Wintering Grounds and Migration Patterns of the Upland Sandpiper

MIGRATION AND DISTRIBUTION

Winteringgrounds andmigration patternsof the UplandSandpiper

Robin P. White

Just now back from the Argentine. Aldo Leopold, A Sand CountyAlmanac (1949)

Photograph/Steveniiolt/ VIREO (h21/ttnacc)

grounds,requested records of specimens teers are an aid to finding some of the rnia Iongicauda)is a shorebirdthat collectedon the winteringgrounds from localities, but some observations have HEfrequentsUPLAND clumpsSANDPIPER of tall and(BARTRA- short museumsin North and South America, had to be omitted because of an uncer- grassin upland grasslands,prairies, and requested unpublished records of Bar- tainty in the date or location of the pastures of the Americas. While the tramia on the wintering groundsfrom sighting.I haveomitted some 87 records conditionof the breedinggrounds is a ornithologists and birders who have of Upland Sandpipersin Latin America critical componentof the reproduction traveled in South America, requested from the wintering distributionand mi- ecologyof Upland Sandpipers(Mitchell sightingsof Upland Sandpipersfrom gration mapsbecause of incompletein- 1967; White 1983), the wintering South American ornithologists and formation. groundsare equally important to a spe- birdersthrough notices in ornithological Second, some reports are detailed, cies'survival (Keast and Morton 1979). newsletters (A. O.U. Ornithological but others give only a general area or This article presentsrecent findings on Newsletter, El Volante Migratorio, time for a sightingwhich makes it dif- thewintering distribution and migration NuestrasAves), met with Argentine or- ficultto placeon a map;e.g.. "abundant patterns of the Upland Sandpiper--a nithologists and birders, and, visited in autumn";"pass through rainy nights speciesof declining numbers and en- potential wintering areas of Bartramia in February and March." I omitted dangeredstatus in some states(Kirsch in Argentina. some 15 records in Latin America from and Higgins 1976; Tate 1981). These methods present severalprob- the wintering distribution and migra- lems. First, published records and un- tion maps becauseof too generala de- publishedobservations may be incom- scription for area and time. Methods plete; records may not have a date or Third, historical changesin range may include1ocational information that must be taken into account. If the dis- To describemore accuratelyand map is not specificas to longitude and lati- tribution of Upland Sandpiperson the the winteringrange and migrationpat- tude. Names given to pinpoint a bird wintering groundshas changed,it may terns of the Upland Sandpiper,I have sighting,for example, may be of minor be inappropriateto map recentsightings searched the literature for published rivers or small villages not found on with records from the late 1800s. After recordsof Bartramia on the wintering many maps. The ornithologicalgazet- organizingthe available records,I des-

Volume 42, Number 5 1247 xgnatedhxstoncal sxghtxngs as thoseprior Phelps1978), "mxgrates to Rio Negro; to the modernizationof farms,drmmng, to 1930, and then marked Mstoncal and accxdentallyto the Malwnas and South and drought.Several shorebird species recentsightings with differentsymbols. ShetlandIslands" (Olrog 1968; and Ol- appearedto be as abundant as in the A fourth problem in preparinga dis- rog 1984). The A.O.U. (1983) describes 1920s but the Upland Sandpiper, de- tribution map is also temporal but on the wintering distribution as "from Su- scribed as one of the most characteristic a shorter time scale. There are records rinam and northern Brazil south to birds of this regionin 1920, was scarce for Upland Sandpipers in thirteen central Argentina and Uruguay." The and "in dangerof becomingvery rare." countries in South America, scattered most detailed descriptionremains that Recent observations from ornithol- throughout most months except June of Bent (1929), although population ogistscontinue, in general, to support and July. Which recordsshould be con- declinesof the bird may render his ac- Olrog's concern. M. Rumboll •ers. sideredto representwintering birds?I count inaccuratetoday. comm.) reports only three sightingsof have plotted sightingsfrom November, Accounts of the abundance of Up- the speciesin Argentina in 14 years December, January, and February, land Sandpipersin SouthAmerica, even (1968-1982); J.P. Myers (pers.comm ), treating thesemonths as the wintering thoseof 60 yearsago, are contradictory. in his 18 months of field work in south- period. Dabbene (1920) and Hudson (1920) easternBuenos Aires province (1973- A final problem is accounting for describedthe bird aswidely and evenly 1974), did not see a single Upland blank spaceson the map. An accurate distributedover the pampasof Argen- Sandpiper;R. Ridsely(pers. comm.) has map of the wintering distribution of tina. In contrast, Wetmore (1927) severalscattered sightings of the b•rd Upland Sandpipersdepends on an in- warned that the bird, "formerly very from his study area in southernBrazil tensivecensus network in the potential abundant," was present in reduced in 1980 and 1981, one sightingfrom range, and because such a data base numbers, and claimed that, due to Paraguayin 1982, severalfrom Bolivia, •s not available, blank spaces on the hunting "Those that remain must seek and only one from northeasternArgen- map representuncertainty--how many the pastures of remote estancias tina in 1982. He calls the winterms blank spacesrepresent birds present but [ranches]in order to survive." groundsof this species"a mystery."In unobserved? Since 1927, only one brief note (O1- contrast,O. S. Pettingill (pers. comm), rog 1967),has been published regarding observedup to 20 birds in one of two population fluctuationsof the Upland different sightingsin northeasternAr- Wintering grounds Sandpiper on its wintering grounds. gentina in 1969 and believesthat "the Publications on the Upland Sand- From 1958 to 1962, Olrog surveyed Upland Sandpiperis among severalof our other North American shoreb•rds piper in South America present a migrant birds in the Buenos Aires sketchypicture of the distribution and Provinceof Argentina. He attributed a that winter in Argentina very com- abundanceof this specieson its winter- declinein the numbersof mostmigrants monly." ing grounds.A descriptionof the wintering rangewas given by Bent (1929) Figure1. Winteringrange of the UplandSandpiper as describedby Bent (1929). (Fig. 1): B 0 L I V I ,:. The main winter range of the speciesis confined to the southern part of South America, north to northern Argentina (Tucaman); and southernBrazil (Irisangaand probably Mattodentro). East to SALTA .• southernBrazil (probablyMatto- dentro); Uruguay (Concepcion, Sta. Elena, and Colonia); and east- ern Argentina (Buenos Aires and Rio Negro). South to Argentina (Rio Negro). West to Argentina (Rio Negro, Mendoza, and Tu- cuman). ATLAN T I C Somefield guidesfor SouthAmerica provide slightly more recent but less detailed range descriptions--"winters OCEAN •nland chiefly in southernBrazil from S•o Paulo south to Uruguay and on

pampas of Paraguay and Argentina CHUBUT south to Rio Negro; a few winter in northern South America" (Meyer de Schauensee1970); "winters principally ß SANTA •n Paraguayand Argentina.Recently found to winter in small numbers in Surinam" (Meyer de Schauenseeand FALKLAND

1248 "Cape Horn present in this area. The only other countries in South America with at least three recordsof Upland Sandpipersin November, December, January, or February are Brazil, Paraguay, and Venezuela. These birds may be over- wintering in these countries,but they may also representlate migrants.

Discussion

The map in Figure 2 portrays two features that deserve discussion: the small number of sightingsand the ap- parentchange in winteringdistribution. Several factors likely contribute to keeping the population low and the rangerestricted. The Upland Sandpiper is protected by law in Argentina, but hunting laws are not strictly enforced either on the national level or among provincesand landowners(Figs. 3 and 4). This lack of protectionled Olrog, as late as 1967, to keep secretthe location of severalUpland Sandpipersto protect them from hunters. Landholdingsare large in some parts of the Argentine Historical: 1 recordo ; pampas and regulation may be based

Recent:5 recordsO 1 record ß; Figure2.Wintering on the discretion of landowners, or of 5 recordsß distributionof the OSouth Shetland Islands local managersfor absenteelandowners. Upland Sandpiper. Some may not care what is hunted on A new map provides the best avail- Sandpipers--in the provinces of Entre their land; others may hunt themselves. able picture of the Upland Sandpiper's Rios, Corrientes and C6rdoba. A total All Argentine ornithologists,however, wintering range(Fig. 2). Out of 318 rec- of 80 birds was counted for the three when questioned,were quick to disre- ords of Upland Sandpipers in Latin locations. gard hunting of this speciesas a possi- America, I used 105 sightingsfrom No- Outside of Argentina and Uruguay, bility; they stated that other problems vember, December, January, and Feb- wintering birds seem rare. Hayer- on the winteringand breedinggrounds ruary to constructthis map. In addition schmidt(1966) describesUpland Sand- (pesticidesand habitat loss)were more to authorscited in the text, other sources pipers wintering in Surinam as "regu- critical factors in Upland Sandpiper for published records include the fol- larly presentin small flocks."Specimens survival. lowing: Ridgway 1919; Beatty 1938; were collectedperiodically from Octo- Predators,other than human, on the Hellmayr and Conover 1948; Howell ber 1965 through April 1966, to docu- wintering groundsdo not seemto pose 1964; Monroe 1968; Land 1970; Karr ment wintering birds, but no thorough a threat. Mammalian predators that 1976; and Spaans 1978. Sightings of study has been conducted of numbers might take a bird 30 centimeterstall are Upland Sandpipersin thesemonths are uncommon, but are strongly concen- Figure 3. Upland Sandpiperhabitat in SouthAmerica: Cattle and windmill on Argentine trated in the area usually identified as estancia. the species'wintering area in Argentina and Uruguay. The records also show that the Upland Sandpiper was observed more often in the Province of Buenos Aires and in Uruguay before 1930; sincethen, the specieshas appar- ently declined in that area. In contrast, records of Upland Sandpipers after 1930 increasenorth and slightlywest of Buenos Aires. In January 1985, I spent one month in northern and central Argentina in searchof Upland Sandpiper wintering sites.I found three areas with Upland

Volume 42, Number 5 not abundant in the grasslands.Rum- Lapwings,however, is not well accepted droughtmentioned earlier (Olrog 1967), boll (pers. comm.) statesthat foxes are amongseveral Argentine ornithologists may account for a decline of Upland not abundant, and that weasels,opos- (R. Fraga, S. Salvador, pers. comm.). Sandpipers in the province of Buenos sums, and other small mammals could First, the lapwingis often found in short Aires. In addition, thesepractices may take a sandpiperonly if it were injured heavily grazed pasturesin contrast to help explain the apparent shift of the or roosting(Fig. 5). Similarly, the num- the mixed short- and tall-grassareas wintering range to areasslightly north- ber of adult Upland Sandpiperstaken most commonly usedby Upland Sand- west and north into the provinces of by birds of prey is not consideredlarge pipers(Fig. 6). Second,food resources Ctrdoba, Santa Fe, Entre Rios, and (R. Fraga, M. Rumboll, S. Salvador, seem abundant on the pampas. Al- Corfientes. Birds may also have moved pers. comm.). though a quantitative study of food farther north to Uruguay, southern Lack of spaceor scarcefood resources abundance is not available, in the Villa Brazil, Paraguay, and Bolivia--areas could be a problem for wintering sand- Maria area, where the largest known that have not been thoroughlysurveyed pipers. Myers (1979) discussesthe lim- concentration of Upland Sandpipers but in which there have been several ited spaceavailable for shorebirdswin- winters in Argentina, the field appeared recent sightings. tering in southern South America, to abound in grasshoppers;insects, in- Favorable changesin thesenorthern comparing land mass at similar lati- cluding grasshoppers,crickets, and and northwesternareas of Argentina tudes on the two continents: "The ex- weevils, are describedas the primary may serve to attract the species. The tent of winter scolopacid habitat in food of this species(McAtee and Beal provinceof Ctrdoba, which hasreports south temperate areas is small relative 1912; Fig. 7). of the most numbers of Upland Sand- to the area used during breeding, and Land-usepractices that affectUpland pipersto date,was originally espinal, or this inducesvery high densitiesin win- Sandpiperson the wintering grounds an area of short, scrubby trees (genus tering populations." He describesag- are not well documented. Several Ar- Prosopis) with a slightly drier climate gressiveencounters between North and gentineornithologists, when questioned and sandiet soils than in the Buenos South American shorebirds and sug- about causes for Upland Sandpipers Aires Province (Cabrera and Willink geststhat there is competition for lim- decline, quickly suggestedthe use of 1973; Fig. 8, this paper). The central ited resources. pesticides. Insecticides were sprayed portion of the espinal belt has been On my visit to Argentina, I observed heavilyin the 1940sto eradicatelocusts, clearedfor dairy farmsand is mainly in only one other shorebird--the Southern and several authors (Barrows 1884; pasture and alfalfa. The Upland Sand- Lapwing(Vanellus chilensis)--feeding Durnford 1877; Dabbene 1920) re- piper is said to prefer the drier climate in closeproximity to Upland Sandpip- ported that the birds ate largequantities and planted grasses,which do well on ers.Myers (1979) describesthis lapwing of theseinsects. While pesticideuse has the sandy soil, to the wetter area with as an extremely aggressivespecies that been correlatedwith eggshellthinning coarsernative grassesfarther south and attacks ferociouslyif other birds come in several carnivorous marine birds and southeast(R. Fraga and S. Salvador, closeto its chicks,probably a response in birds of prey, and with mortality in pers. comm.). All three locations where to limited food resources. On several several songbird populations (Hickey I observedBartramia in January 1985 occasions I did observe the Southern 1961), there is no documentation of were in this espinalregion. Lapwing chase Upland Sandpipers, pesticidedamage to Upland Sandpipers suggestingsupport for Myers' statement on their breedingor winteringgrounds. on aggressivebehavior of the lapwing Changesin agriculturalpractices may Migration patterns and possible competition for food re- be critical to the Upland Sandpiper's sources. limited numbers and to its suggested Important stopoverareas of migrat- A competitive interaction between change in wintering occurrence.The ing Upland Sandpiper have not been Upland Sandpipers and Southern modernizationof farms, draining, and identified. Bent (1929), however, pro-

Figure 4. UplandSandpipers in winteringhabitat: Argentine gaucho Figure5. UplandSandpipers are waryon the winteringgrounds and in background. not easyprey to predators.

1250 American Birds, Winter 1988 Figure6. UplandSandpipers feeding among dumps of tall andshort grass.

vides the most detailed notes available on generalmigration routes of this species. He states that the bird "is not re- corded east of Cuba in the spring, but some birds migrate acrossfrom Yuca- Figure7. Upland tan to Cuba and Florida and then up Sandpiperfeeding the Atlantic Coast." Fall migration is on the wintering describedas "southward, through the grounds. interior and the Atlantic Coast States and through the West Indies to South America." The A.O.U. (1983)provides j,, a more recentsummary of the route-- B 0 t I V "migratessouth through North Amer- ica (rare alongPacific coast from southern Alaska to Washington casually to California, and rare in Arizona, Nova Scotia and the south Atlantic coastal PARANA region), Middle America (not reported northwesternMexico), the West Indies and most of South America (also To- bagoand Trinidad) eastof the Andes." I have constructedmaps of the Up- land Sandpiper's migration patterns usingmethods described earlier (Figs.9 and 10).I used65 sightingsfrom March, April, and May for the Spring map and ATLANTIC 148 sightingsfrom August, September, and October for the fall map. These maps confirm descriptions given by OCEAN Bent (1929) and the A.O.U. (1983) but several additional observations can be made. There are fewer springthan fall recordsfor migratingUpland Sandpipers. espinal Wetmore (1965) describes Upland ß SANTA Sandpipersas found lesscommonly in the spring migration in Panama. pampa FALKLAND Ridgely (1981) alsostates that the bird '• ISLANDS in Panama is an "uncommon to fairly •11an common fall and rather rare spring transient.... "Spring recordsfor the WestIndies are rare.It appearsthat the birds travel south across the Caribbean Figure 8. Biogeographicalprovinces of •4rgentina:Espinal and pampa (after Cabreraand as well as through Central America in Willink 1979)

Volume42, Number5 1251 Figure10. Fall migration patterns of the Upland Sandpiper.

Historical: 1 record o ; 5 records O Recent: 1 record ß ; 5 recordsß

F•gure9, Springmigration patterns of the Upland Sandpipe•

fall but fly north only throughCentral America in spring,although the small number of springrecords from Panama is perplexing.Perhaps there are fewer bird observersand collectors,and thus fewerreports of Upland Sandpipers,in the West Indies and Central America in the springthan in fall. The lengthof the migrationperiod appearsto extend over three or four months in fall and in spring. Records of South American specimens and sightingsare scattered throughout South America from August through May. Historical: 1 record o; 5 records O While some birds may undertake a Recent: 1 record e; 5 records ß fairly rapid flightbetween the breeding and wintering grounds,there also ap- pearto be birdsthat lingerin Central and northern Southern America; mi- gratorystopovers may be lengthyand Conclusion in the four weeks I was in Argentina birdsmay spendas little astwo months than many ornithologistshad seen in on the actual wintering grounds.This Any combinationof the possible the past10 to 15 years.What aboutad- is suggestedby recordsof birdsin Pan- causesdiscussed previously--hunting, jacent ranchesand landholdings?Are ama and Peru in November, and in predators,interspecific competition, Upland Sandpiperspresent there, an Venezuela in November and Decem- pesticides,and land-usechange--may apparenfiysimilar habitat? How many ber. Neither winteringor the year-round accountfor the apparentdecline in Up- of the roadlessor inaccessibleranches presenceof UplandSandpipers has been land Sandpipernumbers. An important are searchedfor upland shoreb•rds? documented in these countries. basicquestion that mustbe addressed, Very fewof the birdinghotspots in Ar- This extendedlength of travel along however,is how rare is Bartrarnia on gentina--IguazfiFalls, Barlioche, Pe- the migration route emphasizesthe its traditional wintering grounds?On ninsula Valdes--include grassland need to preserveinland and upland my brieftrip to Argentina,I found birds habitat and thus are not hotspots for shorebird habitat used by Upland in placesand in numbers(but stillvery Upland Sandpipers.In addition, very Sandpipers,not only on the breeding few)of whichlocal landowners and or- few of the potentialwintering sites an andwintering grounds, but throughthe nithologistswere not aware.In fact, I other South American countries-- Western Hemisphere. found more birds in several locations Uruguay,southern Brazil, Paraguay and

1252 AmericanB•rds, W•nter 1988 Bohvla--are blrded intenselyin poten- Museum of Natural History, Smlthson- KEAST, J A andES MORTON(Eds) Ual Upland Sandpiper wlntenng habi- lan Institution; K. Garrett, Los Angeles 1979. M•grant B•rds •n the Neotrop•cs, Ecology, Behavior, Distribution, and tat County Museum of Natural History;A. Conservation. Smithsonian Inst. Press, Further study on the wintering dis- Jacobberger,Museum of Vertebrate Washington. tnbution of Bartramia in South Amer- Zoology, University of California; J. KIRSCH, L. M. and K. F. HIGGINS. 1976 ma requiresan intensivesystem of cen- Loughlin,Carnegie Museum of Natural Upland Sandpipernesting and manage- suslng. Owners of large landholdings History; J. Navas, Museo Argentino de ment in North Dakota. WildlifeSoc. Bull should be contacted for knowledge of Ciencias Naturales; D. Niles, Delaware 4:16-20. the presenceof the bird, and for per- Museum of Natural History; R. Payn- LAND, H. C. 1970. Birds of Guatemala Livingston Publ. Co., Wynnewood, missionto searchtheir land. The grass- ter, Museum of Comparative Zoology, Pennsylvania. land provinces of Argentina should Harvard University; J. V. Remsen, McATEE, W. L. and F. E. L. BEAL. 1912 serve as the initial search area with ad- Museum of Natural Science,Louisiana Some common game, aquatic and ra- dmonal census work in Uruguay, State University; M. Robbins, Phila- paciousbirds in relation to man. U.S southernBrazil, Paraguayand Bolivia. delphia Academy of Natural Sciences; Dept. Agric. Farmer • Bull. 497:15-16 Only by looking for this bird will we M. Serna, Museo de Historia Natural, MEYER DE SCHAUENSEE, R. 1970. A guideto the birds of South America. Liv- know where it winters. Once major Medellin, Colombia; R. Storer, Mu- ingston Publ. Co., Wynnewood, Penn- w•nteringsites are identifiedthe causes seum of Zoology,University of Mich- sylvania. and extent of wintering mortality can igan; D. Teixeira, Museu Nacional __ and W. H. PHELPS, Jr. 1978. A guide be studied and documented. Universidade Federal do Rio de Ja- to the birds of Venezuela. Princeton neiro; D. Willard, Chicago Field Mu- Univ. Press,Princeton. seum of Natural History; E. Stickney, MITCHELL, G. J. 1967. The Upland Plover and its status in relation to environmental Yale Peabody Museum; B. Webb, ACKNOWLEDGMENTS conditionsand situations,past and pres- Denver Museum of Natural History; ent. Blue Jay 25:58-63. Webber, Florida State Museum, Uni- My trip to Argentina was made pos- MONROE, B. L. 1968. A distributionalsur- versity of Florida. vey of the birds of Honduras. Ornithol sible by a Foreign Travel Fellowship Monogr. 7:112. from the Graduate School at the Uni- MYERS, J.P. 1979. The pampasshorebird versity of Wisconsin-Madison. My community:interactions between breed- travelswere greatly facilitated and en- LITERATURE CITED ing and nonbreedingmembers. Pp. 37- hanced by Bob and Pat Sword and AMERICAN ORNITHOLOGISTS' 49 in J. A. Keastand E. S. Morton (Eds.) Family at Estancia Buena Esperanza, UNION. 1983. Check-list of North Migrant Birdsin the Neotropics;Ecology, Behavior, and Conservation. Smithson- Malcolm and •Anne Marie Pears and American birds, Sixth Ed. American Or- ian Inst. Press,Washington. nithol. Union, Lawrence,Kansas. Family at EstanciaCurupicay, Ana and OLROG, C. C. 1984. Las Aves Argentinas Ennque Castignaniand Family, P. Ca- BARROWS, W. B. 1884. Birds of the Lower Uruguay. Auk 1:315. Administraci6nde ParquesNacionales, Buenos Aires. nevari, R. Fraga, H. Moulin, M. Ojeda, BEATTY, H. A. 1938. Additional records __. 1968. Las Aves Sudamericanas: Una M. de la Pena, M. Rumboll, S. Salvador, for St. Croix, Virgin Islands.Auk 55:131. and D. Wilson. BENT, A. C. 1929. Life Histories of North gula de campo. Univ. Nac. Tucumgn, Inst. Miguel Lillo, Tucumgn,Argentina Unpublished information and help- American Shorebirds. U.S. Nat. Mus. __. 1967. Observaciones sobre aves m•- ful assistancewas providedby B. Abrel, Bull. 146:55-69. CABRERA, A. L. and A. WILLINK. 1973. gratorias del hemisferio norte. Hornero P Alden, K. Anderson, P. Antas, N. .10:292-298. Biogeografiade America Latina. Gen. Banllas, M. Beade, W. Belton, D. RIDGELY, R. S. 1981. Birds of Panama. Sec.Org. Am. States,Washington. Bruning,E. Bucher,F. Cooch,T. Dick, DABBENE, R. 1920. Notas sobrelos chofios PrincetonUniv. Press,Princeton. RIDGWAY, R. 1919. The birds of North P. Donahue, S. Eaton, R. Escalante,T. de Norte America que inviernan en la and Middle America. Bull. U.S. Nat Farr, J. Fitzpatrick, E. Froehlich, S. RepfiblicaArgentina. Hornero 2:99-128. Mus. 50(8):379-385. Furniss,R. Fyfe,A. Gepp, M. Gochfeld, DURNFORD, H. 1877. Notes on the birds of the Province of BuenosAyres. Ibis: SPAANS, A. L. 1978. Statusand numerical M Goodwin, J. Jehl, B. de Jong, R. fluctuations of some North American 166-203. Kantin, C. Leahy, M. Lippsmeyer,J. wadersalong the Surinam coast. Wilson HAVERSCHMIDT, F. 1966.The migration Bull. 90:60-83. Mata, P. Miles, R. I. G. Morrison, C. and wintering of the Upland Plover in Munn, J.P. Myers, T. Narosky, M. No- Surinam. Wilson Bull. 78:319-320. TATE, J. 1981. The Blue List for 1981. Am Birds 35:3-10. res, J. O'Neill, T. Parker, O. Pettingill, HELLMAYR, C. E. and B. CONOVER. WETMORE, A. 1927. Our migrant shore- W. Phelps, M. Pizarro, M. Plenge, I. 1948. Catalogue of the Birds of the birds in southernSouth America. Dept Price, R. Ridgely, S. Robinson, R. Americas. FieM Mus. Natu. Hist. Publi., Zool. Series 13(3):86-90. Agric. Technical.Bull. No. 26, Washing- Rowlett, A. Sada,H. Sick, T. Sherry,F. ton. HICKEY, J. J. 1961. Some effectsof insec- Silva, B. Sorrie, R. Straneck, A. Tarak, ticides on terrestrial birdlife in Middle __. 1965. The birds of the Republic of J. Terborgh, W. Voss, and D. Yzurieta. West. Wilson Bull. 73:398-424. Panama. Part 1. Tinamidae (Tinamous) Data on museum collections used in HOWELL, T. R. 1964. Birds collected in to Rynchopidae(Skimmers). Bull. U.S Nat. Mus. 150. this study were kindly provided by E. Nicaraguaby BernardoPonsol. Condor WHITE, R. P. 1983. Distribution and hab- Alabarce, Fundaci6n Instituto Miguel 66:151-158. itat preferenceof the Upland Sandpiper Llllo; J. Bull and J. DiCostanzo, Amer- HUDSON, J. J. 1920. Birds of La Plata, Vol. 2. E. P. Dutton, London. (Bartramia longicauda) in Wisconsin ican Museum of Natural History; J. Am. Birds 37:16-22. KARR, J. R. 1976. On the relative abun- Dick, Royal Ontario Museum; H. Car- dance of migrants from the north tem- margo, Museu de Zoologia, Universi- perate zone in tropical habitats. Wilson --.Geography Dept., Universityof dadede Sao Paulo; B. Farmer, National Bull. 88:433-458. Wisconsin, Madison, WI 53706

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