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P0248-P0255.Pdf SALT WATER TOLERANCE AND WATER TURNOVER IN THE SNOWY PLOVER JAMES R. PURDUEAND HOWARD HAINES ABSTRACT.--Aseries of water-relatedexperiments showed that Snowy Ploversdo not have any outstandingphysiological capabilities for dealingwith the potentialthermal and osmoticstresses of the Great Salt Plains, Oklahoma. SnowyPlovers and SemipalmatedSandpipers failed to maintain body weight when given 0.3 M NaC1 ad libitum. Killdeer did even more poorly by rapidly losing weight on 0.2 M. Thesesalt water tolerancevalues are limited when comparedto other,particu- larly marine, birds that can drink full strength sea water. The Snowy Plover is no better at conservingwater than other birdsin its weightclass. So ploverswithout drinking water were able to maintaintheir weightwhen furnished mealworm larvae ad libitum, the insectivorousdiet aswell as maintenance behavior of the birds is essentialfor their survival on the salt flats. Department of Zoology, University of Oklahoma, Norman, Oklahoma 73069. Present addressof first author: QuaternaryStudies Center, Illinois StateMuseum, Springfield, Illinois 62706. Accepted3 October 1975. THE Snowy Plover (Charadrius alexandrinus) is a small shorebird (30-40 g) com- mon in habitats of high salinity (Bent 1929, Rittinghaus 1961). One such place is the Great Salt Plains in northwestern Oklahoma, a large, barren, salt-encrustedflat where the plover lives and breeds in abundance. The standing water on the salt plains, ranging in chlorideconcentration from 825.0 to 5156.0 mEq/1, is osmotically stressfulto most organisms,yet the Snowy Plover centersits activities around many of these pools and watercourses(Purdue 1976a). The purpose of this study was to determine the physiologicalcapability of the plover for utilizing hyperosmoticsalt water. The plover's natural history and thermal ecologyon the Great Salt Plains are treated elsewhere(Purdue 1976a, 1976b). For comparison certain tests were also conducted on two other shorebirds, the Killdeer (Charadrius vociferus)and the Semipalmated Sandpiper (Calidris pusillus). The Killdeer breeds in normally vegetated habitats around the salt plains, but is seldom encounteredon the salt flats, while the Sandpiper only migrates through the region on its way to breeding groundsin the north. Individual sandpipersoften spend several days or more foraging around the saline waters on the salt flats. METHODS Birds were mist-netted or trapped on the salt fiats and taken to laboratoriesat the University of Oklahoma in Norman, where they were acclimatedto experimentalconditions for at least 2 weeks, fed groundPurina Cat Chow (protein$0%, fat $%, water 9.5-12%), and providedtap water ad libitum. We conductedfive experimentsto determine tolerance to NaC1 solutions:three on Snowy Plovers, and one each on Killdeer and SemipalmatedSandpipers. The concentrationof NaC1 in drinking water was graduallyincreased by 0.05 or 0.1 M incrementsuntil the birds were unable to maintain body weight. The length of time on each concentrationvaried from $ to 11 days dependingon the conditionof the birds. The lengthsof the regressionlines (Fig. 1-$) with respectto the abscissaindicate the duration of treatmentfor eachconcentration. The plover specieswere allowed two days of tap water betweenincremented increases in NaC1 concentration.Inverted bottlesor graduatedcylinders introduced water into small drinking cups. When salt water was given to the birds, they were taught to userodent watering tubesin order to avoid an upward shift in salt concentration by evaporation from the drinking cups. Environmental conditionsin the test chambersvaried slightly between experimentsbecause of the need to cooperatewith other investigators.Two of the Snowy Plover and the Killdeer experimentswere conductedat $$øC, relative humidity (RH) = 28 to 44%, and a photoperiodof 15 h light and 11 h dark (L:D = 15:11).The other Snowy 248 The Auk 94: 248-255. April 1977 April 1977] Snowy Plover Salt Tolerance 249 IOO .IM y=100.42 +O.74X*,N=4 o m i '•.25 My=99.4-0.38XNs,N=3 _ 9o }04My=9905-•26XNs,N=4' " 0 2 4 6 8 I0 12 TIME (DAYS) Fig. 1. Effectof saltwater ad libitum on body weight of SnowyPlovers. N = samplesize at beginning of Test, * = significantregression coefficient (P < 0.05), NS = nonsignificantregression coefficient. Plovertest--when the NaCI solutionwas 0.25 M--and the SemipalmatedSandpiper experiment took place at 25øC, RH = 30%, and L:D = 11:13. Water deprivation testson SnowyPlovers were conductedat 33øC, RH = 28 to 44%, and L:D = 13:11. Mealworms were fed ad libitum to a seriesof ploversdeprived of any free water and body weight was monitored daily. Tritiated water turnoverwas measuredon SnowyPlovers, Killdeer, and SemipalmatedSandpipers at 25øC(L:D = 11:13, RH = 30%) and 40øC(L:D = 11:13, RH = 13%). The birds, which were in a steady stateas indicatedby constancyof body weight, were injectedintraperitoneally with 50/•1 of 0.52/•Ci//•l tritiated water (THO). At leasttwo bloodsamples (10 to 25/•1 plasma)were taken from the brachialvein within 4 h of injection and subsequentlysingle samples were taken daily. Each samplewas centrifuged; the plasma was transferred to a toluene/butylPBD/PBDO scintillation solutionand counted(Beckman LS-133). Half life (T [5) was determined from the slope of the linear regressionbetween the log of rela- tive concentration(Ct) of plasmaTHO and time in days. Ct is the ratio of cpm/ml at time'(t) divided by cpm/ml observedat the initial blood sampling.The equation for determiningthe fraction of water exchangeddaily (M is: 0.693 T« Total water content was estimated from the following relationship: (ctsi)(vi) VH2ø- CtSo whereVH• o = total watercontent in ml, ctsi= cpm/mlinjected, vi = volumeof THO injectedin ml, ctso= the zerotime valueof the equationctst = ctsoe-•t in whichctst is the plasmacpm/ml at t, and t = time in days. Daily water turnover was determinedby the followingequation: V•2o= V•o'X whereQ.•o = dailywater turnover inml/day. Both V.• o and 'OH• owere relativized bydividing bybody weight. All statisticaltests (significance of regressioncoefficients and analysisof variance)were donein accor- dance with Sokal and Rohlf (1969). 250 PIeRDYEAND HAINES [Auk, Vol. 94 c• 105 m- Wy=99.61 1.98XNs,N=3 • •.,M y=99.68-O.41XNs,N=• o m /'/'• (0.2My:101.35-0.45 XNs,N=3 _ 95 _z o z ,,, 85 o 0.4M y=99.34-2.83 X*,N=2 hi D. 8o I , I , I , I , I 0 2 4 6 8 TIME (DAYS) Fig. 2. Effectof saltwater ad libitumon body weight of SemipalmatedSandpipersß Abbreviations given in Fig. I. RESULTS Salt water tolerance.--The Snowy Plover (Fig. 1) and the Semipalmated Sandpiper(Fig. 2) were able to maintain or gain body weight at salinitiesof 0.2 to 0.25 M NaC1. At 0.3 M significant weight lossoccurred in both species,but weight losswas slowand the birds survivedfor morethan a week. Weight lossin the Snowy Plover (Fig. 1) was precipitous at 0.4 M and death would have resulted if the birds I O.IM y=101.50-0.013 XNS, N=6 /....--4 IOO - TAP WATERy=99.74+O53XNs,N=' 9o •0.2My=9728-3.26 XA- • N=6_ o I 2 3 4 TIME (DAYS) Fig. 3. Effect of salt waterad libitum on bodyweight of Killdeer.Abbreviations same as for Fig. i exceptA = significantdifference between days (P < 0.05), but nonsignificantregression coefficient. April 1977] Snowy Plover Salt Tolerance 25 ! • I00 z n- 50 DW O. IM 0.2M 0.3M 0.4M Fig. 4. Fluid consumptionof SnowyPlovers at concentrationsof NaC1. Rangesare indicatedby length of vertical lines, means by horizontal lines, and 95% confidencelimits either side of mean by darkened block. DW means distilled water. had not been given fresh water. In the sandpiper (Fig. 2) weight loss at 0.35 M and 0.4 M was not as precipitous as in the Snowy Plover, but it was considerablymore rapid than at 0.3 M. The sandpipershad no interruption in their salt water regime, but the plovers did, during the 2-day "tap water break" between molar increments. Thus the sandpipers may have been more fully acclimatized to salt water and lost weight at a slower rate. The Killdeer (Fig. 3) did not maintain its weight at salinities above 0.1 M, and weight loss was precipitous at 0.2 M. Variability was great for the 0.2 M values, and the regressioncoefficient was not significant. There was, however, a significant decrease(P < 0.05) in percent of initial body weight between days. Overall, the sandpiper and the Snowy Plover were about equal in ability to tolerate salt water, with the sandpiperperhaps slightly better. The Killdeer was by far the least capable in this respect. Water content.--Water constituted69.6% and 72.8% of the body weight of Snowy Plovers in two separate tritiated water experiments (one at 25øC and one at 40øC) in which the birds were provided with tap water ad libitum. In a third test a group provided with 0.25 M NaC1 was dehydrated to 62% (Table 1). The latter figure is significantly different (P < 0.05) from the other two according to the Student- TABLE 1 MEAN WATER CONTENT AND DAILY WATER TURNOVER FOR SNOWY PLOVERS1 SEMIPALMATED SANDPIPERS,AND KILLDEER.1 Mean Water Daily Water Temp. Body Wt. Content Turnover N (øC) (g) (% body weight) (% body wt/day) Snowy Plover 4 25 33.7 69.59 (+_ 1.85) 43.32 (+_ 1.05) (Tap water) 4 40 32.2 72.75 (+_ 1.07) 48.45 (+-3.31) Snowy Plover 3 25 34.0 62.26 (+_ 2.56) 39.01 (+_ 1.73) (0.25 M NaC1) SemipalmatedSandpiper 3 25 24.3 58.50 (+ 3.05) 54.45 (+- 4.08) (Tap water) 3 40 22.5 55.93 (+_2.42) 56.55 (+- 7.29) Killdeer 1 40 71.1 67.16 46.25 (Tap water) Parenthetical numbersare standard errors. Drinking fluid was provided ad libitum. 252 PURDUEAND HAINES [Auk, Vol. 94 I I ioo 95 88.4 90.0 89.6 88.0 I I I I 2 :5 4 5 6 7 8 TIME (DAY) Fig.
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