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Telemetric ®Eld Studies of Body Temperature and Activity Rhythms of Acomys Russatus and A Oecologia (1999) 119:484±492 Ó Springer-Verlag 1999 R. Elvert á N. Kronfeld á T. Dayan á A. Haim N. Zisapel á G. Heldmaier Telemetric ®eld studies of body temperature and activity rhythms of Acomys russatus and A. cahirinus in the Judean Desert of Israel Received: 21 December 1998 / Accepted: 15 March 1999 Abstract Two species of the genus Acomys coexist in arid Introduction zones of southern Israel. Acomys russatus is distributed in extremely arid areas, while A. cahirinus is common in both Two species of spiny mice, Acomys russatus and A. ca- Mediterranean and arid regions. Individuals of both hirinus, coexist in arid areas in the south of Israel species from a rodent community in the Ein Gedi Nature (Shkolnik 1971; Kronfeld et al. 1996). While A. cahirinus Reserve were implanted with temperature-sensitive is widely distributed in the semi-arid and even in the transmitters. Body temperature (Tb) rhythms were re- mesic regions of Israel, A. russatus is limited to the ex- corded in free-ranging mice at four dierent seasons of the tremely hot environments of the southern and the cen- year. A. cahirinus (30±45 g) showed a nocturnal rhythm of tral parts of the Rift Valley in Israel (Harrison and Bates Tb throughout the year. In the activity phase during the 1991) and overlaps with A. cahirinus throughout its night Tb increased to 38.2°C. During the day Tb decreased range. The habitat of the spiny mice in the Judean De- to 34°C. This species displayed this pattern in summer sert is characterised by steep rocky mountain escarp- also when ambient temperatures rose above Tb. The Tb of ments running parallel to the Dead Sea. Huge blocks A. russatus (45±65 g) varied between 34.8 and 41°C dur- alternate with small boulders. Spiny mice do not con- ing the hot season, showing a bimodal temperature struct burrows, but seek the shelter of rock crevices. rhythm with maximal values in the morning and in the Trapping data suggest that the common spiny mouse evening. Measurements of activity in this species showed (A. cahirinus) is active during the night, as are most inactivity during the hottest period of a summer day. In desert rodents, while the golden spiny mouse (A. russ- winter A. russatus showed no clearly detectable diurnal or atus) is active during the day (Shkolnik 1971). A. russ- ultradian rhythm in Tb, which remained constant between atus switched to nocturnal activity when the nocturnal narrow limits of 35.2 and 36.8°C. conspeci®c was removed from the habitat. This implies that interspeci®c competition could be the driving force Key words Acomys á Telemetry á Body temperature á for the temporal segregation of activity among the two Activity á Circadian rhythms Acomys species. The golden spiny mouse appears to be forced into diurnal activity by its more aggressive con- gener (Shkolnik 1971). This is supported by laboratory R. Elvert (&) á G. Heldmaier studies demonstrating that A. russatus displays noctur- Department of Biology, Philipps-University, D-35032 Marburg, Germany nal rhythms of body temperature and metabolic rate e-mail: [email protected], Fax: +49-6421-288937 when separated from A. cahirinus (Haim and Zisapel N. Kronfeld á T. Dayan 1992, 1994; Rubal et al. 1992; Fluxman and Haim 1993; Department of Zoology Kronfeld et al. 1994). Chemical cues released by A. ca- Tel Aviv University, hirinus into the habitat may maintain the time separa- Tel Aviv 69978, Israel tion of activity patterns of these two species (Haim and A. Haim Rozenfeld 1993; Friedman et al. 1997). Department of Biology, Trapping data do not allow continuous recording of University of Haifa-Oranim, 24-h rhythms in individual mice and thus detailed ac- Kiryat Tiv'on 36910, Israel tivity time budgets of undisturbed animals cannot be N. Zisapel studied. We applied the technology of body temperature Department of Biochemistry, Tel Aviv University, telemetry to individuals of both species where they co- Tel Aviv 69978, Israel exist at Ein Gedi. This technology enables us to study 485 the (body temperature) Tb rhythm of individual mice, antennae (type Ringo Ranger) placed 30 m apart in the study area. shedding light on their overt temperature rhythms. The Scanning the two antennae improved signal monitoring. Both an- tennae received the signals with dierent strength, and this varied transmitters used were further developed in order to with the position of the mice. Changes in signal ®eld strength be- record also activity patterns of individual animals, tween two successive records were recorded as activity in October enabling us to study the relationship between actual 1996. Constant signal ®eld strength was interpreted as no activity. activity and overt temperature rhythms of free-ranging The tracking receiver (Reichenbach, Freiburg, Germany) was populations of spiny mice. connected via an interface (DACpad-71 B, Datalog. Moenchen- gladbach, Germany) to a low-energy-consuming notebook-com- puter (COMPAQ Contura Aero 4/25) which stored the data on hard disk, and controlled the receiver channels as well as antenna Materials and methods scanning. The entire system was controlled by self-developed software (QB45) which also included ®lter algorithms for supress- ion of noise and interference. Individuals of A. cahirinus and A. russatus were trapped on a rocky Ambient temperatures were measured by thermocouples (cop- hill slope of about 4000 m2 near the Ein Gedi Field School per/constantan) placed in the sun (air temperature, 30 cm above (31°28¢N, 35°23¢E, 100±350 m below sea level) at all seasons of the ground) and 1 m inside a rock crevice. Temperatures were recorded year. The time schedule included three seasons in 1995: February/ from the same positions at all seasons. T , T and activity were March, July/August and October/November; and two in 1996: b air recorded at 4-min intervals and stored in the computer and a data January and October/November. The fall season was repeated logger, THERM 2281-8 (AMR, Holzkirchen, Germany). because of technical problems and loss of data caused by envi- For statistical analysis, data are expressed as the mean SD. ronmental in¯uences, e.g. ¯ood of the study area in November Student's t-test or Mann-Whitney U-test were used for compari- 1995. We trapped the mice with 50 collapsible Sherman box traps sons between mean hourly values of photo- and scotophase. spread randomly in the area. Peanut butter and a small piece of carrot were used as bait. In each study season, traps were set for 25±31 days and they were checked daily just after sunrise and be- fore sunset. In the summer, traps were closed during daytime hours Results and additionally covered with styrofoam for heat protection. The spiny mice were individually marked with toe clipping. Every in- dividual was weighed whenever trapped using Pesola dynamometer A. russatus scales with 0.3% accuracy. Population densities and abundances were calculated using the Jolly-Seber method (Krebs 1989; Suth- Twenty-eight individuals of A. russatus were trapped on erland 1996) which is designed for open populations where all in- dividuals in the ®rst sample are unmarked. This method requires a 377 occasions in March, July and October 1995 and in previous and following trapping period, so it was only possible to January 1996. All were marked when trapped the ®rst estimate population size for two seasons. The densities can be es- time. Two individuals could be retrapped in each of the timated from relationship of ``size of marked population'' to ®rst four seasons (Fig. 1). Mouse no. 5000 was re- ``number of marked animals'' (Krebs 1989). Because of a delay of trapped 7 times in March and July 1995, 8 times in 9 months between the trapping period in January 1996 and Octo- ber/November 1996, the latter study season was not included for October 1995 and 5 times in January 1996. Another population estimates. individual (no. 0200) which was not implanted with a Mice which were trapped repeatedly were implanted with transmitter was retrapped 19 times in March, 11 times in temperature transmitters into the abdominal cavity (Oekokart July, 16 and 17 times in October and January, respec- E&V, Munich, Germany). For the implantation, mice were an- aesthetised with a mixture of ketamine and xylazine. Both species tively. In other individuals, the retrapping incidence required dierent dosages of the anaesthetic (A. cahirinus: ketamine varied between 5 and 13 times during one season. The 80±110 mg kg)1 and xylazine 4 mg kg)1; A. russatus: ketamine density of A. russatus in our study site, based on the 120±150 mg kg)1 and xylazine 5 mg kg)1). Anaesthetics were in- population estimation using the Jolly-Seber method, was jected i.p. and transmitters were implanted in the abdominal cavity. The peritoneum was sutured with resorbable catgut (1.5 metric), about 38 individuals per hectare during July and 35 and the skin with sewing silk (1.5 metric). After a recovery period individuals per hectare during the October season. of 36 h the animals were released again in their natural habitat. High retrapping rates and continuous monitoring of During the measuring period, the implanted mice were retrapped body weight during dierent seasons indicate that the repeatedly and never showed infections. At the end of each re- sedentary behaviour of mice was not disturbed by our cording period, the mice were retrapped to remove the transmitters. The transmitters were of FM type with an integrated internal loop- trapping activities and transmitter implantation (Fig. 1). antenna. They transmitted signals in the frequency range of 150± One individual A.
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