Effects of Raptors on the Activity of Sandgrouse
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j Raptor Res. 28(4):236-241 ¸ 1994 The Raptor ResearchFoundation, Inc. EFFECTS OF RAPTORS ON THE ACTIVITY OF SANDGROUSE PETER N. FERNS AND SHELLEY A. HINSLEY Schoolof Pure and Applied Biology,University of Wales,College of Cardiff, P.O. Box 915, Cardiff,CFI 3TL, UnitedKingdom ABSTR•CT.--Raptorsaccounted for 55% of the disturbanceincurred by two speciesof sandgrouse(Pterocles alchataand P. orientalis)in breedinghabitat in La Mancha, Spain, during springand summer.The main speciesof raptors were the golden eagle (Aquila chrysaetos)in spring and Montagu's harrier (Circus pygargus)in summer.During the breedingseason, Montagu's harriers carried out the majority of their foraging at thosetimes of day when male sandgrousecollected water for their chicks.This presumably enabledthe harriers to locate chicksmore easily. Sandgrouselost no more than 0.5 hr of feedingtime per day as a result of disturbanceby raptors, and yet were still able to rest for at least 3 hr a day at timeswhen they couldhave been feeding. Predators as a whole were probablythe major factorresponsible for the different dispersionpatterns adopted by sandgrouseat different times of year. KEY WORDS: goldeneagle; Montagu's harrier; Pteroclesalchata; Pteroclesorientalis; raptor disturbance; sandgrouse;vigilance. RESUMEN.--Avesrapaces fueron responsablespor 55% de las perturbacionessufridas por dos especies de gangas(Pterocles alchata and P. orientalis)en susareas de reproducci6nen la Mancha, Espafia,durante la primaveray el verano.Las especiesde avesde rapifia mils prominentesfueron el figuila real (Aquila chrysaetos)en la primaveray el aguiluchocenizo (Circus pygargus) en el verano.Durante la estaci6nde reproducci6nlos aguiluchos cenizos 11evaron a cabo la mayorlade su forrajeoen las horasdel dia en que las gangasmachos estaban recogiendoagua para sus pollos. Esto supuestamenteles permiti6 a los aguiluchoslocalizar los pollosmils facilmente.Las gangasperdieron no mils de media hora del tiempo de alimentaci6ndiario debidoalas perturbacionespor avesde rapifia sin embargoestuvieron inactivos por lo menosdurante 3 horasal dia cuandohubieran podido estar alimentando.Las avesde rapifia en generalfueron probablemente el factormils importante en determinarlos distintos partrones de dispersi6n de las gangasen distintas•pocas del afio. [Traducci6n Autores] Raptors can exert a significantinfluence on the rowhawksattacked particular tit (Parusspp.) flocks flocking behaviorof birds, both in the short term about once per day on average,and disturbedthe and the long term. In responseto attacksby spar- birdsfor only about2 min. Longerdisturbances can rowhawks (Accipiternisus), redshanks (Tringa tot- beinferred from detailedstudies of merlinsand spar- anus)deserted their winter feedingterritories for up rowhawks attacking wintering waders (Page and to an hour and formed flocks, while turnstones (Ar- Whitacre 1975, Boyce 1985, Cresswelland Whit- enariainterpres) reduced their individualspacing from field 1994), thoughno actualestimates are available threebird lengthsto onebird length(Whitfield 1988). in these cases. Myers (1984) found that winter feedingterritories In order to derive estimates of the total time lost were abandonedby sanderlings(Calidris alba) in by pin-tailedsandgrouse (Pterocles alchata) and black- years when merlins (Falcocolumbarius) were more bellied sandgrouse(P. orientalis)as a result of dis- abundant. These studies focused on the influence of turbanceby raptors, we recordedthe amount of time raptors on the flocking and vigilance of potential the birds spendbeing vigilant, the number of inci- prey (seealso Metcalfe 1989). The amountof time dentsof disturbanceand the factorsresponsible. This for which target organismsare preventedfrom car- was done as part of the compilationof time and rying out otheractivities has seldom been measured. energy budgetsfor thesetwo speciesof sandgrouse In one early study, Morse (1973) found that spar- in Spain. 236 DECEMBER 1994 RAPTOR DISTURBANCE OF SANDGROUSE 237 STUDY AREA AND METHODS Table 1. Number of raptorsobserved while compiling Observations were made in semi-arid farmland in the sandgrousetime budgets. provinceof La Mancha, Spain, in spring (29 March to 21 April) and summer (22 July to 10 August) 1986. NUMBER OF Sandgrouseoccurred in flocks during the spring, com- SIGHTINGS prisingon average26 (SD = 23, N = 31) individualsin pin-tailed sandgrouse,with a maximum flock size of 75. OF EACH Most daytimeflocks of black-belliedsandgrouse contained RAPTOR 2-7 birds, though slightly larger groups (maximum 28) SUM- roostedtogether overnight. In summer, both speciesoc- SPRING MER curredin scatteredfamily groups.When compilingtime budgets,we scan-sampledbird activity at 1 min intervals. N= iV= In both seasons,the day was divided into 10 equal time RAPTOR SPECIES 23 da 20 da periods.These time periodswere longer in summer (92 Black kite (Milvus migrans) 4 min) than in spring (82 min). The field area and study Red kite (M. milvus) 6 methodsare describedelsewhere (Hinsley 1994, Hinsley and Ferns 1994). A total of over 46 000 bird observations Montagu's harrier (Circuspygargus) 1 23 were made,involving on average29 pin-tailed sandgrouse Common buzzard (Buteobuteo) 3 4 and 6 black-belliedsandgrouse each day in spring, and Imperial eagle (Aquila heliaca) 2 family groupsof 2-5 pin-tailed sandgrousein summer(at Golden eagle (A. chrysaetos) 11 leasttwo different familieswere watchedeach day, and Bootedeagle (Hieraaetuspennatus) 1 over 40 familieswere studiedin total). Every potential Common kestrel (Falco tinnunculus) 1 predatorthat was observedclose enough to the focal birds Peregrine falcon (Falcoperegrinus) 1 2 to disturb them was recorded. If the same source of dis- Little owl (Athene noctua) 1 turbance causedmore than one group of birds to react, this was treated as a single incident, even if the events occurreda considerabletime apart (actual maximum = 7 min). sandgrouse adopted alert upright postures or The observationsin the spring were undertaken at a crouchedclose to the ground (thesebehaviors were rangeof 300-1000 m (typically700 m) to avoiddisturbing alsorecorded as vigilance). the birds, and it was not possibleto distinguishgender at The most common identified source of disturbance this time of year. Observationsin the summer were un- was raptors. During a total of 43 d of observations, dertakenat a range of 200-800 m (typically 400 m). In the summer,groups split up into family partiesin which the number of disturbancesdue to raptors was 60. males,females and juveniles (1-2-mon-old at the time of A further 49 disturbances were attributed to other the observations)were distinguishable. causes.The speciesof raptors involved (Table 1) To estimatethe amountof time that sandgrousespent differed significantlybetween spring and summer, beingvigilant when disturbed,we performedstepwise re- gressionon the percentageof time spent being vigilant though the number of incidentswas very similar. (arcsinetransformed) in eachof the 10 daily time periods The mostimportant species in springwas the golden againstthe number of recordsof disturbancecaused by eagle. The most important speciesin summer was all factorsthat we were ableto identify.These were divided Montagu's harrier (goldeneagles were absentfrom into five categoriescomprising the numbersof the most the area during the summer).Because the samein- commonraptors in eachseason (golden eagle [Aquila chry- saetos]in spring, Montagu's harrier [Circuspygargus] in dividualraptor may havecaused disturbance on more summer);other raptors; other birds (mainly red-legged than oneday, theremay havebeen some unavoidable partridges[Alectoris tufa] and great grey shrikes[Lanius pseudoreplication.However, the proportion of dis- excubitor]);mammals (rabbits, sheep and dogs);and hu- turbancescaused by the mostabundant raptor (com- mans(including noises caused by humanactivities). This method of measuring the affect of disturbancehas the pared with all other birds) in the two seasonswas advantageof taking into accountlosses of feeding time significantlydifferent (Fisher's exact test, P < 0.005). through increasedalertness even when this lastedfor some A greater diversity of raptors was thus responsible time after the disturbance ended. for disturbanceduring the spring (Table 1). Red kites (Milvus migrans) and black kites (M. milvus) RESULTS were recordedin springbut not summer,both being In the absenceof disturbance,sandgrouse vigi- migrantsin the area. A flock of 106 black kites that lancewas typically restrictedto shortbouts with the passedover on migrationon the eveningof 3 August headup. The durationof thesebouts was longestin was not included since it caused no disturbance. male pin-tailed sandgrouseaccompanying chicks (• Golden eagleswere observedcircling quite high = 6.5 sec, SD = 3.9, N = 47). When disturbed, (about30-50 m) overthe fieldsand hill slopes,and 238 PETER N. FERNS AND SHELLEY A. HINSLEY VOL. 28, NO. 4 14 a. SPRING alsoperched on vantagepoints overlooking the fields. Both goldeneagles and red kitesregularly take birds 12 the size of adult sandgrouse(Cramp and Simmons 10 1979). In spring, a juvenile imperial eagle (Aquila heliaca)took a taxidermic mount of a sandgrouse 8 •Pin-tailedS.vigilance Black-bellied /,• (usedto measureoperative temperature) from the S. vigilance // \ \ Numberof raptors 6 groundand was only preventedfrom flying off with it by the attacheddata-logger. Montagu's harriers 4 and red kitesquartered the fieldsat a heightof only // .- \ a few meters,and occasionallycircled