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Breeding Ecology of the Water Pipit

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Breeding Ecology of the Water Pipit BREEDING ECOLOGY OF THE WATER PIPIT NICOLAAS A.M. VERBEEK TaE Water Pipit (Antbus spinoletta) is a holarctic bird breeding in alpine and arctic habitats. Probably becauseof its remote distribution, little informationon the speciesis available. In North America,Johnson (1933) reports observationson one nest of A. s. rubescensin Labrador. Pickwell (1947) givesdetails on adult behaviorand on three nestsof what was probablyA. s. pacificuson Mt. Rainier, Washington. Suttonand Par- melee(1954) givedata on 14 nestsand discusssurvival problems on Baffin Island. Irving (1960a) discussesnutritional conditionsof pipits arriving on their Alaska breedinggrounds. Drury (1961) reportson one nest from Bylot Island, Northwest Territories and discussesecological relationships betweenthe pipit, Horned Lark (Eremophila alpestris), Lapland Longspur ( Calcarius lapponicus) , and Snow Bunting ( Plectrophenaxnivalis ) . In Europe, Gibb (1956) studiedfood and ecologyof A. s. petrosusalong the coast of Cornwall, England, and Ringleben (1957) deals with migration and habitat preferenceof A. s. spinolettain Lower Saxony,Germany. This paper deals with A. s. alticola. The study was conductedfrom 14 June to 21 September1963 and from 12 June to 20 August 1964. The ob- jectives were to obtain details on the species'breeding biology and its adaptationsto breedingin the alpine environment. STUDY A•EA The main study area (Figure 1) was about 116 hectaresof alpine tundra on the BeartoothPlateau, Wyoming, circa 45ø N and 10.9ø 30' W. The averagealtitude is 3,200 m, with a rangeof about 140 m. Barnberg(1961) and Johnsonand Billings (1962) describethe vege- tation; Nim]os and McConnell (1962) the soils; Nimlos et al. (1965) the soil temperature and moisture conditions. For a description of the vege- tation relevantto this study seePattie and Verbeek (1966). The alpine climatetypically has coolair temperatures,intense radiation, great diurnal temperatureextremes, strong winds, and snow almost any month of the year. Table 1 showsclimatic data for the 1963 and 1964 seasons.Because pipits spendmost of the time on the ground,I measured groundlevel temperaturesat 5 cm aboveground at onestation in the valley and one on Pipit Hill in 1964. The 1963 breedingseason was much wetter with more snow, hail, and sleetthan the 1964 season.Snow falling in July and Augustusually stays on the ground only for the duration of the storm or for a short time there- after. Thundershowers,accompanied by hail and occasionalsleet, are 425 The Auk, 87: 425-451. July 1970 426 N•coz^^s A.M. VERBEEK [Auk, Vol. 87 • DRY MEADOW _• MOIST MEADOW -'.':-'• SNOW :.:•ß WET MEADOW ',•':'-Y•-:.. FELLFIELD ACCUMULATION Figure 1. Topography of the study area and distribution of nests of the Water Pipit in 1963 (circles) and 1964 (dots). Contour interval is 30 m. commonthroughout the season.Depth of snowdeposited during the winter varieswith the degreeof exposureand is modifiedby "the prevailingwinds during and immediatelyfollowing the winter snowstorms" (Johnson and July 1970] Water Pipit 427 TABLE 1 MEAN TEi'vIPERATIJREAND TOTAL PRECIPITATION ON Tile STUDY AREA Max. temperature Min. temperature Precipitation (øC) (øC) (mm) Standard Standard shelter Ground shelter Ground Rain Snow Period 1963 1964 1964 1963 1964 1964 1963 1964 1963 1964 15-30 June 11.0 8.8 -1.4 0.2 11.2 21.6 9.7 7.8 1-31 July 15.3 17.0 30.8 3.2 3.9 1.1 54.1 23.7 5.1 0.0 1-31 Aug. 15.1 12.7 27.2 1.7 0.2 -2.2 12.9 18.3 3.1 220.6 1-20 Sept. 11.2 0.6 63.7 6.6 Billings,1962). In 1963 higher temperaturesin late June removedthe snow much more rapidly than in 1964. SPRING ARRIVAL Pipits of both sexeswere presenton the study area when I arrived both years. The road crewstold me that pipits are already on the plateauwhen the passis openedfor traffic the last week of May. Wide variations in their numberssuggest that in the prenestingperiod pipits move from the alpine meadowsto lower altitudeswhenever the weather becomessevere, and return as soonas conditionsimprove. Presenceof severalsingle males duringthis periodindicates that at leastsome males arrive on the breeding grounds unmated. HABITAT AND NEST SITE SELECTION As the birdsarrive before all the snowhas left, the first birdsoccupy snow-freeareas as and wheneverthey appear. More suitablehabitat was availableearlier in 1963, and nestswere found then in placesthat were either too wet or still snow-covered on the same date in 1964. The location of eachnest was determined with the aid of a transit (Figure 1) with ref- erenceto knownbench marks and plotted. In generalthe nestswere con- fined to the faceof Pipit Hill, the Central Ridge,the north and west faces of West Hill, and scattered throughoutthe valley. Within these four generalareas, especially on Pipit Hill, the birds apparentlyselected nodes of territorieseach year, which left much seeminglysuitable nesting terri- tory unoccupied. In 1963 I banded 102 pipits, including 9 adult females. One of these adult femalesreturned in 1964and nestedabout 400 m fromher previous site. Four banded juveniles also returned. Two nests in 1964 were built on top of nestsof the previousyear, and at ]east three nestswere built on top of nestsbuilt prior to 1963. 428 NICOLAASA.M. VERBEEK [Auk, Vol. 87 Territories occupied a great variety of terrain from fellfield to wet meadow,on level to steeplysloping ground, and bare to well-vegetated surface. Two habitat requirementsare apparent: 1) the site must become snow-freeearly enoughin the season,and 2) it must have some rough featuressuch as tussocks,tilted rocks,or erodedspots for nest sites. Soper (1946), Wynne-Edwards (1952), and Sutton and Parmelee (1954) all state that the Water Pipit prefersmossy slopes with southern exposure. In the Riesen Mountains of Czechoslovakia,Hanzgk (1958) found A. s. spinolettais dependenton the presenceof rocky slopesand boulderfields and showsno preferencefor wet areas. This appliesalso to the Beartooth Plateau. TERRITORY In 1963 most territories were already establishedwhen I arrived. In 1964 before continuousgood weather set in, pipits abandonedtheir terri- tories frequently. Those birds that stayed during snowy periods showed little territorial activity except for an occasionalchase. Sometimesbirds flushedfrom what appearedto be their territory and flew away for half a kilometer or more. Throughout these first unfavorable 10 days in 1964, the flight songcould be heard. Once the weather changedfor the better, territorieswere set up rapidly and much chasingoccurred. In 1963 I marked and measuredthe territorial boundariesof five pairs. They varied in extent from 1,580 to 3,355 squaremeters and averaged 1,810 squaremeters or roughly an area of 40 by 45 m. These figuresshow large parts of the study area unoccupied.Many suitablenest sites became snow-freetoo late in the season,after mostbirds had establishedterritories. Also about one-sixth of the 116 hectares consistedof unsuitable nesting terrain, suchas permanentsnow fields and water surfaces.Drury (1961) on Bylot Island, Northwest Territories, gives territorial sizes of "about 100 yards and 200 feet high," or approximately5,600 squaremeters, or roughly three times as large as thoseon the BeartoothPlateau. The number of known nests in 1963 was 47, and in two known terri- tories I did not locate the nests. In 1964 I found 52 nests. The estimated total for each year was 60 nests, or a population density of 0.5 pair per hectare. DISPLAYS OF THE MALE Threat display.--Males often approacheach other almostcasually while feeding,with their wingsdrooped and their tails bobbing. In this and other male displaysthe wings do not vibrate. The approachis not direct. The birds feed back and forth until they are closeto eachother, and one then flies at the other. The attacker may pursue the other bird for some dis- July 197o] Water Pipit 429 tance,often beyondthe attacker'sterritorial limits. Upon returningthe territorial male often settles on an object higher than the surrounding terrain, suchas a boulderor a tussock,where he may preen himself,but usuallyjust sitsand looksaround alertly. Alert display.--Themale standswith his tail elevatedbut not bobbing and his wingsdrooped. He often assumesthis postureon return to his territoryfrom a chase,and alsowhen another pipit flies overhis territory. When the displayis directedto anotherpipit flying overhead,it is ac- companiedby a "pititititi" call. Flight display.--In this displaythe male risesat an angleabout 40 m into the air and then descendsto the groundagain singing "tjwee, tjwee" continuously.During the descenthe raisesand spreadshis tail widelyand extendshis wings fully, so that the innermost(ninth) secondaryis free from the body. In the descentthe phrasesof the songfollow each other more rapidly than during the rise. Often the bird continuessinging for a few secondsafter reachingthe ground,and occasionallyhe singson the ground without a flight display. Throughoutthe early phaseof the breedingcycle when territorial be- havior is strongest,i.e. from the establishmentof the territory until part- way throughincubation, males often walk aboutwith droopedwings and elevatedtail. The tail may or may not bob. This displayis a mixtureof the first two described above. As the female does not behave in this man- ner, it helpsto distinguishthe sexesin the field. NESTS Nests are built on the groundand are more or lessoverhung by rock, sod,or vegetation(Pickwell, 1947; Suttonand Parmelee,1954; Drury, 1961). The 99 nests I located were of four types. Sod nests (32) are placedat the foot of a tussockor inside clumpsof vegetation. Erosion nests(22) are in placeswhere the turf has beenbroken by erosion;some nestsin this categoryare moreexposed than any other type becauseof the absenceof vegetation. Rock nests (28) are situatedunder the edge of a rock and usuallyare sunkso deeplythat the wholenest is overhung.Hum- mock nests(17), placed on the sidesof hummocks,are confinedto the low, wet parts of the valley that remainwet mostof the summer,leaving only the dry exposedhummocks available for nest sites. The six nestsWynne-Edwards (19'52) reportson Baffin Island all faced betweenSW and SE. On my study area 87 per cent of the nests faced betweenW and NE with the highestnumber (28 per cent) facingN.
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