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The Effect of Vegetation Cover on Vigilance and Foraging Tactics in the Fat Sand Rat Psammomys Obesus

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The Effect of Vegetation Cover on Vigilance and Foraging Tactics in the Fat Sand Rat Psammomys Obesus J Ethol (2001) 19:105–113 © Japan Ethological Society and Springer-Verlag Tokyo 2001 ARTICLE Andrei V. Tchabovsky · Boris R. Krasnov Irina S. Khokhlova · Georgy I. Shenbrot The effect of vegetation cover on vigilance and foraging tactics in the fat sand rat Psammomys obesus Received: January 15, 2001 / Accepted: April 17, 2001 Abstract The combination of the visual obstruction and Introduction protection properties of vegetation is considered to be one of the most important factors determining the trade-off between vigilance and foraging in a prey species. In the Vigilance is one of the main antipredator behaviors of prey Negev desert, diurnal fat sand rats construct their burrows species (Edmunds 1974). Vigilance has often been con- in the ephemeral river beds (“wadis”), under dense and tall sidered as an alternative to foraging and was assumed to shrubs of Atriplex halimus, or on the open first fluvial depend on perceived predation risk (Lima 1987a; Houston terrace, covered with scattered low shrubs of Anabasis et al. 1993). The vigilance/foraging trade-off has been articulata. We tested the hypothesis that properties of the reported to be affected by a variety of factors, including vegetation would affect the time budget of female sand rats. individual characteristics (age and sex: Reboreda and Sand rats spent more time aboveground, rested more, were Fernandez 1997; reproductive status: MacWhirter 1991; less vigilant, and moved more slowly under the dense cover body condition: Bachman 1993), social factors (Elgar 1989; in the wadi than at the open terrace. No differences in the Yaber and Herrera 1994), and environmental factors (veg- total foraging time were revealed, but foraging tactics etation cover: Sharpe and Van 1998; proximity to cover: varied between habitats: individuals in the wadi mainly fed Diaz and Asensio 1991; foraging substrate: Lima 1992). aboveground, whereas those at the terrace mainly hoarded. Open habitats allow long-distance monitoring of sur- Our results indicate that sand rats perceive the dense veg- roundings and are often, albeit not always, associated with etation cover as good protection despite its visual obstruc- decreased vigilance (Quenette 1990). Ambivalent proper- tive nature, and that vigilance in Psammomys obesus is ties of cover, namely obstructive versus protective (Sharpe performed at the cost of resting rather than at the cost of and Van 1998), are reflected by controversial results of foraging. empiric studies. Vigilance can increase (Diaz and Asensio 1991; Colagross and Cockburn 1993) or decrease (Metcalfe Key words Vigilance · Psammomys obesus · Vegetation 1984; Lima 1987b) with distance to cover. For example, cover · Antipredator behavior · Obstruction/protection vigilance rate decreased with distance to cover in the trade-off white-crowned sparrow Zonotrichia leucophrys (Slotow and Rothstein 1995). Contrastingly, vigilance of the buffy- headed marmoset Callithrix flaviceps was higher under less extensive leaf cover (Ferrari and Ferrari 1990), whereas the Townsend’s ground squirrel Spermophilus townsendii ap- peared to scan more often in vegetation of medium height and less often in low or tall vegetation (Sharpe and Van A.V. Tchabovsky Severtsov Institute of Ecology and Evolution RAS, Moscow, Russia 1998). In the teal Anas crecca, vigilance first decreased with distance to cover, but then began to increase as distance to B. Krasnov (*) · G.I. Shenbrot Ramon Science Center, Jacob Blaustein Institute for Desert cover increased further (Pöysä 1994). In the yellow baboon Research, Ben-Gurion University of the Negev, P.O. Box 194, Mizpe Papio cynocephalus, openness of the habitat proved not to Ramon 80600, Israel affect vigilance (Alberts 1994). ϩ ϩ Tel. 972-8-6586337; Fax 972-8-6586369 The balance between the protective and obstructive e-mail: [email protected] components of vegetation cover mirrors the trade-off be- I.S. Khokhlova tween the probability of being surprised and being detected Wyler Department of Desert Agriculture, Jacob Blaustein Institute for Desert Research, Ben-Gurion University of the Negev, Sede by a predator (Pöysä 1994) and thus may be reflected by its Boqer Campus, Israel effect on antipredator vigilance (Lima 1987b; Lazarus and 106 Symonds 1992; Arenz and Leger 1999). Vegetation cover densely covered with Atriplex halimus shrubs ranging in size may either be beneficial to a prey species, if it prevents a between 2 and 8m in diameter and up to 2m in height. predator from detecting prey, or it may be harmful, if it There were also a few shrubs of Anabasis articulata, merely obstructs the prey’s view and provides little or no Artemesia sieberi, and Salsola spp. in this habitat. The first protection (Arenz and Leger 1999). fluvial terrace bordering the rocky second terrace was cov- The fat sand rat, Psammomys obesus Cretzschmar, 1828, ered with scattered shrubs of Anabasis articulata that were is a diurnal solitary strictly folivorous gerbil, inhabiting 0.5–1.0m in diameter and up to 0.6m in height. In addition, deserts of North Africa and the Middle East. Animals feed there were also a few small shrubs of Atriplex halimus, both above and below the ground. They cut and take leaves Artemisia spp., and Salsola spp. Apparently because of the of the plants to their burrows to be eaten later (Daly and relatively dry winter of 1998–1999 (54mm total precipita- Daly 1975). Daly and Daly (1975) noted that P. obesus tion), the vegetation of both Atriplex halimus and Anabasis construct their burrows only under their food shrubs and articulata was not especially verdant. Ten of 50 randomly avoid open habitats. However, in the Negev desert P. selected A. halimus shrubs and 13 of 50 randomly selected obesus use burrows not only under the shrubs in the wadi A. articulata shrubs had fresh leaves and stems, whereas the bottom but also those on the first fluvial terrace. This habi- others were dry. tat is much less protected than wadi beds and is covered Within the study area diurnal P. obesus is subjected with scattered and low shrubs of chenopods. Large shrubs mainly to diurnal avian predators, whereas all mammalian of Atriplex halimus in the wadi bottom provide good pro- and reptilian predators in the Negev desert are mainly tection from raptors, but simultaneously obstruct lateral nocturnal. vision, thereby preventing early detection of terrestrial predators, especially those using a creeping predatory tac- Burrow description tic. Scattered small shrubs of Anabasis articulata on the terrace, on the one hand, allow long-distance monitoring, In total, 132 burrows of P. obesus were found at the study but also expose gerbils to birds of prey and provide poor site. A burrow was considered to be active (inhabited or physical protection. visited) either if a sand rat was observed or if fresh feces or We studied the behavior of P. obesus in the Negev High- food remnants were found near the opened and clean bur- lands to understand how they allocate their time for vigi- row entrances. Otherwise, the burrow was classified as non- lance and foraging in relation to the degree of vegetation active (vacant). Burrows were classified according to their cover. We predicted, based on the hypothesis about trade- location as “wadi burrows” and “terrace burrows”; 31 off between foraging and vigilance, that the ratio of pro- active burrows (15 “wadi burrows” and 16 “terrace tective versus obstructive characteristics of the foraging burrows”) were described in detail. A plot 10m ϫ 10m substrate would affect time budgets and especially vigilance with a burrow in its center was established for each burrow. and foraging of the fat sand rats; that is, individuals that live In each plot we measured (a) maximal length and width in the densely covered wadi bottom would allocate more of the burrow area based on the farthest entrances, (b) time to foraging and would be less vigilant than individuals number of entrances, (c) number of shrubs by species, and living on the more exposed fluvial terrace. To test this pre- (d) maximum length, width, and height for each shrub. diction, we recorded time that P. obesus living in two micro- habitats with different vegetation cover allocated to various types of activities. Animals Psammomys obesus were live-trapped using Havahart two- door cage traps (model 1025; Woodstream, Lititz, PA, Materials and methods USA) baited with fresh leaves of Atriplex halimus. Trap- ping was performed under the license 1998/3694 issued by Study area and study site Nature Protection and National Parks Authority of Israel on November 1, 1998. Traps were placed near the entrances The study was carried out on the northern rim of the of the active burrows and were checked continuously during Ramon erosion cirque, Negev Highlands, Israel (30°35Ј N, the day, so that each trapped individual stayed in a trap no 34°45Ј E). The main landscape of the study area is a com- more than 15min. Fat sand rats do not need free water, so plex of hills with a deep loess layer (Յ1m) and wide dry the bait satisfied their water requirements. Trapped animals riverbeds covered with shrubs of Anabasis articulata, were sexed, aged, weighed, and marked individually by fur- Atriplex halimus, Artemisia sieberi, Salsola schweinfurthii, cutting in a symmetrical recognizable pattern. Handling and Noaea mucronata. Data were collected during March time took approximately 7min per individual if it was cap- and April when mean daily temperature ranged between tured for first time. Marked female was handled for 2min to 14° and 29°C. The mean annual rainfall in the area is near check for signs of pregnancy or lactation, whereas a marked 100mm and occurs in winter. male was released immediately. Daily temperature during The study site (800m ϫ 100m) was established along the study period was such that the animals did not need any valley of Wadi Nizzana and included both the bottom of bedding in the traps.
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