA slope (obtained by a Monte Carlo permutation procedure) is a valid criterion to discriminate between (i) communities with directional change in species composition, regardless whether it is caused by directional abundance change of the species or by stochastic change according to a Markov process, and (ii) communities that are composed of species with population sizes oscillating around a constant mean or communities whose species abundances are governed by a white noise process. TLA slopes range between 0.02 and 0.25, depending on the proportions of species with different dynamics; higher proportions of species with constant means imply shallower slopes; and higher proportions of species with stochastic dynamics or directional change imply steeper slopes. These values are broadly in line with TLA slopes from real world data. Caution must be exercised when TLA is used for the comparison of community time series with different lengths since the slope depends on time series length and tends to decrease non-linearly with it.     

Search or relax: the case of bachelor wood bison

Summary Males of gregarious species of mammals often do not associate with female groups during the breeding season. Such males could leave groups either to search for new breeding opportunities, or to restore body condition before rejoining new female groups. To investigate these two competing hypotheses, we studied the chronology of male wood bison (Bison bison athabascae) associating with females, and their aggressive and sexual behaviour over two breeding seasons. With increasing age, males were more often solitary, were more involved in aggressive interactions, investigated the genitals of females more frequently, and tended females more often. Group size did not affect whether or not males joined or left groups. The presence of oestrous females in a group did not affect the number of bulls in that group, and did not affect the joining or leaving of males. Older, dominant males did not exclude younger ones from female groups. Males observed outside female groups spent relatively more time feeding and resting than bulls in groups. Changes in group composition experienced by a male often involved the joining or splitting of female groups, as opposed to actual transfer of males between groups. Our results suggest that males leave female groups in order to recover, not for the purpose of searching for new female groups. We argue that due to the fluidity of wood bison groups, it is possible for a male to encounter new females even when remaining in a female group. Correspondence to: P.E. Komers     

Genetic variability in the marine gastropods Patella coerulea and Patella aspera: patterns and problems

Allozymic variation encoded by 15 gene loci was compared and contrasted in naturally coexisting populations of two marine gastropods (Patella coerulea and Patella aspera) collected along the rocky beaches of the Mediterranean sea of Israel in June, 1979. The genetic variability did not differ consistently or significantly between the two species, but in all the populations studied, P. aspera was more heterozygous than P. coerulea. These results are discussed in terms of the potential ecological factors affecting this genetic differentiation.     

Foraging patterns of yellow-bellied marmosts: role of kinship and individual variability

Summary Two colonies of yellow-bellied marmots (Marmota flaviventris) at an elevation of 2900 m in Colorado were studied to elucidate the role of various behavioral and ecological factors as determinants of spatial foraging patterns. The locations of known individuals were periodically recorded. These locality data were plotted as three-dimensional block diagrams, the peak heights representing the frequency of observation. Predation risk and vegetation distribution influenced the location of foraging areas; kinship was an important factor in the determination of the amount of foraging area shared between individual marmots. Spatial overlap tended to be greater among close kin, but this was modified by individual behavioral characteristics, reproductive state, the existence of separate burrow systems within a colony, and the age of the animal. Mothers and juveniles, and littermates as young and resident yearlings, had nearly identical foraging areas.     

Temporal variability of zooplankton biomass (ATP content and dry weight) in the St. Lawrence Estuary: Advective phenomena during neap tide

In order to assess the temporal variability of living zooplankton in a tidal estuary, the ATP content and dry weight of mixed zooplankton populations (mainly copepods) were measured during a period of 175 h at an anchor station in the Upper St. Lawrence Estuary. Vertical tows were made every 30 min. Hourly vertical profiles of the current speed and direction, temperature and salinity were also obtained during the experiment. A strong tidal influence was found in all series. Maxima and minima of the ATP content (living biomass) and the dry weight (total biomass) were correlated with low and high water slacks. The serial autocorrelation and cross-correlation showed, in both series a 12 to 13-h cycle, and the ATP:dry weight ratio showed a significant 24-h cycle. The cross-correlation with the Kendall τ was used to detect the relationship between biological components and physical indices (stratification and Ri). It is suggested that the proportion of living zooplankton biomass in the Upper St. Lawrence Estuary is most likely the result of a combination of diurnal migration and longitudinal advection.     

Geographical patterns of variability at allozyme loci in the European oyster Ostrea edulis

The variability of 14 enzyme-coding genes has been analysed in samples from 19 populations of the oyster Ostrea edulis L., collected along the Atlantic and Mediterranean coasts of Europe. We found an abundance of clines, which appeared at 8 loci, including the most polymorphic (AP-2 ^*, ARK ^*, EST-4 ^*, MDH-2 ^*, ME-1 ^*, 6PGH ^*, PGI ^* and PGM ^*). Another 6 loci (ALDH ^*, EST-3 ^*, EST-5 ^*, IDH-2 ^*, MDH-1 ^*, ME-2 ^*) exhibited V-shaped patterns of gene-frequency variation, with clines at one or both sides of the Straits of Gibraltar. The observation of coincident clines at many loci can be explained by a model of secondary intergradation. The geographical location of the midpoints of the clines and V-shaped patterns suggests the existence of two ancient Atlantic and Mediterranean oyster stocks which became differentiated in allopatry and subsequently merged. Clines observed along Atlantic and/or Mediterranean coasts at the loci with V-shaped patterns must have arisen independently. The large heterogeneity observed in the levels of gene differentiation (G _ST ) across loci (G _ST ranged from 0.008 to 0.290) and important differences in estimates of gene flow obtained by different methods suggest that the populations of O. edulis are not in genetic equilibrium. Lack of population equilibrium can be due to natural selection and/or restrictions to gene flow. The average among-population variability was higher than in other oyster species that do not show incubatory habits, and represented 8.8% of the total heterozygosity. Levels of intrapopulation variability were lowest in populations from the North Atlantic, suggesting low population sizes in that area.     

Effects of management and climate on elk brucellosis in the Greater Yellowstone Ecosystem.

Every winter, government agencies feed approximately 6000 metric tons (6 x 10(6) kg) of hay to elk in the southern Greater Yellowstone Ecosystem (GYE) to limit transmission of Brucella abortus, the causative agent of brucellosis, from elk to cattle. Supplemental feeding, however, is likely to increase the transmission of brucellosis in elk, and may be affected by climatic factors, such as snowpack. We assessed these possibilities using snowpack and feeding data from 1952 to 2006 and disease testing data from 1993 to 2006. Brucellosis seroprevalence was strongly correlated with the timing of the feeding season. Longer feeding seasons were associated with higher seroprevalence, but elk population size and density had only minor effects. In other words, the duration of host aggregation and whether it coincided with peak transmission periods was more important than just the host population size. Accurate modeling of disease transmission depends upon incorporating information on how host contact rates fluctuate over time relative to peak transmission periods. We also found that supplemental feeding seasons lasted longer during years with deeper snowpack. Therefore, milder winters and/or management strategies that reduce the length of the feeding season may reduce the seroprevalence of brucellosis in the elk populations of the southern GYE.     

Biodiversity consequences of alternative future land use scenarios in Greater Yellowstone.

Land use is rapidly expanding in the Greater Yellowstone Ecosystem, primarily from growth in the number of rural homes. There is a need to project possible future land use and assess impacts on nature reserves as a guide to future management. We assessed the potential biodiversity impacts of alternative future land use scenarios in the Greater Yellowstone Ecosystem. An existing regression-based simulation model was used to project three alternative scenarios of future rural home development. The spatial patterns of forecasted development were then compared to several biodiversity response variables that included cover types, species habitats, and biodiversity indices. We identified the four biodiversity responses most at risk of exurban development, designed growth management policies to protect these areas, and tested their effectiveness in two alternative future scenarios. We found that the measured biodiversity responses, including riparian habitat, elk winter range, migration corridors, and eight other land cover, habitat, and biodiversity indices, are likely to undergo substantial conversion (between 5% and 40%) to exurban development by 2020. Future habitat conversion to exurban development outside the region's nature reserves is likely to impact wildlife populations within the reserves. Existing growth management policies will provide minimal protection to biodiversity in this region. We identified specific growth management policies, including incentives to cluster future growth near towns, that can protect "at risk" habitat types without limiting overall growth in housing.     

On the Origin of Brucellosis in Bison of Yellowstone National Park: A Review

Brucellosis caused by Brucella abortus occurs in the free-ranging bison ( Bison bison ) of Yellowstone and Wood Buffalo National Parks and in elk ( Cervus elaphus ) of the Greater Yellowstone Area. As a result of nationwide bovine brucellosis eradication programs, states and provinces proximate to the national parks are considered free of bovine brucellosis. Thus, increased attention has been focused on the wildlife within these areas as potential reservoirs for transmission to cattle. Because the national parks are mandated as natural areas, the question has been raised as to whether Brucella abortus is endogenous or exogenous to bison, particularly for Yellowstone National Park. We synthesized diverse lines of inquiry, including the evolutionary history of both bison and Brucella , wild animals as Brucella hosts, biochemical and genetic information, behavioral characteristics of host and organism, and area history to develop an evaluation of the question for the National Park Service. All lines of inquiry indicated that the organism was introduced to North America with cattle, and that the introduction into the Yellowstone bison probably was directly from cattle shortly before 1917. Fistulous withers of horses was a less likely possibility. Elk on winter feedgrounds south of Yellowstone National Park apparently acquired the disease directly from cattle. Bison presently using Grand Teton National Park probably acquired brucellosis from feedground elk.     

Temporal patterns in three-dimensional structure and activity of schools of the Atlantic silverside Menidia menidia

Temporal patterns in the structure and activity of schools of the Atlantic silverside Menidia menidia were investigated under laboratory conditions using a new computerized video technique for three-dimensional analysis. Fish were collected by seining at Middle Bridge, Pettaquamscutt River, South Kingstown, Rhode Island, USA. Approximately once a week from May to December, 1979, the behavior of a school was videotape-recorded in the laboratory at 30 min intervals from sunrise to sunset under a temperature and photoperiod regime simulating ambient field conditions. Four measurements of structure and activity were computed: (1) swimming speed (SPEED); (2) distance to nearest neighbor (NN); (3) mean direction of travel (DOT); (4) depth in the tank (DEPTH). Results demonstrated a strong seasonal rhythm in the behavior of schools, but were inconclusive in showing periodicities associated with daily, tidal or lunar cycles. NN and DEPTH were significantly different between spring, summer, fall and winter while SPEED changed significantly across 3 seasons (summer and winter were not significantly different). DOT was bimodally distributed, with peaks in spring and fall. Changes in these variables corresponded to shifts in the behavior of silversides from active, synchronized schools in spring and fall to inactive mills in summer and inactive non-schooling groups in winter. Changes in schooling behavior were only partly related to changes in temperature and photoperiod. Patterns of silverside schooling behavior correlated with their reproductive and migratory cycles and may represent an anti-predator strategy.