Interactions between density,home range behaviors,and contact rates in the Channel Island fox (Urocyon littoralis)

Many of the mechanisms underlying density‐dependent regulation of populations, including contest competition and disease spread, depend on contact among neighboring animals. Understanding how variation in population density influences the frequency of contact among neighboring animals is therefore an important aspect to understanding the mechanisms underlying, and ecological consequences of, density‐dependent regulation. However, contact rates are difficult to measure in the field and may be influenced by density through multiple pathways. This study explored how local density affects contact rates among Channel Island foxes (Urocyon littoralis) through two pathways: changes in home range size and changes in home range overlap. We tracked 40 radio‐collared foxes at four sites on San Clemente Island, California. Fox densities at the four sites ranged from 2.8 ± 1.28 to 42.8 ± 9.43 foxes/km². Higher fox densities were correlated with smaller home ranges (R² = 0.526, F_1,38 = 42.19, P < 0.001). Thirty foxes wore collars that also contained proximity loggers, which recorded the time and duration of occasions when collared foxes were within 5 m of one another. Contact rates between neighboring fox dyads were positively correlated with home range overlap (R² = 0.341, P = 0.008), but not fox density (R² = 0.012, P = 0.976). Individuals at high densities had more collared neighbors with overlapping home ranges (R² = 0.123, P = 0.026) but not an increase in the amount of contact between individual neighbors. This study was the first time contact rates were directly measured and compared to density and home range overlap. Results suggest that foxes exhibit a threshold in their degree of tolerance for neighbors, overlap is a reliable index of the amount of direct contact between island foxes, and disease transmission rates will likely scale with fox density.     

Effects of high density on spacing behaviour and reproduction in Akodon azarae: A fencing experiment

We studied the short term spacing behavioural responses of Pampean grassland mouse (Akodon azarae) with regard to population density in four 0.25 ha enclosures (two control and two experimental) in the 2011 breeding season. Based on the hypothesis that A. azarae breeding females exhibit spacing behaviour, and breeding males show a fusion spatial response, we tested the following predictions: (1) home range size and intrasexual overlap degree of females are independent of population density values; (2) at high population density, home range size of males decreases and the intrasexual home range overlap degree increases. To determine if female reproductive success decreases at high population density, we analyzed pregnancy rate, size and weight of litters, and period until fecundation in both low and high enclosure population density. We found that both males and females varied their home range size in relation to population density. Although male home ranges were always bigger than those of females in populations with high density, home range sizes of both sexes decreased. Females kept exclusive home ranges independent of density values meanwhile males decreased home range overlap in high breeding density populations. Although females produced litters of similar size in both treatments, weight of litter, pregnant rate and period until fecundation varied in relation to population density. Our results did not support the hypothesis that at high density females of A. azarae exhibit spacing behaviour neither that males exhibit a fusion spatial response.     

A mast‐seeding desert shrub regulates population dynamics and behavior of its heteromyid dispersers

Granivorous rodent populations in deserts are primarily regulated through precipitation‐driven resource pulses rather than pulses associated with mast‐seeding, a pattern more common in mesic habitats. We studied heteromyid responses to mast‐seeding in the desert shrub blackbrush (Coleogyne ramosissima), a regionally dominant species in the Mojave–Great Basin Desert transition zone. In a 5‐year study at Arches National Park, Utah, USA, we quantified spatiotemporal variation in seed resources in mast and intermast years in blackbrush‐dominated and mixed desert vegetation and measured responses of Dipodomys ordii (Ord's kangaroo rat) and Perognathus flavescens (plains pocket mouse). In blackbrush‐dominated vegetation, blackbrush seeds comprised >79% of seed production in a mast year, but 0% in the first postmast year. Kangaroo rat abundance in blackbrush‐dominated vegetation was highest in the mast year, declined sharply at the end of the first postmast summer, and then remained at low levels for 3 years. Pocket mouse abundance was not as strongly associated with blackbrush seed production. In mixed desert vegetation, kangaroo rat abundance was higher and more uniform through time. Kangaroo rats excluded the smaller pocket mice from resource‐rich patches including a pipeline disturbance and also moved their home range centers closer to this disturbance in a year of low blackbrush seed production. Home range size for kangaroo rats was unrelated to seed resource density in the mast year, but resource‐poor home ranges were larger (P < 0.001) in the first postmast year, when resources were limiting. Blackbrush seeds are higher in protein and fat but lower in carbohydrates than the more highly preferred seeds of Indian ricegrass (Achnatherum hymenoides) and have similar energy value per unit of handling time. Kangaroo rats cached seeds of these two species in similar spatial configurations, implying that they were equally valued as stored food resources. Blackbrush mast is a key resource regulating populations of kangaroo rats in this ecosystem.     

Spatial ecology of jaguars,pumas, and ocelots: a review of the state of knowledge

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Space use by giant anteaters (Myrmecophaga tridactyla) in a protected area within human‐modified landscape

1. Spatial ecology data are essential for conservation purposes, especially when extinction risk is influenced by anthropogenic actions. Space use can reveal how individuals use the habitat, how they organize in space, and which components are key resources for the species.
2. We evaluated the space use and multiscale habitat selection of giant anteaters (Myrmecophaga tridactyla), a vulnerable Neotropical mammal, in a Cerrado site within a human‐modified landscape in southeastern Brazil.
3. We used GPS transmitters to track eight anteaters in the wild. With the resulting dataset, we estimated home range and core‐area sizes and then used two overlap indexes. We assessed habitat selection by compositional analysis and analyzed events of spatio‐temporal proximity.
4. The average Brownian bridge kernel estimate of home range size was 3.41 km² (0.92–7.9). Regarding home range establishment, five individuals showed resident behavior. Males (n = 4) had larger home ranges and were more active than females (n = 4). Despite the spatial overlap of home range (above 40% in four dyads), maximum temporal space sharing was 18%. Giant anteaters were found in proximity. Habitat selection favored savanna, and exotic timber plantation was always avoided. Roads and built‐up areas were selected secondarily at the landscape level.
5. The selection of anthropogenic sites denotes behavioral plasticity regarding modified habitats. However, the high selectivity for savanna, at all levels, demonstrates a high dependence on natural habitats, which provide the necessary resources for the species. The recurrent proximity of male–to‐female anteaters may indicate reproductive behavior, which is essential for maintaining this isolated population.

     

Spatial ecology and habitat use of giraffe (Giraffa camelopardalis) in South Africa

The lack of long-term studies remains a limiting factor in understanding the home range, spatial ecology and movement of giraffes. We equipped eight giraffes with GPS satellite units and VHF capacity, which were built in to the collars for the remote collection of data on their movements and home ranges over two years on Khamab Kalahari Nature Reserve (KKNR) within the Kalahari region of South Africa. Giraffe numbers in KKNR dropped from 135 individuals to 111 in just five years, revealing the lack of knowledge about their required habitat needs, space use and diet. With over 1000 km² available for roaming within the reserve, habitat selection, principle and preferred food species played a significant role in home range size and overlap between individuals. These giraffes used an average annual home range of 206 km² (20 602 ha) as calculated by a 95% minimum convex polygon (MCP) with a standard deviation core home range calculated by a 50% MCP of 10.1 km² to satisfy their annual needs for survival and reproduction in their preferred vegetation. In the wet, hot season (summer: December–February) when food was abundant, giraffes frequented smaller areas (average 177 km²), while in the dry, cool season (winter: June to August) the mean home range size increased to approximately 245 km². Rainfall influenced spatial distribution since it determined vegetation productivity and leaf phenology. The different seasons influenced giraffe movements, while different vegetation types and season influenced their home range size. Season and food availability also influenced home range overlap between different giraffe herds. Home range overlap occurred when giraffes were forced to roam in overlapping areas during the dryer months when the winter deciduous nature of the majority of the tree species resulted in lower food availability. In winter, the overlap was approximately 31% and in autumn approximately 23%. During the wet and warmer months, overlapping was 15% in summer and 19% in spring, respectively. The percentage of time spent in different vegetation type areas was influenced by the abundance of the principal food species of that plant community. It is thus concluded that the movements of giraffes were primarily influenced by a combination of environmental factors such as season, rainfall and vegetation density.     

Home Range Size of Sympatric Squirrel Species Inhabiting a Lowland Dipterocarp Forest in Malaysia1

Home range sizes and spatial overlap of four sympatric squirrel species were investigated in a lowland dipterocarp forest in Malaysia using a radio‐tracking method. The population density of Callosciurus caniceps was highest and C. notatus was next highest, while C. nigrovittatus and Lariscus insignis were scarce. C. caniceps was larger than C. nigrovittatus and C. notatus while L. insignis was extremely small. For females, home range size was smaller in L. insignis than Callosciurus spp., which may support the body weight hypothesis: larger species have larger home ranges. Among the three Callosciurus species, female C. caniceps had the smallest home range. These differences were accounted for by habitat characteristics rather than by density or body weight; C. caniceps was dominant in bushy areas and used crowded small trees while C. notatus and C. nigrovittatus used large trees in the forest. In this study, home range size did not change seasonally; this differs from studies in temperate regions, possibly because food availability is much less variable among seasons in tropical rain forest. Home range overlap among heterospecific individuals was common but different species seemed to partition space by using different vertical levels of the forest. Consequently, the home range size and spatial overlap of sympatric squirrel species may be affected by habitat diversity in tropical rain forest.     

Intraspecific Variation in Home Range Overlap with Habitat Quality: A Comparison among Brown Bear Populations

We developed a conceptual model of spatial organization in vertebrates based upon changes in home range overlap with habitat quality. We tested the model using estimates of annual home ranges of adult females and densities for 30 populations of brown bears (Ursus arctos) in North America. We used seasonality as a surrogate of habitat quality, measured as the coefficient of variation among monthly actual evapotranspiration values for areas in which study populations were located. We calculated home range overlap for each population as the product of the average home range size for adult females and the estimated population density of adult females. Home range size varied positively with seasonality; however, home range overlap varied with seasonality in a nonlinear manner. Areas of low and high seasonality supported brown bears with considerable home range overlap, but areas of moderate seasonality supported brown bears with low home range overlap. These results are consistent with behavioural theory predicting a nonlinear relationship between food availability and territoriality.     

Home‐range size of an Andean bird: Assessing the role of physical condition

Because space‐use patterns are a key aspect of the ecology and distribution of species, identifying factors associated with variation in size of territories and home ranges has been central to studies on population ecology. Space use might vary in response to extrinsic factors like habitat quality and to intrinsic factors like physical condition and individual aggressiveness. However, the role of these factors has been poorly documented in the tropics, particularly in high‐elevation bird species. We report the home‐range size of a Neotropical Andean bird, the gray‐browed brush finch (Arremon assimilis), and evaluate the role of physical condition in explaining variation in home‐range size among individuals. We performed spot mapping to estimate the home ranges of 14 territorial males in Bogotá, Colombia, using minimum convex polygons (MCP) and 95% kernel density estimators (KDE). The mean home‐range size estimated for the 100% MCP was 0.522 ± 0.305 ha (range = 0.15–1.18 ha), whereas the 95% KDE estimation was 0.504 ± 0.471 ha (range = 0.13–1.88). We calculated the real mass index of each bird as a proxy of physical condition to assess whether individuals in better physical condition had larger home ranges. Because we found no relation between our estimations of physical condition and home‐range size, we conclude that space use in this species might depend more on ecological factors such as habitat quality or neighbor density than on individual traits. Abstract in French is available with online material.     

Inter‐seasonal persistence and size‐structuring of two minnow species within headwater streams in the Eastern Cape,South Africa

This study examined temporal variation in population dynamics and size structuring of two cyprinid minnows, Pseudobarbus afer and Barbus anoplus, in relation to their proximate physical habitats. Population estimates were determined using three‐pass depletion sampling during both summer and winter. The habitats were characterised by seasonal variation in all physico‐chemical conditions and spatial variation in substrata compositions. Whereas significant differences in population size were noted between seasons for B. anoplus, no differences were found between seasons for density and capture probability for either species. An increase in boulders was associated with increase in population size and density for P. afer; for B. anoplus, increased percentages of bedrock and bank vegetation were associated with an increase in population size and probability of capture, respectively. According to Canonical Correspondence Analysis, size structuring in P. afer was explained predominantly by seasonality, with smaller length classes associated with the seasonal variable of summer, while larger length classes were associated with pH that was higher in winter. By comparison, for B. anoplus, the habitat variables – bank vegetation and bedrock – accounted for much of the explained variance for size structuring. Recruitment appeared to be the major driver of size structuring for the two species; refugia, especially boulders and bank vegetation, also appeared to be important. Overall, the two species were adapted to the headwater streams that were generally variable in environmental conditions. Potential invasions by non‐native invasive fishes that occur within the mainstream habitats threaten these two species. Efforts should continue to protect these minnows from such invasions by constructing barriers to upstream migration of non‐native fishes into these headwater habitats.     

Passive acoustic telemetry reveals highly variable home range and movement patterns among unicornfish within a marine reserve

Marine reserves are the primary management tool for Guam’s reef fish fishery. While a build-up of fish biomass has occurred inside reserve boundaries, it is unknown whether reserve size matches the scale of movement of target species. Using passive acoustic telemetry, we quantified movement patterns and home range size of two heavily exploited unicornfish Naso unicornis and Naso lituratus. Fifteen fish (N. unicornis: n = 7; N. lituratus: n = 4 male, n = 4 female) were fitted with internal acoustic tags and tracked continuously over four months within a remote acoustic receiver array located in a decade-old marine reserve. This approach provided robust estimates of unicornfish movement patterns and home range size. The mean home range of 3.2 ha for N. unicornis was almost ten times larger than that previously recorded from a three-week tracking study of the species in Hawaii. While N. lituratus were smaller in body size, their mean home range (6.8 ha) was over twice that of N. unicornis. Both species displayed strong site fidelity, particularly during nocturnal and crepuscular periods. Although there was some overlap, individual movement patterns and home range size were highly variable within species and between sexes. N. unicornis home range increased with body size, and only the three largest fish home ranges extended into the deeper outer reef slope beyond the shallow reef flat. Both Naso species favoured habitat dominated by corals. Some individuals made predictable daily crepuscular migrations between different locations or habitat types. There was no evidence of significant spillover from the marine reserve into adjacent fished areas. Strong site fidelity coupled with negligible spillover suggests that small-scale reserves, with natural habitat boundaries to emigration, are effective in protecting localized unicornfish populations.     

Fall movements of Red‐headed Woodpeckers in South Carolina

Fall migration of Red‐headed Woodpeckers (Melanerpes erythrocephalus) can be erratic, with departure rates, directions, and distances varying among populations and individuals. We report fall migration departure dates, rates, and routes, and the size of fall home ranges of 62 radio‐tagged Red‐headed Woodpeckers in western South Carolina. Rates of fall migration differed among years; all radio‐tagged woodpeckers migrated in 2005 (15 of 15), none (0 of 23) migrated in 2006, and 54.2% (13 of 24) migrated in 2007. Of 28 woodpeckers that left their breeding territories, we relocated eight either en route or on their fall home ranges. These woodpeckers migrated short distances (4.3–22.2 km) south along the floodplain forest of a large creek. The variable migration patterns we observed indicate that Red‐headed Woodpeckers may best be described as facultative migrants. We determined the home range sizes of 13 woodpeckers in both seasons, regardless of whether they migrated, and fall home ranges were smaller (mean = 1.12 ha) than summer home ranges (mean = 3.23 ha). Fall‐winter movements of Red‐headed Woodpeckers were concentrated on mast‐producing oak (Quercus spp.) trees, which may have restricted home range sizes. The partial migration we observed in 2007 suggests that factors other than mast crop variability may also influence migration patterns because woodpeckers that year responded to the same annual mast crop in different ways, with some migrating and some remaining on breeding season home ranges during the fall.     

Is there a higher risk for herbivore outhreaks after cold mast years? An analysis of two plant/herbivore series from southern Norway

Historical data on two plant‐herbivore interactions from southern Norway were used to test the hypothesis that the degree of herbivore outbreaks in post‐mast years is negatively related to summer temperatures in the mast year, because plants are more depressed after a high seed production if temperatures and thus the photosynthetic activity is low. The plant species were the sessile oak Quercus petraea and the bilberry Vaceinium myrtillus. For the former species post‐mast years were identified from reports given by the local forest authorities for the period 1930–48, and from acorn export curves for the period 1949–98, For the latter species, post‐mast years were identified mainly from bilberry export curves for the period 1920–31, from game reports for the period 1932–78. and from diary notes for the period 1979–87. The herbivore species used were the green oak leaf roller moth Tortrix viridana and the capercaillie Tetrao urogallus. Eight moth outbreaks on oak forests were reported by the forest authorities in the period 1930–98, and they all started in a post‐mast year of the sessile oak. There were however also eleven post‐mast years without moth outbreaks. According to game reports, observations and autumn counts, all increases in the autumn population size of capercaillie during 1920 88 occurred in or after a year with high bilberry production. Among i8 post‐mast years, there were seven with strong increase, seven with slight or moderate increase, and four with no increase. For both herbivore species, post‐mast years with marked population increases had significantly lower summer temperatures in the preceding (mast) year than had post‐mast years with no or slight increases. For moth populations there also was a negative effect of high temperatures in April, possibly because moth eggs tend to hatch too early relative lo budburst if spring temperatures are high. For the capercaillie, high amount of precipitation in June – July seemed to have some negative impact on the autumn population sizes, as also found in previous studies.     

Variation in the population structure between a natural and a human‐modified forest for a pioneer tropical tree species not restricted to large gaps

The distribution of tree species in tropical forests is generally related to the occurrence of disturbances and shifts in the local environmental conditions such as light, temperature, and biotic factors. Thus, the distribution of pioneer tree species is expected to vary according to the gap characteristics and with human disturbances. We asked whether there was variation in the distribution of a pioneer species under different environmental conditions generated by natural disturbances, and between two forests with contrasting levels of human disturbance. To answer this question, we studied the distribution patterns and population persistence of the pioneer tree species Croton floribundus in the size and age gap range of a primary Brazilian forest. Additionally, we compared the plant density of two size‐classes between a primary and an early successional human‐disturbed forest. Croton floribundus was found to be widespread and equally distributed along the gap‐size gradient in the primary forest. Overall density did not vary with gap size or age (F‐ratio = 0.062, P = 0.941), and while juveniles were found to have a higher density in the early successional forest (P = 0.021), tree density was found to be similar between forests (P = 0.058). Our results indicate that the population structure of a pioneer tree species with long life span and a broad gap‐size niche preference varied between natural and human‐disturbed forests, but not with the level of natural disturbance. We believe this can be explained by the extreme environmental changes that occur after human disturbance. The ecological processes that affect the distribution of pioneer species in natural and human‐modified forests may be similar, but our results suggest they act differently under the contrasting environmental conditions generated by natural and human disturbances.     

Home on the range: spatial ecology of loggerhead turtles in Atlantic waters of the USA

Aim Although satellite tracking has yielded much information regarding the migrations and habitat use of threatened marine species, relatively little has been published about the environmental niche for loggerhead sea turtles Caretta caretta in north‐west Atlantic waters. Location North Carolina, South Carolina and Georgia, USA. Methods We tracked 68 adult female turtles between 1998 and 2008, one of the largest sample sizes to date, for 372.2 ± 210.4 days (mean ± SD). Results We identified two strategies: (1) ‘seasonal’ migrations between summer and winter coastal areas (n = 47), although some turtles made oceanic excursions (n = 4) and (2) occupation of more southerly ‘year‐round’ ranges (n = 18). Seasonal turtles occupied summer home ranges of 645.1 km² (median, n = 42; using α‐hulls) predominantly north of 35 ° latitude and winter home ranges of 339.0 km² (n = 24) in a relatively small area on the narrow shelf off North Carolina. We tracked some of these turtles through successive summer (n = 8) and winter (n = 3) seasons, showing inter‐annual home range repeatability to within 14.5 km of summer areas and 10.3 km of winter areas. For year‐round turtles, home ranges were 1889.9 km². Turtles should be tracked for at least 80 days to reliably estimate the home range size in seasonal habitats. The equivalent minimum duration for ‘year‐round’ turtles is more complex to derive. We define an environmental envelope of the distribution of North American loggerhead turtles: warm waters (between 18.2 and 29.2 °C) on the coastal shelf (in depths of 3.0–89.0 m). Main conclusions Our findings show that adult female loggerhead turtles show predictable, repeatable home range behaviour and do not generally leave waters of the USA, nor the continental shelf (< 200m depth). These data offer insights for future marine management, particularly if they were combined with those from the other management units in the USA.     

Spatial ecology of arboreal snakes (Hoplocephalus stephensii,Elapidae) in an eastern Australian forest

Abstract Stephens' Banded Snakes (Hoplocephalus stephensii Krefft 1869) are large (to 1 m), highly arboreal elapid snakes, restricted to mesic forested areas along the eastern coast of Australia. Radiotelemetric monitoring of 16 individuals at Whian Whian State Forest in north‐eastern New South Wales over 25 months provided the first data on spatial ecology of this threatened taxon. Two major influences on movements by Stephens' Banded Snakes were identified: the distribution of large hollow‐bearing trees, and the avoidance of conspecifics. Radiotracked snakes were sedentary inside tree hollows for extended periods (mean = 8 days) during their active season, interrupted by occasional long (mean = 124 m) nocturnal movements to another shelter tree. Snakes travelled on the ground rather than within the canopy, and thus were potentially exposed to terrestrial predators. Although the home ranges of the radiotracked snakes overlapped substantially (mean = 27%), simultaneous occupancy of ‘shared’ shelter trees was less common than expected by chance. Hence, we conclude that adult Stephens' Banded Snakes generally avoid the presence of conspecifics. Snakes used from five to 30 shelter trees and home ranges of male snakes were larger than those of females (mean = 20.2 vs 5.4 ha). The large spatial scale of these movements, and limited overlap among individuals, means that a viable population of this taxon requires a large area of contiguous forest. This requirement may explain why the species has not persisted in small forest fragments.     

Space use of common genets<Emphasis Type="Italic">Genetta genetta</Emphasis> in a Mediterranean habitat of northeastern Spain: differences between sexes and seasons

The seasonal home range size and spatial relationships of 16 adult genetsGenetta genetta Linnaeus, 1758 (6 males and 10 females) were estimated in a Mediterranean habitat of northeastern Spain. Genets minimum density was estimated as 0.98/km². Mean annual home range was 113.1 ha in males and of 72.0 ha in females. Males had larger home ranges than females in all seasons, but differences were only significant in winter. Home range size changed seasonally and showed a similar pattern in both sexes, with lower values in summer (males — 41.2 ha, females — 29.0 ha) and maximum ones in spring (males — 78.8 ha, females — 56.1 ha). Animals displayed spatial fidelity throughout the year. Core areas (MCP50) represented 27% and 19% of total home range size for males and females, respectively. Resting home ranges (based on locations of inactive animals) were 9 times lower than overall home range size. Individuals of the same sex overlapped less than individuals of different sexes, especially with regard to core areas, which showed almost no overlap. The results obtained suggest that (1) different factors are likely to affect the space use of genets, such as body mass, food abundance and reproductive cycle; (2) genets use space in a heterogeneous way, with areas of greater activity than others within their home range; (3) there was intrasexual segregation with regard to space use.     

Living in the suburbs: Space use by vervet monkeys (Chlorocebus pygerythrus) in an eco‐estate,South Africa

We examined vervet monkey (Chlorocebus pygerythrus) space use using GPS/UHF telemetry data from 10 vervet monkeys across six troops over 9 months within a 420 ha suburban eco‐estate. We documented a mean home range of 0.99 km² (95% MCP) and 1.07 km² (95% KDE) for females (n = 6), 1 km² (95% MCP) and 1.50 km² (95% KDE) for males (n = 4) and 0.87 km² (95% MCP) and 1.12 km² (95% KDE) for troops (n = 6), respectively, indicating that males and larger troops had larger home ranges. These relatively small home ranges included shared territorial boundaries and high home range overlap. Vervet monkey movements indicated higher morning activity levels, and habitat selection indicated significantly more use of golf course, urban residential and forest, thicket and woodland areas, and avoidance of wetland, grassland and shrub, and urban built‐up areas. Our results suggest that modified habitat use by vervet monkeys is a consequence of behavioural facilitation to access highly available food resources, thereby facilitating their persistence in green spaces in urban areas of South Africa. Conflict management is dependent on the conservation of sufficient natural habitats and food resources, to minimise their dependence on anthropogenic supplementary food resources and consequently reduce human–monkey conflict.     

Null model analyses of temporal patterns of bird assemblages and their foraging guilds revealed the predominance of positive and random associations

Patterns of species associations have been commonly used to infer interactions among species. If species positively co‐occur, they may form predominantly neutral assemblages, and such patterns suggest a relatively weak role for compensatory dynamics. The main objective of this study was to test this prediction on temporal samples of bird assemblages (n = 19, 10–57 years) by the presence/absence and quantitative null models on assemblage and guild levels. These null model outcomes were further analyzed to evaluate the effects of various data set characteristics on the outcomes of the null models. The analysis of two binary null models in combination with three association indices revealed 20% with significant aggregations, 61% with random associations, and only 19% with significant segregations (n = 95 simulations). The results of the quantitative null model simulations detected more none‐random associations: 61% aggregations, 6% random associations, and 33% segregations (n = 114 simulations). Similarly, quantitative analyses on guild levels showed 58% aggregations, 20% segregations, and 22% random associations (n = 450 simulations). Bayesian GLMs detected that the outcomes of the binary and quantitative null models applied to the assemblage analyses were significantly related to census plot size, whereas the outcomes of the quantitative analyses were also related to the mean population densities of species in the data matrices. In guild‐level analyses, only 9% of the GLMs showed a significant influence of matrix properties (plot size, matrix size, species richness, and mean species population densities) on the null model outcomes. The results did not show the prevalence of negative associations that would have supported compensatory dynamics. Instead, we assume that a similar response of the majority of species to climate‐driven and stochastic factors may be responsible for the revealed predominance of positive associations.