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The Water Economy of the Sage Sparrow, Amphispiza Belli

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The Water Economy of the Sage Sparrow, Amphispiza Belli THE WATER ECONOMY OF THE SAGE SPARROW, AMPHZSZZZA’ BELLZ NEVADENSZS RALPH R. MOLDENHAUERl AND JOHN A. WIENS Department of Zoology Oregon State University Corvallis, Oregon 97331 The water balance of desert birds has only distribution somewhat more northerly than recently received intense study. Chew ( 1961)) the Black-throated Sparrow, extending from Bartholomew and Cade ( lQ63), Cade’ ( 1964)) the high sagebrush deserts of eastern Washing- Schmidt-Nielsen ( lQ64), and Dawson and ton south through eastern Oregon and south- Bartholomew ( 1968) have reviewed the early western Idaho to southern Nevada and north- literature on the subject, and, based on these em Arizona (A.O.U. 1957). The breeding early works, the ecological and evolutionary ranges of the two species broadly overlap in aspects of avian water economy in deserts have Nevada. Both species are considered graniv- been discussed (see, e.g., Bartholomew and orous and commonly occur far from surface Cade 1963). In general, it appears that most water. Because of the similarities in distribu- species are dependent on surface water or tion, habitat, and diet of these two closely-re- succulent foods to satisfy their physiological lated species, a comparison of their water requirements for water. However, more recent economies may provide further insight into investigations have shown that certain ,desert the patterns of avian water economy in desert species are sufficiently able to reduce ;water environments. This study provides an analysis losses to survive long periods on a dry diet of of several physiological aspects of the water seeds without drinking. These species i elude economy of the Sage Sparrow. the Budgerygah, Melopsittacus un ulatus ( Cade and Dybas 1962)) and the Zebra inch, METHODS AND MATERIALS :: Taeniopygia castanotis (Calder 1964; C de et EXPERIMENTAL ANIMALS al. lQ%), of Australia; the Cutthroat k inch, Of 35 Sage Sparrows used in this study, 27 were cap- Amadina fa.sciata (Edmonds 1968); Starks’ tured with Japanese mist nets during the summers of Lark, Spizocoys starki, and the Grey-backed 1967 and 1968 in the vicinitv of Cabin Lake Guard Finch-Lark, Eremopterix verticdis (Willoughby Station, 9 mi. N of Fort Rock, Lake County, Oregon, and eight were collected in a similar manner near lQ&3), of Africa. Unionville. Humboldt Countv. Nevada. in November Of the North American desert specjes in- 1966. At the time of capture; the‘ birdswere’ weighed vestigated, only the Black-throated Sparrow, to the nearest 0.1 g with an Ohaus triple beam balance. Amphispiza bilineata, has such capabiIities.2 The birds were transported to Oregon State Univer- sity where all laboratory tests were conducted. An It can survive indefinitely either withoutwater’ additional 47 birds were shot in the vicinity of Fort or on 0.40 M NaCl drinking solution, an d mea- Rock between April 1967 and April 1968 for informa- surements of urine Cl- concentrations indicate tion on body weight and stomach contents. levels higher than for most avian s ties MAINTENANCE ( Smyth and Bartholomew 1966a). The lack- $” Except during salt water discrimination tests, all birds throated Sparrow inhabits some of the most were housed individually in small cages (22 x 24 x extreme desert conditions in North A erica, 41 cm) located in a windowless room with a 12-hr breeding in sagebrush and creosote bus1 des- photoperiod (QQ:45-21:45). Room temperatures gen- erally remained constant at 20°C but occasionally erts from northeast California and southwest- fluctuated from 18.5 to 22°C. Humidity was not con- ern Wyoming south to northern Mexicol trolled and varied between 10 and 30 per cent. Al- The Sage Sparrow, Amphispiza belli, khich though the birds remained in excellent condition on is the only congener of the Black-throated a diet of only vitamin-enriched chick-starter mash, they were occasionally given millet seed, freshly- Sparrow, inhabits similar desert region. One chopped lettuce and Tenebrio larvae. Unless otherwise subspecies, A. belli nevadensis, has a breeding stated, only the chick-starter mash was used during actual experimentation. Water was provided ad libi- Present‘ address: De artment of Biology, Sam Houston tum. The birds were weighed to the nearest 0.1 g State University, Huntsvll .P e, Texas 77341. with a Mettler top loading balance or an Ohaus triple 20hmart and Smith (Auk, in press) have recently shown beam balance either daily or every other day, at the that the Brewers’ Sparrow, Spizella brew&, is also capable of surviving long periods on a dry diet without drinking. end of the dark period. 12651 The Condor, 72:265-275, 1970 266 RALPH R. MOLDENHAUER AND JOHN A. WIENS WATER CONSUMPTION immediately melted shut at one end and centrifuged for 3-5 min at 5000 rpm. The plasma was separated Tap water consumption was measured with inverted by cutting through the collecting tube at the boundary 25-ml graduated cylinders fitted with L-shaped drink- between the clear plasma and the packed red blood ing tubes similar to those described by Bartholomew cells. The urine was separated from any oil and uric and Dawson ( 1954). Two control drinking devices acid residue in a similar manner. Both the plasma were used to measure evaporation, and urine samples were transferred individually to WATER DEPRIVATION capped microfuge tubes and frozen or stored at 4°C until analyzed. Osmolar concentrations were deter- The ability of Sage‘ Sparrows to survive without drink- mined with a Mechrolab Vapor Pressure Osmometer. ing was investigated using six well-hydrated birds An Aminco-Cotlove Chloride Titrator was used for which were deprived of water and provided with only Cl- determinations and a Coleman Flame Photometer a dry diet of mash ( ca. 9.0 per cent water) until death was used for determinations of Na+ and K+. or near death. In order to ascertain the effects of Salt water discrimination tests were carried out with water deprivation on food consumption and activity, groups of four or five birds in a circular cage (61 cm the total energy intake and perch activity of six addi- diam x 61 cm high) made of one-half inch hardware tional birds were measured for five days prior to cloth. Four drinkmg devices were located outside the water deprivation and during four days of water cage at 90” intervals to one another. The two solu- deprivation. Daily gross energy intake was determined tions to be tested were placed in drinking devices by measuring the difference in the dry weight of the opposite one another. Each day at the end of the food provided at the start of the photoperiod and that dark period the drinking devices were rotated 90”. remaining at the end of the 24-hr cycle plus any Measurements were taken for at least six days. Four spillage, which was separated from the fecal drop- NaCl concentrations were tested: 0.1.I 0.15.I 0.20. and pings. Alumniurn trays with sides 12.5 cm high 0.30 M. placed under the cages were used to catch any spilled food. The mash (mean caloric value = 3.944 kcal/g ESTIMATES OF METABOLIZABLE ENERGY dry wt) was equilibrated to room moisture condi- AND EXCRETORY WATER LOSS tions, and samples were taken for moisture determina- tions prior to each feeding. The residual food was Estimates of metabolizable energy intake and excre- dried at 98°C for 24 hr prior to weighing. All weigh- tory water loss were made under three regimens of ings were made to the nearest 0.05 g with a Mettler fresh water intake: ad libitum. restricted (2.0 ml/ top loading balance. The motor activity of each bird day), and no drinking water but with TenebAo larvae was registered on an Esterline-Angus event recorder supplied (40 larvae/day), Metabolizable energy was each time the bird depressed a mechanical perch taken as the gross energy intake minus the excretory located transversely in the cage. energy output (see King 1961). The daily gross energy intake was determined, as previously described, UTILIZATION OF SUCCULENT FOODS for each group for at least five days. Excretory energy was determined for each bird from caloric In order to determine the ability of Sage Sparrows to measurements of two or three pooled 24-hr collections utilize succulent foods as a water source, two groups of fecal material. Collections were made from cages of birds (n = 6 and 10, respectively) were deprived of fitted with metal perches, one-half inch hardware drinking water and given lettuce (95 per cent water) cloth floors, and cork legs about 1.9 cm long. Voided or Tenebtio larvae (60 per cent water) along with fecal material was collected on a sheet of preweighed the regular mash diet. Approximately 15 g of freshly- aluminum foil beneath each cage. The cages and chopped lettuce or about 40 Tenebrio larvae were foil were located on aluminum trays to contain the provided each day. Each group was maintained on dried fecal droppings and spilled food. At the end this regime for at least six days. of each 24-hr period, the foil, wire mesh floor, perches and the trays were removed and replaced. SALT WATER UTILIZATION The fecal material adhering to the floor and perches Sodium chloride drinking solutions were used to was scraped free and combined with the loose fecal ascertain the ability of Sage Sparrows to utilize saline material separated from the spilled mash and not water and to test its effect on the ionic and osmotic adhering to the foil. The gathered fecal droppings composition of the blood and urine. Three concentra- and the foil were dried for 24 hr at 98°C and then tions were used: 0.20, 0.25, and 0.30 M.
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