Factors Influencing Numbers of Syntopic House Sparrows and Eurasian Tree Sparrows on Farms
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382 ShortCommunications and Commentaries [Auk, Vol. 110 The Auk 110(2):382-385, 1993 Factors Influencing Numbers of Syntopic House Sparrows and Eurasian Tree Sparrows on Farms PEDROJ. CORDERO InstitutoPirenaico de Ecologœa,Consejo Superior de InvestigacionesCient•ficas, Apartado202, 50080 Zaragoza,Spain The House Sparrow(Passer domesticus) and the Eur- was mappedand classifiedinto agriculturalland, in- asianTree Sparrow(P. montanus)show differencesin cluding: (FORAGE)orchards, cereals and associated habitat use. The former is predominantly an urban natural vegetationwhere sparrowsforaged; (FOR- and suburbanspecies, while the EurasianTree Spar- EST)woodlands, including Mediterranean oak, pine row is more rural (Summers-Smith 1963, Pinowski forestand Mediterranean scrub;and (GARDEN) gar- 1967, Lack 1971, Cody 1974, Dyer et al. 1977). How- dens, including groves of several tree speciesand ever, they often coexistalong suburban-ruralgradi- ornamentalshrubs where the sparrowsdid not forage. ents (Cody 1974), where extensivediet overlap (An- Open fields supportedagricultural activities, mostly derson1978, 1984) and nest-sitesegregation (Anderson intensivevegetable growing (89%),with the rest in 1978,Cordero and Rodriguez-Teijeiro1990) have been alfalfa, wheat, barley and maize. found. Some ecological(Pinowski 1967, Anderson To evaluate livestock I considered cattle equiva- 1978)and geographical(Summers-Smith 1963, 1988) lents of domestic animal biomass (LIVESIND; Table evidence suggeststhat habitat differences,in part, 1). To obtain this, total masswas evaluatedusing the may be modeled by competitive interactionsduring following estimatedvalues: 0.325 kg for a domestic which the House Sparrow is dominant. Assuming pigeon(Columba livia); 2.5 kg for a domesticfowl; 3.5 that EurasianTree Sparrowsare nest-sitelimited (An- kg for a rabbit;27 kg for a dog; 100kg for a pig; 450 derson 1978), I have analyzed the habitat character- kg for a cowor bull; and 850kg for a horse.Livestock isticsof House and Eurasian Tree sparrowson farms exceptpigeons, dogs, and occasionallydomestic fowl where nest boxes were introduced to increase the were in enclosedbuildings; with the presenceof live- available nest sites. stock,an abundantfood supplywas providedfor the My objectivewas to determine the effectsof dif- sparrows(both vegetablesand insects;Dyer et al. 1977, ferent habitat variableson the breeding numbers of Lowther 1983). both House and Eurasian Tree sparrows on farms I usedthe possiblepredictor variables summarized and at rural residences. I have examined the rela- in Table 1. Other possible variables, such as year of tionships between variables related to food, nest site, investigationin the plot (in which nest boxeswere and the number of individuals of each species,both introduced), elevation, distance to coast, distance to in natural holes and introduced nest boxes. nearestfarm or rural house,nest box support(tree or I studied 14 farms or houses in rural areas of Bar- wall), and climatic features were excluded because celona Province in northeastern Spain. All siteswere the range of variation was minimal, or earlier de- used by man and were isolated, surroundedby cul- scriptiveanalyses indicated that the variable was not tivatedfields or forest.Other hole-nestingspecies were important for this analysis. absent except for a few pairs of European Starlings Linear-multiple-regressionanalyses (Norusis 1985) (Sturnusvulgaris) at one of the farms. I put up 10 to were used to determine which of the factors best ex- 131nest boxes (Table 1) on buildingsor on the nearest plained the observed numbers of House Sparrows trees adjacent to them. In the breeding season I and EurasianTree Sparrowsin the plots. Before the searchedintensively for occupiednests (as in Cordero analyses,I transformedthe variableslogarithmically and Rodriguez-Teijeiro1990) and checkednest boxes (log) sothat they were normallydistributed and had at three- to four-day intervals to determine whether homoscedasticvariances (e.g. Zar 1984). The factors they containedcompleted clutches. The number of that could affect the numbers of birds (Table 1) were sparrowswas determined basedon the total number entered into the analyseswith forward stepwise in- of nest sites occupiedand for which there was evi- clusion of variables (Norusis 1985). Four regression denceof breeding.Also, cavities potentially available models were developed. In two, the dependent vari- for nesting were noted. ableswere the total numberof House•parrow nests Variablesassociated with food availability were re- (ALL-HS) and the total number of Eurasian Tree lated to vegetationcover and livestock(Dyer et al. Sparrownests (ALL-ETS). In the other two models, 1977, Lowther 1983). The vegetationalcover in the the dependent variableswere the number of House 100 m around eachplot was measured,assuming that, Sparrownests in nestboxes (HSBOX) and the number normally, foraging trips were within that distance of EurasianTree Sparrow nests in nest boxes (ETS- (Summers-Smith1963, pers. observ.).The vegetation BOX).For eachfinal equation,the regressionresiduals April 1993] ShortCommunications andCommentaries 383 T^n•,E1. Designation and descriptionfor variables included in multiple-regressionanalyses, with means, standard deviations and ranges observed at 14 sites. Code Description œ+ SD Range FORAGE Percent cultivated land and 62.43 + 22.72 30-99 associatedvegetation cover FOREST Percent forest + scrub cover 29.28 + 24.25 0-60 GARDEN Percent grove and scrub 4.29 + 7.28 0-20 garden cover TREES Percent FOREST + GARDEN 33.57 + 22.57 0-60 LIVE$IND Livestock index a 11.86 + 26.93 0-98 CAVITY No natural cavities 22.00 + 21.72 3-70 BOXES No nest boxes 31.21 + 29.47 10-131 HSCAV No House Sparrow nests in 12.71 + 15.05 0-43 natural cavities ETSCAV No EurasianTree Sparrow nest in 1.14 + 1.23 0-4 natural cavities HSBOX No House Sparrow nestsin 3.78 + 6.09 0-22 nest boxes ETSBOX No Eurasian Tree Sparrow nests in 5.35 + 7.99 0-31 nest boxes RHSCAV. Ratio HSCAV/CAVITY 0.45 + 0.76 0.0-0.9 RETSCAV Ratio ETSCAV/CAVITY 0.13 + 0.17 0.0-0.5 RPSCAV Ratio (HSCAV + ETSCAV)/CAVITY 0.57 + 0.23 0.0-0.9 ALL-HS No House Sparrow nestsb 18.00 + 18.34 0-50 ALL-ETS No Eurasian Tree Sparrow nestsb 6.71 + 9.23 0-36 • Cattle equivalentsof domesticanimal biomasscalculated as the sum of estimatedmean massand total numbersof all domesticanimal species in plot divided by estimated mean massof cattle (450 kg). • Nests included that were in open (i.e. not in cavities or nest boxes). met the assumptionsfor linear-multiple-regression explained 57% of total variance (Table 2). The nega- analysis(e.g. Norusis 1985). Results are given as )• + tive association with TREES is related to food avail- SD. ability. Food overlap between House and Eurasian Both House and EurasianTree sparrowsoccurred Tree sparrowsmay be considerablein the breeding in 12 (86%)of the plots (n = 14), but the HouseSpar- season (Anderson 1984). However, some differences row was more numerousthan the EurasianTree Spar- in foraging ecology (a consequenceof greater attach- row (Table 1; Wilcoxon matched-pair signed-ranks ment of House Sparrowsto humans)should exist be- test, z = -2.118, two tailed, P = 0.034). The number tween the two species.The House Sparrow could ob- of available natural cavities (CAVITY) was the most tain food both from agricultural areas (including important factor affecting the total number of breed- associatednatural vegetation)and from areasclose to ing House Sparrows(ALL-HS) explaining 54%of the human dwellings (e.g. food waste, livestock),while variation (Table 2). Variables related to food avail- the EurasianTree Sparrowmay obtain mostof its food ability (i.e. livestockindex, percent of foraging areas) from agricultural areasin open fields where natural were not determinantsfor ALL-HS or, at least,they vegetation is common. An increaseof woodland and were not so within the range of variation found in gardens with tree plantation reduces the potential the plots investigated(Table 1). LIVESIND, although foraging areas (r = -0.99, P < 0.001), and this may approaching significance,did not enter in the re- limit the carrying capacityof the habitat for the Eur- gression equation. Although the relationship be- asian Tree Sparrow. This result does not contradict tween the House Sparrow and presenceof livestock the findings of another study on habitat selectionby is well documenteddue to the additional food sup- the Eurasian Tree Sparrow, in which forests repre- plies (e.g. Summers-Smith 1963, Dyer et al. 1977, sented a secondarybreeding area for the species(Pi- Lowther 1983),agricultural land alsosupported high nowski 1967). In Pinowski's study, nest boxes were densitiesof sparrows.The sparrowsconcentrated in put up in villagesand forests,while in mine the nest isolatedbuildings amid orchardsand croplands;old boxeswere placed only near buildings. In both sit- roofs (with pottery tiles) in these areasprovided nu- uations, EurasianTree Sparrows needed to increase merous nest sites. foragingdistance from the nestswhen forestcoverage The total numberof EurasianTree Sparrows(ALL- was extensive. ETS)was positively related to the number of available Although the numbers of nest boxesoccupied by nest boxes(BOXES) and negatively to the percent of the two specieswere similar (Wilcoxon test,z = - !.27, total tree and scrub cover (TREES),which together two tailed,P = 0.20),proportionally, the EurasianTree 384 ShortCommunications and Commentaries [Auk, Vol. 110 TABLE2. Coefficients for multiple-regression