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Insubation Behavior of the Dead Sea Sparrow

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Insubation Behavior of the Dead Sea Sparrow 340 SHORT COMMUNICATIONS W. Weller for their helpful suggestions in revising JOHNSGARD,P. A., AND J. KEAR. 1968. A review of the manuscript. parental carrying of young by waterfowl. Living Bird 7:89-102. LITERATURE CITED KEAR, J. 1970. The adaptive radiation of paren- tal care in waterfowl, p. 357-392. In J. H. Crook BERGMAN, R. D., AND D. V. DERKSEN. 1977. Ob- Led.], Social behaviour in birds and mammals__I. servations on Arctic and Red-throated Loons at Academic Press, London. Storkersen Point, Alaska. Arctic 30:41-51. KLOPFER. P. H. 1959. The develonment of sotmA-ALU COLLIAS, N. E., A~TII E. C. COLLIAS. 1956. Some signal preferences in ducks. Wilson Bull. 71: mechanisms of family integration in ducks. Auk 262-266. 73 :378-400. LENSINK, C. J. 1967. Arctic Loon predation on GOTTLIED, G. 1965. Imprinting in relation to pa- ducklings. Murrelet 48:41. rental and species identification by avian neo- nates. J. Comp. Physiol. Psychol. 59:345-356. Department of Biology, Queens’ University, Kingston, H~~HN, E. 0. 1972. Arctic Loon breeding in Al- Ontario K7L 3N6, Canada. Accepted for publication berta. Can. Field-Nat. 86:372. 16 February 1978. Condor, 80:340-343 @ The Cooper Ornithological Society 1978 INCUBATION BEHAVIOR OF THE temperatures (Ta) during the incubation period DEAD SEA SPARROW (April-August) can exceed 45°C at noon, and rela- tive humidity (RH) may fall to less than 10% (Ros- nan 1956, Mendelssohn 1974). Their large, covered Y. YOM-TOV nest is generally built on dead branches of tamarisk A. AR trees, and its exterior is totally exposed to solar radia- AND tion. Occupied nests are always within 100 m of H. MENDELSSOHN fresh water. The nest is a massive oval structure of twigs (25 x 35 cm), weighing up to 1 kg. An The importance of precise regulation of egg tempera- S-shaped tunnel leads from the top of the nest to an ture during incubation was demonstrated by Lundy incubation chamber lined with soft materials in its ( 1969) in studies of the domestic fowl, in which no lower part. Hence the eggs are largely insulated embryo survived continuous incubation above 40.5”C from direct sunlight while the nest is still well ven- or below 35°C. Similar results were reported for tilated. The eggs are incubated almost exchrsively by pheasant, duck, quail and domestic fowl by Roman- the female. During the day she leaves the nest for off and Romanoff ( 1972). A nest temperature of short periods about six times per hour. When am- 34.O”C c 2.3”C (mean -C SD) was reported by Hug- bient temperatures are high (Ta > 39”(Z), females gins ( 1941) for 37 bird species representing 11 or- sometimes stand at the nest opening and pant. ders, which suggests that most birds have similar Egg temperatures were recorded by using fresh or thermal requirements for successful incubation. The dummy gypsum eggs in which a thermistor was in- optimal nest temperature is primarily achieved by the serted. Preliminary observations indicated no differ-.-- incubation behavior of the parents, and according to ences in temperatures of the real and dummy eggs, White and Kinney ( 1974), the nest attentiveness of and the female incubated the experimental egg nor- some birds is a rectangular hyperbolic function of mally. We recorded egg temperatures (Ti) in four ambient temperature, where the horizontal asymptote occupied nests and six abandoned nests in the Jor- crosses the ordinate at maximal attentiveness time dan Vallev. 25 km S of the Sea of Galilee: in two “L-no- (min/h) in the nest, and the vertical asymptote in- cupied nests in Ein Yahav, 40 km S of the Dead Sea; tersects the air temperature abscissa roughly at and in one occupied nest at the Tel Aviv University optimum incubation temperature. However, they Wildlife Research Centre (WRC). At the same time mentioned only briefly the behavioral responses when we watched and recorded indirectly (from egg tem- ambient temperature exceeds optimal incubation perature records ) the sessions and recesses of the___” temperatures, although heating above optimal tem- incubating females. Daily water losses of the eggs perature may be much more critical for embryonic in the nests and eggshell water conductances were survival than a similar exposure to cold (Romanoff also measured, using the method suggested by Ar et et al. 1938, Drent 1976). al. ( 1974 ) . The aim of this paper is to report how the Dead Sea Sparrow (Passer moabiticus) regulates the tempera- RESULTS AND DISCUSSION ture of its eggs over a range of ambient temperatures. Mean daily temperature of incubated eggs (4 nests, 24 observations) was 33.7”C & 0.5”C (SD). This MATERIALS AND METHODS temperature is well within the range reported for The Dead Sea Sparrow, a small passerine of about 14 other passerines (Huggins 1941, Drent 1976). Rela- g, breeds along the Rift Valley in Israel, where air tive humidity was calculated as follows: mean daily SHORT COMMUNICATIONS 321 5 Ambient Temoerature (Ta 1°C FIGURE 1. The relationship between shaded am- FIGURE 2. The relationship between shaded am- bient temperature (Ta) and the temperature of an bient temperature (Ta) and attentiveness in high egg in the nest (Ti) in three occupied nests and six (Yh) and low (Yc) temperatures in occupied nests. nests where the birds were driven out. Measurements Each point represents the mean of 2-21 one-hour for deserted nests have been omitted (see text). Each interval observations (see Table 1). The hyperbolic point represents a mean of 2-21 one-hour observa- dashed line is calculated for low ambient tempera- tions. ture according to White and Kinney ( 1974). The equations predict attentiveness in percent. water loss of two eggs was 16.52 mg HnO/day in one nest and 16.15 in another (mean egg weight was 1.64 180.4 (where Yh is percent attentiveness at air tem- g). Eggshell water vapor conductance measured in peratures above 36°C; Fig. 1). the laboratory was 0.49 and 0.44 mg H,O/day. We feel that our linear model for Yc is a better mmHg, respectively. Thus, a mean water vapor expression of real incubation behavior of the Dead pressure difference between the inside of the egg Sea Sparrow than White and Kinneys’ ( 1974) rect- and the nest atmosphere of 35.1 torr was calculated. angular hyperbolic model which describes incubation At mean nest temperature of these two nests behavior of Temperate Zone birds and assumes zero (335”C), the RH in the nest is 10.5%, which, as attention time at Ta = Ti. Our calculated curves, proposed previously, is a reasonable result at this besides fitting the data better (F = 24.5; P < O.Ol), incubating temperature ( Ar et al. 1974). The mini- intersect at Ta = 35.1”C, close to the mean Ti, and mal egg temperature measured was 25°C and the predict a minimal attentiveness of 37.3% (22.4 min/ maximum never exceeded 39°C over an air tem- h). This approximates the observed value. The fact perature range of 15-41” (Fig. 1). Ti in deserted that minimal attentiveness above zero suggests either nests followed Ta with a lag of a few minutes, but that the birds enter the nests in order to “measure” they did not differ in hourly mean temperatures. egg temperatures or that other factors are involved. The Dead Sea Sparrow is able to maintain mean egg Assuming that our equations might be extrapolated temperatures below high ambient temperatures, as to 100% attentiveness, then the limits of our model are the larger White-winged Doves (Zenaidu asiatica) are at air temperatures approximately between 1°C in Arizona (Russell 1969). and 45°C (Fig. Z), which correspond to the known Attentiveness and ambient temperature. Figure 2 breeding distribution of this species-between south- and Table 1 summarize mean attentiveness calculated ern Turkey, where they have been seen nesting over from 326 h of observation. Attentiveness (defined snotv-covered ground (Kumerloeve 1965, air tem- by min. h-l or percent) is only a crude estimate of real peratures were not given), and north of the Gulf of thermoregulation because it does not describe how Aqaba where the mean maximum monthly air tem- tightly the brood patch is applied to eggs and the perature during part of the incubation period is 43°C amount of heat transferred to the eggs. For air tem- and daily maximum rarely exceeds 46°C (Rosnan peratures below 35°C we found that the best fit to 1956 ) . our data ( P < 0.01) is a linear model in the form of Since avian embryos tolerate overheating less than Yc = -1.8.Ta + 101.2 (where Yc is the attentive- cooling, it is not surprising that a small rise in Ta ness in percent at air temperatures of 35°C and be- above 36°C is followed by a pronounced increase in low ) . attentiveness. The slope of Yh is 3.4-fold steeper A simple way to avoid overheating eggs is to in- than that of Yc. Attentiveness is of minimum vari- cubate them continuouslv. The eggs-- should then ability at 40°C and 41°C (Table 1). We attribute approximate the body temperature of the parent this to the parents’ urgent need to attend the nest (Russell 1969). As Ta rises, more intimate contact at these high Ta, in order to prevent overheating of and attention are needed to keep egg temperature the eggs. close to body temperature. Thus, at air temperatures Heat and water budget. High ambient tempera- above 35.1”C, our data take the form Yh = 6.2.Ta - tures require a strict heat budget, which may be ap- 342 SHORT COMMUNICATIONS TABLE 1.
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