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Fat Reserves of an Opportunist and of a Specialist Species in the Negev Desert

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Fat Reserves of an Opportunist and of a Specialist Species in the Negev Desert FAT RESERVES OF AN OPPORTUNIST AND OF A SPECIALIST SPECIES IN THE NEGEV DESERT YEHOSHUA $I-IKEDY AND URIEL N. $AFRIEL Departmentof Zoology,The Hebrew University of Jerusalem,Jerusalem 91904, Israel, andthe Mitrani Centerfor DesertEcology, Jacob Blaustein Institute for DesertResearch, Sede Boqer 84990, Israel ABSTRACT.--Weextracted fat from samplesof two sympatric lark speciesresident in the Negev Desertof Israel.The CrestedLark (Galeridacristata), which fed on a wide range of seed sizes,exhibited within-day but not seasonalfluctuations in fat content and carried on average12.9% fat (of lean dry mass).The Desert Lark (Ammomanesdeserti), which fed on a narrowerrange of seedsizes, exhibited smaller within-day fluctuationsin the amountof fat, and carried on average 8.9% fat. We proposethat although the two larks occupythe same environment and are of the same feeding guild, these differencesare adaptive. When food becomesperiodically scarce, feeding specializationsof the Desert Lark may often sufficeto overcomeshortage, but their absencein the generalistCrested Lark may make it more dependableon fat reserves.Received 9 July 1990,accepted 20 December1990. SPECIESthat utilize few food types are ex- within the Negev Desert of Israel, the Crested pected to exploit each of them more efficiently Lark consumedseeds that rangefrom 0.3 to 53.5 than speciesthat utilize a wide range of re- mg in mass,whereas most of the seedscon- sources(Pianka 1981a).If high efficiencyis ex- sumedby the DesertLark were lighter than 1.0 pressedin the ability to exploit effectively a mg. The CrestedLark generally was lesstena- resource even when it becomes rare, then food ciousand, dependingon the time of year, moved specialistsshould be less susceptiblethan food between regions and habitats;but the Desert generaliststo a simultaneousreduction in the Lark was much more sedentary($hkedy 1990). level of most resources. Thus, with respectto both food resourcesand Fat storage,however, can be usedas a buffer habitat utilization where both speciesoverlap, againstfluctuations in food availability (King the Desert Lark is a specialistand behavesas if 1972),but the size of the fat reserveis expected its environment is rather stable. The Crested to be determinedby a trade-offbetween benefit Lark is a generalist, frequently changesfood (reducedprobability of starvation)and cost(in- types and habitats, and behaves as if this same creasedrisk of predation)(King and Farner1966; environment is rather variable. Specialization King 1972;Blem 1975, 1976;$tuebe and Ket- vs. generalization are regarded as adaptations terson 1982; Nolan and Ketterson 1983; Pien- to stable vs. unstable environment (Pianka kowskiet al. 1984;Lima 1986,1988; Rogers 1987). 1981b,Futuyma and Moreno 1988).Because both It can be predictedthat specialistsand species species occur in the same environment, their of stable environments should deposit smaller different adaptationsmay reflect differencesin fat reservesthan generalistsand speciesof vari- the "interpretation" of environmental stability. able environments. We explore whether this implied difference in We comparedfat storagein two speciesof "interpreted" environmental stability and in nonmigratorylarks (Alaudidae).The distribu- mode of resourceutilization by co-occurring tion of the Crested Lark (Galerida cristata) ex- speciesis reflected in differencesbetween them tends from southern Sweden in the north to in fat storagetactics. northern Kenya in the South (Hall and Moreau 1970). Most of this range is nondesert,hence METHODS desertmight be marginalhabitat for the Crested The study site was on Halouqim Ridge in the cen- Lark. The distribution of the Desert Lark (Am- tral Negev Desertof Israel near Sede-Boqer(30ø52'N, toomanesdesertl) is confinedto arid regions(Hall 34ø57'E; 475 m above mean sea level). The terrain and Moreau 1970). Both speciesare ground consistsof stonyhillsides, with sparseshrubs (mostly dwelling, and their diets compriseseedlings, Artemisiaherba alba, Hammada scoparia, and Zygophyl- insects,and a large proportionof seeds($hkedy lum dumosum)(Evenari et al. 1982). Mean annual rain- 1990). We found that in a region of sympatry fall is 91 mm. Rainfalloccurs only during winter (Sep- 556 The Auk 108: 556-561. July 1991 July1991] FatReserves in Larks 557 TABLE1. Mean (+SD) wet mass(WM), lean dry mass(LDM), fat content(F), and fat ratio (FR) of all birds. Level of significance:*** = P < 0.001. Species n WM LDM F FR (%) Crested Lark 118 39.0 _+ 2.7 11.5 + 1.2 1.48 + 0.71 12.9 _+ 7.2 Desert Lark 103 22.0 _+ 2.5 6.6 -+ 0.8 0.59 _+ 0.22 8.9 _+ 3.2 Statistics 59.0'**a 38.9'**a 12.7'*'4 5.47'**b values, value, Mann-Whitney U test. tember to May). Mean monthly air temperaturesvary but in CrestedLarks it was lower in September from 9.8øCin January(the coldestmonth) to 25.5øC than in December (Table 2). On average, fat in July (the hottestmonth). During January,air tem- ratio in both specieswas lower in September peraturesmight drop to -IøC (Zangvil and Druian than in other seasons, but this trend was not 1983;data from meteorologicalstation located 2 km significant(Table 2). There were no significant from the study site). seasonal differences in fat ratio. We obtained our data from 118 Crested Larks and 103 DesertLarks collected from February 1987to July Fat ratio calculated over the entire study pe- 1989.Samples of both specieswere securedfour times riod increasedduring the day in CrestedLarks. a year (hereafter seasons):before the breeding season The highest fat ratio was between 1,500 and (February through March, hereafter February),just 1,700,and the ratio later decreased(Fig. 1). The after the breedingseason (July), when CrestedLarks mean increaseof fat ratio over the entire study return to the study site (September),and December. period was 0.5% per hour. In the pooled data Eachshooting expedition lasted approx. 2 weeks.Right of birds secured in December of 1987 and 1988, aftercollection, we weighedeach bird (wet mass,WM) fat ratio increaseduring the day was higher to the nearest0.5 g (50-gPesola spring balance), mea- than in all other seasonscombined, and higher suredits wing length (WL) to the nearest1 mm (from than that of the Desert Lark (Fig. 1, Table 3). carpaljoint to wing tip; Newton 1969) and sexedit The correlation between fat ratio and time of in the field by dissection.Because the gonadsare small during nonbreedingseasons and there was damage day waslow and not significantin Februaryand causedby the shotgunpellets, we sexedonly 56 Crest- July (Table 3). ed and 49 Desert larks. The birds were deep-frozen The daily increasein fat ratio of Desert Larks 1-4 h after death. We used Moreau and Dolp's (1970) wasmuch slower and not significant(only 0.03% methodsfor determiningdry mass(DM), the amount per hour; Fig. 1, Table 3). of fat (F, extracted with petroleum ether), and lean In February, lean dry mass in both species dry mass(LDM). We then calculatedthe fat ratio by increasedsignificantly during the day (r• = 0.37, calibratingextracted fat with the bird'slean dry mass n = 28, P < 0.05, in the Desert Lark; and rs = (FR = F/LDM) and with wing length (FR1 = F/WL). 0.36, n = 37, P < 0.05, the Crested Lark). Wing The resultsof the statisticalanalysis concerning both measuresof fat ratio (FR and FR1) were similar, thus length also increasedin the Desert Lark (rs = we only present the resultswith fat ratio of fat cali- 0.33, n = 26, P < 0.05) and in the Crested Lark bratedwith lean dry mass. (rs = 0.29, n = 36, P < 0.05). In both species male wet masswas greater than female wet mass, RESULTS and mean lean dry massof Crested Lark males was greater than females(Table 4). There were On average,Crested Larks were heavier, they no significantdifferences between sexesin any stored more fat, and their fat ratio (fat to lean other index. dry mass)was higher than DesertLarks' (Table 1). In July lean dry massof Desert Larks was DISCUSSION 8.7%lower than average;in CrestedLarks, wet masswas 4.4%lower than averageand lean dry There are only a few studiesof fat deposition masswas 9.2%lower (Table 2). This is probably in nonmigratorybirds of nontemperatelati- becausejuveniles (which we did not alwaysdis- tudes (Grant 1965, Ward 1969). The amount of tinguish from adults) were presumably part storedfat (ca. 13%of the lean dry mass)as well of the July samples,and juveniles are lighter as its seasonal fluctuations (with extreme de- than adults (PJtzold 1986). Desert Larks' mean viation of a monthly mean from an annual mean lean dry masswas similar in all other seasons, of <0.3 g fat) in the tropicalYellow-vented Bul- 558 SHKEDYAND SAFRIEL [Auk, Vol. 108 TABLE2. Seasonalvariation (• + SE) in lean dry mass(LDM), fat ratio (FR), and wet mass(WM). • LDM FR (%) WM Desert Lark February 6.76 + 0.12 9.2 + 0.6 21.60 _+0.36 July 6.06 + 0.11b 9.3 + 0.4 21.44 -+ 0.37 September 6.81 + 0.15 7.2 + 0.6 21.80 + 0.48 December 7.18 + 0.17 9.5 + 0.6 22.54 _+ 0.49 Crested Lark c February 12.01 + 0.12 13.4 + 0.9 39.24 _+0.29 July 10.46 + 0.16b 13.8 + 1.5 37.29 + 0.52 September 11.59 + 0.21a 10.2 + 1.0 38.69 _+0.66 December 12.24 + 0.10 a 13.1 + 1.2 40.26 _+ 0.50 aSeasonal variation analyzedby two-way ANOVA. The year of study and the interactionsbetween the year and the seasondid not affect significantlyany of the independentvariables. Comparisons between seasons were evaluatedby Duncan'smultiple range test only when the F valuesof the seasoneffect were found significantby the two-wayANOVA. bDifferent from all other seasons(P < 0.05). • Wet massin July was significantlylower than in Februaryand in December(P < 0.05).
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