WESTERN BIRDS ■ Vol. 10, No. 3, 1979 WESTERN BIRDS f JuarterEy Journal inf Western Field Otltllholcr^b'i fttWwiertf John S LLathnr V- r Jcc -Prtf sided . Ttwntfl H Wsh .1 ffi'jituri'i* Phvl Schaeffer Ate m bershrp Secretory Schfurfftu Seerafury PhyJlia Layman *%cr*4ary Linda Delaney Dkedon: Laurence C Banfafdi, Jianh* A. Corvry. t^Slfld F. DffSanpa, Stephen „A 7 Layman,. Jylifi S Luther,. Guy McCaikle. Richard W, SfalKcup, Tw*fV« R Wahl, Jdrtfft Wibvnirvn Edftffl- Alan M Craig Editor No ix-q A Moore FdlfocHaJ Buerrd Robert AndfttfWL Alan BiMrldg* William H. 33ehlc. Andrew J Berger Laurent* C Eiiniord fChnirruftni ileann* A Cnnry-, David h DeSanEu, Richard ElKhsOrt, Jowph DlltnWl^. Joseph R X:h\ Ji , Ned K Johnson Virginia P. Johnson , Brin a KgswI, Charles S Lawn^ti, Stephen A i_aymon, John S Luther, Tbn Mandli Brian J. MeCaff«j|[i Guy McC-aiki*, M, lipnolhy Myr** Harry B Nehcs ThomaE L Rodgers Stephen M Russell Olsver K Sco!!. P David Skftftr Rkrhsrd W Stnlhrup, David Stirling. G. 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I iliun.in, Cnlii-Ts.lR ^49^ [1 Make checks pa uahk 1 tu Western Field OtnllholagwU A 15% commission sa nll-o-wed Ecu agembti. WESTERN BIRDS Volume 10, Number 3, 1979 OCCURRENCE AND DISTRIBUTION OF THE MOTTLED PETREL DAVID G. AINLEY and BILL MANOLIS, Point Reyes Bird Observatory, 4990 Shoreline Highway, Stinson Beach, California 94970 The Mottled Petrel ( Pterodroma inexpectata) is much reduced from its former numbers. Due to the clearing of forests and predation by introduced mammals, it no longer breeds on the two main islands of New Zealand but is now restricted to a few islands south and east of the South Island, principally the little ones near Stewart Island and the Snares Islands (Warham et al. 1977). Its confinement to so few sites led these authors to express concern for the future of the species. They went on to summarize what is known of the species’ terrestrial (i.e., breeding) biology. Here we summarize information on its oceanic distribution. We report for the first time several occurrences of Mottled Petrels in California and Washington, and summarize some recent records from other areas along the North American Pacific Coast. We also summarize the diffuse literature and unpublished observations on its oceanic occurrence, and from this establish its oceanic range and seasonal movements. “NORMAL” RANGE Reports summarized by Watson et al. (1971) and Warham et al. (1977), plus more recent observations (Ainley, unpubl. data), in- dicate that the Mottled Petrel, during its late OctoJ^df-40-^arJv; June breeding season, feeds in Antarctic waters justjtfbrch jcLtfatJ jaabkdce from 95°E to 140°W (to 75°30 'S), with sca^r^iiKh"VT9ctaIs ; t^jSr - ring even farther east (Figure 1). Breeding^birds-“could easily’ -make the 2200 to 4000 km trip between nestifig and feeding grounds, given the 12-14 day spells of incubation (Warham et al. 1977)\ Since the species is abundant in southern wafers, during "the Ibreediog ‘ sjii Western Birds 10: 113-123, 1979 i u 1 1 1 < i > , , , MOTTLED PETREL season, most of the population must be present. A few individuals do occur elsewhere at that time {see below) , but substantial occurrence near South America, as implied in Palmer (1962), is not consistent with data presently available. Mottled Petrels are not present at breeding localities from early June through September (Warham et al. 1977). Neither are they very often, if at all, reported as beach-cast specimens in New Zealand during those months, as compared to breeding months (Roberts 1975, Veitch 1975). Szijj (1967), one of the few ornithologists to have censused pelagic birds in Antarctic waters of the South Pacific during winter, found no Mottled Petrels in areas where they occur commonly during summer. These several clues suggest that the species is absent from South Pacific waters during the non-breeding season, and North Pacific records support this conclusion. Gabrielson and Lincoln (1959) listed four specimen records for coastal Alaska between mid-May and early August 1882-1911, and on that basis considered it to be a “straggler” to Alaskan waters. Kessel and Gibson (1978), summarizing the above plus many recent unpublished records, considered it to be a “very rare visitant” in southern Alaska coastal waters and an “uncommon visitant” in off- shore waters from May through October. As for pelagic waters of the northern North Pacific region, sufficient observations are now available to describe its status. Kuroda (1955), on a cruise from Japan to the western Aleutians and Bering Sea in June and July 1954, encountered a number of Mottled Petrels on several days when east of 162 °E (at 50 °N) and South of 53 °N (at 164 °E), thus in the north-central Pacific. Wahl (1978), on a cruise track very similar to Kuroda’s (except that he also went much farther east on the extreme southern Bering Sea during June and July 1975), observed many Mottled Petrels between 45°N, 166°E and 50°N, 180°W, again in the north-central Pacific, and saw only four scattered in- dividuals in the southern Bering Sea. Hamilton (1958), on a cruise between Japan and Seattle during June 1955, a more southerly route than the above, observed many individuals between 41 °N, 180°W and 44 °N, 155 °W, a few hundred kilometers south of the Aleutians. Sanger (1972), working in eastern and central North Pacific waters during 1955-1967, considered the species to be “com- mon” in w$J,g£§ 4 J60 km NW of Vancouver Island and northwestward into^l^Gu’i cj Alaska during June-August, and considered it to be ppasejjt Bjjt^re^utip^February-March. In July 1969, he (in Gibson ^9^).- again fourif tH% species to be common between 52°N, il60?W (320 km south the Shumagin Islands, Alaska) and 50°N, r 14Q°W (760'knr WSW c§ the Queen Charlotte Islands, British Col- - urubia) rHdis^rtrbe track Tan from Seattle to Adak Island (western MOTTLED PETREL Aleutians) and back. When the cruise was repeated in early October, he observed only two Mottled Petrels, near 54 °N, 144 °W. Wiens et al. {1978), on a cruise from Alaska to Hawaii along 158 °W during late October-early November 1976, considered the Mottled Petrel to be “common” north of 45 °N and less abundant as far south as 36 °N. For several cruises in coastal waters of southeast and western Alaska and the eastern Aleutian Islands in May-October 1975 and 1976, they did not report this species (see also Bartonek and Gibson 1972). Most recently, DeGange and Ainley (unpubl. data) con- ducted censuses on four summertime cruises criss-crossing almost all of the region just reviewed, as well as the Bering Sea from the Aleu- tians to Bristol Bay and Nunivak Island west to 175°E (at 57 °N). Their observations agree with the above but they also found many Mottled Petrels (flocks of up to 22 birds) in a narrow corridor extend- ing north from Adak Island, or just east of Bower’s Bank, to about 57 °N, 179°W in the Bering Sea (see also Kessel and Gibson 1978). These records establish the Mottled Petrel as a common May- October resident in the northern and eastern North Pacific Ocean, principally in association with the Transitional, Central and Western Subarctic and Alaskan Stream domains of surface waters (Dodimead et al. 1963; Figure 1). Noteworthy facts consistent with this associa- tion are 1) the lack of records in warmer waters of the far western North Pacific (Austin and Kuroda 1954, Kuroda 1957, Dement’ev and Gladkov 1968, Ornithol. Soc. Japan 1974, Nakamura and Tenaka 1977) and 2) the scattered summer records in the Bering Sea (except in the corridor east of Bower’s Bank where the species is abundant) and in the most western part of the northern North Pacific and adjacent Okhotsk Sea (Kenyon and Phillips 1965, Bartonek and Gibson 1972, Shuntov 1972, Nakamura and Tenaka 1977, Wahl 1978, Wiens et al. 1978, DeGange and Ainley unpubl, data). P. inexpectata’s movements through the tropical Pacific, established by records independent of those cited above, provide still more clues about the periods of residence in South and North Pacific waters. King (1967, 1970) reported the species’ rapid migration through the Hawaiian Island area (175°E to 150°W), flying north in April and May and south in October and November (Figure 1). It appears now, based on new data, that the northward migration begins much earlier. Ainley (unpubl. data), on a cruise from Samoa to Los Angeles, noted many northward flying Mottled Petrels southeast of Hawaii between 24 and 30 March 1979. So abundant were they that the movement probably began at least a few weeks earlier and, based on the breeding season detailed by Warham et al. (1977), probably involved non-breeders or failed breeders. The route seems to be a diagonal one between New Zealand and the Gulf of Alaska, and 115 MOTTLED PETREL there appears to be no evidence for the circular route clockwise around the Pacific as proposed in Palmer (1962). The many birds observed by Beck (in Loomis 1918), 640 to 1600 km west of central California between 19 and 26 November 1906, must have been at the eastward edge of their migratory route and among the last of the southward migrants that year. The 540 birds counted by Mobberley (in Bourne and Dixon 1975) 112 km WNW of Cape Flattery, Washington, on 28 April 1972 must also have been at the eastward edge of the route but in the vanguard of the northward movement. Many of the birds observed by Wiens et al. (1978) were probably migrants, particularly those observed south of the Subarctic Front (at ca. 40 °N; see Discussion). Figure 1. The approximate oceanic range of the Mottled Petrel ( Pterodroma inexpectata) . 116 MOTTLED PETREL “EXTRALIMITAL” RECORDS Few records of the Mottled Petrel exist for areas outside the above described range. The first record for North America was a very unusual one as it was from New York (Wallace 1961). The bird perhaps flew north in the Atlantic, having probably been one of the individuals that occasionally fly as far east as the Drake Passage in the South Pacific (Watson et al. 1971). Szijj (1967) and Palmer (1962) are the only persons of whom we are aware who have reported this species in South American waters except in the Drake Passage; they reported it in Chilean waters and near the Galapagos, respectively. The species’ usual migratory route brings it near the Americas only in Alaska, thus the scarcity of North American records from British Columbia south is not too surprising. In the last several years though several have been reported. Their timing is rather in- consistent with the species expected occurrence. These records are as follows: British Columbia. A bird captured aboard ship, photographed and released 46 km SW of Estevan Point, Vancouver Island, on 24 February 1971 was the first record for inshore waters of this province (Campbell and Shepard 1973). The day before, one was seen about 280 km west of this locality (Crowell and Nehls 1971). During the next year, on 17 March 1972, one flew aboard another ship 480 km SW of the Queen Charlotte Islands. The specimen is now at the Vertebrate Museum, University of British Columbia (Campbell and Shepard 1973). Washington. Three inshore occurrences have recently been recorded. One live bird was sighted by Glen and Wanda Hoge (pers. comm.) on 28 February 1976 at Ocean Shores, the north jetty of Gray’s Harbor, Gray’s Harbor County. The sighting is still being con- sidered by rare bird authorities in Washington, but appears to be ac- ceptable (T. Wahl pers. comm.). This would be the first state record. Two dead individuals were then found in Gray’s Harbor County by Jack L. Smith (pers. comm.) while conducting beached bird cen- suses for the Washington Department of Game. The first was found on 2 March 1976 between Westport Lighthouse and the Twin. Har- bor access, and the second was found on 5 March 1976 just north of Moclips. The first is now a specimen at the University of Puget Sound; the second was badly decomposed, indicating that it prob- ably washed ashore a week or more earlier. Oregon. The first record for this state, and for the Pacific Coast south of Alaska, was a bird found dead 3 km north of Alsea Bay, Lincoln County, on 25 July 1959 (Wallace 1961). Its late stage of decomposition and the fact that it was in the “winter high tide line” 117 MOTTLED PETREL suggest that it probably washed ashore long before it was found. The incomplete skeleton is now at the Museum of Vertebrate Zoology, Berkeley. More recently, on 16 February 1971, Narca Moore {pers. comm.) observed one 175 km west of Tillamook, Oregon, at RV YAQUINA station CP2A; and on 18 March 1972, Wayne Hoffman found two dead ones while conducting a beached bird census, 7 and 10 km south of Newport, Lincoln County (Crowell and Nehls 1972). Both of the latter birds are now in the collection at Oregon State University, Corvallis. California. One of us (BM with Ane Rovetta), during a Point Reyes Bird Observatory beached bird census, found a Mottled Petrel at Point Reyes Beach, Marin County, on 25 February 1976. This was the first record for California. The study skin is now in the California Academy of Sciences (Figure 2). A few days later, on 28 February, another was found at Cayucos Beach, San Luis Obispo County, by Dana Tryde (pers. comm.). This specimen is now at California Polytechnic State University, San Luis Obispo. A third was found during another PRBO beach census at San Simeon Beach, San Luis Obispo County, on 13 March 1976 (DGA, Louise Squibb and Lois Felmlee). The skeleton is now at PRBO (Figure 3). Later that year, on 11 August, a freshly dead individual was found on a beached bird census at the mouth of the Mad River, Humboldt County (Winter and Erickson 1977a; specimen now at Humboldt Figure 2. Mottled Petrel found on Point Reyes Beach, Marin County, California, on 25 February 1976; the first record for California. 118 MOTTLED PETREL State University, Areata). No more Mottled Petrels were recorded until 1 May 1977, when one was found alive but weak at Bolinas Lagoon, Marin County. The date and its extensive feather wear in- dicated it was at least 1 year old. It died on 8 May and is now a specimen at California Academy of Sciences. DISCUSSION Mottled Petrels breed at sites where waters are about 5-13°C, but during the breeding season they apparently prefer to feed at the northern edge of the Antarctic pack ice where temperatures are 1-3°C (Figure 1). A great many icebergs are a part of this preferred environment (Ainley unpubl. data). During the non-breeding season, after rapidly crossing tropical waters, they frequent subarctic waters in the central and eastern North Pacific that are about 5-13°C. Mottled Petrels conceivably have the opportunity to fly far- ther north to colder waters, and even to the edge of the Arctic pack ice. To do so, though, they would have to leave their preferred oceanic water and fly over the shallow shelf water of the northern Bering Sea. At the northern edge of the Antarctic pack ice in sum- mer, the Mottled Petrel is virtually the only Pterodroma present, and Figure 3. Mottled Petrel found on San Simeon Beach. San Luis Obispo County. California, on 13 March 1976; the third record for California. 119 MOTTLED PETREL is among the most abundant of avian species. In waters around its breeding islands at that time, many species of petrels occur abun- dantly including several Pterodroma. In North Pacific waters during the northern summer, the Mottled Petrel is the only Pterodroma (in those waters where the species regularly summers) and ranks among the most abundant of seabirds present. By flying farther north it would encounter many other seabird species in great numbers. If the long-distance movements of seabirds can be influenced by the ex- istence of unexploited resources in certain areas, then the Mottled Petrel provides a good example. The increased number of North American Pacific Coast records since 1971, compared to earlier years, is probably a result of inten- sified activity among bird watchers. For example, of the 10 P. inex- pectata reported within 100 km of the coast south of central British Columbia since 1971, 7 were found on organized beached bird cen- suses. Before 1971 a few people made occasional beach censuses along this coast. Since then the number has increased dramatically, reaching a peak of about 80 beaches regularly censused by 1975. Most of the Mottled Petrels (12 of 14) found from British Columbia south since January 1971 have occurred during the short period be- tween 16 February and 18 March. The timing is outside the species’ usual peak occurrence period in the North Pacific. The birds involved must have been non-breeders. Do non-breeders or failed breeders move away from nesting sites and into the North Pacific before breeders and juveniles do so in April and May? The close timing of records regardless of year does suggest a regular, though minor migratory movement. Ainley’s recent observations of many Mottled Petrels crossing the tropical Pacific in March further support this. Another pattern was visible in the “extralimital” occurrence of this species. Three birds occurred in Oregon and British Columbia within 8 days of one another in February 1971; two birds occurred a great distance apart off British Columbia but within a day of each other in late February 1972; three birds occurred in Oregon and Washington, a great distance apart, again within a day of one another, in mid- March 1972; and 6 birds occurred in California and Washington within 14 days of one another in February-March 1976. The close dates of occurrence in widely spaced localities is further evidence for migratory movement to the North Pacific earlier in the year than previously expected. It thus appears that few truly extralimital records exist for this species. In fact, the only ones of which we are aware are those from New York, the Galapagos and northern Chile. The fact that 11 of 14 records (mid-February through mid-March) since 1971 were in two of a possible seven winters may provide some clues about the reasons for the more inshore occurrence during 120 MOTTLED PETREL some years. Using northern California as an index, the two outstand- ing winters for inshore Mottled Petrels, 1971-72 and 1975-76, also happened to be periods when Northern Fulmars ( Fulmarus glacialis ) were far more abundant than usual in coastal waters (cf. Ainley 1976; Stallcup et al. 1975; Stallcup and Winter 1975, 1976a, b; Winter and Erickson 1977a, b; Erickson and Morlan 1978; Winter and Manoiis 1978). The latter species is abundant in the Gulf of Alaska during winter and moves southward and shoreward in con- junction with cold waters of high salinity (i.e., the characteristics of central Subarctic waters in the Gulf of Alaska; Dodimead et al. 1963, Ainley 1976). Possibly the same conditions brought the Mottled Petrels shoreward as well, since they too seem to prefer oceanic cen- tral Subarctic waters. SUMMARY Records in the literature are summarized to determine the oceanic range of the Mottled Petrel throughout the year and to establish the limits of its usual occurrence, especially in the North Pacific. The species is very infrequently reported within 100 km of shore from southern British Columbia south to California, but since January 1971 “first records” for the species were established in each coastal province and state, with the exception of Oregon. These records in- volved individuals not in the breeding population. They were the result of increased activities among birders, were probably in the forefront of the species’ northward migration, and were probably in- fluenced by fluctuations in oceanographic conditions. ACKNOWLEDGMENTS We wish to thank the several persons who contributed details of their observations and other information: John Butler, Wayne Campbell, George Divoky, Glen and Wanda Hoge, John Luther, Guy McCaskie, Narca Moore, Jack Smith and Dana Tryde. DGA’s observations in the polar and tropical South Pacific were gathered on cruises funded by the National Science Foundation, Division of Polar Programs; cruises in the western Pacific and Bering Sea were funded by the U.S. Fish and Wildlife Service. Terry Wahl and Alan Baldridge improved the paper through their comments. This is Con- tribution 190 of the Point Reyes Bird Observatory. 121 MOTTLED PETREL LITERATURE CITED Ainley, D.G. 1976. The occurrence of seabirds in the coastal region of California. West. Birds 7:33-68. Austin, O.L., Jr. and N. Kuroda. 1954. The birds of Japan, their status and distribu- tion. Mus. Comp. Zool. Bull. 109:279-637. Bartonek, J.C. and D.D. Gibson. 1972. Summer distribution of pelagic birds in Bristol Bay, Alaska. Condor 74:416-422. Bourne, W.R.P. and T.J. Dixon, 1975. Observations of seabirds 1970-1972. Sea Swallow 24:65-88. Campbell, R.W. and M.G. Shepard. 1973. Laysan Albatross, Scaled Petrel, Parakeet Auklet: additions to the list of Canadian birds. Can. Field-Nat. 87:179-180. Crowell, J.B., Jr. and H.B. Nehls. 1971. The winter season. Northern Pacific Coast region. Am. Birds 25:615-619. Crowell, J.B., Jr. and H.B. Nehls. 1972. The winter season. Northern Pacific Coast region. Am. Birds 26:644-648. Dement’ev, G.P. and N.A. Gladkov. 1968. Birds of the Soviet Union, vol. II (Transl. from Russian). Israel Prog. Sci. Transl., Smithsonian Inst., Washington, D.C. Dodimead, A.J., F. Favorite and. T. Hirano. 1963. Review of oceanography of the subarctic Pacific region. Int. N. Pac. Fish. Comm. Bull. 13:1-195. Erickson, D. and J. Morlan. 1978. The autumn migration. Middle Pacific Coast region. Am. Birds 32: 250-255. Gibson, D.D. 1970. The fall migration. Alaska region. Am. Birds 24:79-82. Hamilton, W. J., Ill, 1958. Pelagic birds observed on a North Pacific crossing. Condor 60:159-164. Kenyon, K W. and R.E. Phillips. 1965. Birds from the Pribilof Islands and vicinity. Auk 82:624-635. Kessel, B. and D.D. Gibson. 1978. Status and distribution of Alaska birds. Studies Avian Biol. No, 1. King, W.B. 1967. Seabirds of the tropical Pacific Ocean. Preliminary Identification Manual. Smithsonian Inst., Washington, D.C. King. W.B. 1970. 1 he trade wind zone oceanography pilot study, Part Vll: observa- tions of sea birds March 1964 to June 1965. U.S. Fish Wildl, Serv. Spec. Sci. Rep. Fish. No. 586. Kuroda, N. 1955. Observations of pelagic birds in the northwest Pacific. Condor 57:290-300. Kuroda, N. 1957. A brief note on the pelagic migration of the Tubinares. Yamashina’s Inst. Ornithol. Zool., Misc. Rep. No. 11:436-449. Loomis, L.M. 1918. A review of the albatrosses, petrels and diving petrels. Calif. Acad. Sci., Proc., 4th Ser., 2:1-187. Nakamura, K, and Y. Tanaka. 1977. Distribution and migration of two species of the genus Pterodroma in the North Pacific. Yamashina Inst. Ornithol., Misc. Rep. 9:112-120. Ornithological Society of Japan. 1974. Check-list of Japanese birds. Gakken Co., Tokyo. Palmer, R.S. ed. 1962. Handbook of North American birds, vol. 1. Yale Univ. Press, New Haven, CT. Roberts, P.E. 1975. Sea birds found dead in New Zealand in 1965 and 1966. Notor- nis 22:151-161. Sanger, G.A. 1972. Checklist of bird observations from the eastern North Pacific Ocean, 1955-1967. Murrelet 53:16-21, Shuntov, V.P. 1972. Seabirds and the biological structure of the ocean (Transl. from Russian). Natl. Tech. Inf. Serv., Springfield, V A. 122 MOTTLED PETREL Stallcup, R., D. DeSante and R. Greenberg. 1975. The fall migration. Middle Pacific Coast region. Am. Birds 29:112-119. Stallcup, R. and J. Winter. 1975. The winter season. Middle Pacific Coast region. Am. Birds 29:735-740. Stallcup, R. and J. Winter. 1976a. The fall migration. Middle Pacific Coast region. Am. Birds 30:118-124. Stallcup, R and J. Winter. 1976b. The winter season. Middle Pacific Coast region. Am. Birds 30:760-774. Szijj, L.J. 1967. Notes on the winter distribution of birds in the western Antarctic and adjacent Pacific waters. Auk 84:366-378. Veitch, C.R 1975. Seabirds found dead in New Zealand in 1973. Notornis 22:231-240. Wahl, T.R. 1978. Seabirds in the northwestern Pacific Ocean and south central Ber- ing Sea in June 1975. West. Birds 9:45-66. Wallace, W.M. 1961. Scaled Petrel in Oregon. Condor 63:417. Warham, J., B.R. Keeley and G.J. Wilson. 1977. Breeding of the Mottled Petrel. Auk 94:1-17, Watson, G.E., J.P. Angle, P.C. Harper, M.A. Bridge, R.P. Schlatter, W.L.N. Tickell, J.C. Boyd and M.M. Boyd. 1971. Birds of the Antarctic and Subantarc- tic. Folio 14, Antarctic Map Folio Ser. Am. Geograph. Union, New York. Wiens, J.A., D. Heinemann and W. Hoffman, 1978. Community structure, distribu- tion, and interrelationships of marine birds in the Gulf of Alaska. Final Rep., Envir. Asses. Alaskan Cont. Shelf, OCSEAP, NOAA, Boulder, CO. Vol. 3:1-178. Winter, J. and D. Erickson. 1977a. The fall migration. Middle Pacific Coast region. Am. Birds 31:216-221. Winter, J, and D, Erickson. 1977b. The winter season. Middle Pacific Coast region. Am. Birds 31:367-372. Winter, J. and T Manolis. 1978. The winter season. Middle Pacific Coast region. Am. Birds 32:394-397. Accepted 7 September 1979 123 124 Sketch by Narca Moore MORNING AND EVENING ROOSTS OF TURKEY VULTURES AT MALHEUR REFUGE, OREGON DEBORAH DAVIS, 1812 Center Street, Walla Walla, Washington 99362 The tendency of Turkey Vultures ( Cathartes aura) to perch at some particular site other than the overnight roost, both in morning and evening, appears to be predictable and universal within the species. As with many components of Turkey Vulture behavior, very similar behavior is shown by the California Condor (Gymnogyps calif ornianus; Koford 1953) and the Andean Condor ( Vultur gryphus; McGahan 1972). At a large roost in southeastern Oregon, I observed the arrival and departure times of Turkey Vultures, and their movements to and from different perching sites. STUDY AREA AND METHODS The roost was located at the southern end of Malheur National Wildlife Refuge, Harney County, Oregon. The vultures roosted primarily in a row of cottonwoods { Populus sp.) at P Ranch Station. Before and after roosting, many perched about 100 m NW of the trees on a metal observation tower approximately 30 m high (Figure 1) . North of P Ranch were the irrigated meadows of the Blitzen River Valley. Further north lay Malheur Lake, a vast marsh fluctuating from 200 to 20,000 ha and rich in avian and mammalian fauna. About 3 km SE of the roost were the lower slopes of Steens Moun- tain, characterized by Western Juniper ( Juniperus occidentalis) and Quaking Aspen ( Populus tremuloides) . The general region was characterized by Great Basin sagebush ( Artemisia sp.) — Greasewood ( Sarcobatus uermiculatus) association. 1 observed vultures for 330 hours during late spring and mid- summer 1973. I watched the birds from the time of their arrival until the cessation of activity in the evening, and in the morning from twilight until their departure. Every 15 minutes I recorded time, temperature, wind and precipitation. Five Turkey Vultures were captured in a trap, such as that used by Coles (1938), baited with carrion, primarily carp ( Cyprinus carpio). A captive vulture was left in the trap as a decoy. I marked each bird with a 3 cm x 18 cm colored leg-streamer of plasticized fabric (Safety Flag Company of America, Pawtucket, Rhode Island), riveted to a USFWS size 7b band. Western Birds 10: 125-130, 1979 125 TURKEY VULTURE ROOSTS RESULTS AND DISCUSSION The number of Turkey Vultures roosting at P Ranch each night averaged 104 (68-151; n = 28). Although Nauman (1965) concluded that individual Turkey Vultures have specific preferred perching sites, marked vultures in this study showed no such preference. Fur- ther, data from sightings of marked birds suggest that at least some individual birds did not spend every night at the roost. For each of the four marked vultures that used the roost, I calculated the ratio of the number of days sighted to the total number of observation days. The mean percentage of the frequency of roost usage for the four vultures was 68% (58%-79%). Koford (1953) found that California Condors often did not roost at the same place each night, and that they were likely to roost near a carcass until the carcass was consumed. The time at which vultures began arriving at the roosting area varied. In late spring, vultures might not arrive until 1.5 hours before sunset, while in early and mid-summer there were vultures on the tower by 2.5 hours before sunset. Arrival times also varied from day to day. When a storm front approached, the birds returned to the roost earlier than when the weather was clear and calm. In strong wind the birds formed a “wind-flock,” all oriented at the same angle to the wind, their bodies parallel to one another. The flock glided laterally without making headway, so that it appeared to be “hang- ing” over the roost. Koford (1953) observed the same behavior in California Condors. On 12 June, when skies were completely over- cast and the wind was strong and gusting, I saw a wind-flock near the roost about 6 hours before sunset. On the same day, 2.5 hours before sunset, 34 birds were at P Ranch. Although many Turkey Vultures perched on the tower, the main roost at night was the trees. Nauman (1965) observed Turkey Vultures near Columbus, Ohio, coming into a roosting area and perching as early as 3 to 4 hours before sunset then moving to their final roost approximately 45 minutes before sunset. Koford (1953) found that California Condors first roosted high on a cliff or in trees high on the side of a mountain, and later, shortly before or after sunset, moved to lower elevations. There are likely several advantages in this pre-roosting behavior. In Coles’ (1938) study, Turkey Vultures pre-roosted in what he called “sunning trees.” These were dead trees without foliage or shade, which allowed the vultures to arrive and depart with ease. He noted that after long periods of “preening, resting and wing-spreading,” the birds moved from these trees to their final roost. At P Ranch, the tower played the same role as Coles’ sunning trees. The tower, con- structed of widely spaced bars and located in an open meadow, allowed unobstructed landing and departure. As with sunning trees, 126 TURKEY VULTURE ROOSTS the tower was not shaded and thus provided good conditions for carrying out the preening, sunning and stretching activities characteristic of Turkey Vultures. The steel tower also provided stable perches for the sometimes vigorous movements required in r\ Figure 1. Tower at P Ranch, Malheur NWR, Harney County, Oregon, used by Turkey Vultures as a pre- and post-roost. 127 TURKEY VULTURE ROOSTS preening. Finally, the pre-roost served as a site for what were often intense agonistic encounters. After sunset, the nature of activity at the pre-roost began to change. Preening ceased and many vultures flew, hopped or walked to new perching sites. Some movements were a result of agonistic encounters, while others appeared to be spontaneous. During this period of increased movement, many vultures left the tower for the trees. Those that stayed on the tower generally moved to the upper- most levels. By 45 minutes after sunset there was little or no visible movement on the tower. Those vultures that flew directly to the trees rather than the tower demonstrated a period of preening and agonistic encounters similar to that seen on the tower, with all visible activity ceasing shortly before darkness. In the morning, many vultures returned from the trees to the tower, in this case using the tower as a post-roost. On 23 of 34 morn- ings (68%), over half of the birds to use the post-roost that day were at the tower by 15 minutes after sunrise. On the remaining 32% of the mornings, the vultures flew to the tower gradually, either singly or in small groups, the number on the tower reaching a peak shortly before the birds began to depart the roosting area. For each of the mornings when the birds were not disturbed at the roost, I used the peak number of birds on the tower just before departure as an estimate of the number of vultures using the tower that morning. Of those 35 mornings when the birds were not disturbed, the mean number of vultures post-roosting on the tower was 59. Thus, each day roughly half of the vultures roosting in the trees used the tower as a post-roost. Observation of marked vultures indicated that in- dividual vultures did not use the pre-roost and post-roost every day. I assume that most vultures used the tower at least occasionally. Coles (1938) described Turkey Vultures in Ohio flying after sunrise to favorite sunning areas where they remained until soaring conditions developed. Koford (1953) reported that California Condors usually changed their perching sites at least once before departing in the morning. In McGahan’s (1972) study, Andean Condors flew from shaded roosting ledges to sunny perches before they began foraging. These descriptions of cathartine behavior, together with my observa- tions, indicate that preening, stretching and sunning are the primary activities of post-roosting vultures, and that the outstanding require- ment of a post-roost is sunshine. The vultures’ departure time from the roosting area was taken to be that time (in minutes after sunrise) when the number of vultures on the tower had decreased to one half the peak number of vultures on the tower that morning. On over half (55%) of the mornings the vultures departed the roosting area between 3 and 4 hours after sunrise. When a breeze was blowing, the mean departure time (146 128 TURKEY VULTURE ROOSTS minutes after sunrise) was significantly lower than the mean depar- ture time when there was no breeze (191 minutes after sunrise; group comparison t-test, p<0.05). On 3 mornings in April there were periods of rain, snow and persistent wind. Then, with skies generally overcast and temperatures below freezing, the vultures left earlier than in the summer. If rain or snow was falling, some vultures would not leave the roost all day. Generally, soaring conditions are poor during periods of rain (Pennycuick 1972). The Turkey Vulture is capable of surviving several days without food (Hatch 1970) , prob- ably as an adaptation to an opportunistic feeding niche. Thus, if flight conditions are poor, a vulture should be able to remain at the roost for 1 or 2 days or more, until flight conditions improve. Presumably Turkey Vultures left the roost when there were suffi- cient thermals for soaring. But social facilitation apparently affected departure time. The first birds to depart usually flew directly off without circling. However, when one or more vultures began circling nearby after flying from the roost, other vultures often took off and joined the circle. Several birds in succession would then leave the roost or post-roost. This was likely a response to a visual cue, which is substantiated by the presence of vultures in the trees that would not join a circle formed by birds from the tower, presumably because the tree vultures did not see the circling behavior. When a bird circled in front of the trees, out of sight of birds on the tower, only vultures in the trees joined the flight. SUMMARY During spring and summer of 1973, I observed movements of roosting Turkey Vultures at a large roost in southeastern Oregon. About 100 vultures perched in a row of cottonwood trees each night, and used a tall observation tower as a pre- and post-roost. Daily variation in arrival times was related to fluctuations in weather. After sunset, most vultures on the tower flew to the trees to spend the night; about half of the roosting vultures returned to the tower after sunrise the following morning. The tower provided a sunny site for preening, sunning and agonism. The requirement for such sites is shared by other cathartine vultures, most notably condors. On over half of the mornings the Turkey Vultures left the roosting area be- tween 3 and 4 hours after sunrise. The time of departure was most influenced by wind and perhaps social facilitation. The widespread occurrence of social roosting and the associated pre- and post- roosting phenomena among cathartine vultures suggests that these behaviors are a vital component of these species’ survival strategies. 129 TURKEY VULTURE ROOSTS ACKNOWLEDGMENTS I wish to thank the foilowing persons: Charles H. Trost of Idaho State University, who was my advisor on this project and who reviewed the manuscript, John O. Sullivan, who suggested the topic of research and who helped design the study, and my parents, Frederic and Jean Managhan Davis, for their continued support. I am also grateful to Carl B. Koford and Sartor O. Williams, III, for their comments on an earlier draft. LITERATURE CITED Coles, V.E. 1938. Studies in the life history of the Turkey Vulture (Cathartes aura septentrionalis) . Ph.D. Thesis. Cornell Univ., Ithaca, NY. Hatch, D.E. 1970. Energy conserving and heat dissipating mechanisms of the Turkey Vulture. Auk 87:111-124. Koford, C.B, 1953, The California Condor. Natl. Audubon Soc., Res. Rep. 4, McGahan, J. 1972, Behavior and ecology of the Andean Condor. Ph.D. Thesis. Univ. Wisconsin. 304 p. Univ. Microfilms, Ann Arbor, MI. Nauman, L.E. 1965. Spatial distribution in a Turkey Vulture roost. M.S. Thesis. Ohio State Univ., Columbus, OH. Pennycuick, C.J. 1972. Soaring behavior and performance of some East African birds, observed from a motor-glider. Ibis 114:178-218. Accepted 18 October 1979 130 Sketch by Tim Manolis DISTRIBUTION, BIOLOGY, AND STATUS OF A RELICT POPULATION OF BROWN TOWHEE (Plpllo fuscus eremophllus) BART CORD and JOSEPH R. JEHL, JR., Hubbs/Sea World Research Institute, 1700 South Shores Road, San Diego, California 92109 The Inyo Brown Towhee ( Pipilo fuscus eremophilus) is a relict population of a species that was formerly widespread in the southwestern United States and northern Mexico (Davis 1951). A member of the Crissalis group of subspecies, eremophilus became restricted to mountain areas in the northern Mojave Desert as a result of climatic changes beginning in the Pliocene (Davis 1951). Current- ly it is known only from the Argus Range of Inyo County, California (Cord and Jehl 1978, contra AOU 1957) (Figure 1). The nearest neighboring population, P. /. carolae (formerly kernensis) occurs 65 km due west in the Walker Basin of the southern Sierra Nevada. Like all other races of P. fuscus, eremophilus is considered resi- dent (Davis 1951). It was described by Van Rossem (1935) on the basis of specimens he collected at Mountain Spring (1400 m) and Lang Spring (1830 m) in the southern Argus Range, and from a single specimen taken by F. Stephens at “Searle’s Garden” (Fisher 1893). 1 Because of its limited and largely inaccessible range, which is rare- ly visited by ornithologists, very little is known about the biology, re- quirements, or population size of this isolated desert dwelling race. We attempted to gather such data at the request of the Desert Land Plan staff of the U.S. Bureau of Land Management. METHODS In May-June 1978 Cord visited water sources in the Argus Range between Indian Joe Spring and Stone Canyon, with the exception of La Motte Spring (Figure 2). Surveys were restricted to riparian habitats because they provide the only towhee nesting habitat in the Argus Range; the surrounding area is high desert. Extensive hiking was required, usually through trailless canyons with steep slopes and rugged rock formations. 'The exact location of “Searle's Garden" has been disputed. We (Cord and Jehl 1978) have argued that it is immediately adjacent to Indian Joe Spring, which is in southern Inyo County, and not in San Bernardino County as suspected by some (e.g., AOU 1957). Western Birds 10: 131-156, 1979 131 INYO BROWN TOWHEE Whenever a towhee was observed, a 50 pace toe-point transect was taken to the nearest riparian habitat to provide precise data on vegetative components. Other data collected were: legal description of site, description of locality, water flow (gpm), stream length, area of free surface water, mean water depth, major plant species, area of riparian growth, degree of recreational use, and evidence of use by burros. Photographs were taken at each water source and at each locality where towhees were found. In late 1978 Cord discovered extensive riparian growth in the Quail Spring-Benko Canyon area, in the center of the apparent Figure 1. The range of Brown Towhee populations in California (stippled). The range of the Inyo Brown Towhee (solid) is confined to the southern Argus Range. 132 INYO BROWN TOWHEE range of eremophilus. Supplementary field work was carried out in that area in April-May 1979, as well as in a few areas surveyed in 1978. In both years, the extent of riparian habitat was carefully mapped. Field work in fall and winter 1978-79 was aimed at determining eremophilus ’ status and requirements at those seasons and in- vestigating the possibility of seasonal dispersal or emigration. Detailed descriptions of the habitat and of the itinerary are available in Cord’s field notes; copies are on file at the Bureau of Land Management, Riverside, California, and Hubbs/Sea World Research Institute. RESULTS DISTRIBUTION In May-June 1978, towhees were found at only 6 of 24 areas in the southern Argus Range (Table 1): Indian Joe Spring, Great Falls Basin, Crow Canyon, North Homewood Canyon, Ruby Spring, and Mountain Spring Canyon (Figure 3). None were found in 10 canyon areas (20 water sites) in the northern Argus. In April-May 1979, towhees were present in 8 of 10 areas in the southern Argus, in- cluding the following additional localities: Shelf Canyon, Rusty Can- yon, Green Canyon, Benko Canyon, Layne Canyon, and the Bob- cat Canyon-Water Canyon complex (Figure 4). No further studies were made in the northern Argus. However, in response to the report of a possible sighting in the Coso Mountains, just to the north of the Argus Range, Cord visited Black Spring on 8 May; he found a lone Green-tailed Towhee ( Pipilo chlorurus) . In both years, all 12 sites combined, a total of 75 birds (including three nestlings in one nest) was recorded. This figure is conservative. Desert towhees are difficult to census in the rugged canyon country, because they have large home ranges and often remain silent and hidden in dense riparian cover. P. f. eremophilus is evidently confined entirely to the southern Argus Range of Inyo County. Thirty-six percent of the sightings were made within a circle of 3 miles diameter centered at Benko Canyon, and 100% within an 11-mile circle. The vast majority of the sightings (85%) were made within a 6-mile circle; most of this land is within the confines of China Lake Naval Weapons Center (Figure 2). Towhees are more common on the east side of the Argus, ap- parently because of the larger riparian habitats there. Their absence from the northern Argus seems due to lack of habitat; soil formation is poor, water sources are few and widely scattered, and riparian vegetation is scanty. There were, and still are, active mines at almost every water source in the northern Argus, which has further depleted riparian growth. In some areas only Squaw Waterweed ( Baccharis sergiloides ) remains; it does not provide nesting habitat (see below). 133 o 5 io 15 MILES Figure 2, The Argus Range area. Numbers refer to the major spring or canyon areas visited in this study. 1) Indian Joe Spring (+ Searle’s Garden); 2) Great Falls Basin; 3) Crow Canyon; 4) Moscow Canyon; 5) Benko Canyon complex (includes Shelf Canyon, South Homewood Can- yon, Rusty Canyon, Benko Canyon, Benko Spring and Green Canyon); 6) North Homewood Canyon, Ruby Spring, Layne Canyon; 7) Mountain Spring Canyon; 8) Water Canyon-Bobcat Canyon complex (includes Coyote Spring); 9) Shepherd Canyon; 10) Onyx Mine area (3 springs); 11) Revenue Canyon; 12) Snow Canyon; 13) Thompson Canyon; 14) Stone Canyon (French Madam Spring, Jack Gunn Spring); 15) Black Spring, Coso Mountains. The Inyo Brown Towhee is confined to the area within the circle. 134 INYO BROWN TOWHEE 135 Figure 3. Mountain Spring Canyon, site 5. Two pairs of towhees are estimated to inhabit this area INYO BROWN TOWHEE 136 Figure 4. Coyote Spring, at the head of Water Canyon. One pair of towhees is estimated to occur here. Table 1. Distribution, habitat parameters, and population size of Inyo Brown Towhees, Argus Range, Inyo County, California 1978-1979. INYO BROWN TOWHEE a a in o w a o C c 3 £ g 2 "D C 3 O - cn co c o IS o3 CD CD g .9- 03 — , 1- (1) > a 3 flj c o - cu *— ■ > c o i-. ro fO V, 9- CT 3n a> JC CQ c. 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c fl u c cu CD c >— iCMcostinot^oo lUipiDQ>Q)Q> ddddw^cocoyoco c o P\ c U T3 O O 3 cis E o X J0 t o Z co c ’JCC a co Pn JO D DC Layne Canyon Site 1 450 A 21,000 350 Site 2 100 AT 4,000 100 Site 3 0 TIF 8,000 200 Site 4 50 TA 5,000 120 Table 1. (Cont.) INYO BROWN TOWHEE — a. (/> Q LU £ O C c 5 £ g 2 ■* -a c 3 o '-i-i to T3 S— s T3 C 3 O M— ( 1/5 T3 s— 55 at t''- at rH cn c a m oo r"~ at 7 —i cn c *c Q. to o C o ■£ «c * ^ Q) . 2 - on : u d) > c *3 t u E- > < E- < < < <<£ s o o o o o o o o co to o o o o o o in in to in cm co c o 3s c a U V-. 0) CD to co c /5 st

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