Interspecific competition in tree squirrels: do introduced grey squirrels (<Emphasis Type="Italic">Sciurus carolinensis</Emphasis>) deplete tree seeds hoarded by red squirrels (<Emphasis Type="Italic">S. vulgaris</Emphasis>)?

Red squirrels (Sciurus vulgaris) and introduced eastern grey squirrels (S. carolinensis) scatterhoard seeds of broadleaf trees. Scatterhoarded seeds are an essential resource in spring and their consumption increases red-squirrel fitness. We examined whether grey squirrels partly deplete the high-energy food resources cached by red squirrels, reducing their consumption, in two ways: (1) at the population level, comparing energy intake of feeding on cached seeds between a study site with red and grey squirrels and one with only red squirrels present; and (2) at the individual level, in the study site where species co-exist, relating hoard recovery of red squirrels to the amount of core-area overlap with grey squirrels. There were no significant site differences in the mean daily energy intake of red squirrels feeding on seeds recovered from caches. However, in the red-grey site, during spring, red squirrels that had a high percentage of their home-range core area overlapped by grey squirrels had a lower daily energy intake than low-overlap red squirrels. Body mass of red squirrels in spring was negatively correlated with the percentage of interspecific core-area overlap, but not with core-area overlap with other red squirrels. Our data suggest that interspecific competition for scatterhoarded seeds, with grey squirrels pilfering red squirrels' food caches, caused a reduced energy intake in red squirrels with a high degree of interspecific core-area overlap, and reduced body mass in spring. Therefore, cache pilfering is likely to reduce reproductive output in red squirrels, and thus play a role in the replacement of red by grey squirrels. Electronic Publication     

Gap-Crossing Decisions by the Red Squirrel, a Forest-Dependent Small Mammal

Abstract:  Forest-associated species in fragmented landscapes must traverse potentially inhospitable gaps to move between habitat patches. Although conservation biologists advocate connecting patches with corridors or improving the matrix to make it suitable for movement, little is known about the factors influencing gap-crossing decisions for most species. We investigated gap crossing by the red squirrel ( Tamiasciurus hudsonicus ) in logged landscapes in southeastern Alaska, where the species avoids microhabitats associated with gaps created by clearcutting. We released individuals across clearcuts and determined the routes they took home with tracking spools and radio telemetry. Of 36 adult red squirrels translocated across six clearcuts, 14 crossed clearcuts to reach home. Squirrels were more likely to cross clearcuts if the detour efficiency (distance to home crossing gap divided by distance of forested detour) was low, indicating an ability to compare distances along alternate routes and travel costs or risks in different habitats. No other landscape metrics, such as gap size or crossing distance, predicted crossing behavior. Red squirrels of low body mass were more likely to cross clearcuts, where the probability of encountering conspecifics is low. Distance predicted route choice for squirrels detouring around clearcuts. Indirect evidence suggests that perceived predation risk, energetic costs, or both are higher in clearcuts. Detour efficiency reportedly influences the gap-crossing decisions of some forest-associated birds, but this is the first demonstration of its role in gap-crossing decisions by a mammal.     

Brood neglect and contingent foraging behavior in a pelagic seabird

Among species where there is a risk to leaving offspring unattended, parents usually take alternating shifts guarding their young. However, they may occasionally exhibit brood neglect by leaving their offspring unattended at the nest. To investigate this phenomenon further, we examined the foraging behavior of the northern gannet (Morus bassanus) during chick-rearing. This species has a prolonged nestling period (13 weeks) and the single chick is usually guarded by one or other of its parents, because unattended chicks are frequently attacked by conspecifics. We tested the prediction that the foraging behavior of adults when they left their offspring alone at the nest (unattended trips) would differ in character to when adults left offspring with their partner (attended trips). Brood neglect typically occurred after a longer-than-average attendance period at the nest. Unattended trips were, on average, about half the duration of attended trips, and therefore much closer to the colony. There was also a difference in departure direction between attended and unattended trips, with unattended trips tending to be northeast of the colony. Chicks were fed by parents after both attended and unattended trips, but the frequency and the duration of unattended trips increased as chicks grew older whereas the duration of attended trips decreased as chicks grew. These results indicate that parents may be making a trade-off between risk of attack to their offspring and food-intake rate, and that the solution to this trade-off is dependent on chick age.Communicated by C. Brown     

Pigeon homing: new airbag experiments to assess the role of olfactory information for pigeon navigation

Summary In three series of experiments we assessed the effects of olfactory and non-olfactory information collected en route or at the release site on the initial orientation of homing pigeons. In the first experiment, pigeons were transported in open crates to two sites located in opposite directions from the home loft. They were left at the site for 1 h, then put into airtight containers filled with air from that site and brought back to the loft. From there, controls were transported back to the original site. Experimentals were transported to the opposite site. Upon arrival at the site, the olfactory mucosae of both groups were anesthetized with Gingicain. Thus in this experiment, control and experimental pigeons were exposed to different olfactory as well as to different non-olfactory information during displacement and at the site. In the second series, controls and experimentals were treated as in the first experiment, except that they were enclosed in the airtight containers at the very beginning of the experiment and were ventilated with synthetic air until arrival at the final release site. This treatment excluded the possibility to perceive olfactory information en route or at the site. In this series, the two groups differed only with respect to non-olfactory information perceived during displacement and/or at the release site. In the third series, we exposed pigeons at the loft to air collected either at the later release site (controls) or to air collected at a site located in opposite direction of the home loft (experimentals). Here the two groups differed only with respect to their exposure to air of different origin. In all three series, the pooled controls showed a directional preference that was statistically indistinguishable from the home direction. All three experimental groups were disoriented. In the first two series, the differences in the initial orientation of control and experimental pigeons were highly significant. In the third experiment, there was only very weak statistical evidence for a difference between controls and experimentals. These results suggest that more than one factor is involved in the pigeons' navigation system. According to the present experiment with synthetic air, pigeons probably gather and process non-olfactory information during the first part of their transport from the home loft to the release site. Thus, some kind of, in this case non-olfactory, route reversal seems to be involved in the homing process. In addition, the experiment involving only manipulations of airborne information indicated some olfactory component. Previous experiments at our loft did not result in disorientation of the pigeons if either only the access to airborne information had been removed or when otherwise unmanipulated pigeons had been transported in detours to the final release site. Therefore, we assume that our pigeons' navigation system relies on several cues. Deletion of one cue can be compensated by other information. Often the initial orientation of our pigeons is disturbed only when at least one cue is removed and another one provides false information.     

Movements of dispersing flying squirrels in relation to siblings and parents

Knowledge of kin interactions can be informative in explaining the processes underlying dispersal. By dispersing, relatives can avoid kin competition for resources or mates and prevent inbreeding. We investigated sibling movements in relation to each other and parents before and after dispersal in radio-collared juvenile flying squirrels (Pteromys volans L.). Before dispersal, most siblings moved in different areas and were not in contact while exploring the surroundings of natal area. After dispersal, all siblings settled far away from each other, as they dispersed different distances and to random directions compared to each other. No clear effect of litter size or presence of same-sex siblings was observed on dispersal. Time spent in the same nest with the mother did not differ between dispersing sons and daughters. Mother did not force dispersers to leave the natal nest; instead, the mother often moved away from the nest before offspring. Father and offspring were not observed to be in contact before dispersal. Dispersal direction was unaffected by the location of fathers’ home range. We conclude that after dispersal interaction between relatives is limited in flying squirrels, but we did not find any indication that interaction between relatives before dispersal is determining dispersal decision. Siblings’ settlement far away from each other was already indicated by the movement within the natal home range, but, interestingly, the latter could not be used to predict dispersal patterns of sexes.     

Oceanographic characteristics of areas used by Cory’s shearwaters during short and long foraging trips in the North Atlantic

Many breeding seabird species are central-place foragers and constrained to find productive prey patches within their foraging ranges. We assessed how different populations of a pelagic seabird species, the Cory’s shearwater Calonectris diomedea, breeding in oceanic and neritic conditions, cope with these constraints in the North Atlantic, during both incubation and chick-rearing periods. We analysed 237 foraging trips to study the movements and oceanographic characteristics of foraging habitats of seven different populations of Cory’s shearwaters. Generally, oceanic populations exhibited higher foraging effort, by travelling more time and to more distant areas, and larger home ranges and feeding areas, than the neritic population (i.e. breeding on an island within the Portuguese continental platform). On their short trips (i.e. ≤4 days), birds from the different populations fed mostly in shallower waters around the colony. During long trips (i.e. ≥5 days), feeding areas of both oceanic and neritic populations were characterized by high concentration values of chlorophyll-a, low sea-surface temperature and shallower habitats, with oceanic populations of the Azores exploiting areas north of the islands over known seamounts and frontal regions. Birds from other oceanic population (Selvagens) also exploited the African continental shelf system on their long trips. The home ranges of the different populations overlapped widely, but there was a general spatial segregation in terms of the core feeding areas at the population level. Core feeding areas and areas of foraging overlap between different populations should be important to inform conservation management measures, such as the definition of Marine Important Bird Areas for seabirds over the North Atlantic.     

Alarm calling behavior of the thirteen-lined ground squirrel,Spermophilus tridecemlineatus

Summary Alarm calling in a population of thirteenlined ground squirrels, Spermophilus tridecemlineatus, was studied over a three-year period. Data on ground squirrel reactions to human and canine approaches and to the approach or presence of avian predators were used to quantify alarm calling behavior.The results support the hypothesis that alarm calling in this species functions to warn genetic relatives. Human and canine approach-elicited calls were most frequently given by mothers and their recently emerged young; adult males and nonparous females rarely called. The onset of maternal calling coincided with the aboveground appearance of a mother's own litter, and both juvenile and maternal calling were terminated at approximately three weeks post-emergence. Alarm calls were rarely emitted during encounters with avian predators.Alarm calling behavior in S. tridecemlineatus thus appeared to be dependent upon the presence of newly emerged juvenile relatives. To investigate whether the population structure of S. tridecemlineatus was perhaps incompatible with the evolution of alarm calling directed toward adult relatives, the distance between the home ranges of adult relatives and the distance over which the alarm vocalization is audible to ground squirrels were measured. The results revealed that females were likely to have adult relatives relatives residing within audible range of the call.     

Site fidelity and homing behavior in juvenile loggerhead sea turtles (<Emphasis Type="Italic">Caretta caretta</Emphasis>)

To investigate site fidelity and homing behavior in juvenile loggerheads ( Caretta caretta, L.), a mark-recapture study spanning four years (1998–2001) was conducted in Core Sound, N.C., USA. Each year of the study, approximately half of the turtles captured were tagged and released near the capture sites ( n=207), while the remaining turtles were displaced 15–20 km and released ( n=198). Loggerheads in both groups were recaptured in equal proportions near the original capture sites and many individuals were also recaptured in subsequent years. These data imply that juvenile loggerheads often returned to their capture sites following displacement, because if turtles dispersed randomly or remained near their release sites, then fewer displaced turtles should have been caught again. Moreover, because turtles migrate out of North Carolina sounds each winter, turtles recaptured at the same locations in different years evidently returned to specific sites following long migrations. To further investigate homing behavior, a small number of displaced turtles ( n=28) were tracked using radio telemetry following their release. Although transmitters detached from most turtles within a few days, analyses of initial headings showed strong orientation in the direction of the capture site. In addition, four turtles successfully tracked for longer periods of time all returned rapidly to the vicinity of the capture location and remained in the area. Taken together, the results of this study indicate that juvenile loggerheads exhibit fidelity to specific areas during summer months and possess the navigational abilities to home to these areas following forced displacements and long-distance migrations.     

Why do female Belding's ground squirrels disperse away from food resources?

We examined the effects of food provisioning on the natal dispersal behavior of Belding's ground squirrels (Spermophilus beldingi). We provided extra food to adult and yearling females in their maternal territories during pregnancy and lactation, and to offspring of these females in their natal areas for 6 weeks after weaning. We used unprovisioned young of unprovisioned mothers as controls. Provisioning influenced the probability of dispersal from the natal area by female but not male S. beldingi. All surviving male S.␣beldingi dispersed by 55 weeks of age, regardless of whether they and their mothers received extra food. By contrast, we observed a significant trend, beginning 3 weeks after weaning and continuing through the yearling year, for a greater proportion of provisioned than control female S. beldingi to emigrate from the natal area. Competition for food did not appear to influence natal dispersal of females. However, overall population density, density of females weaning litters, and rates of aggression and vigilance among these females, were higher in provisioned than control areas, suggesting that competition for non-food resources was unusually intense in provisioned areas. We propose that juvenile female, but not juvenile male, S. beldingi may emigrate from the natal site to increase access to areas with low densities of conspecifics. Together with findings of earlier workers, our results suggest that spatial and temporal distributions of environmental resources are important influences on the dispersal behavior of female ground squirrels. Received: 28 February 1996 / Accepted after revision: 16 October 1996     

Influence of oceanic factors on long-distance movements of loggerhead sea turtles displaced in the southwest Indian Ocean

The routes of five satellite-tracked loggerhead turtles (Caretta caretta), subjected to an experimental translocation away from their usual migratory routes, have been analysed in relation to the concurrent oceanographic conditions. Remote sensing data on sea surface temperature and height anomalies, as well as trajectories of surface drifters were used, to get simultaneous information on the currents encountered by the turtles during their long-range oceanic movements. Turtles mostly turned out to move in the same direction as the main currents, and their routes were often influenced by circulation features they encountered. A comparison between turtle ground speeds with that of drifters shows that in several instances, the turtles did not drift passively with the currents but contributed actively to the overall movement. Two turtles embarked on an oceanic crossing, probably induced by seasonal changes in surface temperatures, a crossing that was largely determined by the main currents existing in the area.     

Testing the navigational abilities of ocean migrants: displacement experiments on green sea turtles (Chelonia mydas)

Like many animals migrating through the oceans, sea turtles face difficult navigational tasks when they have to reach distant, specific sites. The paradigmatic case of Brazilian green turtles (Chelonia mydas), which nest on the tiny Ascension Island in the middle of the Atlantic Ocean, has often been the subject of hypotheses concerning their navigational mechanisms. To investigate their nature, we displaced 18 females from Ascension and tracked them by satellite after release from eight different points in the ocean, 60-450 km away from the island. Four turtles moved to Brazil soon after the release, 4 moved in various directions before heading to Brazil, and 10 reached the island. All the successful trips, bar 1, were winding but ended with a final straight segment of variable length, as if the turtles were searching for a sensory contact with the island which they obtained at various distances. The approach to Ascension mostly occurred from the direction opposite to the trade wind, suggesting a navigational role of wind-borne information originating from the island.     

Correct and false olfactory orientation of homing pigeons as depending on geographical relationships between release site and home site

Summary Pigeons from two German home sites were released at a site near Mantua in northern Italy. The home sites, Andechs and Würzburg, are 303 and 508 km north of the release site, respectively. Not only the initial bearings but still more the distributions of recoveries after a longer flight distance (median 65 km) were very different in pigeons from these two lofts. While the majority of the Wurzburg birds were found north of the release site, almost all birds from Andechs were found south of it (Fig. 1). Pigeons from both lofts, if made anosmic by sectioning the olfactory nerves, showed no average tendency towards change of latitude. These findings strongly suggest that both correct and false positional information were deduced by the birds from olfactory inputs. A coherent (though very hypothetical) interpretation of these and earlier results is based on regularly varying proportions of chemical tract compounds in the atmospheric boundary layer over the Alps and adjacent regions (Fig. 4).     

Desert ants adjust their approach to a foraging site according to experience

In the habitat of desert ants, Cataglyphis fortis, a constant wind is usually blowing during the daytime. When visiting a familiar food source, the ants steer some distance downwind of the feeder, rather than attempting a direct approach that might miss small food sources, in particular. In the downwind area, the ants pick up the odor plume emanating from the food and follow it upwind to the prey. This strategy saves considerable walking distance and time. The additional path necessitated by the downwind strategy is only about 0.75 to 2 m, depending on nest–feeder distance, while missing the food on the upwind side results in much longer search trajectories. During the initial three to five visits to a feeding site, downwind distance and length of the approach path are shortened notably, and the approach trajectory is straightened. Desert ants further exhibit considerable short-term flexibility in their approach. Experienced individuals are evidently able to decide upon leaving the nest which direction to choose toward the feeder, depending on current wind direction (that fluctuates slightly during the day). Notable changes in wind direction occur primarily overnight. For larger nest–feeder distances, the animals adjust their approach en route to the altered wind direction during their first foraging trip in the morning.     

Nest site selection in the open-nesting honeybee Apis florea

We studied nest site selection by swarms of the red dwarf honeybee, Apis florea. By video recording and decoding all dances of four swarms, we were able to determine the direction and distances indicated by 1,239 dances performed by the bees. The bees also performed a total of 715 nondirectional dances; dances that were so brief that no directional information could be extracted. Even though dances converged over time to a smaller number of areas, in none of the swarms did dances converge to one site. As a result, even prior to lift off, bees performed dances indicating nest sites in several different directions. Two of four swarms traveled directly in what seemed to be the general direction indicated by the majority of dances in the half hour prior to swarm lift off. The other two traveled along circuitous routes in the general direction indicated by the dances. We suggest that nest site selection in A. florea has similar elements to nest site selection in the better-studied Apis mellifera. However, the observation that many more locations are indicated by dances prior to lift off also shows that there are fundamental differences between the two species.     

Can pigeons be fooled about the actual release site position by presenting them information from another site?

Summary The aim of the experiment was to test the hypothesis that pigeons depend on route- and/or site-specific airborne parameters to establish their position relative to the loft. Pigeons were transported to the release site with free access to the environmental air. They were then enclosed in large airtight containers filled with air from the release site and either transported via the loft to a release site in the opposite direction with respect to the loft (experimental birds) or via the loft back to the same site (control birds). Before release the pigeons (still inside the containers) were made anosmic by a local anesthetic applied to the olfactory membranes through the nostrils. Vanishing bearings of experimental birds were on the average in a direction opposite to home while the vanishing bearings of control birds were homeward oriented. For this initial orientation the pigeons seem to rely on airborne cues obtained at a site where they last had access to ambient air or cues obtained from the air inside the sealed containers. Irrespective of the treatment the bearings pooled with respect to north still show a residual orientation to the NE. There was no difference either in the homing speeds or in the homing times of anosmic control and experimental pigeons. We therefore have to assume further mechanisms guiding the pigeons home in addition to a possibly olfactory one.     

The association of female kin in the arctic ground squirrel Spermophilus parryii

Summary The association of kin in arctic ground squirrels (Spermophilus parryii) was studied near Haines Junction, Yukon Territory, during the summers of 1977, 1978, and 1979. Males dispersed in this population, thus only females were likely to live near relatives.Close female kin (sisters, mothers/daughters) had greater overlap of home ranges and interacted more amicably and less agonistically than did less closely related females. Closely related females clumped their young at emergence, whereas more distantly related females did not; there was little indication of clumping of pre-emerged young. The overlap of home ranges of distant relatives (known genetic relatives that had not associated in a natal burrow) was intermediate to that of close relatives and non-relatives. The types of interactions between distant relatives were more similar to those between non-relatives than between close relatives.I conclude that female arctic ground squirrels exhibit nepotism. Females may benefit from associations with relatives by sharing watching for predators once juveniles become conspicuous. Male arctic ground squirrels commit infanticide and several females may be more effective at protecting their young from infanticidal males than females living alone. I suggest that clumping of young by close relatives may provide a mechanism allowing distantly related females to identify each other.Address for offprint requests     

Sibling recognition in thirteen-lined ground squirrels: effects of genetic relatedness,rearing association,and olfaction

Summary I investigated sibling-sibling recognition in captive thirteen-lined ground squirrels (Spermophilus tridecemlineatus) by cross-fostering lab-born pups shortly after birth. When young reached about 45 days of age, I observed dyadic interactions in a test arena of pairs from four relatedness X rearing groups, and recorded the frequency of exploratory encounters between individuals. Sibs-reared together and nonsibs-reared together exhibited significantly fewer exploratory encounters than either sibs-reared apart or nonsibsreared apart. Young reared together were equally exploratory, regadless of relatedness; similarly, young reared apart, whether they were sibs or nonsibs, showed similar levels of exploration. Thus, the differential treatment of siblings in the lab appears to be based on rearing association and not genetic relatedness per se. I interpret this recognition based on association (rearing familiarity) in the context of the species' social organization and compare my results on S. tridecemlineatus with similar studies on S. beldingi, S. parryii, and S. richardsonii. I also used an olfactory impairment technique (zinc sulfate) and found that differential treatment in thirteen-lined ground squirrels was influenced by olfactory cues.     

Changes of movement patterns from early dispersal to settlement

Moving and spatial learning are two intertwined processes: (a) changes in movement behavior determine the learning of the spatial environment, and (b) information plays a crucial role in several animal decision-making processes like movement decisions. A useful way to explore the interactions between movement decisions and learning of the spatial environment is by comparing individual behaviors during the different phases of natal dispersal (when individuals move across more or less unknown habitats) with movements and choices of breeders (who repeatedly move within fixed home ranges), that is, by comparing behaviors between individuals who are still acquiring information vs. individuals with a more complete knowledge of their surroundings. When analyzing movement patterns of eagle owls, Bubo bubo, belonging to three status classes (floaters wandering across unknown environments, floaters already settled in temporary settlement areas, and territory owners with a well-established home range), we found that: (1) wandering individuals move faster than when established in a more stable or fixed settlement area, traveling larger and straighter paths with longer move steps; and (2) when floaters settle in a permanent area, then they show movement behavior similar to territory owners. Thus, movement patterns show a transition from exploratory strategies, when animals have incomplete environmental information, to a more familiar way to exploit their activity areas as they get to know the environment better.     

Spacing of leks in relation to female home ranges,habitat requirements and male attractiveness in the great snipe (Gallinago media)

Summary In an attempt to examine three main hypotheses on the evolution of leks, data on female home ranges, distance between leks and male site fidelity in the great snipe (Gallinago media) were obtained. In a 30 × 40 km area in central Sweden, the locations of 12 leks were identified. Six were within the study area (7 × 11 km) in which probably all leks were known. In the study area, nearest neighbor distances between leks were longer than the predicted drawing area (diameter of female home ranges plus the detection range of leks). Furthermore, in only 1 of 13 radio-tracked females did the estimated home range enclose many leks. These data do not fit the prediction from the hotspot hypothesis on the dispersion of leks, namely, that the distance between leks should be less than the drawing area and hence an average female home range should enclose more than one lek. The observed distance between leks was about the same as predicted by the female preference hypothesis. This hypothesis also predicts females visit mainly one lek; however, we found they sometimes visit two and nest close to a third. We propose that males may first settle according to hotspot rules, but females will resettle according to preferences for certain males and/or larger leks. In this way some hotspot leks are abandoned, and the distance between leks is increased as males become more tightly clumped. However, when all males and females have settled, the leks still existing would be on hotspots. In the female preference hypothesis, males are assumed to aggregate on leks because females prefer clustered males as they can be more easily compared than if they are dispersed. Alternatively, in the attractiveness hypothesis, it is suggested that females prefer certain males, and hence unattractive males surround the attractive are forced to join leks in order to come close to females. Our data on male site fidelity support the attractiveness hypothesis because successful and dominant males return to the same lek and territory, both within and between years, whereas unsuccessful males move to other leks. Attractiveness of certain males may explain why according to hotspot rules in this species males are more clumped than expected. Offprint requests to: J. Höglund     

The behavior of heterotypic resting schools of juvenile grunts (Pomadasyidae)

Resting schools of juvenile grunts (less than 12 cm length), composed primarily of Haemulon flavolineatum and H. plumieri, were studied from 1972 to 1976 on a series of patch reefs surrounded by seagrass beds on the northeast coast of St. Croix, U.S. Virgin Islands. Juvenile grunts form large inactive multispecies schools in reef areas by day. Repeated censuses over several years reveal that certain parts of the reef, particularly those with large formations of the corals Porites porites and Acropora palmata, are traditional schooling areas. When the fishes reach a size of about 15 cm, the H. plumieri form homotypic schools which roam the patch reefs but are no longer associated with particular formations of coral. In contrast, larger H. flavolineatum are uncommon on the patch reefs and they may move offshore to deeper water. The schools of juvenile grunts migrate some distance away (often 100 to 300 m) into seagrass-covered areas to feed on invertebrates at dusk each day. The routes which are followed into the seagrass beds are precise and persist over years. At dawn the grunts return to the reef on the same routes. Mass marking and release experiments show that juvenile grunts will move over long distances (approximately 3 km) to reach a home reef.