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HomeMy WebLinkAboutAPA3340SUMMER DISTRIBUTION, NUMBERS, AND FOOD HABITS OF THE GYRFALCON (Falco rusticolus L.) ON THE SEWARD PENINSULA, ALASKA APPROVED: APPROVED: ) - ·-< 1_/ (i .. ,_ " .) /)). }.·: (.\_ {lf>Q/,]_::__.< DATE I 0 ///.'!.:(., /9 ?-;; __ _ =n-ea_n_o---:r:;--·-the College of Biological {J Sciences and Renewable Resources ~ // C::..---CL~--------------Vice President for Research and Advanced Study SUMMER DISTRIBUTION, NUMBERS, AND FOOD HABITS OF THE GYRFALCON (Falco rusticolus L.) ON THE ---------- SEWARD PENINSULA, ALASKA A THESIS Presented to the Faculty of the University of Alaska in Partial Fulfillment of the Requirements for the Degree of MASTER 0~ SCIENCE By David Gale Roseneau, B.S. College, Alaska Hay 1972 ABS'l'RAC'I' During the summers of 1968, 1969, and 1910 aerial and ground surveys were made of approximately 17,000 square miles of the Seward Peninsula, Alaska, to locate nesting Gyrfalcons (Falco rusticolus L.). During the three summers 131 nestings were observed and populations remained both high and relatively stable on a region-wide basis. He-utilization of specific nest cliffs was low, however, and cliff-shifting was conspicuous. In smaller areas of the peninsula the numbers of pairs utilizing a given area changed from year to year in a manner correlat8d with prey availability. Prey remains and pellets were collected from 37 nests over the course of the study and 1,483 kills were identified. A minimum of 40 speci.es was represented includ- ing 32 bird species and eight mammal spec~es. This is a considerably longer list of prey species than has been reported previously. Four species dominate the food sample (Rock Ptarmigan, Lag onus mutus, 1di llo1·r Ptarmigan, L. la;;:;opus, the Arctic Ground Squirrel, Sp~rmophilus undulatus, and the Long-tailed Jaeger, Stercorarius longicaudus), making up 81 per cent by number and 92 per cent by weight. Prey utiliza- tion va~ied with availability both from year to year and from habitat type to habitat type beti'Teen the hunting ranges of nesting pairs. iii ACKNOVJLEDGiviENTS I would like to express my appreciation to the organizations and persons who supported and encouraged this study. The Alaska Department of Fish and Game supported the project during 1968 and 1969 (under Project No. W-13-r-3 and W-17-l, Work Plan B, Job No. ll, and under Project No. W-17-l and W-17-2, Study Plan B and R, Job No. B-ll and R-10.4). In 1970 this agency supported the analysis of food habits data (under Project No. W-17-3, Job ~o. B-11 and R-10.4). Dr. Robert B. Weeden, formerly a biologist with the Alaska Department of Fish and GE~me and Project Leader, deserves special mention for his encouragement and valuable comments. The Nome office of the Alaska Department of Fish and Game, under the leadership of John Burns in 1968 and 1969 and Robert Pegau in 1970, provided outstanding cooperation and assistance throughout the course of the study. Edward Muktoyuk, Carl Yanagawa, and Peter Kashiverof lent valuable assis- tance and information. The National Audubon Society funded a major portion of the study during 1970. Their financial support and encouragement during 1970 made it possible to complete this thesis. The University of Alaska and the College of Biologi- iv cal Sciences and Renewable Resources provided additional necessary support and funds throu~hout the course of the study. My special thanks go to Dr. L. G. Swartz, academic advisor and teacher. His encouragement, advice, and friendship were of the best throughout my academic career. William Griffin, Alaska Department of Fish and G~me biologist, Neal W. Foster, Foster Aviation, and George Johnson, Foster Aviation, piloted various portions of the aerial surveys throu~hout the study--a sometimes dangerous and tryins job. Their combined expertise and cooperation made Lr1e uerial survey possible and lent much towards its success. Dr. C. L. Sainsbury and Travis Hudson of the United States Geological Survey contributed valuable cooperation and assistance throughout the course of the field work. Wayman E. Halker, fellow graduate student at the University of Alaska, assisted in field operations during 1970. As an 2blc co-worker and excellent conpanion, he shared my enthusiasm for birds of prey. William R. Tilton assisted with field work during 1968. He contributed a high level of enthusiasm, undamped spirits, and a keen eye. :Patrick Wright worked as my field assistant du~ing a'port:lon-of the_1969 field season. Sam Stoker donated valuable time during a memorable v exploration of the peninsular coast between Nome and Unalakleet in 1969. vi TABLE OF CO~TENTS :!:NTRODUC'l'ION. OBJEC'l'IVES. f!IETHODS . . Study Area . . . . Numbers and Distribution . Food Habits. RESULTS . Numhers and Distribution . Population Changes . Food fiabits. Prey utilizat5on on the Yearly variations in prey utilization . Variation in prey species utilization between different pairs . SUMMARY AND CONCLUSIONS . LITERATURE CITED. vii l 3 4 4 6 8 13 13 23 1!1 it 1+ 59 121 123 'l'able l. 2. 3 . lj . 5 . 6 . 7. LIS'l' OF ':::'J\BLES Numbers and densities of breeding Gyrfalcon pairs on the Seward Peninsula, Alaska ..... Successive use of nesting cliffs by predatory birds on a 20-sauare-mile nortion of the Seward Peninsul~, 1968-1970. . . . .. Soecies utilizing the 34 active Gyrfalcon nesting cliffs observed on the Seward Peninsula, Alaska, 196R-1a70 .. Species utilizing the 48 active 1969 Gyrfalcon nesting cliffs observed on the Seward Penin- sula, ~laska, 1968-lq7o. . . . . . .. Collection dates of Gyrfalcon prey remains Total 1968-1970 Gyrfalcon food remains The three major Gyrfal~on food items .. 8. Total migratory bird species identified from the 1968-1970 Gyrfalcon prey remains and listed in various important gro~ps . . 9. 1968-1970 Gyrfalcon food rem8ins on a yearly basis .... corn-quted 10. Migratory species identified from the 1968- 1970 Gyrfalcon prey remains compared on a yearly basis and listed in various important groups . . . . 11. Comparisons of the occurrence of nine prey groups in Seward Peninsula Gyrfalcon prey remains computed on a yearly basis . . . -. -12a.-Gyrfalcon eyrie no~ 1, Seward Peninsula, -.A-laska . . . . . . . . . . . . . . . . . 12b~ ·GYrfalcon food_remains from ~able l2a grouped . irt various major food categories ...... . vij_i 13 19 32 45 46 48 52 62 tSB 85 86 Tab1e 13a. Gyrfa1con eyrie no. 2, Seward Peninsula, AJ.aska ..........• 13b. Gyrfalcon food remains from Table l3a grouped in various major food cate~ories. 14a. Gyrfa1con eyrie no. Alaska ...•. 3, Seward Peninsula, lJ.tb. Gyrfalcon .food remains fran rrable J.lta grouped in various major food categories. 15a. Gyrfalcon eyrie no. 4, Seward Peninsula, Alaska. . . . . . . . . . . . . . 15b. Gyrfalcon food remains from Table 15a grouped 16a. in various major food categories ...... . Gyrfalcon eyrie no. Alaska ..... 5, Seward Peninsula, 16b. Gyrfalcon food remains from Table 16a grouped in various major food categories. 17a. Gyrfalcon eyrie Alaska ...•. no. 6, Seward Peninsula, 17b. Gyrfalcon food remains from Table 17a grouped in various major food categories. 18a. Gyrfalcon eyrie no. 7, Seward Peninsula, Alaska. . . . . . . . . . . . 18b. Gyrfalcon food rer:1ains from 'l'able 18a grouped in various major food categories. 19a. Gyrfalcon eyrie no. 8, Seward Peninsula, Alaska. . . . . . . . . . . . . . l9b. Gyrfalcon food remains from Table 19a grouped in _various major food. categories. . . . -. . 20a. .. Gy-rfalcon eyrie ·no. ·9, Sevrard Peninsula, Ala sleet. ·, . . . 0 • • • • • • 20b; ·Gyrfalcon food remains from Table 20a grouped in various major food categories. . ... ix 87 88 89 90 91 92 93 95 97 98 99 100 101 102 Table 21~. Gyrfalcon eyrie no. 10, Seward Peninsula, Alaska. . . . . . . . 103 2lb. Gyrfalcon food remains from Table 2la grouped in various major food cate~ories. 104 22a. Gyrfalcon eyrie no. 11, Alaska ..... Seward Peninsula, 22b. Gyrfalcon food remains from ~able 22a grouped 105 in various major food cateGories. 106 23a. Gyrfalcon eyrie no. Alaska ..... 12, Seward Peninsula, 23b. Gyrfalcon food remains from Table 23a grouped 107 in various major food categories. lOB 24a. Gyrfalcon eyrie no. 13, Seward Peninsula, Alaska .. 109 24b. Gyrfalcon food remains from Table 24a grouped in various major food categories. 110 25a. Gyrfalcon eyrie no. 14, Seward Peninsula, Aln.ska. . . . . . . . lll 25b. Gyrfalcon food remains from Table 25a grouped in various major food categories. 26a. Gyrfalcon eyrie no. 15, Seward Peninsula, 112 Alaska. . . . . 1J.3 26b. Gyrfalcon food remains from Table 26a groured in vario 1,lS ma:j or food categories. 1111 27a. Gyrfalcon ~yrie no. 16, Seward Peninsula, 27b. 28. Alaska. . . . . 115 Gyrfalcon food remains from Table 27a grouped in various major food categories. . . . -· The range in the per cent occurrence and ~numbers of the· three ma,j or prey :categories, ~igratory bird species, and resident species at 23 SeNard Peninsula Gyrfalcon nestings . ~ X 116 ll'T Fir;ure 1. The Seward Peninsula, Alaska, showing the boundaries of the 1968-1970 substudy area, the approximate tree line and the 162°00' Vl. meridian. . . . . . . . . . . . . . . . . . 2. Percentage occurrence of selected ~roups of prey species in summer Gyrfalcon nrey remains from the Seward Peninsula, Alaska, 1968-1970. . . . . . . . . . .... 3. The percentage occurPence of ptarmigan in the 1968-1970 Gyrfalcon prey remains com- pared with an observed subjective estimate of ptarmigan population trends from the Seward Peninsula, Alaska ..... . 4. The percentage occurrence of jaegers in the 1968-1970 Gyrfalcon prey remains com- pared with an observed subjective estimate of jaeger population trends from the Seward Peninsula, Alaska. . . . . . . .... 5. The percentage occurrence of microtines in the 1968-1970 Gyrfalcon prey remains com- pared with an observed ~;ubj ecti ve estimate of microtine population trends from the Seward Peninsula, Alaska . . . . . . . . . 6. The percentage occurrence of Arctic . . Ground Squirrels in the 1968-1970 Gyrfalcon prey remains compared with an observed subjective estimate of the ground squirrel population trends from the Seward Peninsula, Alaska . . . . . . . . . . . . . . 7 .. (A) The estimated biomass o~ ~tarmigan and Long-tailed Jaegers calculated as a per- ._~entage by weight of the avian remains,-and (B) the estimated biomass of Arctic Ground ,Squirrels c-alculated as a ~}ercentage by -weight of the mammalian remains identified -·rrorn-the' 1968-19'{0-Sevrard r·eninsula· Gyrfalcon PTeY remains . . . . . . . ... . .. . . . xi 15 69 '{l 73 75 79a Figure 8 . Major Gyrfalcon prey iters expressed as a percentage of the total of yearly kills by weight: (A) ptarmigan, jaegers, ground squirrels; (B) ptnrmi~an, ground squirrels; (C) ptarmigan, jaeGers; (D) ground squirrels, j aegers. . . . ·-_..., xii 82 INTRODUCTION The Gyrfalcon (Falco rusticolus L.), largest of the falcons, is found north of 59° N. Latitude around the world. It is the Arctic counterpart of the desert falcons, considered by Otto Kleinschmidt (as cited in Voous, 1960) "as the arctic form of the semi-cosmopolitan group of the Saker (Falco cherrug Gray) and Lanner (Falco biarmicus Temmincl:c). 11 This species is closely assocj_ated with tree- less arctic and alpine terrain, and is adapted for catch- ing birds and small mammals on or near the ground. In Alaska, the Gyrfalcon lives and breeds through- out the foot-hills of the Brooks Range, the De LA nO' ....... ....., ...... D Mountains, the Baird Mountains, the Bering Sea coast and Seward Peninsula, the high uplands between the Yukon and Tanana Rivers, the foot-hills of the Alaska Range, the hills between th~ Kuskokwim River and Bristol Bay, parts of the Alaska Peninsula, and (almost certainly) some of the larger islands in the Aleutian Chain. They also have been ~aid to breed sparingly on Kodiak and Nunivak Islands and .. possibly on St. Lawrence Island ( Cade, 196-o). · Cade .(1960} mentions the Talkeetna and Chugach Mountains as . . . - add~tl.~nal possible habitats. I have kno't'l-ledge of at . . least; three -su:ccessful nestings· (Roseneau, ~unpublishe-d) ----o.·-_· ·-:J· . ~--... -- ... ,_ . : --. ----· --- 1 2 ~~, ------------------------------------~~ - ------------------------------- 1971) in the Chugach Mount~ins. Breeding may also occur in portions of the Chigmit Mountaihs (pers. observation). Prior to the present study, little was known about Alaskan Gyrfalcon populations, their numbers~ and food habits, except from Cade's (1960) research. -Cade collected data from portions of the Arctic Slope from the Colville River in the Brooks Range and relied heavily on Alaska Range data collected by Murie (1946, unpublished), M. w. Nelson, and J. H. Doyle. He further attempted to piece together all breeding records of Gyrfalcons in Alaska and to summarize what little was known about food h~bits. Cade' s data indicated that the highest densi_ties of Gyr- falcons. were likely to be found in western Alaska, adjacent to the Bering Sea (including the Seward Peninsula). He Estimated the entire Alask~n population to be.between 200 .and 300 pairs. ._._c'4 ---_j OBJECTIVES This study was formulated to pursue the following specific objectives: 1) to determine the summer distri- bution and numbers of Gyrfalcons on the Seward Peninsula, Alaska; and 2) to determine the food habits of breeding Gyrfalcons. Often studies of predatory animals are confined to relatively small areas by logistic and other problems, and I feel that this is unfortunate. Predators in general tend to be more thinly· distributed than the mem- bers ofrlower trophic levels and for this reason, studies of small areas are likely to be representative of highly local conditions and to suffer from small sample sizes. Accordingly, I have attempted to study Gyrfalcons over a very large area, an area which also possesses the advan- tage of bein_g well circumscribed geographically. This study was intended to be broadly applicable to the breed- ing range of the Gyrfalcon and thus to permit extrapolation and perspectives on Gyrfalcons in the whole circumpolar ·zone.· ·-.;:·.;.._ . ~ METHODS Study Area The Seward Peninsula is the westernmost extension of mainland Alaska .and, because most of it seemed likely to be comprised of good Gyrfalcon habitat, it was chosen as the major study area. Although one aerial search was made east of Golovin, the formal study was conducted west . I of 162° W. Although·potential Gyrfalcon habitat exists east of-162° W., only one aerial survey was made east of that line due.to logistical factors. A portion of the major study area was selected as ~ sub-study area for in- .~-- \"_) tensive investigation near Nome and is described later (see Figure 1). Extensive g~ourid and aerial work was done during the periods 24 May through 30 August 1968, 22 May through 15 August 1969, and 23 May thr9ugh 25 August 1970. The Seward Peninsula is situated between 64° 30' N~ and 66° ·30' N. Although the region lies at approximately · the same latitude as Fairbanks, vlhich is in the center of typi·car·sub-arct·ic boreal forest, maritime influences help- create biotic conditions on the Seward P~ninsula th~t_are ·arctic-1ri nature. The tree-line stops at a north-south line_ rep~¥sent_ing _?.pproximately 162° · 32' i'J., and the ·-·-·...... ·-. peniri~t!J~a-i-s ~practically t-reeless Nes·t of this line. Hov-1...., ··-ever? some _·sprue~ forest extends Nest along the· southern _-{)~·-lJ .. ' l j l ' 1 ~ 1 coast to Golovnin Bay and northwest up the Nuikluk River valley tri a ~oint approximately 15 miles upriver f~o~- Coundil. Scattered spruce continue almost to the North Fork of the Nuikluk River and reach the southern and eastern portions of McCarthy'~ Marsh. The topographic profile is characterized by low rolling hills, generally under 1,500 feet elevation. The land is drained by a complex network of shallow rivers and tributa~y creeks, many of which usually become dry by mi~­ July. Four small mountain rang~s exist . . . The York Mountains, near the western tip of the peninsula, are barren and rugged, rising abruptly from the 5 -~ -0 sea to become high rolling tundra-covered hills a few miles riorth and offer few suitable outcrops but afford some sea- cliff sites. The Darby Mountains northeast of Golovnin Ba.y e~tend north to merge with the more centrally located Bendeleben range. ~he Darby Mountains contain a large section of granite intrusives in the southeast section pro- viding -many spires and "blocks:' suitable for -·large cliff ·nesting raptors. The Bendeleben Mountains are extensiVely -weathered and rounded. The Kigluiak r11ountai11s, just south·· ·~ .. of the·kuZ'itrin River_ and Imuruk .. Basin, are-rugged and rise abruptly~:f:rom -relatively lor! e·levat·ion. A large section _of the· J;_~iij_n_sula riO'rth of'-the .. Be-ndele ben rl!ounta1ns contains · numel"cOU:s -.-c.inder .. cones and is covered by geologically recent -'~,3 ~ava ·flows. : Cinder cones and kett.le lakes cover much of -. :"'·.;;:.··-··:-=.·.:,-: -·· 6 .. the Cape Espenberg sector north of 66° N . . Rock outcrops, often of limestone or its meta- morphic products, occur commonly throughout the peninsula. These outcroppings tend to face south. River bluffs occur on som~ of the larger rivers and ~orne small canyons exist along deep-cutting streams. Sea-cliffs are limited to a few important stretches of coast east of Nome, the.Grantley Harbor and Tuksuk Channel area, the York Mountain section ·between Lost River and Tin City, and some sections of the northern coast between the Goodhope River and the Buckland River. Numbers and Distribution In 1968, exploratory work began on the outskirts of Nome and continued along the three major roads (to Teller, Taylor~ and Council) as theY opened to vehicular i· traffic. r1ost of_ tbe intensive ~1o·rk on study eyries was . -· done on foot and with road vehicles. Snow machines were utili2ed from arrival through mid-Jun~ to explore areas well off of the roads. Rivers were crossed on foot and occasionally by canoe. A skiff and a 38 foot inboard " patrol-vessel (-lent to the Alaska Department of Fish and -pame by-. th.e Department-of the Army) were used for coastal··. ·-trave·l~eastl.,rard and westward from Nome. A Cessna 180 and . -----· -a PA...:.l8~:Super Cub. were used f-or the aerial surveys. My· - 0 7 previous familiarity with Gyrfalcon nesting requirements and s~tuations often allowed me to travel on the ground or in aircraft directly to likely nest sites. Aerial survey te6hniques were developed which allowed me to 16cate nesting cliffs, nest sites, riount young in the nests, and, in some instances, observe eggs. Ratcliffe (1962a) has defined the.concepts of nnesting cliff, !t "nest site, n and "eyrie. •! His definitions will be followed-throughout the text of this thesis. Low level flights wer~ cqnducted during June and early July and on favorable days lasted up to 6 hours~ Optimum flight dura- tion was 3 td 4 hours; beyond this time, fatigue and eye- .) strain affected efficiency. PA-18 Super Cub flights were --~_./ manned by a pilot experienced in aerial· survey operations and one observer; an additional observer covld be accommo- dated on C~ssna 180 flights. Total search hours flown wer_~; PA:....r8 Super.Gub, 32.2 hours; Cessna 180, 13.1 hours. Efficiency in the Cessna 180 was initially mar- ginal because of higher speeds and reduced visibility from the aircraft-windows. -· With experience, efficie·ncy using the Ces_sna 180 · i.ncreased reaching a. level· comparable vd th the -s~l-ewer PA-18 Super Cub by May· 1970. United States· .. : .Geo.l6gfcQ.l S1lrvey topographic maps of scales ~1: 63000 and 1:2500-"QO'""'.were-.:used to .record poidtions.-A Sony. Model· ··-· --~ -" .... _ .. -:"-::·: .. :-;._:_·--_:;:_._ -.. · . T_C-lQO_:-cassette ·tape recorder was used_ to record aerial ./""~ ~.~obs_e~~a-~io~~~ .. Access to nests visited on foot was gained ·(~ ·.~) _c:J a by routine rock-climbing techniques. Methods used in 1969 and 1970 were identical to those described for 1968 with a few exceptions. Snow machines were not employed i~ 1969 or'l970 bec~use of early snow-melt. In 1969 26.9 search hours were spent in a PA-18, 8.8 in a Cessna 180. In 1970, the aerial survey was con- ducted solely from the Cessna 180 aircraft (28.4 hours). One helicopter trip to a remote sectcir was provide~ by Standard Oil Company geologists in late July 1970. During all three years of the study, various men from the. U. S. Geological Survey, particularly Dr. C. L. Sainsbury, pro- vided valuable information and assistance. Food Habits Collection· of the uneate.n portions of prey i terns (hereinafter referred to as "prey remains"), along with reg~rgitated pellets, were made from all accessible occu- pied nesting sites. Prey remains were analyzed tb deter- mine the prey species utilized for food by nesting Gy:rf.a],_cons. Although an .analysis of pellets vias made for -c-omparative purposes, prey remains received the major emphasis-··· Pellets, with the· ex·ce-ption of t_beir m.icrotine c_onten.ts, were_ less informative' due to the qualitative .. - nature of-the-contents ··anti to the fi:rct that· upon-arialysfs the pellets rarely off~~ed evidence of prey specie~ other ·--. . . . than .tbos;e .already represented in the prey remains (with -..:.:. "'" _. -. _.;:.:,. ·,:r •·· \ . .. ' 1 ''· 1.· ~ .C-.'::;-" ~- ·~ ---- the exception of microtines). Regurgitated pellets and prey remains were collected from 10 nesting sites in 1968, 14 nesting sites in 1969, and 16 nesting sites in 1970 {see Table 5, 9 page 115). On the first visit to a nest site, a 1'c1ean-up:' of the nest and its sur~oundings was made. Weathe~ed pellets and remains. were discarded, and only fresh speci- mens were co1l~cted, leaving the places of accumulation cl~an of food residues. On subsequent visits only those resid~es ahcumulated by the breeding pair would be present· in the sample·. Outcrops, ridge tops, and ·grassy hummocks in the immediate area were search~d todetermine the loca,.. (~ tions of favorite perching places. Pellets fo~nd away from the known perches or the nest were discarded since usually they could not be attributed to any one species of raptor (Weir, 1967). Throughout the summer, the.nearby slop~~ were criss-dressed o~ foot; additional kills usually in the form o-f-.. a ring of plucked ptarmigan feathers were noted. The.m&jority of the samples came from the occupied nests,-the· slopes directly beneath them, ahd the two or three pr~mary perching piaces nearby. --: .~...; ... ~ Tne tables were constructe.d using data from .the. . . . -·: .. remailJ.S collected at these locations. These remains ·. ·l:lS1laiJ;i:.,G,Rn.si_!;3ted of actual skeletal material with adhering 3.. . fle.slr:,=rur) or feathers. r ... . . . . \ were u':S7j~J.i,y: ~mitted, since almos.t all such feather-rings Feather-rings on the nearby slopes ·. --.-----~---~~----~-.-~--;_:_:_: .... ---_:_=-:-· ~:. ;-·--- ... ___ ·_-_...:::------.. .-/ .. ::-_··-:- :~ I I I 1 1 .. ,·, 10 represented ptarmigan (Lagopus lagopus and Lagonus mutus) in winter plumage and were obviously from the previous winter or early spring. It was gen~rally impossible to determine if feather-rings .from a prey species represented other kills of that species or if they belonged to the skeletal remains of that species collected from the nest area. When searching·a nest for ~neaten remains a careful search was made for sig-nificant small feathers or skeletal material, in recognit~on of the fact that.the larger avian and mammal remains would be the most conspi- /~ cuous and, therefore, most often recorded species J (Errington~ 1932). This search usually produced passerine and/or microtine remains hidden in the nest litter. Collect~ons made at .each riest sit~ were placed in-labeled plastic bags. It \'las common to find portions (feet' feathers-, etc.) from two or three separate kills or the same· species at the perching -and -plucking places. Each -set was·pla6ed in its individual plastic bag before being added to-the total collection. The collection from any one eyrie on any one date ------. was treated as.follows: ·each labeled pl~stic bag w~s vented to ~llow drying and to prevent mold and decomposition -. ·''-·""" --. -and-p.l'ace-{r-fn -a labeled paper bag; with a liberal quantity -. . . -. -0 of na~·ttrti""c~rystal:s. -These bags l-lere stored in a cool, dry place whenever possible and, once remains were safely dry, _ .·..-:··· I . .: 11 ...---,.. C .. -· •. ~ ~--'1_ .. ' -~:J-------- .y • .. J the sealed paper bags were packed in cartons for shipment to Fairbanks. Identification of the skeletal material, feathers, and other uneaten portions was carried out during 1970 and 1971 at the bird and mammal collection of the University of Alaska .. Whole pellets were broken apart and examined for content. In the majority of cases, the pellets were com- ·-.. po&ed.of mammal hair, small bones~ and feathers from the heads, necks, and breasts of birds (as described by Bond, 1936). Pellets resulting from a meal of· ptarmiga-n commonly contained the antebra,chium and the manus and often the humerus of a wing, or the tibio-t~rsus, tarso-metatarsus, phalanges, and often the femur of a leg. Rarely did the examination of a pellet reveal signs of a species not alrea<;ly-present among the uneaten prey remains. In almost &11 ca~e~ sp~cies represerited in p~llet contents were pro- portioriaL to those. found in the corresponding prey remains. Evidence of microtines was more often found in pellets than in prey remains (by a factor of about five). This indicates ;··-~ that rriicrotiile rodents are ta."ken to a greater degree than t.·muld·-be·assumed· if only prey remains .are examined . .. c ~-·· .::.-. 'Pe.ilet examination tends to reveal qualitative, but-not ~.<I£antita~~ve, information (Erringt.on, 1930,. i·932L . ·,~ · 'rheref_ol'e; only the uneaten portions of prey l"emains were used to~:pev(:?lop a quantitative picture· of Gyrfalcon diet in l!!lll!lll ••• A&Mi,W.Ai'"•.ww . · =· -~ I t j I ! i ·I ! I j I 12 northwest~rn Alaska. .. ·• .-.. •.o;,_:_-":]. -_·· . .:c·_---~--~ -J RESULTS Numbers and Distribution The major sttidy area comprises approximately 17,000 square miles of potential Gyrfalcon habitat, _and includes essentially the whole Sevrard Peninsula. Virtually all of this large area was examined from aircraft or on the ground. The first survey flights conducted in 1968 served to familiarize me with the topog~aphy and terrain of the Seward Peninsula. By aerial observation, it became clear that because_of geologic and altitudinal-factors some-areas were essentially devoid of nesting sites. The Kigluiak .'!f" () t•lountains are steep,· with crests averaging about · 3, 000 feet above sea level. Valleys are narrow, barren, and do not ·o ~upport an abundance of prey species. More important, out- crops suitable as nest sites are rare. The surrountling hills are rounde~d-~nd are. also not ch~racterized b~ rock outcrops. ~In the eourse of thi_:.3_ study; no cliff...;nesting raptors have been observed in this region. In contrast to the rugged Kigluiaks much of the --. Bendele ben range is vvell weathered and-rounded. Some scatt~red·rock outcrops do occur, most in the form of tors, but f-:m,r:offer suitable ledges for nest construction. A few su1"tab1e out~croppi-ngs exist~ primarily along· the v1ater --._, . ,• ~: -·--. . ' co-urses,-generally not far above the valley floors. A rela- .. ·.-· 13 14 tively small number of nestings occurred in the Bendeleben Mountains; the few outcrops available were not heavily utilized. Rock outcrops are rarely encountered in the marshes, poastal plains, lava flows, and broad low basins associated with the larger iriland river systems; c6nse- quently, few nesting sites are available. Rock outcrops do occur in upland 11 hilly" regions, ln faulted areas, on some upland benches, on the shoulders of some hills, and along some water courses. ·Granitic intrusions in some wel.l-fau1~ed zol}es have produced spires and.tors well suited as nesting sites. A relat~vely small proportion of '~ '"_) the coast line offers sea-cliffs suitable for nesting . .. The sub-study area (Figure l) covers about 2,400 square miles and includes the rugged Kugluiak Mountains, coastal marshes and plains along the southv-1estern coast, and t~o relatiVely minor stretch~s of s~a cliffs. ·The remaining po_rtion of th~ · s.ub-study area; primarily. s-outh of the Kigl~i~k Mountains, is good nesting habitat and is charact-erized by---high . ( 1, 000. to 2, 000 ·reet} rolling hills- -. -·. -. · interlaced by 12 relat·i vely short--( 16 to 40 -mfies) ·drainage syst~fus. A-l~rge part of this-hi~ly regiort consists of . metamorph).(J schist: and limestone. These tl'io basic rock types are,_.interspersed and lend themselves \'lell to the formati-on of numerous outcrops and tors through faulting, --··-..,_ . . ' ·. . ' . . ···- .-t) _foldj_ng,: .. thrust.ing, and erosion. These outcrops and tors . "-==' . . . . , ... ----· .· ... ,... . ··.'···_; .... .. ·-, --..... 1 I ·' 'i i; ' . , N '. Bo!lndary of substudy area Serg.Hot . 0Spgs. 'f0.0,· T. ayior v ' ~ 0 -~ ? Tree· line . (approximate) .. '! Figure L • The Set-rard Peninsula; Alaska, sho~Ting the boundaries of the 1968 -1970 substudy ~rca, the approximate tree line and the 162°06 1 .W meridian. 16 are heavily utilized by Gyrfalcons and other cliff nesting "· spec.ies. This area includes about 1, 000 square iitiles and has been designated as Area I. The remaining approximately 14,600 square miles of the Sew~rd Peninsula includes the Darby Mountains, the Bendeleben Mountains, and the rqgged York Mountains. These ranges interrupt the general pattern of rolling 1,000 to 2,000 foot hills and are interlaced by numerous ~tream systems. , .This general pattern of tundra covered hills has a few exceptions; a large lava and cinder cone-covered area .lies between the Bendeleben Mountains and Imuruk Lake an« extends west to a point near the junction of the Kuzitrin ,· C) River and the Noxapaga River. A.large marshy lowland basin contains the extensive Kuzitrin and Noxapaga River systems and Imur~k Basin. These river systems and surrounding pattern of tundra ponds drain into Imuruk Basin north of the .Ki·gluia:k-P.fountains. An additional large area (about 2,500 square miles) extending from Wales to the mouth of _ the Goodhope River consists of wet lowland tundra with ext-ensi-ve systems of sloughs,. ponds' and rivers flm-iing ·into large --·coitstallagoons • F-inally, a Sp·ruce. forest .. .extends ·ove·r .an eastern portion of the peninsula· (see Figure :~11". In all, about four to five thousand square miles of the Seward ·Peninsula outside of the sub-study area are rarely utilized by nesting Gyrfalcons because of the lack of suitable. nesting sites. . I I ( i ~ I l i 17 In the remaining 9,600 to 10,600 square miles, the occurrence of suitable outcrop~ i~ generally low, though these-outcrops are important to Gyrfalcon pairs as nesting cliffs. These outcrops tend to be widely scattered or concentrated in small unevenly distributed clusters, with one important exception--an area of about 1,200 square miles in the central-western portion of the perrin- sula where numerous _outcrops occur. This area has been desi~nated Area II for purposes of discussion. Area II is simila~ to Area I in that it consists mainly of schist and limestone~ --·In a:ddi tion, Area> II has ·a.' maJor faulted zone running through its northern half. Igneous r.ock in t.he ,c-~ ···-J form Of granitic intrusions ITO'i'T existing as large granitic tors occurs where a second important fault trenp intersects the first. These large complex formations are contained _ within a relatively small 20 square mile portion of Area ~I ~hd-offer many suita~le fiestin~ sites (~ee Table 2). .Ruril)g 1,968, 1969, and_ 1970, the sub-study area (Figure 1) was given comprehensive aerial and ground cover- age. -This region of---about -2,4 00 square mi:les contains about _140· outcrops suitable for nesting by at least one of three srecile~ of cliff-nesting raptors-~Gyrfalcons, Rough--fegged HaNks (Buteo lagopus), and Golden Eagles -- (Aquila chrysaetos}. These outcrops also serve as potential -nesting c~ltffs for Ravens -(Corvus cor ax) . Thirty-eight of Q the 140 -q_uterops are known to be suitable as Gyrfalcon- w I ' . I I !' . l I I I 1 1 l i i Table 1. Nur.tbers and densities of brE>edins Gy1•falc~n pairs on the Scw:~rd Penlr.sula, Alaska Area ~· Sub-study 2 .~oo Area_ r 1,000 Ar.ea ;u -1-,200 ==-=T=: ~ 1968 1969 1970 3 years 1968 1969 1970 3 years -1968. 1969 1970 3 years ~Uctivc Nes~ 19 1 1~2 1.2. 52 x = 17.3 16 12 16 ~~ X "' 111.7 42 126.3 171.4 !._?£.,.)_ 62.5 83.3 62.5 X = 171.4 ·75.0 63.16 18 I I I /~ X = 14.0 x = io3.2 ,I I }Area II 1,200 Good coverage years . (1969, 1970) "'../ Entire Seward Peninsula (West o:r 162° W.) ·· Entire Seward Peninsula (West of 162° W.) _ 17,000 1968 1969 1970 3 years Good coverage years (1969, 1970) 35 131 97 X:= 17.5 113.7 i .. 48.5 500.0 354.2 3116.9 i = 401.3 R = 350.5 I f ~ i· I ~-I lrn add1 tion, one other n:ti r '~-"'-~ l<"c"t~':l th'lt -=i\;;~._.'!,.;;;~-to~<t1re-l<'st.::ct'fJ.::c.;:t ,:;-iil.JI:r.h-=-ffi--;-t="=t;~E'1 fluf!rlc:...-:._-=--~ ~ =-=--=--=-----=-=jjj to frequent the nesting cliff until about mid-July. _ ~; ?-one additional single bird was foun~ defending an outcrop where only an empty nest was .i ~epeatedly obse~~ed from-the-a~rcraft. · ~ 3rh1s area received poor aerial coverage durin~ 1968, which resulted in what is certainly a ~ lower number of nestings that actually occurred. The total number listed for 19E3 reflects ~ this. -I I I r: ~ ~ i I 0 .. ., ... ~.~.--,._. -~-_/ '. Table 2~ S~ccesslve use or nestin; clirrs 1 by predatory birds on a 20-squnre-mlle rortion or. the Sol;:ll'.d Peninsula, 196:3-1970 19 _ .. ______ "'="'* ..,.. ~~--=- Site No. Use in 1~68 Use in lCJ~O llse ~!1 1970 l Gyrfalcon 2 Gyrfalcon 3 Rough-Legged Hawk Rough-Leg~ed Hawk Rough-Legged Hawk lj Rough-Ler:ged Eawk Rour.;h-I.er;ged Hawk 5 Golden Eagle Golden Eagle prer;ent 2 6 Raven 7 Gyrfalcon Gyrfaleon3 8 Gyrfalcon 9 Gyrfalcon 10 Gyrfalcon· 11 Rou,;h-Legt;ed Hawk 12 Rough-Ler;ged Hawk !\ 13 r.aven __ _) 14 Raven 15 Gyrfalcon 16 Rough-Lege;ed 'Ha~1k 17 Rough-Legged Hawk 18 Rou~;_h-T.cgr;ed F.awk 19 Rough-Legged Hawk 20 Ro:~t;h-Le::r;e<i ~at·!~-':' 21 -Rough-Legged ·Hawk 22 Raven 23 Unidentified NOT ·usED-BY ANY-SPECIEs· DURING 1968 -1970 36 ··-,AU these cUffs have at least -one old stick nest Ol' an eyri.e ledge fprr.1ed. by the rem-'l.ins or a ·stick nest on them. . _ . · · · · · · . 2At 1east one tllcd...spent cons.l.cie-rable time here and was observed·. a nu:nbe:-of times b:Jt no evidence of breeding was found. . 3Th is t<~as-·the same pa1r based o'n color and behavioral characteristics. / 1 f-- \ t' ~ ....... --- -. -.-. 20 nesting cliffs. Gyrfalcons were observed utilj_zing these outcrops over the three-year study period; reliable local residents reported that Gyrfalcons had used six additional outcrops prior to 1968. About 30 more outcrops appear to be useable based on comparisons of cliff height, ledge size, and the presence of old, large stick nests. Table 1 lists the numbers of breeding Gyrfalcon pairs.and their relative densities for the Seward Peninsula including three sub-areas. During the 1969 survey a decrease in the number of breeding pairs was observed in the sub-study area and in its sub-unit, Arei I. This decline amounted to about 25 per cent and occurr~d in a . . . year when some important prey p6pulations declined and 1vill be discussed further in the Food Habits section. In 1970 the number of breeding ~airs·was found to have returned to . 'th~"l968 leveL. :This ~ccurr~q in a 'year when SOITle impor-. . -. -. ·-- -·_ -. -. ' -. . . . --- --- tant prey species populations appeared to have recovered. to a level somewhat below the 1968-"high" but· markedly above that of. the 1~.69 "low". (see Food Habits section) • ·· Duririg. the course of the study,' 52, or 39.7 per cerit of the 131 total Gyrfalcon nestings observed on the pehinsul~, occurred within the appro-ximately 2 ,lfOO square mile sub-study area. Forty-four, or 33.6 per cent of--the 131 tbtal Gyrfalcon nestings, occurred within the approxi- (-~ mately 1,009 square mile Area I. Of the 52 total Gyrfal_con \_~; . . nest~ngs~pbserved within the sub-study area, ?4.6 per cent I . . . ' -~ :-' ;~---,..Jc ---.., __ -' 21 occurred in Area I. Area II was discovered during the last survey flight conducted in 1968. Thorough coverage was not possi- ble that season. In succeeding years (1969 and 1970) Area II was given thorough coverage. The mean density figure from 1969 and 1970, omitting 1968, is probably a reasonable representation of Area II although it lacks the full per- spective of all three years of the study. -During the course of the study, 42, or 32.1 per cent of the total 131 .Gyrf~lcon nestings observed, occurred within this approxi- mately l, 200 squa-re mile area. Combining both areas of /"~--good nesting .habitat {Area I and Area II), 86, or 65.7 per ',_J cent of all observed nestings, occurred \td thin only approxi- mately 2,200 square miles or about 13 per cent of the Seward Peninsula. Within the ~ntire 17,000 square miles of the -- Seward Peninsula (west of 162° W.) a total of 131 breeding pairs of Gyrfalcons with a mean density of about one pair per 40i-square miles was observed over the course of the study. -Because I was unf~miliar wit!! the terrain and a larger portion of the 1968 search time (about one half) h'as uti.lize-<i to locate areas of concentration and re-check- -- poasible riesting situations, I consider the 1968 total of ·--:-:-.. -::-:.· -··· 34 hre_eding Gyrfalcon pairs to be low and not representative @ of the Sewar~ Peninsula as a \'.Jhole for that year. In 1969, . -a t-0tai e£:--48 breeding pairs of Gyrfalcons \'las found, and in ·.---::-:-·" ----'-"'-"'2-~--:.:;. _ .. -·--=-~-_,_ •.. .; •. _____ .,__ 22 1970 a t9tal of 49 breeding pairs (Table 1). The mean density for these two years is aboat one pair per 350 square miles. This figure, in my judgment, is minimal for the Seward Peninsula, but may be representative of a few other areas of upland coastal habitat in western Alaska. From the present data it appears that Cade (1960) was ~orrect in attributing high densities of Gyrfalcons to · the Seward Peninsula and the western coast of Alaska -ad~acent t6 the Bering Sea. He reported from Eskimo in- formation (the reliability of whl9h he was uncertain) that there were 11 30 known nesting areas· on the Seward Peninsula." It is the Seward Peninsula supports more breedirig Gyrfalcons than any other area of comparable size in Alaska when prey levels are high. I have p~eviously estimated (Roseneau, 1970) the total S·evmrd -Peninsula Gyrfalcon popul'ation during abunda.nt prey years at ·about 70 pairs. This e~timate is based on aerial views of.the eritire peninsula and knowledge of the availability of nesting site~ in some areas that were not .intensively surveyed. ·Assuming the estimate to be accurate, i t .. ls pos_si~.le t_hat densities of Gyrfalcons on the · SeNard -. . -. . Pentnsti~?. -IJ1ay-.be as higp as one pair per 24 3 square miles ·· .for the~s:e -T7-j.ooo -square mile?.• - -tade (1960) estimated the total Alaskan Gyr- ,3 falcon P.ORI!lation including both breeders and non-breeders to be 2o:o t.o. 3.00 pairs. Because of the lack of Alaskan " ' '-23 s.~-:--.:'"2.1con data, he felt that an "adequate conception of :the z.::. z:::: of the Gyrfalcon population in Alaska" could not be :> ... _::--_:_·J.lated. The Seward Peninsula generally has an unusually ..:_~-;:;-:: number of Gyrfalcons. While it is doubtful that many l~€'= areas of habitat as good as the Seward Peninsula ez~st in Alaska, very little of inland Alaska has been e::.:~ined, with the exception of some river courses. Aerial ~~~~~naissance is desirable before reliable population esti- ~~~es can be made, but it is possible th~t the total popu- :~~:on ~h Alaska during good years may exceed Cade's /-~ .. \J :?r:..::~..:: ~at ion Changes Little information on changes in Gyrfalrion popu- ~~~:ons is available. Cade (1960) has summarized the =c~~~Y work on Alaskan populations and best sums up'the 2 ~::~-=-~tion Nhen he states: "Mount McKinley Park and the cr:..:~!lle River are the only two-regions of Alaska where any -z::.:-~:: of' continuous record of the numbers of Gyrfalcons has b~~~ kept." Iri Mount McKinley Nati6nal Park data are 3 -r-,~:::-'se. Cade (from Dixon, 1938, and Murie, _1946) cites ~:;_-:_7 :"·i-V-e known nesting cliffs_ supporting a· maximum _of --~---·. ·. ··-----: .. :~- -"::?~:-o'=~ pai~s"_in _any single year. The Col ville River data .. ..:-· -~:·:::-::-:=::..:::at·e--a· population of three pairs of Gyrfalcons inhabi- _-~~~!~g_1~~-~pper~and middle reaches of the river in 1952. ~@.':•::--::?-59;'--:~~bl~ sanie stretch of river -supported 12 pairs. '-1 . I /·~{· ~-ecce=) · ........ _______ ./ 24 During those two years peregrine numbers remained rela-. tively constant (17 pairs in 1952 and ~5 pa~rs in 1959). In another example from the Colville, a maximum of six pairs of Gyrfalcons have bred in any one year (1959), with an average over five years of oniy 2.6 pairs (Cade, 1960). Apparent numerical conflicts in Cade's data make the total counts for 1959 uncertain (pages 177 and 256, Cade, 1960). On the Seward Peninsula during good survey cover- age years (1969 and 1970), a relatively constant number of breeding pairs was found. In 1968, only 34 breeding pairs wer~ located, but this almost certainly reflects the poor .. survey coverage. In addition) 1968 was vihen the abundance of prey species (including two major food species) was the greatest (see Food Habits section). So far my data suggest that on a regional basis Gyrfalcon populations, within. these large areas of nesting habitat, remain rela- -tively constant, at least on the Seward Peninsula. Fluctuations in numbers did occur within some areas of the Seward Peninsula. These fluctuations in numbers of breeding pairs are similar to those commonly mentioned throughout most of the Gyrfalcon literature, e.g., ·cade (1960) and Hagen (1952). ~ln one 20 square mile porti6n of A~ea II offering abundant bUtcrops, careful ground and aerial reconnaissance located a minimum of 36 potential nesting cliffs showing signs of' }:f;t'evious use by at lea~t one of the four large . :;-·;.·.· .· . ' ~~-.'J-~ -- ,-,_ -\ 25 cliff-nesters found on the Seward Peninsula (Gyrfalcons, Rough-legged Hawks, Golden Eagles, and Ravens). Of these 36 potential nesting cliffs, about half appea~ed to off~r nesting conditions similar to most known Gyrfalcon eyries. Twenty-two of the 36 cliffs wer~ observed to be utilized by the four cliff-nesting species over the period 1968 to 1970. Table 2 presents the history-of occupation of the nesting cliffs in this small section of Gyrfalcon habitat. It would appear that this relatively small and isolated ~ -. . .. . - 11 community 11 of large cliff-nesters, including Gyrfalcons, is in-a constant state of flux. In another area, an approximately 50 mile section one pair of Gyrfalcons breeding on it in 1968, two pairs irt 1969, and five pairs in 1970. On another large inland river (over 100 miles long) only five nesting cliffs occur. In 1968, one was occupied by Ravens .and fourwere unus~d. Iri 1969, the cliff used by the Ravens -· was occupied by Gyrfalcons. Of the four unused cliffs occurring on the same bank of a one-half-mile stretch of the river, one was unused, one was occupied by Gyrfalcons, one was g~eupied ·by Rough-legged Ha11,rks, ··and one was occupiedby Ravens. In 1970 Ravens were nesting again at -. .. - ·thei:r 1968 cliff. Of the other four cliffs, Gyr.falcons again use·d?·their 1969 site. Ravens used the 1969 Rough- i I I l . l l i ·.,'\~··.-.--··· !;;,.:;::.-------~ ..---. 26 legged Haw]< cliff and Rough-legged Ha~'lks used the 1969 Raven site. The one nesting cliff with a well-constructed large stick nest situated under an overhang offering excel- lent shelter had remained unused all three years. In one five~mile stretch of creek in 1968 I found eight nesting cliffs. Three of these were occupied by bree~ing Gyrfalcons, three by breeding Rough~legged Hawks, and two were empty. In 1969, this same section of creek was utilized by three pairs of Gyrfalcons. A pair of RoUgh~legged Hawks inhabited on~ of the remaining five nesting cliffs. In 1970, only one Gyrfalcon pair was found nesting in the vicinity of the creek (four miles no~th), and one of the 1968 Gyrfalcon eyries was being utilized by a pair of breeding Lesser Canada Geese (Branta canadensis). In two successively larger areas of habitat, Ar~a I arid the_sub-study area, variation in yearly riumbe~s of breeding Gyrfalcons occurred (Table 1). Area I, sub~ stu4y area, lost four pairs in 1969. The larger sub-study area lost five pairs this same year. Considering both areas separately, both-experienced a decline in breeding Gyrfalcon.pairs of about 25 per cent in 1969. The 1969 .·decline ~can be attributed in part to a decline in some of the important. food species (see Food Habits section). ''.rhe yearly rilimbers of Rough-legged Hawks and Golden Eagles . ··11 .. ~ ' _..c-~ 27 (i.e., breeding pairs) support this hypothesis. In the sub-study area, 28 pairs of Rough-legged Hawks bred in 1968, only 8 pairs in 1969, and 35 pairs in 1970. In 1968 seven pairs of breeding Golden Eagles were present, in 1969 only tv10 pairs, and in 1970 again seven pairs. The Rough-legged Hawk numbers, in particular, clearly reflect the 1969 microtine "crash." It is possible that the decline in breeding numbers of Gyrfalcons in the sub-study area also refle6fi'the microtine crash as well as a simultaneous decline in the ptarmigan population (see Figure 3 and Figure -5). Frequent references are~made in th~ literature '_J to the apparent response of Gyrfalcons to microtine and @ - - ptarmigan cycles (Dementiev,and Gortchakovskaya, 1945; Hagen, 1952; Cade, 1960; Gudmundsson, 1970; Bengtson, 1971). Gyrfalcon~ are reported to be nonexistent cr scarce breeders if! many areas during microtine or ptarmigan "lows."· It should be noted that these areas tend to rep~esent situa~ ti6ns in which Gyrfalcons are d~pendent primarily on these two ~ategories of food. Dementiev and Gortchakovskaya (1945) did not_ find an interdependence betv1een Gyrfalcons and lemniiiigs-and-'suggests Gyrfalcon utili-zation of a sea bird-·colb:riy as. an explanation. =.-..;. :-~There are indications that some· food species, notably,~tarmigan, Long-tailed Jaege~s, and microtine~, did I I I I I I I I I 1 l l I _j i . I l ' -~~J---- I ' not decline to the same degree ·in the northern half' of' the Seward Peninsula during 1969 as they did in the southern -half. --If t-hese indications ·are correct, one _ !1Jight speculate that numbers of raptors, including sorrie Gyrfa~cons, shifted breeding activities to the northern parts of the peninsula in response to this situation. 28 Although Rough-legged Hawks were not of primary concern, nest locations were noted. Only six occupied nest ~it~s were observed in 1968 in the northern half of the peninstila; About six more pairs were observed from a distance and were_ probably breeding in this sector. A higher concentration (28 pairs) occurred in the southern ·(: --~ -'\; __ .__) sub-study area. In 1969 34 breeding pairs of Rough-legged Hawks were found in the northern half of the peninsula, but only eight breeding pairs were evident in the sub-study area. 1970 appeared to be a year of general abundance for Rough-legged Hawks over th~ entire Seward Peninsula •. - 'J.lhirt_y-five pqirs were found nesting -in the sub-study area and_38·pairs ih the northern half 6f the p~ninsula. In 1969-, when 14 breeding pairs of Gyrfalcons inhabited the sub-st~dy areai more nesting p~i~s ~er~ evi~ent in a strip -north of· t-he sub-study area" but south of Area II;-.. - --_In a very large region of habitat,_ such as the };7_, OO __ p _§q)J.a~e miies-of the Seward Peninsula conside;red in __ this ·study,-· the numbers of Gyrfalcons probably remain rela- (J tively c~c)nstant unless a major portion (or all) of the /~ region is affected by drastic changes in prey population size, availability, or vulnerability. Cade (1960), speaking of Gyrfalcons, states: "One pair may have several alternate aerie areas, or, as 29 appears to be true on the Colville, pairs actually breeding from year to year· fluctuate greatly in any given region; he~nce' there is no meaning to a static estimate of breeding population size." I contend that estimates of the breeding popula- ti~n size of Gyrfalcons ha~e meaning only when applied to !!regions" of an isolated nature and/or of large_size (such as the Seward Peninsula). From my observations, it is ,~) . possible that the changes in cliff occupancy or the flue- tuations in numbers of breeding pairs in areas reported in the present Gyrfalcon literature may be relatively local in nature. These changes probably. reflect a population. shifi io~other·areas of a large ~egion rather than a general non-breeding of Gyrfalcon pairs. Few authors . . . ·~. . .r . . ·- state whether non-br~eding Gyrfalcons were observed attached to·nestin~ cliffs ~r if they were in the vicinity of the cliffs. The implication is usually that they were ·_ not. Hagen ( 1952) claimed the 11 • • • existence of non- breedines· y·ears in the life of a gyr-falcon pair. 11 and is based on-what is, in_ my opinion, insufficient. data. Cade ,_ ...... ; . .... :; .•. :., .. ·". . ~ ... · -·.ao .. ..:,.. ----· ( 1960-)-trnP-J.ted that a Gyrfalcon pair,' if not· breeding at (~ the sameL.ne:::?t;ing cliff each year, v1as not breecting~ .. at al·l- I I I 30 ---~~: .. ------------------ ' ·,_ --- that·year. Within a large region of abundant nesting sites, th-e-possibilities of local population shifts or at least breeding attempts in other areas would seem more reasonable than actual non-breeding. On the Seward Peninsula during 1968-1970 only two pairs and four other individual Gyr- falcons were observed that could h~ termed possible non- breeders~ Further research will be· required to gain a better u~derst~nding of_ the possible non-breeding of Gyr- falcon pairs and how such occurrences may relate to the population as a whole. The theme that Gy~falcons not breeding in one area or region may in fact breed in another area or region /~ \_J that was· not investigated is often understated. The only circumpolar region where there are definite indications that Gyrfalcons actually stay in an area~ or actually ·occupy an eyrie and do not attempt to breed, is Iceland. . . Gudmundsson (pers-. comm .-, 1969) mentioned-such observations in th~ Lake Myvatn region. Gudmundsscn (1970) states: . uAs the ptarmigan is the staple diet of the Gyrfalcon in Ic~land it is not surprising that the Gyrfalcon t1u6tuates ·w±trr·the..,ptarmigan·and that ·during years-of ptarmigan sca:rcity many Gyrfalcons do not nest at all. 11 ···- Region-wide Gyrfalcon population fluctuations probapl_y_ ct·o, in fact, occ~1r in Alaska (and throughout~ the _·--Gyr-falcon range)_; but· further ,study is needed to document @ -~--~~~-. I . . . ··-·---~--~- . ~ .l ' i ' -· . '· I ' . ' ; ' . I I I i . ' ' ' I I "' f) ·, _ _} 3-l the scope and degree of these changes in population numbers, and to document non-breeding. Gyrfalcon nesting cliff tenacity was low oh the Seward Peninsuia. Of the 34 active Gyrfalcon nesting ·cliffs located in 1968 (Table 3), 18 (53 ~er ~ent) were found inactive and unused by any species of cliff-nester in 1969. Eight (24 per cent) were used again by breeding Gyrfalcons. Of the remaining seven, three were occupied by Ravens 1 two b~ .Rough-legged Hawks, one by a pair of Lesser Canada Geese, one by an unidentified species, and one was not checked~ In 1970, 22 (65 per c~nt) of the 1968 Gyr- faicon. nesting clif-fs were found to be inactive. Foul"' (12 ~~-___ ,_' .... ·----e· }'C:J._· ~,.;o;:;uv; wer utilized by Gyrfalcons. Of the remain-· ing eight, one was occupied by Ravens, five were occupied by Roug}1._-legged Hawks, one by Golden Eagles, and one by an ·unidentified species. Twenty-three (48 per cent) of the 48 ·active Gyrfalcon nesting cliffs located in 1969 (see Table 14) --. { .. were found-unoccupied by any species in 1970. Fourteen (about 30 per cent) were occupied again by ne~ting Gyr- falcons .. Of the remaining 11, th~ee were occup~ed by Ravens) on~·by Reugh-legged Hawks, ·one by Golden Eagles, and .sX~ we_re not. checked. In about five to six per cent o·f the· cas-es where the nesting ciiff vms unoccupied, the .. . - nests wer~_: g"one' making the· nesting cliffs at least _ .tempo:r(lr-i;I.y_ unusable. -·'-~~A total of 131 Gyrfalcon nestings v.rere observed t I I I i I -~:-9--------------~----. - - - - - - - - - - - - - -- --32-- -- -.. - - L·~ ···._~) Table 3. Species u_tilizing the 34 active Gyrfalcon nesting cliffs observed on the Seward Peninsula, Alaska, 1968-1970 1968 1969 197-0. Species Number Number Number Gyrfalcon 34 8 4 Unoccupied 18 22 Rough-legged Hawk 2 5 Raven 3 1 Golden Eagle l Lesser·Canada Goose· 1 Unidentified Species· 1 ,. 1 Unknown 1 Total 34 34 311 ·-· . - ?./~ --~ /'\ \_~o..7) 33 Table 4: Spec~es utilizing the 48 active 1969 Gyrfalcon nesting cliffs observed on the Seward Peninsula, Alaska, 1968-1970 1968 1969 1970 Species Number Number Number --- Gyrfalcon 9 14 8 14 Unoccupied 22 23 Rough-legged Hawk 11 1 -Raveri 1 3 Golden Eagle 2 1 Unidentified 1 -Unknown 10 6 Total 48 i48 }~ 8 -----=. ·d 34 over the cour~e of the study. These nestings occurred ··at 107-nesting cliffs. Seventeen nesting cliffs (16 per cent) were involved in Gyrfalcon nestings during two successive years (1968 and 1969, or 1969 and 1970). Only 20 (19 per cent), including the 17 mentioned above, were utilized two out of every three years. Only four other nesting-cliffs (four per cent) 1·1ere occupied by breeding Gyrfalcons during all three years (1968, 1969, and 1970). Eighty-three (about 77_ per cent) of the 107 nesting cliffs were utilized by Gyrfalcons only one during the three years. Twenty-seven (about 25 per cent) of the 107 nesting cliffs were utilized once during,the three years by other /-----.... r-- \ ·--~) · cliff-nesting species. Sixteen of' these cases involved .-.-~ Rough-legged Hawks, five involved Ra~ens, four involved Golden Eagles, one·involved Lesser Canada Geese,,and one involv~d an unidentified -species. .. Most nesting cliffs had at least one and often two (or more) alternate nest sites that had been utilized by_ the cliff--nesting species at some time in the past. Of the 20 h~sting cliffs ~h~re Gyrfalcon nestings occurred in -- two out of the thr~~ years, the same nest site was occupied iri at -least-ten cases. In the other four_instances Where the _nestj;p_g ·cliff l'ras occupied all three years by Gyr- falcons, _.there are a total of eight possible nest sites (two a:t; each nesting cliff). -THo of these nest sites were- utilized-~ll three years~ two were utilized two out of the -.:..__. --~-~-,---·~·. 35 three years, two were utilized only once, and two were never used. In general, all existing nest cliffs on the Seward Peninsula that showed signs of having had at least one stick nest constructed on th~m sofueti~e in their his- tory (estimated to be 400-500) had one nest site that appeared from signs of useage to play a predominant role in nestfngs by any of the cliff-nesting species. In subjective terms the situation appeared to follow that of the nest site useage observed in the sub-study area and I believe that these predominant nest sites are involved in about 50 per .~ cent of the nestings occurring on these nesting cliffs and \~J' therefore might be considered as "traditional" nest sites. This low nest site and nesting cliff tenacity (comoared to that attributed to Gyrfalcons and some of the eth~~ large ~alcons [Cade, 1960]) is somewhat opposed to the limited Alaska Range data. In one case, Gyrfalcon pairs h~ve occupied a nesting cliff with only one nest site for the l~st nine years and probably longer~ Prior to 1~63, t.bis .ne.st was occasionally visited by successful egg collectors a.r1d j.n at least t'l'lO years (Hr. Vern Seifert, -~ pers. cornin., 1969) the pairs were shot. Cade ( 1960) dis- cusses~ __ t.r.~:tli~tional nesting ciiffs and hm'l nesting -cliffs -such as thi:S one may develop relatively long histories of . - use by ~fa} cons . . · ~-:i__-Tt. is clear that. although the total Gyrfalcon ~--w~: ., . I . I I r I t I l l ~.--_·-_:;·"'". -- "-._ @ -~- - 36 population p~obably remained constant the population under- went conside~a~le shifting from area to area, nesting cliff to nesting cliff, and nest site to nest site. It appears that in a small area of habitat or along a short stretch of creek Nhere nesting cliffs are common considerable 11 cliff-shifting" occurs (i.e., the utilization of different nesting cliffs by what may be the Gyrfalcon pairs). It is also common for the number of . . . . . breeding Gyrfalcon pairs inhabiting the area to change from year to ~ear, and for some vacated nesting cliffs to func- tion as nesting cliffs for other large cliff-nesting '.t spec_~.es. tha deg~e~ to which these phenomena occur may vary locally due to changes in local prey population~, reduction of nest sites (nests falling off of cliffs or be~ng d~stroyed by rock-falls), creation of nest site& (the construction of a stick nest on some cliff ledge by Ro~gh-l~gged Hawks, Ravens, or Golden Eagles), or even the behavior-of Gyrfalcons themselves. This cliff shifting and alternation of nesting ··cliffs-a~d.nes~ sites probably doe~ not ~eflect the implica- _tions of the comment made by Dementiev and Gortchakovskaya .(i9LI5).when they stated: r'Thus it may be seen that-the Gyrfal~on used t6 breed on the same territory constantly . . - ehough~ though not every successive year, having probably been_ chased off by the Raven.!! I believe it is unlikely that Gyr:r:alcons are "chased 11 from or forced to abandon __---,_ i~_:) -,__-.!-:_J"=~ j--- -, 37 nesting cliffs and nest sites by any of the three other important species of cliff-nesters. Cade (1960) comments on this same statement and our conclus~ons coincide. Cade (19~0) also reports that he found a dead.uneaten Raven, presumably killed by Gyrfalcons, below ~rt eyrie on the Oolamnagavik River in 1956 and he mentions one other simi- lar instance recorded j_n the literature (Brull, 1938). At . one i96~ Gyrfalcon eyrie I found a freshly killed but_ uneaten body of a-Raven "cached il beside the nest--an obvious .loser in some aerial encounter. . . Information on nesting relationships between -~ GyrfaTcons and Ravens, Golden Eagles, Rough-legged Hav1ks, · arid Peregrines is summarized by Cade ( 1960). In 1968 I observed the successful nesting of a Raven pair and a Gyrfalcon pair on the same hillside about 300 yards apart. In 1969 and 1970 a pair·of Ravens successfully ·nested about 100 yards Trom an active Gyrfalcon eyrie ln the ~ub-study area. The nests were at approximately the same level and in plain sight of one another. On one occasion in 1969, a 45~minU:te battle between the :pairs-\'Tas observed (the birds '\tier~ unaw~_:r·e of my. presence both before and durlng the conflict). The initial encounter was not witnessed, btit from that. "Point on the Gyrfalcons were the aggressors. The male Gyr:t::_alcon as initial combatant against both Ravens \'las joined at th~ end of the first ten minutes by the female -· ·--·~-·-"·.· . which ha·J"-~b~en brooding eggs or nevdy hatched young at the 38 eyrie. The male immediately stepped up the force and frequency of his attacks. The female Gyrfalcon retired to her eyrie after 35 minutes and her mate broke off the attack moments afterward. On many occasions one or both of the Ravens succeeded in returning to their nest to perch or brood the newly-hatched young and were forced to leave the nest because of extremely powerful close oasses by the male Gyrfalcon. The female Gyrfalcon also was successful in driving the Ravens from their nest a few times, but she did not put as much effort into her attacks, often breaking off her passes before an actual encounter occurred. However, she tended to be more vocal. The Ravens were completely successful in avoiding actual con- tact, ~sually by side-slipping at the last moment. At each encounter of a Gyrfalcon and Raven, the Gyrfalcon . presented lts feet as if to strike, while the Raven coun- tered by rolling and presenting its feet toward the falcon (usually while in the act of side-slipping). In two cases, one of the Ravens actually flew upwards to meet the male Gyrfalcon on a direct collision course, forcing the f~lcon to flare off, but ~ithout a doubt the Gyrfalcons demon- strated a clear advantage over the Ravens. The termination of the aerial battle appeared to result from fatigue and loss of interest on the part of the falcons. The Ravens did not appear to withstand the physical demands of the battle as-~ell as the falcons. During other observations l . l I 39 at this eyrie, including two three-day periods, no other encounters or conflicts were observed between the two species as they both attended to the feeding of their young. From 1968 to 1970 Raven pairs nested in close proximity (100-400 yards) to Gyrfalcon in seven instances. no encounters between G8ldcn !~agles and Gyrfalcons were observed, though they must occur (Cade, 1960). In one instance Golden Eagles nested about 400 yards away from a Gyrfalcon eyrie on the same hillside and at nearly the same elevation. On one occasion, both Gyrfalcons were observed to watch one of the eagles from their nesting cliff as it soared briefly over the hill top. The nests .,,;ere hidden fr·om each other by a small series of outcrops and the falcon nesting cliff faced the back side of the eagle nesting cliff. Rough-legged Hawk encounters with Gyrfalcons were numerous and paralleled the observations of Cade (1960). In one instance, a male Gyrfalcon was observed stooping at a male Rough-legged Hawk that had its nest about 200 feet away on the same cliff. Both birds were unaware of my presence. The conflict began when the Rough-legged Hawk; soaring parallel to the cliff, passed directly in front of the falcon eyrie (and about 100 feet out from it). The male Gryfalcon was resting on a perch about 100 feet beyond his eyrie (where his mate was brooding eggs) and as the Rough- .c•, --- l_ _t_t:W to intercept lj 0 the intruder. The Rough-legged Hawk side-slipped to avoid contact and both birds began loudly vocalizing. During the next ten minutes t~e Rough-legged Hawk continued to circle about on a course that brought him closer to his own nest while the Gyrfalcon repeatedly stooned at him from above. I was impressed with the apparent ease with which the Rough-legged Hawk avoided the Gyrfalcon's intent stoops by employing a combination of watchfulness, abrupt hoverin~ and side-slipping. The intensity of the battle slowed down after about the first five minutes and continued to do so until both birds drifterl back to their respective portions of the cliff to perch. In another instance, where Rough-legged Hawks nested on the same cliff and about 100 yards away in direct vtew from nesting Gyrfalcons, n1y approach flushed a newly fledged Gyrfalcon. The young .falcon flevJ' up the river and under the Rough-legged H<:nvk nest (which contained five young about to fledge). Both Rough-l~gged Hawk adultB dove on the young Gyrfalcon almost .forcing it into the river. The .femal~ Gyrfalcon, soaring overhead, responded immediately and nkekking, 11 stooping hard at fhe Rough--legged-Hawks, narrowly-missing· -one before they could continue their harassment of the young Gyrfalcon. The Rough-legged Hawks retreated and the female Gyr.falcon turned her attention to.me. Duri~g the course of the study, Rough-legged Hawks are ~nown to have shared the same cliff with Gyr- falcons in at least nine instances. The nroximity of the nests varied frorn 200 feet in one case to about 400 yards. Many· additional ca~~s existed in which Rough-legged Hawks nested on another, often smaller outcrop, within 200 to 400 yards from active Gyrfalcon nesting cliffs. Food Habits No attempt is made in this paper to condense the circumpolar data pertaining to species known to have been killed by Gyrf~lcons. Hagen (1952) and Cade (1960) have already summarized a considerable portion of this informa- tion; those and three other papers (Murie, 1946; Dcmentiev and Go::-'tchakovskaya, 191!5; and Bengtson, 1971) are of rn·i-- mary importance because they involve: (1) reasonably larsc sample sizes; (2) observations over more than one year 1 s time; and/or (3) ojse~vations at more than one eyrie. The general characterist:cs of Gyrfalcon diets, based on these papers, can be summarized into the followjng points: l. Gyrfalcons rely heavily on resident species of birds and mammals in the Arctic and subarctic. a. Gyrfalcons prey heavily on V!illow rtarmi·- _._.,_· Ptarmi~an-ean be considered as the single ~ost important food item· (Cade, 1960; Hagen, 1952; Bengtson, 1971) on a circumpol~r basis. 42 b. Small mammals are known to be an important food source on a weight basis in Alaska (Murie, 1946; Cade, 1960). The Arctic Ground Squirrel is particularly important and can surpass ptarmigan as the most utilized food species in some areas of Alaska (Murie, 1946; Cade, 1960). The lemmings (Lemmus lemmus, the Norwegian Lemming, !::· trimucrona tus, the Brovm Lemndng, and Di_cro~C2.!_1Y x £_;roen- landicus, the ColJared Lemming) on a Horld wide basis see:n to play more variable roles and usually are not as imnor- tant as food (Dementiev and Gortchakovskaya, 1945; Hagen, 1952; Cade, 1960). In J\L1sJ~a the Hro1·m Lemming, the Collared Lemming, the Tundra Vole (Microtus o~conomus), the Singing Vole (~. miurus), ·anrl the Red-backed Vole (91ethr_~_C?_r:!?mV~ rutilus) appear to play an important but indirect role in relation to Gyrfalcon fooc<. habits ( Cade, 1960). Large numbers of microtines commonly a.ttract laqr,e numbers of rodent-eating birds, many of which are utilized by Gyrfalcons for food probably in greater proportion (at least by weight) than the microtines themselves. 2. In Iceland a relatively stable and large population of ducks is heavily utilized as an important source of food by Gyrfalcons nestin8 in, or frequenting, the Myv_a:tn area. Utilization of Rock Ptarmigan increased as the ptarmigan population increased suggesting a prefer- ence for this food species over the ducks (Bengtson, 1971). lj 3 Salomonsen (1951) considers waterfowl to be one of two principal ~ources of food to Gyrfalcons during the nesting season in Greenland. 3. Dementiev and Gortchakovskaya (1945) docu- mented a high utilization of seabirds (Alcids, LariJs, and Anatids) by Gyrfalcons nesting near seabird colonies on Kharlov Island. Salomonsen (1951) considers seabirds to be a principal source of food for nesting Gyrfalcons in Greenland and indicates that Gyrfalcons usually nest near seabird colonies. 4. Gyrfalcons, on a circumpolar basis, take a relatively wide range of prey other than the species listed above, ranging from smaJl passerines to other raptors. Gyrfalcons, however, tend to take a narrower range (in terms of numbers of species) of prey than rto Peregrine Falcons (Falc~~ per_Qr;rinus) (Cade, 1960). ·5: In general, it appears that Gyrfalcons rely principally on two or three prey species as a source of food (Cade, 1960), and that coastal-nesting pairs feed chiefly on Alcids, Larids, and Anatids while inland-nesting bir~s feed chiefly on ptarmigan and sDall mammals (Dementiev. and Gortchakovskaya, 19115; Cade, 1960) ._ 6. Gyrfalcons also appear to be opportunistic, capable of taking a wide range of sizes of prey, and are able to shift to other food sources when ptarmigan become scarce (Bengtson, 1971; Cade, 1960). 411 7. In some parts of the world Gyrfal.cons probably fluctuate with ptarmigan cycles (G11dmundsson, 1970) and in some instances appear to breed in lower numbers in response to ptarmigan lows. The degree of this relationship to ptarmigan poptllations probably varies from region to region and is poorly understood. Fluctuations of Gyrfalcon numbers have been reported by Salomonsen (1951) in connection with lemming nur:1bers in Northeast Greenland. Prey utilization on the Seward Peninsula During the course of this study a total of 1,1133 Gyrfalcon kills were identified from the immediate area around ~0 Gyrfalcon nestings (see ~able 5). Avia~ snec10s constituted 85 per cent by nurher and approximately 87 oer cent by weight·of the total kill. Mammalian species consti- tuted the remaininr.; 15 per cer..t by number and 13 per cent by weight of the prey remains (see Table 6). Cade (1960) reports lower mammal subtotals for ten Colville River eyries (13.6 per cent of total/6.6 per cent of total/1.2 per cent by weight based on 142 kills), but a much higher mammal subtotal (76.2 per cent of total/80.1 per cent by weight based on 429 kills) for three Alaska Ran~e eyries (data obtained from Murie, 1946, and unpublished; M. W . . Nelson; and J. H. Doyle). This marked difference in the proportion of mammal remains can be attributed almost entirely to the differences in frequency of occurrence of Table 5. Collection dates of Gyrf::tlcon jlrcy rer~ains J-:yrj_£ 1 .2 3 5 8 9 10 17 18 19 Total eyries 2 3 4 5 6 7 8 12 ,c .LU 20 21 22 23 2 Vern-69-1 Total eyries 1 2 5 9 11 12 13 14 15 16 21 24 25 26 27 28 Total _ey:ries 'I.'otal eyries Total nest-:l,r.gs Year 1968 1968 1968 1968 1968 1968 1968 1968 1968 1968 10 1969 1969 1969 1969 1969 1969 1969 1969 1969 1969 1969 1969 1969 1969 14 1970 1970 1970 1970 1970 1970 1970 197'0 1970 1970 1970 1970 1970 1970 1970 1970 16 29 = 40 22 June; 16 Aur,ust 6 June; 11 July 16 June; 6 July; 23 August 4, 9, 14, 25 June; 5, 23 July 21 July 31 Nay; 9 July · 9, 14, 23 Jun0; 6 July 13 June; 12 July 21 June 19 July Number of visits = 24 19 June; 3, 13 July; 12 August 211 May; 16, 23, 24 June; ·r, 11, 17 July 20 July 16 June 21, 26 June; 8, 12 July; 5 August 11 Aup;ust 13 June 5, 13 July 27 r~ray 13 June 13 June 20 June 16 July July (date unknown) Number of visits ~ 26 25 July 1 June; 1 July 29 Nay; 4 June 8 June 7, 8, 13, 14 July 6 June; 7, 8, 13 July 27 July 28 May; 20 June 24, 25 June - 31 May; 5, 12, 22, 27 June; 9 July 26 July 23 June August 15 July 15 June 27 July Number of visits = 29 Grand total number of visits = 79 lNot included in the total. 2This cy~±~ ~as located by Mr. Vern Seifert, who collected ~;hat remains he could find for donation to this project. T:tbl~ 6. Tot-:\1 106~-1070 Cyrrnlcon food reM~iM TOTAL BIRDS Aretie Ground Squ1r~~~ (Sae~~on~!1us un8ul2tu~) Collar~d I,ei..rn1ng Cp!.::-~stc~-'X.. z.ot~~}j.;o.~1co.:s) Unlct~nt1t1ed le~n1c.~> Drown Lcl""' . .'lling ((.l:' .... r.,u-: t:"'!mucronntu:~) Un1dent1r1ed voi::?s--• -- Red-bac~ed '/ole (r.le~:-"_-:onor."-;5 r'•tilus) Un1dent1fled r.ic~o~ices --- Tundra Vole _q~t-:~ :--"'~2.!!2~) Short-tailed ~:cas~l (:--·-.::.tel3. cr!"lin~a) Mink (~~~ y~) -------------- TO:JJ. I·'J,X.':t.LS Un1dent1t1ed kills "'l'OTAL KILLS Total categories -49 1'ote1 identified op~~:-< • -39 (31 ev1en, 8 r.:ammal) 'l'ot~l miri1mun spcc1eo ""-~0 (32 av!a:'l, 8 roa"'-"'al) "'Oata rrom Cade, 196C ~·ota1 ~ 876 155 30 29 23 1~ 1~ 13 12 11 11 8 7 7 6 5 5 lj 3 3 3 2 2 2 2 1 1 1 1 1 1 1 l 1 1 1 1 ~ 1,260 170 24 8 6 5 ~4 2 1 1 _!. 262 _! 1,~83 Pe!> cent ~--~ 59.1 10.5 2.0 1.5 1.6 0.9 0.9 0.9 o.a 0.7 0.7 0.5 0.5 0.5 o.~ 0.3 0.3 0.3 0.2 0.2 0.2 0.1 0.1 0.1 0.1 0.1 83.5 llS 1.6 0.5 0,4 0.3 0,3 0.1 0.1 0.1 1~.9 .Jl.d 100.0 ! .02 r(\'r ~<""nt F!r:1-···:1:-~ 6'l.5 12.3 2.4 2.3 1.8 1.1 1.1 1.0 l.O 0.3 0.9 0.6 0.6 0,6 0.5 0.4 0.4 0.3 0,2 0.2 Ow2 0.2 0.2 0.2 0.2 0.1 -- 97.6 61.1 l0.8 3.6 2.7 2.3 16.8 0.9 o.s o.s 107.7 A~pl•c-rr1.J:te rc r· cent \-H'h"':ht cl:tss or tot.1l !!l.L:::"':.~-~ !,:;. W~1,•M •<so 71.6 ·3~0 7.4 '135 0.6 • 30 0.1 100 0.2 380 o.s 100 ~.2 ·~ao 0 -, 370 0.6 Sc 0.1 sco C.1 1 90ii 0.9 tOO 0.6 275 0.3 •goo 0.7 § 30 0.1 500 0.3 320 0.1 500 0.2 30 0.1 110 0.1 12 0.1 17 0.1 35 0.1 1.~50 0,2 300 O.l 45 C'.l O.l 100 O.l " 30 0.1 .. .-.... -0.1 -.;JU 200 O.l 200 0.1 ).0 0.1 HO 0.1 20 0.1 ---.12 .. ~: .. .!. 86.9 •soo 1"2. 6 50 o.a so 0.1 50 0.1 30 0.1 30 0.1 30 0.1 •o 0.1 200 0.1 ill _Qd 12.9 lCO.O the Arctic Ground Squirrel in the prey remains reported for these different regions, and probably reflects this species' . availability at the times the.data were collected. On the Seward Peninsula, three categories of prey stand out as major sources of food to the Gyrfalcons, both by number and weight (see Table 6). These Dajor prey species are shown separately in ~able 7. Ptarmigan clearly appear as the primary prey. Though no attempt was made to separate the two species (~. laGonus and~· mutus), it is known that both were taken. Ptarmigan co~prised 59.1 ner cent of the total prey reJ'lains ancl 71. G per cent by body weight. That the various species of ptarmigan form a major portion of Gyrfalcon prey throughout the Gyrfalcon's cir- cumpolar distribution is generally well known (Dementiev and Gortchakovskaya, 1945; Hagen, 1952; Cade, 1960; Drown and_Amadon, 1968; White and Weeden, 1966; Ben~tson, 1971). Ca.de._ .(1960) noted a preference 1'even in a year when they (i.e., ptarmigan) vv-ere scarce!! from his Colville River data. Hagen (1952) comments on this preference when microtines were abundant and available to Gyrfalcons in Dovre, Norway, and Bengtson (1971) states that in the Myvatn area of Ice- land, trptarmigan, when abundant, a1'e the preferr.ed prey even in areas containing. excellent habitat for vraterfov.rl. :r Bengtson -(1971) reports an interesting increase in pta~migart utiliiation by Gyrfalcons as otarmi~an increased in numbers I . I 'l'ab1<: .7.. 'l'h~ ttlr<::e maJor Gyrfalcon food items Per cent Per cent ~ Humber 1968 Total flircl->•:a;:~mal ?tarnica~ \!911 G4.~ 7~.8 Ground :;quirrels 43 9.4 68 .'3 Long-tailed Jae~.;ers 42 ...2.:2 lCi. 7 TOTAL 379 83.1 Per cent 1968-1970 ?tar~.1JO;an 876 59.1 69.5 Grour.t'J ::;gu1rreln 170 11.5 76.6 I,rmr:-tn11crl Jaer:crs l~ ~ l1.:J. T0'1't\r1 1,201 ill.O Per cent Per cent ~~ 1969 Tot't1 B1 rd-!·lammal lit] 48.6 62.7 63 21.11 95.5 .Ji ..2.:.!!. ...1...:.!2. 222 75.4 Numt>er 439 64 -21. 600 Per cent 1970 Total 59·9 8.7 l.U 81.8 Per cent Bi:--d-~!3:7.l~al 63.7 63.8 15.2 ~:ote: \.lillow Ptr.trm1p;:tn <k· laro:ous) and Rock Pt!lrnir.;an (J.. r:111tu0) ~:ere concl.dcrcd as one category. Xo attcm~t was made to separate these two species. Three additional jaer,cr kills could not be identified to specie5, but were probnbly Lnng-tailcd Jae~crs (~. lonricnuclu!)• A fourth kill was n Pnras!tic Jaeger(~. pnrn~ttleus). CJ in an :=trea whe·re thous:=tnds of cluc-ks forr.1ed a readily available and heavily utilized fc6d ~ource. lj 9 It appears that these observations of ptarmigan "preference" are generally valid. Such a nreference for ptarmigan by hunting Gyrfalcons seems logical and expected for an avian predator that is almost completely dependent upon them for about two-thirds of the year. From the extensive literature of falconry, it is generally known that trained falcons and hawks can readily develop specific interests in a particular prey species or type. Although almost no data are available to axnlain what young Gyr- falcons do during the winter months, it is probable that they too are strongly dependent upon ptarmigan for food as winter sets in. It seems likely that ptarmigan are strongly fixed as a nprey imagen in Gyrfalcons; that this occurs early in the life of a Gyrfalcon; and that the i'preference :I could be. termed one of necessity. Cada (1960) reported that ''On the arctic ~lope ptarmigan usuallymake up nearly 90 ner cent by weight of the total consumed. 11 Certainly the hir;h availabili.ty of Long~t~iled Jaegers and Arctic Ground Squirrels on the Seward Peninsula accounts for a large portion of the approx-· imately l8_per cent difference between Cade's data and nine. Arctic Ground Squirrels were the second most fre- quently ·bbdu~rfng prey species in the Seward Peninsula prey .t'ema:Lns . (see '.L'ab le 7) and compr'i sed about 13 per cent by 50 weight of the total Gyrfalcon prey remains (see Table 6). Cade (1960) re~orded only six eround squirrels constituting abou~ 5.8 per cent of the total prey remains and about six per cent by weight from ten Gyrfalcon eyries on the Colville River in 1959. Eieht arctic slope eyries where prey remains were picked up from 1952-1957 produced only one ground squirrel which contributed a small 0.7 per cent by number and 0.8 per cent by body weight of the total prey remains. Murie 1 s (1946, unpublished) Alaska Range datfi (reported in Cade, 1960) from three eyries reports 259 ground squirrels totalling 60.5 per cent by number and 78.9 per cent by weight of the total orey remains. One Sheenjek River eyr;e visited by George B. Schaller (reported in Cade, 1960) pro- three of which were ground squirrels. In my opinion these differences in ground squirrel useage are due to differences in their availability to Gyrfalcons at the times and places these collections were made. Cade's (1960) comment that "Gyrfalcons are much more specialized than peregrine falcons, Falco peregrinus, on a pop~lation-wide basis in-their food habits~ depending primarily upon one or tHo resident species of prey, !I is supported by the fact that ptarmigan and Arctic Ground Squirrels accounted for 1,046 (about 70 per cent) of my total li483 identified Gyrfalcon kills. By body weight these resident species (Willow Ptarmigan, Rock Ptarmigan, and J\.rctic Ground Squirrels) comprise about 81! per cent of J I 51 the total biomass (see Table 6). Cade (1960) found a comparable situation when he analyzed Murie's Alaska Range data and his own Arctic slope and Colville River data. Of the three data sets, Cade found ground squirrels oredominant in the Alaska Range data and ptarDigan predominant in the Arctic slope and the Colville River data. My own observa- tions indicate that Gyrfalcons in the Alaska Range do indeed take large numbers of ground squirrels. Ground squirrels, unavailable to Gyrfalcons from about 1 October to 1 May, contributed almost twice the bio- mass on a Height basis than Long-tailed Jaee;ers cUd in the summer on the Seward Peninsula. However, it should be made clear that Gyrfalcons do not normally consume the digestive tract of ground squirrels and this portion plus the heaviet> mammalian bones constitute a relatively large percentage of the weight of this species. Conversely, Gyrfalcons often appea~ to con~u~6 the digestive tracts of jaegers and this probably results in a more equal ratio of consumed biomass for these two important species. Two caotj.ve birds were also observed to follm'l this feeding nhabi t., Long-tailed Jaegers were the third most frequently occurring prey species in the Seward Peninsula Gyrfalcon prey rem~!ns (see Table 6 and Table· 7). This species accounted for 10~5 per cent by number an~ 7.4 per cent by vleight of the t-otal prey remains. On a 1vei~ht basis all jaegers 6bmbined (Long-tailed Jaegers, Parasitic Jaegers ':'.l!~.:le 8. 'rotal ~1~r:;.to~y 't!.rd !!p~cieo 'lC.entific<! !'ror-:. th~ 1968-1970 ()j'rfalcon p:-cy rc:nri1n5 anc': 11:;;':-ed in Vtt.:'"i("'U3 f~r:-o~t~nt ~;roui)~ ----~--------------~·~-~~ Chnr<Jh1rd!l ~n~~~~dn c~ncltJ~1n~ jcc~cr= ~nd rJucko) Un11~~t1f1cd n1~rato~y t.! rd ::pp. 159 79 12 58 30 ?c:" cc~t of totnl %ill_ 0.8 2.0 25.5 -------·~---·-· ---------- l'c:-cl!nt by ·(rc~~.ht of tc~~:l~~ 2.2 0.5 0.~ 17.6 fer ccr.t. c!' t'>t::t1 :::1.o;r~:tory ~ :-~.1,_!1'----- 20.6 7.3 F~r c~r:.~ tly w~~ir:ht of ~~~.:'ll ::.1._i;o.~":<':_::' ~:'..:-d :~-:.1 ~ 12.5 !;.6 2.2 ' 18.3 99.9 '~ '-'' [\) 53 [0._. parasi_ticus], and three unidentified Stercorarius spp.) accounted for about 7.5 per cent of the total biomass. Cnde (1960) recorded only three Long-tailed Jaegers in his Col- ville data and two Parasitic Jaegers from his Arctic slope data. White and Springer (1965) reported only one Long- tailed Jaeger from among 38 kills they identified from a Gyrfalcon eyrie near Hooper Bay, Alaska. It is possible that the Sevrard Peninsula, juttim~ out into the Bering S.::a i~ a more desirable jaeger habitat. It c .. r)pears that large lernuinr;-vole por,ula.t ic)11s influ~nce Gyrfalcon utilization of some nigratory avian species,. notably Long-tailed Jaegers, Short-eared Owls legged Hawks, since the availability of these species to Gyrfalcons depends strongly on the presence of microtines. Cacle (1960) states that nThe sit:;nificant fact for the gyrfalcons on the Colville was not their predation on microtines, but the presence of rodent-eating birds which had -be-en drawn into the region-by -the microt ines. :r Cade found that "on a weight basis more than 20 per cent of the 1959 sample of gyrfalcon food consists of rodent-eating birds." Of the total 1,483 Gyrfalcon ld1ls i_dentLfied from the Seward Peninsula, 171 or about 11.5 per cent were rodent- eatinp; birds (j <w_gers and Short-eClred Owls). On a wei13:ht basis this food source totaled about 8.3 per cent of the biomass (see Table 8). 'l'hes-e dat-a :1ppear to indicate th~lt this food source is important to Gyrfalcons whenever it ls availible to them. Cade (1960) found the.t ~he S'~_;:·,p·ei' foods of Gyr- falcons in Alaska characteristically contained two or three species that contributed "more t~12.n 70 pPrcent of the total sample. 11 From the literature, it appears that thj s car; be considered a general circumpolar characteristic of Gyr- falcons, and that it is not unusual for the percentage to become as high as 90 ner cent in certain regions 1f we con- sider ducl{S i.n general as one cater;ory of 1Jrey (Sen[!tson, 1971). My data agree with this ~eneralization and with Cade's statement. In order of occurrence (see Table 6), ptarmigan, Arctic Ground Squirrels, and Long-tailed Jaegers constituted about 81 per cent (1,201 kills) of the total 1,483 Gyrfalcon kills from the Seward Peninsula. By body weight, this small group of species, including three resi- dent species, accounted for nearly 92 per cent of the total biomass. The remaining prey species constitute a small propo~tlcin of the total prey remains and are generally com- parable-to the data of others (e.g., f'-1urie, 194 6; Hagen, 1952; Cad~, 1960; Bengtson, 1971). Cade (1960) states in reference to Gyrfalcons that, J!In Alaska the data on food -hab~ts of interior populations support the conclusion of Dementiev and Gortchakovskaya (1?45) that gyrfalcons, i11 distinction from peregrines, are primarily oriented toward predation on permanently resident arctic species of prey whereas peregrines prey on migratory species.11 IV!y data further support the fact that Gyrfalcons do indeed prey heavily on permanent resident specie~, particularly those Gyrfalcons nesting at some point removed from an actual coastline but that these resident species play an important part in the food habits of Gyrfalcons more closely asso- ciated with the coast. The occurrence of one Mink (Mustela vison) in 1968 must be considered unusual. This mammal is a rare species over the western portion of the Seward Peninsula (John Burns, pers. comm., 1968), and it is doubtful that Mink are often in a position in which Gyrfalcons may take them. The occurrence of one Short-tailed \{easel (Mus~ela erminea)-in 1970 is also unusual, thout;h this specie·s is _fairly common on the Seward Peninsula particularly in high lemming years, and is often observed hunting on the open. tundra where it is exposed to aerial attack. Dementiev and Gortchakovskaya (1945) identified the only other specimen of this species reported in the GyrfaJ con literature fro:n an eyri~ on Kharlov Island. The role of migratory birds in the food of Seward Peninsula Gyrfalcons is of considerable intrinsic interest since it is ~his component that lends the majority of di~er- sity to the spectrum of prey taken. The migratory species have been divided into various grouos for comparative pur- poses. As a whole, migratory species comprised about 26 per cent by number and approximately 10 oer cent by weizht of the total prey remains. These data indlcate that Gyrfalcons do take a significantly smaller r~orortion of migratory species than arctic peregrines tend to take (based in part on Cade's 1960 data). It is interesting, in addition, that this food source is of a large enouch size to serve as a relatively steady and probably significant source of pesti- cide and PCB (polychlorinated biphenyl compounds) contami- nation to Gyrfalcons if in fact these migratory species are contaminated themselves.1 It is interesting to compare Cade's (1960) Arctic slope and Colville River data with my total list of orey species (s~e Table 6). By combining his data with that of Murie (1946, unpublished), Cade (1960) summarized most or the available Alaskan information on the summer food haJits of Gyrfalcons. These combined Gyrfalcon data contain '' .. ~ohly twenty-one species of prey as compared to fifty- seven species that are Us ted for peregrines. . . '' ( Cade, .1960). Three of these 21 categories are comprised of uniden- tified ducks·, 'un-identified passerj_nes and vole species that _ 1 Prel:Lminary inves ticat :Lons by I:Jayr::an E. vialker, University -of Alaskay indicate that all specimens of Gyrfalcon prey specie,s -t~Ollected_ in 1970 analy2~d so far co:1tained detect- able levels of both chlorinated hydrocarbons and PBC. This study ·will be published separately. 57 may or may not have been different from those similar species listed. The arctic slope data for 1952-1957 com- prise a total 6f 142 kills collected from eight eyries, and. consist of only 12 categories includin~ unknown vole spe- cies and unidentified ducks. A similar situation exists with the Colville data--103 kills consisting of 11 cate- gories of prey species including that of unidentified passerines. Murie's Alaska Range data (reported in Cade, 1960) are even less diverse--429 kills consisting of seven categories of prey species including a known total of at least six species. My data for the Seward Peninsula, pre- sented in Table 6, list a total minimum of 32 avian and eight mammalian species. Cade (1960) stated that, orr have had to utilize pellet analysis to a great extent because of the infrequency with which other types of remains have been found at most of the aeries during the time of my visits. n In my expel"- ience it is unusual not to find numerous skeletal material with attached feathers or fur in or below eyries .. Since Cade was primarily studying eyrie.s situated on river cliffs,. river break-up and changes in l'later levels may explain his difficulty in finding prey remains. The striking differences between cade's (1960) data and mine may reflect: ( 1) a difference in d~versity and availability of prey·species betwee~~he regions (i.e., at the time of study) and (2) a difference in sample sizes. It is possible to extract a 58 number of combinations of eyrie collections (see Tables J2a,b to 27a,b) from my data that will construct data sets of similar numbers of eyries and numbers of kills and thus give results comparable to the low diversity of species found by Cade. Bengtson's (1971) Icelandic data are more compar- able to mine. He lists 24 identified avian soecies (ll of which are duck species) and three a~ditional prey catecories (Anas spn., untdentified dudes, ~mi :::;;--1:111 ducklings) from a region lacking small mammals. The data that best compare with mine are those of White and Sprin~er (1965). Their data are from only one Gyrfalcon eyrie near Hooper Bay, Alaska, and consist of 38 kills collected in a brief 15-day segment of time during the Gyrfalcons' nesting period. White and Springer (1965) list 18 categories involving 2 minimum of 14 spec~es. Eleven (28.9 per cent) of the total kills were shorebirds, nine (23.8 per .cent) were ntarmisan, six (15.7 per cent) were Larids, six (15.7 per cent) were passerines, and four (10.5 per cent) were Anatids. While Cade's (1960) data n ••• pertain mostly to eyries in the more inland tundra and mountainous ar·eas. n (as stated by White_and Sprihger, 1965), my data and that of White and Springer (1965) are from coastal areas or peninsular areas strongly influenced by the coast. Yearly va-riations in prey utilization Gyrfalcon prey re~ains from the Seward Peninsula ':rere segregated by year and the yearly compositions of the 1968-1970 prey remain~ are listed in ~able 9. In attemot~ ing to compare the 1968, 1969, and 1970 collections, con- sidcration must be given to: (1) the consistency of the search effort; (2) the consistency of the tir.dng of the collection visits; (3) the nunbe:t' of eyries visited dur>ir.g each year; and ( i!) the chanp:es in orey soecies numbers and their availability to Gyrfalcons. 1. Search effort: SeClTch effort '.-Jas generally consistent throu,c:;hcut· the study. f~o1;1ever,_ in 1969, ~3om~ nests were not climbed to ti1ough they were visually observed from close range. Such nests were not climbed to because they appeared essentially devoid of prey remains. In l9[q mor>e effort via.s DX!Wnded in search belm·v-ncstinc; cliffs and at nearby perching: places. For unknm,m reasons, pre:,r remains were scarcer both in the nests and below them in 1969. Dur- ing that year only· 2911 kills i'Jere collected--a much lower figure compared to those of 1968 Rnd 19'70 • ·· · " --2 .. --'l'iminp; of visits: During 19GB, visits to Gyrfalcon eyries were spread fa5rly evenly over the period 31 f.[ay-tcr-·23 July. In 1969, the uajor'ity of the visits occurred between 15 June anrl 20 July. It is oossihle that some remains, accumulated prior to 15 June, were scattered and thus unrecoverable. In 1970, v~sits corresponded 60 closely with those of 1968, occurring evenly over the period 28 May to 27 July. 3. Number of eyri~_s visited: In 1968, ten Gyrfalcon eyries were visited to collect prey remains and pellets~ In 1969, 14 eyries were visited, representing an increase of 30 per cent in the eyrie sample size over that of 1968. In 1970, 16 eyries were visited, representing an increase of 60 per cent in the eyrie samole size over thnt of 1968 and 33 per cent over 1969. li. Prey __ abundal]_ce and avai lability: Prey ~pecies' numbers changed markedly in some instances from T9GB to-j_970 .. ~The scope of this study did not permit detailed censu~ of pPey species, but general day to day observations were made and are discussed later. It is evident from Table 9 that ptarmigan, ground squirrels, and Long-tailed Jaegers stand out both by nuJ'1her and weight as the Ilro~inent prey species durin~ each summer (also see Table 7). It is significant that, by weight, these three prey categories together comprised 92.4 per cent, 85~~ ~er c~nt, and 93.5 pe~ cent, respectively, of the 1968, 1969, and 1970 prey remains. It is also interesting to note that the -two avian categories· (ptarmigan and Long-tailed Jaeger~~ tbgether accounted for a large proportiori (82.3, -_.o--- 62.0, and 83.7 per cent, respectively) by weight of the yearly total kills,. and a very large proportion by Vleight (92.6, 80.9, and 93.5 per ce~t, respectively) of the avian species taken. By comparison, ground squirrels constituted a relati~ely ~mall, though obviously important, segment of the yearly t6tal kill by weieht (10.1, 23.3, and 9.8 per cent,-rer;pectively), but comprise .q very large portion (93 .I~, 99.5, and 95.7 per cent, respectively) by wejght of the yearly Mammal kill. Further, it is note-worthy that the tvw major resident specles' categories (ptarmir:;an and ground squirrels} together ·by ~'Ieight co-nstlb.i'ted a large and relativeli stable portion (86.1, 81.5, and 84.0 per cent, respectivcily) of the yearly total kill. The remainder of tile yearly 'prey remains consisted primarily of small numbers of a variety ~f migratory bird species and micro- tines, and constituted a relatively small portion by weight (7.6, 14.7, and 6.5 per cent, respectively) each summer of the yearly total kill. During each summer of the study, Gyrfalcons on the :3e\·Jard Peninsula took a significant pro- por>t ion o-f·-migra tory bird species. These l'la:ta are presented in 'l'able 10. -It should be emphasized, in vie~.., of potential · pestici-de and -PCB contamination, that a relatively large proportion of these migrat.u1·y blrds are top level consumers in their own right. The yearly data generally follow.the overall data from the £eward Peninsula listed in Table 6 and pTeviously discusse~ in this paper. In seneral, the same conclusions as those obtained from the' data ln rrable 6 could be drawn from any nne year's data. The most interesting aspects of 62 ~·r,ble 9. I ~63-1 ~i ~ C:;rfslcoiJ rood rf!o:--e ~ n~ CC1ipUt('d 0:1 • yculy bo.~', .. ------------~---------------~-------~· .... -........ _... .. _..-.....--_ ....... ~------- --·--J~~~------------____ l.c_~---~-------___ _:1_ .. !1. ('!._ ___ ------------- ~·ot> l Per C't" r\''~ r:""r c l nt 'r~tlll Per ce:".t ;\·:· .-r: :·t 7c!:.ll :e~ CCI'.~ !(':' .'let. r.llls £L~.f.l f<. ~ '!" -~-... '! :--.'ll !!._11 ~ ~ t~ '"-~ r:_t_:-.::~:.·:_·.21. f_!];~ f'_::._~t~l_ ~~.:-.:~: ... :~:..:~.:~:_ ?tt~r=.1 r::~n !jJt·~!(':S 29b 6~. 5 74.8 1~3 ~3. (, C?.7 bJ9 ~9-9 (.::. 7 !.onr.--t.1llf'.! ;; .7!t ~-.er ~2 9.2 10.7 16 5.b ; .~ 97 13 -~ 4.~.2 J.r:;crie.:~.-, G~-l::c~ .~ l "': ·.·~:· ~ 0.9 l.O 11 3. 7 ~.E 15 ~.n ?~3 Un1CI!'ht~!"1t".j r·•-~~-~:--lne 5 1.1 1.3 3 2.7 3.~ 16 2.~ 2 .. 5 l'nl~c~.t! !'! cd bi: -:~ 5 l.l 1.3 ;> 0.7 o.~ 1(. 2.9 i. 5 v~~'(ac-nt1~!ed s~,)P:t ~r'\.!.:S 3 0.7 o.e 5 1.7 ;<,.2 6 o.o 0.9 !'-l~<'%-)~f""e:1 l\ 1 t t 1w::~.c 3 0. 7 0.8 11 J. :· l.7 Cc;;":.ou ~r.! re 7 1.5 l.S 6 o.e o.~ St'J."Jrt-~<·:-e-1 c~l 6 1.5 2.0 2 0.7 0.9 2 0.3 ~. 3 \.";1!::~·r-cl ~ 0.9 1.0 2 0.7 0~9 5 0./ O.R F.o~:.1a l 0.2 0.3 8 2.7 3.S 2 (1.~ 0.3 0:1!ci,:ont!ntd ... &.tcr!'c\11 l 0.2 0.3 3 1.0 1.3 ~ 1).5 o.G F! .~t:t 11 3 0.7 o.e 3 l.O 1.3 l C·.l 0.2 '!'.; rt.t":.: h:l~ln 7 2.-3.1. '£'.:,r-t.?.~ >:.j. Goo ... 1 t 0.1 0.8 3 (•.~ 0.5 O}.r.:-q•..-:.1.." ~ 1.- l.3 1 ').l 0.2 L:,p~ ;..";.i J.(l.~f: ~-i ..l':" l 0,2 0.3 2 0.7 C.9 2 t·.:. 0.3 r i ~~·cr. G ::J.:.!"··. ::. il l.~ !.f. u~ .... :!~c-&:t~ rJ .. r;J s?·;:e-=-~ ·-2 o .. ~ o.r; l 0.1 0.2 ~:-.1:>::r~t~!2.:.-~ <! 1::-!r.:-3 0.7 0.6 Cor r.y-C ~;co:-~:"-.~ ·::~~'-' l~: 1.0 1.3 1.11-::<!t 'i\:;:. ~ ~:: t:r '= 0.2 (1.3 1 C.l 0.2 ~-C'.r! r-_:>]1 !. ~;('..:'! "·' 2 0.7 0.9 ":.;·::.' ~ . ._, ..• r ~ ,.;: 1 0.3 (, .. ~ (· ·'-C.?. ~nc-u r. ·; .~. f, ~ ~' .. <: f"J..) ? . ~ };) ~~;k fc'.CO 1 (..). VoL c~·t r:l:··• 1 n ._,. '! ·:, \l (!.~ 0.3 St:!:l!f-!··c-.?.t·.::! f~r--~~~ l 0.3 0~ ,.l it.'er.d~r ~:-_:.-:'.t ~:-.lE:t" o.:-o;3 .. - -~· . . . C.l c.~ .l•t";:! :'>~1 11:: .;..'l,.'•'lt:t· ~ tia~ J! 1::1 r! :..·-: ::·.:·:1r:.f~~;-., 1 c.; 0.3 Patasllic: . :. -~ ~· .·.•.• ;-1 O.l. 0.~ P.:: rr~:·:·( ~ t:.;:; , .. , 1 c-., (\.~ Unhh:.u.r:(d Ol~ll~ ts 0.3 o.~ ~ ..... -~·o;.·~·-· 1 (\.l. o.,' \'z..:-JcJ ·::,;-·.t:'ih 1 (J.l 0.2 a·.j.t.(.;· 1': Co 1 ~ 1 0.! ':.1.2' Fot sr·~:-;·.:.k __ l_ _;:.~_, _ -..c.~? 'i"(fJ"}.!,.. ~It:>~ 393 SG.~ lCO.O ~2e 77 .(. 10/).': 639 e-: .2 !(('. ~ l••ct! • Cl't';;··•l g~1 .1 J .... ~el ~l >"·.( 6~.) 0 ~] ·' S5.5 (; L1 ,-'I .· ... ·. Collel'c'.i I ,_··:::r'b 11 :t.l! !'( .s 3 1.0 ~.t lO 1.( 1.) ,, Ur.l~.!nl! ~ ' . ..:j ! I';~.:'!!;;~;~. ~ 1.1 t.~ r.ro~:l Lt:::.:-.1 ~~.:; 5 l.l 7.9 1 ~·.1 C.2 VnJ-:\('nl;! r: :--1 V\•1(·~ 1 ·o.2 1.6 • C.5 t.) f.'!d-t.a~i:cJ -IC:l-l' ~ Ci.5 Z:,J Cr.idl"n~! r! !'•.! r.~!-: ~~~~! f\C~ 2 0.\ 3.2 ~undr:i \'ole 1 (1.1 0.? !'f·,C"rt.-t . .,' ~ •.:.1 \.'~t.St·l 1 t.l c .. ~ l:lnl: ..1 _.£.:1. _!;(. T!"J:'}.!, }:J.~"..vJ,:..'t I;] n.e }(:0.0 CG 22.5 }(•,.0 9J 1?.1 \(;C.~ Cnldcr.t!rlcG k1ll• _! ~:!,- ronL ~:1!..f.'; ~~6 ~,-733 To.Olc'-lO. '. 1,) ., J nce:cr:J !:;ho~~birr!:; .s:~ort-~arcd ?a;jz~.r!nc:J ::":·"A~ ! r·J ~ (~':::c: 1 ,:vj1 rifj Du·~kr. Migratory op~cieo 1 1clcnt1f1c1· froM the listed 1n yarioun 1~po~tant ~~ciupo I 19(;3· ?cr cent of of total r.;i;:_:r:1tcry Nu~tH'"r k111(1~55~ ::3-:l:.'Ci~::; ij~ 9.7 ~1L9 .,., <.) s.o 23.5 Owl:~ a 1.8 8.2 7 1.5 7.1 j~Cf:CI'3) G 1.3 6.1 5 l.l 5.1 Un! •!r:-nt 1 ilcrl r:,1;:-;:-::to~y ~p~c~.e~ -~ .ld hl 'i'07ili-9fl 21.3 100 t:ur.:t-cr- 16 19 2 21.J 12 , .... ... u 2 ~~ ')) ),~~ l?70 Fer CC:1~ or ?~'r" CC:lt or of tot~l ~lr:r-atory or tot.l\1 :::;.sr~t. .. y:·y !{~ ll (~2~ • 81'f~·~~C!3 Nt:~be::--~ill \7;<) ~~~ 5.~ 18.3 99 13.5 49.5 6.5 22 ,ll 37 5.0 1(). 5 0.1 2.~ 2 0.3 1.0 8.2 2S.2 .,.,. 3.7 13.5 ,, 1;,1 14.1 12 1.6 6.0 3.4 11.8 7 1.0 3.;3 0,11 2 .}t 15 2.:! _8_,_g_ 28.9 100 2CO 2ll. 3 lCO 1 ~h~;~·n~c ~v1~~ ~p~c1c~ ~nd it in ~~~um~G th.!t rn1grat~~~y ~oec!es e~ a w~ole are the ~ajor ~ou~c~ ~r p~~t1e1ee ~~G ?~~ con~~m!~~t~n~ tr· Grrfnlcon~o ==::::::;:====-- the yearly comparisons made in Tabl~s 7, 8, 9, and 10 are those gf th~ changes in the frequency of occurrence of some species in the Gyrfalcon diet over the three-year period~ To clarify the significance of these chan~es a brief introduction to some of the changes in prey species levels ·on the Sev:ard· Peninsula is needed. It is typical of the far north that both migra- tory snecies and resident species may vary greatly in numbers from year to year. As a consequence, the avail- ability of Gyrfalcon prey species (particularly microtjnes, rodent-e~ting~birds, and ptarmigan) may change from·year to year. Gyrfalcons appear to be well adaoted to cope with this situation, at least in the Alaskan Arctic and sub- arctic. Cade (1960) notes that on the Colville River in 1959, ptarmigan became scarce by mid-June and l!that the final gbod fl~dgirg success of these gyrfalcons was made possible thr-oug-h the fortunate co1ncldence of a sui table alternate population of large-sized prey ani~als'' (Short- eared O~ls, Marsh Hawks, Long-tailed Jaegers, and Parasitic Jaegers attracted to the. resion by ;r.q. moderately high den-· si ty ·of microtines :r). It appears that a similar situation occurred on the Seward Peninsula in 1968 and 1970 when microtin_e ·populations attracted numbers of rodent-eating birds ta th~ area, particularly the Lon~-tailed Jaeger which played an important part_in the Gyrfalcons' diet these tv10 _yc=ars. 65 In 1968, considerable amounts of snow remained unt.i.l late June on the Seward Peninsula. f-1any migratory bird species were concentrated along the road system and in other scattered areas free of snow until early June. Lemmings, particularly Collared Lemmings, were olentiful. This species had even appeared in numbers on the snow sur- face in mid-winter (John Burns, pers. comm., 1971). Hed- backed Voles were also quite common in grassy 2reas. Con- sequently, L~ng-tailed Jaegers and Short-eared Owls were found in large numbers over the tundra. Both Rock Ptarmigan .. and Will9w PtArmitari~ere-found'in large numbers throughout the peninsula. Rock, Ptarmi~an Here often located at lm·1 elevations in habitat more characteristic of Willow Ptarmi- can, and in many instances both species were found nesting in the same locations~ __ Ground squ~rrels apoeared plent1~ul and were observed over a wide variety of habitat types. -lt!aterfm,rl vrcre gcncraJ.ly abunc.1ant ." -Pfntails ~Anas ~lcut__r3_), in pa:rt:!cular, frequently nested in valley bottoms and marshei, and occasionally under willows at considerable distan~e~ fro~ streams. In 1969, light winte~ snowtaiis and high May temperatures resulted in a rapid snow-melt which was completed approximately 30 days ahead of the 1368 melt. Relurnins pD_;,~.erines anCl. shorebirds Here observed scatte.·red~-across the tundra inste2.d of concentrated alone the cleared road system. During the 1969 summer, microtines -. -.··-.-· -. appeared to be virtually non-existent. Only nine Short- eaPed Owls were observed during the entire summer. Long- tailed Jaegers were thinly scattered across the peninsula and the majority of those present were non-breeders ranging about in small groups and feeding on insects and berries. Ptarmigan numbers were reduced from the 1968 level and species segregation was more noticeable. Ground squirrels were present in large numbers co~parable to the 1968 levels but also appeared to be more restricted to oarticular locales. Waterfowl were less coMmon than they had been ·196·8. : }fo du-eks vJerea.iscove'red ne"sting outside-of large ~ "'"' .. LJl marshes or river syatems. In 1970, generally light snow- falls and warm May temperatures were again the cause of a rapid snow-melt. Winter conditions (1969-1970) were such that a major portion of the peninsula's road system was open to the public (the 71 mile Nome-Teller road) almost all winte~. A~ in 1969, snow-melt was completed approximately 30 day~~6ea~ of the 1968 melt. Returning migratory bird -species, particularly passerines and ~horebirds agajn did not concent~ate along the road system but were scattered across the:relatively snow-free tundra. During the 1970 summer considerable microtine activity 1;1as observed. Bodies of Tundra.Yoles were commonly found in active Rough-legged Hawk .nests, but Red-backed Voles 2.nd lemminr;s v1ere rare1y observed_-~· Long-tailed Jaegers were common breeders aJ.tho'1P:;11 they did n¢t_appear to be as numerous as they did in 1968. 67 Short...,earecl Owls Here also comr:10n, but vrere not as numerous as in 1968. Ptarmigan· mimbers apneared to be slir;htly hisher than in 1969. However, almost all ptarmigan observed were Willow Ptarmi~an; very few Rock Ptarmigan were evident and ntarmle;an species segref~at lon ap;ain occurred. Ground squirrels were presen~ in large numbers but, as in 1969, appeared more restricted to particular locales. Waterfowl numbers did not appear to change from the 1969 level. 'l1 ahle 11 compares by year the occurrence of nine major ~foups of Gyrfalcon prey snecies in the Seward Penin- .. esula Gyri'a.±con prey remafns. rrhese prey r;roups include tl1e three orimary categories of Gyrfalcon prey remains, ptarmi- gan, ground squirrels, 2nd jaegers and six nd~itional groupings that include the total numbers of shorebirds, passerines, microtines, seabird_s-· (Alcids and Larids), cFicks, and Short-eared Owls. The yearly changes in the occurrence of ·some--Gyrfalcon l"lrey species on the Set·rard Peninsula are- shown in Figure 2. The following discussion will certain primar-ily t~o these data. ~he occurrence of ptarmi~an remains fell sharply in 1969, and increased to near the 1968 level in 1970 (see 'l'able 11· and Figure 2). 1968 vras a :rhighn ptarmigan year, and both ·Rock Ptarmigan and Hi1low Ptarmigan were common along tb~ road system (three to five pairs per mile) and vrere const;antly flushed while ·hiking ·across· ·c-ount-ry. In 1999, _ fe:n Hock Ptarmigan were observed ar-ter mid-June. 68 1Tab]e 11: Corrlp3.r1sons of thr;-oc:cu:·rence r:i.' J'I~Jl.~'" 'Drey groups in Se1·rard Pc!1:l.nsula G2n•L'.~C()n nrey l~e·nJ·, .. l•nc· C0~1fJU·+-·,.,(, u-" ..., .,,.., .. lrly h'J"'~'-.. C-U /, l.1 \__.. .J. ;. .L Ll .) ._.. ( .... Cl•J -L ,_) ____ ! 9 f)=~---__ _] 969 ____ -·-... 1 ~]_():_ _____ Per c!ent Per cent Per cent Groun -----··- 1l'Ot<?-J:_(!; 56)_ T~t_~~~~L Total ( ·q :() I .•. ' l>t ... r aJ:rnJ_r;a.n 64. ~5 lid. G 59.9 Ground squirrels 9. l.j 21.11 8.7 Jaeger:.; 9.G 5. l.j 13.5 Shorebirds 5.0 6.5 5.0 Pat>scrines , r: -'-•.J 8.2 3.8 - T!iicrotincs 4.2 1.0 3.8 Seabirds 1·3 lj • 1 1.6 Short -e;,~red ONl s 1.8 0. '1 0.3 65 . - . 60 55 ~ 50 GJ 1-1 >.. 11' 1-1 f.\. ~ •.-1 '/15 . CJ ~ ~ •.l) J.f "".:"" ~I B :>5 ~I 0 C) to.? o• u t:: GJ r.J· ~! CJ p. 20 l I 1 15 10 5 ------·---~---:..._ _________________________ ---~·~---····-.. ·····------- ptarmigan jacgers gioun~ squirrels ..shorebirds. n;i•::ro tines 1968 1969 1970 Y e a r Figure 2. Pcrce~t~ge occurrenc~ of selected groups of prey spc.:i.cs .i~~ ,;u!~n,er Gyrfalcon pl"t!.y rcinair.s fro:n t:t·H~ Se\o.'ard T~t:!!il:!:.ul~·i, .-\las.J-~a, 19h~1-1970. Prey grcups that incraas2d during 1969 are represented by dashed li!1~~s .. 70 Willow Ptarmigan observations averaged about one pair per ·thr~e to fi~~ ~il~s of road in some of the optimal Willow Ptarmigan habitat. In 1970, Rock Ptarmigan, never observed to be numerous that summer, were not seen after about the . first week of June with the exception of three specimens collected in August. Willow Ptarmigan were commonly observed all summer but remained below the observed 1968 level. Family groups and small flocks of this species became common in mid-July and _August. Local hunters comr.1ented that otar- mig~n se~med ·scarce in 1969, hut that in 1970 hunting was . better and-t11at-th.ere were again !r lots of birds. ;r These observations suggest that the entire nt~rmigan population declined in 1969 but rose again in 1970 due primarily to an increase in Willow Ptarmigan. However, the 1970 level. did not attain that of the. 1968 high. Figure 3 s11ows that the f~equency of ptarmigan in the Gyrfalcon prey remaJns agrees vli th my subjective observations of the ptar- mi"gan population. The occurrence of jaegers in the Gyrfalcon prey remains fell in 1969, but increased in 1970 to a level above_that found in 1968 (see Table 11 and Figure 2). Nine- teen sixty-eight was a high jaege~ year: All three species t._rere common breeders' especially in the southern half of the peninsula. Long-tailed Jaegers were by far the most abUrtdantf Pomarine Jaegers were the least abundant. Long- tailed Jaeg.ers Nere commonly observed catching and eating 65 60 55 . 50 45 . 0 ""L-- 'l. A------~ o-----<> Ptarruican p0pulaiion esti~ate Oc~urrencc of ptarmigan in prey remainG I _j L ______ _L ____ ~_j _________ j _________ .......J 1963 1969 1970 Y C! a r Figure 3. Th~ percentage occurrrnce of pt&rmisan in ~:'!lc 196( -1970 Gyrfalc>on pr.~y r0mains co;;!p;;red \ilth an cbscn·ed st<Ljcctive est5~:;tc o£ ptc;rini_;:;an · poptllction Lrend:.; from the Scuard Peninsula, Alaska. 100 75 50 O) r:..;:; ('j C\ rl r: 0 ~ r.~ '"d (l ~ ~ !I) ~ ~ -':l ••• ,, <:.> r, lJ t1l c C) t;-o~ !I) r; --~ t' "'> .;:t h; ., ..... ('j r--i t~ :.:! C:J c ... C...l 0..-l A. () :> >. •rl '~ .,LJ ~ C.' H Ci .1.J 25 :r.,• :11 ::; }; Vl;,; I) ·-1 / .. 72 Collared Lem~ings, B~ciwn Lemmings, and voles. Counts as hir;h as-100 birds were common on one 35-r:lile stretch of the Teller Road. In 1969, only one observation of a small group of Pomarine Jae[:::c:r·s \'!2S r'lade, anrl that occuT'Y'r:.: ;;'X!'ing spring migration. A few Parasitic Jaegers were observed along the ~outhern coast and a few pairs of this species apparently bred near Safety La~oon. Long-tailed Jaegers were thinly scattered across the peninsula. A few nestinG pairs of this species were found but the majority of those observed .did not appea~ to breed and were in small groups of 10 to ~0 that wer~ Nandering aeross the tundra eating insects and berries. In 19'10, Po::1arine Jaee;ers, though present,_were scarce. Parasitic Jaegers were again present, though not in the numbers observed in 1968. Long-tailed Jaegers were agai~ common although somewhat below their 1968 level. Long-tailed Jaegers were observed catching and eatir1g vol~-$ ( almos_t _certainly Tundra Voles). Observations th~refore suggest that the jaeger populations, and in particular Long-tailed Jaegers, declined in 1~69 and ro~~ again ~n 1970 probably due to an increase in Tundra Vole numbers, -to a level approachine; but belm'l the 1968 high. In Figure-ll the frequency plot of 5 aeger 2 . remains also -. ·-. · 2 The t.Qt_,.'ll-j aer?;er remains are almost ent.trely Long-tailed -Jaegers. It is interesting to note that for unknown rea- sons·this species took the brunt of Gyrfalcon predatJon, while th~ other two jaeger species were almost entirely untouchE?\1· 0 eo ('j 30 ~ 15 Gl u ~~ Q) p.. l. JO L '~ I -------------------------- -4t \ \ \ \ \ \ \ Ct-----1;. Jaeger population estimate c-----o Occ.urrence of jaeeers .in prey rern~i.ns. \ .¢. \ , \ , \ I \ / \ / ' I \ I \ / \ / -~ ·/ . \ I \ I \ // / 0'--.. \VI -~ 'd ~~ j I o L_. ____ _: ___ l ______ j __ _ __ _j _____________ _j 19GB 1969 1970 Y e a r Figure 4. The perccuLage occurrence of Jaegers in the 1968 -1970 Gyrfalcon prey .~s~ains co~parcd with an observed subjective estirn3tC of j&eger -·papulc:llorl tre:1ds !ro~u the Se~ .. "ard I·c-~ninsula, Al3.sk;, lCO . (.C -.D C'. .; t:! H 0 <! Col "\1 :--. !lJ ~~ l!} r.: .~~ .D .u 75 ~ ~ 0 c,:) t:~ u ul -~ Q) r.: > 0 <!) -rl r-1 <J 74 appears to follow the trend of my subjective observation --of the jaeger population. The percentage of microtine kills in the Gyr- falcon prey remains fell in 1969 and rose again in 1970 to a level comparable to that of 1968 (see Table 11 and Fig11res 2 and 5). The 1968 season was a year of abundance for microtine species in general, particularly Collared Lemmings and Red-backed Voles; Brown Lemmings and Tundra Voles were also cor.1mon. _In marry areas Collared Lemminss were found on the tundra in quantities that allowed a collector to cap- ture numbers of them by hand. Prior to the 1968 summer a build-up in microtine numbers had apparently occurred. Microtine trapping data from John Burns (oers. comm., 1971) indicate that. in the vicinity of Nome trapning success increased T~om about 11 per cent in July 1967 (1~3 trap- nights) _to about e. 0 per cent in October ( iW9. trap-nights) . Duri~g mid~October one area produced 41 animals in 144 trap- nigh~s--a success of about 28 per cent. Following this general build-up, primarily of Collared Lemmings and Red- backed V6les, Collared Lemmings were observed moving about on top of the snow durin~ the winter 1967-1968--an appar- en_tly un_usual o"ccurrence (John Burns, pers. comm., 1971). puring the summer of 1968 Burns reported a trapping suc6ess -.. of about 40 oer cent in early summer with an increase to . . . high as··Gs per cent by late Aucus t. These trapping data indicate that of the four species discussed, Collared -~.,·----··-· ... _______ -------------~--·--·--·-"'1 30 ~ ~ \ \ . ··•!' J.I}C"I 4-·----~.:~ !--r.~~c"('::_;jr~~ pnp·.i.l~~-ti0n (>,, ll ... ,.t..., C!------·------o <?cc.urrt:1r:~ cf :Pit,::0tint.~S ~ .. ... .: c::: ...-:: hC' C.-i 1!1 25 {) r. Q) 1-< r ~~ :: 20 -~ N1 ·~ .., r:: ~ (..' I< () 1'4 15 -· 5 \ ' 10 prey remaJns ~l) \ \ \ \ \ \ \ I \ I ,. I \ /L l \ ! 1 \ /; I \ I ~ \ I I \/ __..-o j -c..___-....__________ '>r. _________ __..___...... I " -I -,.,)____ I v l---..----.. --------J-... ~------·-·--··-·--...... 1 ..... --~--·--·-.. --_ _L_ ....... _ .... ___ , ____ ~I 1%8 l%9 1970 'l a r Figure 5-TLil~ pereen t.::tVJE ,~o~:c~· rr2nc!!. (. ~­ microrin(S i~ tl1e 196S--l97C-Gyrfalcon prey remains co;~rpared vit~; ?;1 0b;;erved ftubjcct:ive l?stin:Htc cf r;}_c·t·otl.:·:e ~-"-""ru1...:•.-­ tion trend~-: frc·m th;..: Se!,.JGr1 Fenii!sula 1 Alaska. (j) f-t'.1J . .:t ::i!lci.Lys J :; :i: <0 i Ciltf:'J microtinc~ occ~!!. ~Lc~i~ f!~2 t1mr.~; ;J.::-.. :.:~tt~'r:. ;~~ ;:~·~·-prL~y 1·t:··~ P.l[~lft~~ p~.O ~ t etJ CF t h5. ~: gr~. t·h l .. .. ,:: ~. ;jJ '"· (/! ;:l c;: ~' ,.C; 75 c -~ -·-l .u r.1 ). ... ~~ v <;; ~ Q -: ~-1 G.· ._J :, C) (:} 50~~ d :..~. ~.t ~~ .~: •• J d.: > :.,-.., ·~·i ~-· ~J ~; 0 ~ '.J "-: T) ••i 25 .!""I·~ ~'J ~ Lemmi-ngs were the dominant species c:md the rrundra Vole 'il3.S the least abundant. In 1969, all microtine species appeared sc~~c~; however, a few Tundra Voles were observed crossing . . roads and small numbers of this species were present in some areas, at least on the southern half of the nen:Lnsu1a. Three hundred fifty trap-nights in three good 1968 ~icro- tine habitats near the active Gyrf:.l1con eyrie::> nrodnced only one Red-backed Vole. Burns (pers. comm., 1971) re- ported a catch -o·f only· 25 anim::J_l-s ·of the· ass6rted species in 1 3 070 trap-nights in July 1969. During 1970, a few Collared LeE1mings and Red-backed Voles were observed lng roads ann were r·arely encountered on the tunclro.. Tund:ea Voles Her~. freauently o_b~erved. _ .!~ouc;h-legro:ed j ae;~;ers ,. and .Short-e;:~u'"'ed ·O~·Jls \·Jere observed cat chine and e2.ting; them~ they \·Jere commonly f"ound. in Rour,h-legsec~ 1 l'1\·ri< nests~ The percentage of nicrotine kills found in the prey remains generally followed the estimate of the th~e~-y~ar trend in the microtine populations (see Figure 5). -The general high pooulation of microtines in 1968 attracted a large number of rodent-eating birds~ including, besides the jaegers, a few Marsh Hawks and·many Short-eared Owls. In 1968 up to 12 Short-eared Owls were commonly db- served along many one-mile sections of road. By P!Tay 1969, the micro-tines had undergone a c;eneral population "crash!! and were-scarce. This condition drastically reduced the numbers of·rodent-entin~ birds on the Seward Peninsula th:::tt 77 season. Only nine Short~eared Owls were observed during the entir~-1969 summer and eight of these birds were in two family groups observed hunting along the Kuzitrin River during mid-August. In 1970, an increase in the numbers of the mi~rotine~, prima~ily the Tundra Vole, again attracted ·. large_ .numbers of rodent-eating birds to-the Sev:a~d Penin- sul~ (again including a few Marsh Hawks), alth~ugh below the numbe~s observed in 1968. Short-eared Owls were again common. ~aunts alo0g a 3Q-mile ~tretch of road in tha- -'-Kou·garot-:-:Kuzitrln.:.1>iTg-rim--R.iver systems ranged from 25-30 individuals throughout the summer--an average of about one ·Sho~t~~ared Owl per mile. The percentage of Short-eared /"--""'\ _ ONJ.-s in the prey re1rtains fell in 1969 and continued to fall _\·--:-_! __ in_~_l9TO ir1 spite gf an -observed-increase ·in number in- -----·- 197:0-(s·ee T9,bTe.c.lO and 'Figure 2}. 'i1he significance of this case, inconsistent Ni th the general picture~ ·cannot be ··- asses·sed. The foregoing outlines a situation in which 1969 -·--population deciines in three groups of prey species, ptar- migah, rodent-eating birds; and microtines, were correlate_d -·- with a l.ov..rer percentage of these _species in coll~cted Gyr,.... falconJir.~Y: ___ re_m_a;i._ns _. The remaining Gyr-falcon prey groups - ducks_ (s~e~-~'J':iible 11 and Figure 2) rose in proportionate-·.------.;: .. :·. ----~ . ' --. -. representa.t'ion in Gyrfalcon food iri 1969. ·:"' Tfie crccurrence of ground squirrels in Gyrfalcon --.,... __ •...!,.. •• ~ .. ..-;;. •• -:·.-·-1 ·------- . ___ ._,. ~ ~ . -------.-·.:~-;.: .. :-~::.-: .. .. food remains, while rising in 1969, declined to slightly be~ow~he 1968 level in 1970 (see Table 11 and Figure 2). Gro~nd ~quirrels were abundant oyer most of the Se~a~d Peninsula du~5ng all three years and were commonly observed' the entire lerigth of the road system and on every over:....land hike.~ Actual numbers ot ground squirrels did not really appear-changed fro~ those found in 1968 and certainly any change that may have occurred was relatively small and not easily detected. Figure 6 shows the sharp increase in the · .. · .. ·----__ ; ___ --~-----·· ·.·:.- per qqnt of ground squirrels in the Gyrfalcon prey remains in 1969_, vlhi1e the ~stimated ground squirrel population indicates little change over the three-year period. The remaining prey specles, consisting_ -o:f. passer·:tne::r;-shorebirds, ducks, and se-abirds, a!'l show the-same tlpward tendency ·in their occurrence in the 1969 Gyrfalcon prey remains. These groups did not appear to exhibit any-note-worthy changes in_population numbers over . -___ ---.-.. --·· -·-· the ye_-~:rs .. Tbese data indicate that the 1969 declines in numbers a~d availability of some Gyrfalcon prey species, primariX_y: .. ptarmigan and Long-ta.iled Jaegers, on the Sevmrd Peninsu1-a-resulted in t·hes-e species being taken .to-a les·ser . ,__; -.. · .. deg.ree hy Gyrfalcons, and that other species, specifically .the, Arct_:L¢=·c~Ground -Squirre-l-,--were taken ·to a-hi.gher-·degree as a meq.g_s_ of compensation. Because ground squirrels are tc3 an .impot~aQ~ Gyrfalcon prey species on the Se\1ard Peninsula, '79 both by number and by weight, and were relatively plentiful, these data suggest that Gyrfalcons relied heavily on this species in 1969 to compensate for the lack of other prey Bpecies. $ince ground squirrel numbers did not appear to ~change to any important extent over the three-year period, I believe that the increase in the occurrence of thi~ spe~ cies in the Gyrfalcon prey remains indicates a higher degree of utilization, not because ground squirrels were more abund~nt but because other prey species were less abundant. VJh-ether 'ground 'squlr-;rels-~rere -·hurited-"sele-ctively,· or wh~ther - they were simply killed more frequently because they were -· . ~-. - there and available is unknown. In either case, how·ever, it app-ear3 that the increase in utilization of the Arctic Ground· Squj_t>-rel vva-s an--important form· of compensation in 1969:, wh-en ptarmigan,· Jaegers·, and microtines declined. It appears that, like ground squirrels, the passerines, shorebirds, ducks-, ·all:d seabirds played-a rol--e-in-the 1969 Gyrfalcon·· .. · , .. diet as_ c __ ?mpensation for the dec1ine of the other prey popu- lation, though to a much Jesser extent. Though important prey compensation apparently took place in 1969, there was an aprr~9Ximate 25 per cent decline of the Gyrfalcon popula- tion Nit l-it.!} the sub-:-s tudy area and ·its .. su_b-uni t, Area I, ·that year.:·{see·page 20). It is possible that the prey became available too ~at~ in the Gyrfaldon breeding cycle to prevent some pairs from abandoning the area. Ground o. ~~· "-J· Ql 25 0 ~ C!) ,.., B u 0 5 A--....... .:...:_ . . . • ----~-t:X-~ ..... ---------.6. 100 • 75 A-----t. Ground squirrel population estimate 25 0._..._,_~-o OccurrE!nce of ground squirrels in prey re;aains. 1 . J~=---· __ .~_r ____ ...;._ _ _:._.;.-.:..~..;__ _____ .l..r _____ --J 0 1968 1969 1970 y·· e a r Figure 6. The percentage occurrence of Arctic Ground Squirrels }.n the 1968:-.1970 Gyffalcon ·prey remains co~pared >Ji.th an observed subjective estimate of the ground squi-rreJ:-population trends from the Seward ·Peo,insuia,· Alaska. . t::(O rJ \,Q. C\ ...... -~ ... 0 i. •. .·~ \-.Y 80 _squirvels are unavailable io Gyrfalcons from about 1 October to l May ·.due :to their vlinter hibernation.- Tp eJnphas.iz_e the apparent importance of ground .squirrels to Gyrfalcons nesting on the Seward Peninsula in 1969, a plot of the calculated per cent by weight of the yearly ~urns of avian species and mammalian species ideriti~ .fied Erom the prey remains has been made (see Figure 7). This figur~ indicates that, by weight, avian species declined ~bout 13 per cent in 1969, while in this same year ground. squirrels; by ~Ieight, increased about 13 per cent ln ·the Gyrfalcon pr~y remains. Table 6 (fr~m which the plot vras obtained) sho'trs that during all three years ground squirrels, by weight, constituted almost all of the mammalian .... ·· contr.J.bution -to the prey remains. Simj_larly, the approxi- mately 13 per cent decrease in the 1969 avian kill can be ... attributed to a J.arge extent to the decline, by weight, of ptarmigan and jaeger~. Figure 8 is an attempt to show the compen~atory role. played by ground sql,l.ir_rels and the rel.~.tionships of the three .. important categories· (ptarmigan, j aegers, ·and ground squirrels). PtarMigan and_jaegers, together by ~~ight, rleclined sharply in 1969 (Curve C). On the other hanq,-"f>he-;three primary categories including ground . sc:quirrels::~curve A),-· together by Neight, . did not decline as sharpl~ ~n~969. Grouping ptarmigan and ground squirrel~ \~~----(Cur-v-~-.:-:BJ:~~: thes.e tvw categories by Neight. res.ulted in the. --:~---~--~ -. ·.--. 95. 90 85 80· "-' ..c: eo •.-I ~ ~ >. .a ~ eo <11 .j..l· 25 ~ ~ C) H -~ ll:< 20 ~~ \ _ _) 15 10 5 ~ \J_· ,.-..------------------~-----------·---·-·--~-----------, (A) (B) 1968 1969 1970 Y e a r ·rigure 7. (A) The estimated biomass of ptarmigan and ... Long-tailed Jaegers calculated as a p~t;c::entage hy · · ·l~teight of" the avian rem.:lins, and. (B) the estimated . · biomass or· Arctic Ground. Squirrels calculated -a£ .a · ~.percentage. bY tveight of the rr.aa~~lian· remains identi..:.. , :,fled .from the .1968 -1970 Seward Peninsula Gyrfalcon -:;~·:PI"~ remains. 100 95 90 . . 85 80 75 u ~ 70 .... •rl Q) .:?. 65 .>. . ,0 20 15 io ·s ·o- (A)'-.._. '--__ ___L_ ______ _J..i ------''_;__----1 1968 196_9 1970 . y e a r Figure 8. Major Gyrfalcon pr~y item~-e~pressed as a percent- --_ ag~-of the tot.:;.l of yearly kills by· weigtt: {A) ptarniigan, j ;~-~g-ers, ground squirrels; (B) ptarmigai1, ground squirrels'; (cf'ptarmigan, jaegers; (D) ground squirrels, jaegers • . : -~----·:__,_ __ . -- --.: ____ _ ._:_·~-)~--. .. ~ 83 flattest curve. This indicates a nore stable yearly utili- zation of these two resident species in changing combina- tions. However, grouping ground squirrels and jaegers (Curve D) shows that these two categories together by w~ight, increa&ed in 1969, even though jaegers contributed the lea~t amoun~ by weight and, in fact, declined irt numbe~ of occurrences-that year. The increased occurrence of ground squirrels indicates them to be an jmportant source of food that helped "buffera the decrines in ptarmigan and jaeg~r biomass. it ~ust be reiterated that based on p~llet exam~ ination,· it was estimated that microtines occurred up to 5 times more frequently than was evident from the prey ·remains~· rn·terms of calculated biomass and percentage by weighti-this fact-has very little effect on the calculated Values presented on the basis of the prey remains alone (one per· cent--or less)~-In._fact, i'lere -microtines to occur up to fen--times the number they represent in terms of prey remains, -the results· in t_er-~s of percentage by \'Ieight. cal- culations are still affected only to a minor degree; Because of the differences in the ye-arly sample _ .sizes o~ _the Gyrfalcon prey remains, a _2 X 2 contingency table ~as~constructed and chi-iquare values were obtained - to he:J..p-0:0.lnpare by year the prey groups _listed in Table 11. In all cg,_ses, \'lith the exception of the Short-eared 01-..rls, (-"' the chi-~-qlJ.Stre value followed a pattern that is consistent '~ --·"-'""· -. . &J ". ',· ... '! .. ,. (_-_) vii th the picture presented by Figure 2 and the hypothesis that utilization of some prey groups increased in 1969 when other prey groups became less available to Gyrfalcons. Variatfon in prey species utilization betv1een different pairs Tables 12a~b to 27a~b list prey remains fr6m a series of Seward Peninsula Gyrfalcon eyri~s where 25 or more. kills v1ere found .. 'l'hese tables describe a range of variability of prey vlithin ·a s·ample of 23 Gy!'falcon nestings in differ-ing hal?itats and situations. T1,v0 factors may account-·· for this variation: ·( l) the location of the eyries in relation to prey species habitat and numbers, and (2) .~ pair or individual preferences on the part of the Gyrfalcons. To· demonstrate the variation in prey remains that occurred betw~~ri the Gyrfalcon eyries described in Tables 12a,b through 27aib, the three major prey categories (ptarmigan, j aege:rs_, ~nd ground·. squirrels), migratory bird species and resident species were ex:tracte.d. and are presented. in Table 28. It·· may p.1s:o .be _noted that of the 23 nestings ·listed where prey remains were col-lected, ptarmigan occur in every iristanc~, ground squirrels occur in 21, and jaegers occur . "·-'---"• ·Tn: l4>:·::o:he or ·more of the. resident species. occurred in all the nes.~iYrgs and migratory bird species occurred in 22. ·:.:"':It has been previously mentioned that Gyrfalcons· _::-_:~_-:_....__:. __ , ·ne~rting "in insular situations feed primarily on Alclds, --_-.. ···' .. _,_ .. ~-~ \--1 . "..___/ ;~ -~ .'-:~-,-=~ 85 Tabl~ 12a. Gyrfalcon ~yrie no. 1 1 , Seward Peninsula, Alaska 1968 1970 Per cent Per cent Name Number of kill Number of kill --- Long-tailed Jaeger 15 29.4 81 55.9 Ptarmig·an species 18 35.3 214 16.6 Bl~ck-legged ~ittiwake 3 5.9 11 7.6 Unidet1tified birds 3 5.9 5 3.4 American Golden Plover 1 2.0 3 2.1 v1himbre1. · 1 2.0 3 2.1 Pintail 2 3.9 1" 0.7 Unidentif-ied waterfowl 3 2.1 U:nident_ified shorebirds 3 2.1 Bar~ tailed Godt.fit_· 1 .2.0. 2 1.4 Unidentified Alcids 3 5.9 ·Black ·sra:nt ·· · 1 0.7 Green~winged Teal 1 2.-0 Ruddy_ T1:1:rnstone 1 0.7 Parasitic Jaeger 1 0.7 Short~eare_<i .Owl 1 2.0 LapHlrid Longspu·r 1 2.0 Unidentified passerines 1 ~ .. TOTAL BIRDS 50 98 .J 140 95.7 Arctic Ground _Squ:!-rrels .. 1 2.0 ll 2 . .s Vole sp~·gies 1 -0. 7. TOTAL MAMf>'IALS 1 2.0 5 3.5 TOTAL KILLS 51 145 1 Eyrie. no·:· 1. is located overlooking the sea above a narroH coastal: plain. -_:..:· __ . __ ..,_ ---· ~-·-··· ................. ____ ..__... __ .. ____ ........ ...-~ ·-86 .-'rab·le 12b. ·Gyrfalcon foo·d ·remains from Table 12a. grouped in various major food categories C~ategory Ducks Shorebirds Passerines Jaegers Seabirds Short..:..eared Owl unidentified birds Migratory-b-irds · subtotal·· Ptarmigan. ..· .. Grouncf -s-quirrels l\Ucrotines ·· . . . Resident species -,sub't;ot;§:l __ _ -~ta~migan~ jaegers, an:dground squirrels P~armigan and j aef;ers -------- Pt_armigan and ground squirr~J§ lF· __ l_gu:ses -_f:or -1968 are 2F· _ 1:-gures . for 1970 are ·······- --: ~-. .::..:..:: --=-----. -· -0.. .--''....0::..:.. 1968 1 19702 Per cent Per cent Number of kill Number Of kill 3 3 1 15 6 1 _l 32 18 1 19 33 19 .. calculated 5.9 5.9 2.0 29.4 11.7 2.0 . 5. 9 61.5 34.6 1.9 -- 36.5 65 .1~ 63.5 36.5 on the calculated-on-the 5 3 . lt 12 8.3 1 0.7 82 56.5 11 7.6 _2 ~ 116 80.0 211 16.6 4 2.8 1 ~ .. 29 20.0 109 75.2 105 72-.4 28 19.3 basis of 51 ld.lls. basi-s ·of 145-kill~ • 1 Table l3a; Qyrfalcon eyrie rio~ 2 1 , S~ward Peninsula, Alaska ' ij j'·· .N· :. :.:. :\: ·/.: • 'j_; ·= arnet .:\. -i\ •· -~··!;;:·!, P~anmi~an s~ecies ' Long-tailed ,jaegers ; Jae~er species • · , Unidentified shorebirds Americ~n Golden Pl6ver 1t!himbre 1 ·' . Par~keet: Auklet ·· Unidentified passerjnes Unidentified birds ' TOTAL ·BIRDS Ar,ct.ic Ground Squirrel Brbwh ·Lemming ' . - po:I1ared Lemming I •• ' TOTAL MAMMALS ' . TOTAL KILLS ,•j ' ' ·~ ••• ••• r J \, : Number ;2lf '2( 1 + 53 1. 1 2 55 I ' : .. 1:968 P~r:cent of kill L13. 6 4q,l i.B 1.8 96.4 1.8 1.8 3~6 ..... Number fe,r .rent· iQf kill 37 ·74.0' 6 J-2.0 I 1 2.0 1 2.0 1~8 96.0 2 1!. 0 2 l!. 0 ,50 I Number. 74 4 1 1 1 1 82 1 1 2 84 Per cent· of kill ;. ·88.1 4,. 8 1.2 1.2 1.2 1.2 97.7 1.2 1.2 2 ,l! 'iT? '• 2 .. .Gyrle no.. . ls located at an inland river bluff in a narrow valley, 10 ~ir-miles from the coast. 2This'eyrie was ·probably r"970. I. I . : ---r .• ) I occupied py the same pair of g~rf~lcon~ during 1968 and 1969, and . ! i . ~ . • ! • ! ' ' ' ! I: I·' ! Cl].' ... · ... L I ~ ! . Table l3b. G~rfalc6n fobd remain~ from Tabl~ 13a g~ouped in various major categories .••,, • : .. i, ·:, ' . . ' i . i ' ~ :) .. l >i ·JI,: : ii \. .: ! :1 ~.~.· ' ,. ., ) . Ga't'e·gotv · '\i .I :·! ·; .;-_. '.'_v,.: ..•. ' ::; ; ·nucks:' · $hqr~birds Passerines Jaeg~rs: ·seabirds .Unidentified birds i· 19681 __ .._ 'I i ' ' · Per cent Nti~b~r· of kill ··r i 1·. 8 j8 ! 50.9 -:--·-- Migrato~y birds subtotal 29 52.7 Ptarmigan Grdurid squi~rels :.1ic:rotines 24 1 1 Reiident ~pecies subtotal . 26 ; j ! Ptarmigan~ jaegers, a~d grou~d .sqtifrrels ~tarmigan a~d j~egers Ptarmigan and :ground · . squirrels !Figures for 1968 are ?Figures for 1969 are 3Figures for 1970 are 53 52 25 caJ!culated calculated calculated 43.6 1.8 1 .. 8 47.3 96.4 94.5 l{ 5 . li on the basis on the basis on the bas:i.s of 55 kills. of 50 kills. of 81.t. kills. ·--.-..... '~ '~ _ _) Table i4a. .. . ... ·1. Gyrfalcon eyrie no. 3 , Seward Peninsula, [\_laska · 1968 1969 2 89 Per cent Per cent Name Number· of kill Number of kill - Ptarmigan species 10 32.3 16 38.1 Robins 3 7.1 Unidentified passerines 3 9.7 2 4.8 Bar-tailed Godwits 2 6.5 American Golden Plover 2 4.8 Long-tailed Jaegers 2 1L8 -RedpoJ.ls. 2 4.8 Jaeger species 1 3.2 Whimbrel 1 2.4 Ruddy Turnstone 1 2. lj .. Tr-ee Sparrow. __ 1 . 2. 4 GraY:-che-eked Thrush 1 2. 1+ Unidentified shorebird-1 2 ~ Ll Unidentified bird .: 1 2.4 -TOTAL BIRDS 16 51.7 33 61L 5 Ground squirrels 15 48.4 _2_ 21.4 TOTAL M.f\.l'lfr1f!.LS 15.: 48.4 9 2l.l.J ---ToTAL KILLS 31 42 · 1 Eyrle .110 ~-3 is located on the lm'ler slopes of a narrow · river va·lley 8 air-miles from the coast. 2This ··eyrie "~>Tas occupied by the same pair of Gyrfalcons -1968-:19.69. , ... .,....· . .. ,·.,_:-.; ._·. - I j I l I i .! I 90 Tab~e l4b. Gyrfalcon food remain~ from Table l4a g~buped in various major categories Per cent Per cent Category Ducks Shor-ebirds Passerines Number of kill Number of kill _Jaegers Seabird Unidentified birds fliigratOJ?Y -bir_cts sub~otal Ptarmigan __ Ground squirrels Nicrotines Resident species subtotal Ptarmigan, jaegers, and _ ground squirrels Ptarmigari and jaegers Ptarmigan and ground squirrels 2 3 1 6 10 15 25 26 11 ?I') '-./ 1 Figures for.l968 are calculated 2--. __ -_ -----Figures-:for--1969 are -calculated ,..~,-·,L.,::-._· ... 6.5 9-7 3.2 19.4 32.3 }~ 8 . lj -so. 7 83.9 35.5 80.7 on the on the 5 9 2 1 17 16 9 -25 25 basis of --- basis of 11.9 21.4-- 4.8 2 .1-J 4-o .-s 23.8 21.1~ 59~5 -59.5 31 kills. 42 kills. () Tabie t5a. Gyrfalcon eyrie no. 41 , Sevmrd J;eninsula, /\ \ J '·._/ Alaska· Name ---- Ptarmigan species Tufted Puffins Pigeon Guillemots Unidentified v.raterfowl Unidentified passerines Unidentified birds h'himbrel Unidentified shorebirds Long-tailed .Jaeg·er's- _Robins TOT AT_, BIRDS Ground squirrels 1 .- -TOTAL r1Ar.'IMALS TO'rAt. KILLS Eyrie no.-4 is located a seabird- ---. ~-~ ,--.:--·- colony. ··· .,. -. . . <.~ 1969 ?er cent Number of kill ll 26.8 7 17.1 4 9.8 3 7.3 2 1L9 2 4.9 l 2 • 1~ 1 2,11 l 2.4 l 2.4 33 80.4 8 . 19.5 8 19.5 1!1 on a, sea-cliff on the periphery of 92 Table 15b. Gyrfalcon food remains from Table 15a grouped in_ various-major categories Categor~ Du-cks Shorebirds Passerines Jaegers Seabirds Unidentified birds Migra~ory birds subtotal Ptarmigan f.1icrotines Resident speci~s subtotal Ptarmj_p;an ,-.) aeg-ers, and -- Ptar~igan and jaegers Ptarmigan and grJund squirrels Number 3 2 3 1 11 2 22 ll 0 C) 19 20 12 19 19691 Per cent of kill 7.3 4. 9- 7.3 2.4 26.8 ~.__2. 53.7 26.8 19.5 46.3 I.,-, Lj o . ·r 29.3 46.3 1 Figures for 1969 are calculated on the basis of 41 kills. '(' .~\ Table 16a. Gyrfalcon eyrie no~ 5 1 , sekard Peninstila, Alaska i 'Ji:'" . I Name Ptarmigan specie~ Common snipe , American Golden Plover Jaeger species Short-eared Ovll Robin Unidentified passerines 'l10TAL BIRDS Arctic Ground Squirrels· Collared Lemmin~ Brown Lemming Vole species · TOTAL MAMr·1ALS :,l,OT AL KILLS · Number· .8 5 2 1 1 17 8. 8 }~ 1. 21 38 1968 ' " Per cent' of kill 21.1 13.2 5.3 2.6 2.6 :4 4. 8 21.1 21.1 10.5 2.6 55.3 .. 1969' Number · Per cent or kill 8 1 "! 1 10 1 1 11 72.7 ' 9.1 ' 9.1 90.9 g,l 9.1 1970 Per cent Number of.kill 29 67.4 1 2. 3 3 7. 0 1 2. 3 1 ~ 35 81.3 ' 5 11.6 2 ' 1+ • 7 1 2.3 8 18.6 )~ 3 1 Eyri~ no. 5 is lo6ated on a hillside in a narrow valley system 23 air-miles:from the coast. '1 ., . I \ ! {0 r;, ') 111' [),, I {~ ~I ) '---······· Table 16b. Gyrfalcon food remains from Table· l6i grouped in var~ous major categories .1968 1 I ' ' :;·. ,i l'::i i>Cate€;or.lf:' ·, I ·., Ducks . Shorebirds· Passerines ,Jaegers Seabirds Short-eare<;i 0111 Number 7 1' Migratory birds subtotal 1 9 8 8 Ptarmigan Ground squirrels Microtines .u_ Resi~ent species subtdtal· 29 ' Ptarmigan, jaeg~~s,, and ground ~quirr~ls Ptarmigan and Jaegers Ptarmigan and ground squirrels 16 ' i ~Figures for 1968 are calculated Figures.for 1969 are calculated 3Figures.for 1970 are calculated Per cent 'f' 0~ kill Number 42.1 9 on the basis of 38 on the basis of 11 on the basis .of 43 19692 · ., :P'er cent •' .· i ; of kill : 9.1 ': 9.1 . 18.9 72.7 9.1 81.8 I 81.8 kills• kills:. kills. :Number 4 1 1 6 29 5 _l 37 .35' 30 3L1 19703 I Pe.r cent df kili 14.0 67.4 11.6 ~ 86.0 81. 3~ 69.7 79.0 95 Table 17a~ Gyrfalcon eyrie ~o. 61 , Seward Peninsula, Alaska Name Ptarmigan species American Golden Plover Robin Gray~cheeked Thrush Long-tailed Jaeger Semipalmated Plover Unidentified shorebirds Lapland Longspur Unidentified p~sser~nes . TO'J:IAL BIRDS Ground squirrels Collared Lern-ming TOTAL rJIAMr·iALS 7 TOTAL KILLS Number 8 6 3 2 1 1 1 1 1 24 38 1 39 63 1969 Per cent of kill 12.7 9.5 4.8 3.2 1.6 1.6 1.6 1.6 1·. 6 38.2 60.3 1.6 61.9 1 Eyri~ no.-6 is located on a hilltop in a wide river valley 11 air-miles from the coast. - ---- - - - Table ~7b. Gyrfalcon food remains from Table 17a grouped in various major categories Category Ducks Shorebirds Passerines Jaegers Seabirds Migratory bird5 subtotal Ptarmigan Ground squirrels Microtines Resident species subtotal Ptarmigan; jaegers, and ground squirrels Ptarmigan and jaegers Ptarmigan and ground squlrrels Nurr:ber 8 7 1 16 8 -:<R ..J" 1 Jn 117 9 116 1969 1 Per cent of kill 12.7 11.1 1.6 25.4 12.7 C:.r. ..., uv • .) , c:. ...1... • 1._} 74.6 71!. 6 11~. 3 73.0 1 Figures for 1969 are calculated on the basis of 63 kills. 97 Table 18a~ Gyrfalcon eyrie no. 7 1 , Seward Peninsula, Alaska Name Ptarmigan species Unidentified waterfowl Pintails TOTAL BIRDS Ground squirrels 'l'OTAL MAMMALS TOTAL KILLS Number 36 ll _l 43 _l- 3 46 Per cent of kill 78.3 8.7 __§__0_ 93.5 1 Eyrie no. 7 is located on a river bluff in a broad, marshy valley 36 air~miles from the nearest coast. 'l'able 18b ·-. Gyrfalcon food remains from Table 18a grouped in various major cRtegories categ_?ry Ducks Shorebirds Passerines Jaegers Seabirds Migratory birds subtotal Ptarmigan Ground squirrels Microtines Resident ipecies subtotal t'tarmj,gan, .i aegers ,-and ( ground sq~irrels Ptarmigan and ground sql!i:t:_I'e 1~ Number 7 7 36 3 39 39 Per cent of kill 15.2 15.2 78.3 6.5 81~ • 8 _ _g 4 • 8. 1 Figur~s f~r 1969 are calculated on the basis of 46 kills. I l I l .I Table 1.9a-. -·Gyrfalcon eyrie no. 8 1 , Se~-mrd Peninsula, Alaska -1968 99 Name Number Per cent of kill Ptarmigan species hfhimbrel Short-eared Owl Unidentified passerines Unidentified bird TO'l'AL BIRDS -Ground squirrel -TOTAL r·1AMMALS TOTAL KILLS 54 2 2 l l 60 4 4 6ll 84.4 3.1 3.1 1.6 1.6 93.8 6.3 1 Eyrie no. 8 is located on the shoulder of a ridge in a -hig~ ba~ren valley system 32 air-miles from the near~st coas.t .. ;.;·,H>f!l<'i~TY ilF .~ ....... ,. .............. .-.. ------- u--"NIV. Oli' -AtASKA .. LlliRAH.Y ~ ~ i I I ~ f. I i I l I I i i i I i 100 Table 1.9b. Gyrfalcon food remains from Table 19a grouped in various major categories Category Ducks· Shorebirds Passerines Jaegers Seabirds Short-eared Owl ·- Migrator~ bir~s subtotal Ptarmigan Ground squirrels l'l!icrotines Resirt~nt species sybtotal Ptarmigan, jaegers, and ground squirrels ~.P_tarJnigan and j aegers Ptarmigan and ground squirrels Number 2 1 2 6 511 lj 58 58 Per cent of kill 3.1 1.6 3.1 9 . 1~ 8 4. lj 81! • l! 90.6 . 90.6 1 Figures-for 1968 are calculated on~the basis of 64 kills. ---· ... ---·· 101 Table 20a. Gyrfalcon eyrie no. 9 1 , Seward Peninsula, Alaska Name Ptarmigan species Short-eared OvJl Unidentified shorebirds Pintail Long-tailed Jaeger TOTAL BIRDS Arctic Ground Squirrel Unidentified microtine Mink TOrr AL MAI\1MALS TOTAL KILLS 1968 Per cent Number of kill 153 93.9 3 1.8 2 1.2 1 0.6 159 97.5 2 1.2 1 0.6 1 0.6 4 2.4 163 1970 Pe-r cent Number of kill 21 65.6 , 3.1 ..L 8 25.0 -·- 30 93.7 2 6.3 2 6.3 32 1 Eyrie no. 9 is located on a hillside in a broad vaJ.ley system, 16 air-miles from the nearest coast. 102 Table 20b. Gyrfalcon foo0 ~~~~~~s ~rom Table 20a grouped in various major categories -------- Cater:;ory -Ducks Shorebirds Passerines Jaegers Seabirds S~10rt-t=;ared _Owl I·Ur;rator,y b:Lrds subtotal Ptarmlgan Ground squirrels !:Jicrotlnes Resident species subtotal P tarm]_gan, j aegers, and ground squirrels Ptarmigan and jaegers Ptarmigan and ground squirrels Number ---- 1 2 3 6 153 2 l --'- 157 155 Per cent of klll 0.6 1.2 1.8 3.7 93.9 1.2 0.6 96.3 Number J. 8 9 21 2 23 31 29 23 Per cent of ldll 3.1 25.0 28.1 65.6 6.3 71.9 96.9 90.7 '71. 9 1 Fig~res for 1968 are calculated on the basis of l63 kills. 2--. -Flgures for 1970 are calculated on the basis of 32 kills. 103 'rabl-e 2la. Gyrfalcon eyrie no. 101 , Seward Peninsula, Alaska Name Ptarmigan species Unidentified waterfowl species Unidentified sandpiper species Unidentified passerines TQ'r.AL BJ:RDS Arctic Ground Squirrel ·collated Lemming· · Unidentified microtine · TUr AL l'-1M!JivrALS · · 'fOTAL KILLS ____ 1968 Number 11 l 1 1 3 2 1 11 25 Per cent of klll 44.0 11. 0 4.0 4.0 56.0 32.0 8.0 11 • 0 ljll • 0 1 ~Eyr~e no. 10 is located high on the side of a river valley· 15 air~miles from the nearest coast. . . ( Table 2lb. Gyrfalcon food remains from Table 2la grouped ~n various major categories Ducks Shorebirds Passerines Jaegers Seabirds Mi~ratory birds subtotal Ptarmigan Ground squirrels rucrotj_nes Hesj_dent species subtotal Pta:r'migan, j aeg:,ers" and ground squirrels Ptarmigan and jaegers · Ptor·rnigan and ground squirrels ------ 1968 1 ----------~· ----------- Number 1 1 1 3 11 8 __]_ 22 19 Per cent of kill 4.0 4.0 4.0 12.0 44.0 32.0 12.0 88.0 76.0 1 Figures for l963 are calculated on the basis of 25 kills. ( I \ 105 Table 22a. Gyrfalcon eyrie no. 11 1 , Seward Peninsula, .Alaska Natne Number Ptarmigan species TOTAL BIRDS TOTAL KILLS 45 1970 Per cent of ;.:J.ll 100.0 100.0 100.0 1 Eyrie no. 11 is located on a river bluff at the edge of the central lowlands in a large drainage system 42 air-miles from the nearest coast. 106 Table 22·b. Gyrfalcon food remains from Table 22a grouped in various major cate~ories Category Duc:Ks . Shor.ebirds Passerines ·Ja~gers Seabirds Migratory birds subtotal Ptarmigan. Ground.squirrels f·Hcrotines 'Resident species subtotal Ptarmigan, jaegers\ and ~round squirrels Pta~migan and jaegers Ptarmigan and. gr.ound squirrels Number Per cent of kill 100.0 100.0 1 Fig.ures for· 1970 are calculated on the basis of 45 kills. /-~, ~~~ -~----5 107 1 Table 2Ja. Gyrfalcon eyrie no. 12 , Seward Peninsula, Alaska Name · Ptarmlgan species Short~eared Ov1l Long-ta~led Jaeger TOTAL BIRDS -Red-backed Vole Lemming species TOTAL MAMMALS TOTAL KILLS Number 62 2 1 3 l 11 69 1970 Per cent of kj_ll -89.9. 2.9 l. li 9 !J • 2 11_ 3 1.4 ~ '7 -' • I 1 Eyrie no. 12 is lb6at~d ori a river bluff at the edge of ·-the central lowlands in a large drainage system 42 air- miles from the nearest coast. < ••• • 108 'l'able · 2-3b. · Gyrfalc·on · food renains from 'rable 2 3a p.;rouped in various major categories Category ·Ducks Shorebirds Passerines Jaegers Seabirds Sr1ort-eared 0\vl f'ligratory birds subtotal Ptarmigan Ground squ~rrels Microt·ines Resident species subtotal Ptarmigan, j~egers and ground squirrels . Ptarmigan and jaegers Ptarmigan and grouod squirrels Number 1 2 3 62 lj 66 67 Per cent of kill 1. lj 2.9 --· 4.3· 8'.L 9 -~ 95. '[ 97.1· lFigures for 1970 are calculated on the bas'is of 69 k:!.lls ·?"..·-.:. ··-;· .. :-·· ... 'l'able .~I~ a~. Gyrfalcon eyrie no. 13 1 , Set'Jard Peninsula, A1flska 1970 109 Per cent Name - Ptarmi~an species · Unident::tf1.ed birds OldsquaN . - American Clo}rlcr: Plover common sn:l.pc. Lesser Ycllowlegs Lon€_~-ta:llr:d Jaer;er •rree f>parr·ovr SriOW Bun!~ tnr;-- Unidentified passerine 'l'O'f!\L BIRDS Arctic Ground Squirrel .'l'O'l'A L fv1Af1UiiALS~ Number 81~ 10 1 1 1 1 1 1 1 1 102 _]_ 7 109 of kill 77.1 9.2 0.9 0.9 0.9 0.9 0.9 0.9 0-.9-- __Q_:_.2_ 93.5 6.4 \ 6. lf lEyri~ no.-13 is located high on the edge of a plateau ~ecion falling off to the coastal lowlands and 27 a1~~ miies from the rtearest coast • . - - - - - - - - ( i \ .110 Table 24b. Gyrfalcon food remains from Table 24a grouped in various major categories Ducks· Shorebirds Passerines Jaegers Seabirds Unidentified birds Migratory birds subtotal Ptarmigan Ground squirrels Microtines Hesident specie.s subtotal Ptarmigan, jaegers, and ground squirrels Ptarmigan_and jaegers Ptarmigari and ground squirrels Number 1 3 3 ·1 10 18 811 7 91 92 85 91 1970 1 Per cent of kiJ.l 0.9 2.8 2.8 0.9 __ 9 -~ 16.5 - 77.1 6.4 83.5 8 ~~ • 4 78.0 83.5 1 Figtires for 1970 are calculated on the basis of 109 kills. -..:.; . -·~· 111 4 1 . Table ~5a. Gy~falcon eyrie no. 1 , Seward Penjnsula, Alaska 1970 Per cent Name Number of kill ----- Ptarmigan species 11 37.9 Unidentified passerines 3 10.3 Golden Plover 2 6.9 Common snipe 1 3.4 l.rlhimbrel 1 3. li TOTAL BIRDS 18 61.9 Arctic Ground Squirrel 8 27.6 Collared Lemming 2 6.9 Lemming species l 3.4 TO'rAL MAMMALS ll 3'7. 8 TorrAL KILLS < 29 I \ l __ . llj located creek bluff in broad rela-t;yrie no. is on a a · tively open valley system 17.5 air-miles from the nearest coast. ( -==------·--=~~ -------~ .~~ 112 Table 25b. Gyrfalcon food remains from Table 25a grouped in various major cate~ories Categorv. Ducl-;:s Shorebirds Passerines _ .raegers Seabirds Migratory birds subtotal Ptarmigan Ground squirrels i\Ucrotines Resident species subtotal Ptarmigan, jaegers, and ground squirrels Ptarr:;igan and j aegers Ptarmigan and ground squi:rrels Number 3 3 7 ll Q u _]_ 22 19 Per cent of kill 10.3 10.3 21-t .l '15. 9 2'7. 6 10.3 75.9 65.5 1 F'igures for 197 0 are calculated on the basis of 29 kiJ.ls. ---- Table 26a. Gyrfalcon eyrie no. . Alaska Name Ptarmigan species Urtidentified passerines ADerican Golden Plover Unidentified shorebirds Lon~-tailed Jaeger Snow Bunting Unidentified birds TO'l'AL DIRDS Arctic Ground Squirrel Lemming species Collared Lemming Vole species TOTAL J'.1M/iMALS Unidentified kill TOTAL_UNIDENTIFIED KILLS Tor.r AL KILLS 113 1 15 , Seward Peninsula, Number 17 5 I[ 1 1 J. 1 30 6 2 1 1 10 1 1 1970 Per cent of kill 41.5 12.2 9.8 2 .li 2. ll 2 . L~ 73.1 lil. 6 4.9 2,11 2 . )~ 211 • 3 2 .11 2.4 1 Eyrie no. 15 is located hi~h on the side of a narrow creek valley surro~nded by high ~ills 15 air-miles from the coast. ---~- / ' ( \ llLJ Table 26b. Gyrfalcon food remains from Table 26a grouped in various major categories ·category -Ducks Shorebirds Passerines Jaegers Seabirds Unidentified birds IV!igratory birds-subtotal _Ptarmigan Ground squirrels f':Iicrotines Resident species s~btotal -Ptarmigan, jaegers, and :_ ground sguirrels Ptarmigan and jaegers Ptarmigan and ground squirrels Number 5 6 1 1 13 17 6 4 27 2lJ 18 23 Per cent of kill 12.2 14.6 2. l! 2.4 31.7 Jn .5 14.6 2~~ 65.9 31.5 43.9 29.1 1 Figure& for 1970 are calculated on the basis-of 41 kills. - --- "" ----..--·---· 115 Table 27a. Gyrfalcon eyrie no. 161 , Seward Peninsula, Alaska Name Ptarmigan species Common snipe Unidentified passerines Robin Lapland Longspur Bar-tailed Godwit Unidentifi~d shorebirds Say's Phobe Varied Thrush Water Pipit Fox Sparrow TO'r AL BIRDS Arctic Ground Squirrel Collared Lemming _L_ernming species Vole species TOTAL l\1AMMALS 'l,OTAL KILLS Number 27 3 3 2 2 ]_ 1 1 1 l 1 IJ3 25 lt 2 1 32 75 1970 Per cent of kill 36.0 ILO ILO 2.7 2.7 1.3 1.3 1.3 1.3 1.3 1.3 57.2 3 ? -. .J •. 5 ).3 2.7 ~ 112. 6 1 Eyrie rio. 16 is located hi~h on the side of a broad river valley 12 air-miles from the coast. ' -~ 116 Table 27b. ·Gyrfalcon food reDains from Table 27a grouped in various major categories Cate~o~ .· IJucks Shor·ebi rds Passerines .Jaegers Seabirds Migratory hirds subtotal Ptarmigan Ground squirrels Microtj_nes Resident species subtotal Ptarmigan, jaegers; and ground squirrels Ptarmigan and ground squirrels Number 5 ll 16 27 25 _]_ 59 52 ----- Per cent of kill 21.3 36.0 33.3 9. 3_ "(8. 7 69 .11 1 Pi;-::_;ure-s f~r 1970 are calculated on the basis of 7·5 kills. " 'The ran~;e 1 in the per cen·t occurrence and numbers of the three' ma,j or prey categories, migratory · bird species, and resident sp~cies at 23 Seward Peninsula Gyrfalcon ne~tings 1968 1969 1970 Pr-ey Ca';;egory Per cent Numbers Fer cent Numbers Per cent Numbers _ _2.£_ G ro :1 12__ Ra:1:;;e Ranl\e Rans;e Ran~e Ran~e Ra:;'lr;e Ptnrmi~a.n 21.1 j 93.9 8; 153 12.7; '(8.3 8· 37 16.5; 100.0 • 1 • 84 ' ~...~..., Jat~·;_;,:?rs 8.0; 50.9 0. 28 0. 0; 12.0 0. 6 0. 0; 56.5 o· ,......, ' ' ·' C>c: G:rouncl squirrels 1.2; 118. 4 1; 15 4. 0; (j 0 .. 3 1; 38 0.0; 33.3 0; 25 :'<Ugro. tory bird species 3. 7; 63.5 3; 33 15.2; ~53 .. 7 2; 22 0. 0; 80.0 0; llG Resident species 35.5; 96.3 19; 157 1!6.3~ ()4 .. 8 9• .47 20.0~ 100.0 22; Ql .. ~ 'l'OTAL NESTINGS 7 6 10 1 n11 arithmetic figures rep~esent the minimum and maximum values obtained from tables 12b through 27b. =======================================================·--===================================== 1113 Larids, and Anatids, and that pairs nesting inland feed primarily on ptarmigan (Dementiev and Gortchakovskaya, 1945; Cade, 1960). This suggests that Gyrfalcons will tend to utilize those snecies that are the most abundant (i.e., available and therefore possibly the easiest to catch) in the general vicinity of the Gyrfalcon nesting cliff. From my data on the food habits of Gyrfalcon nesting on the Seward Peninsula, it is apparent that pairs nesting on the coast or in an area strongly influenced by the coastal environment pr~y ~~on species associated with both the coastal and the inland habitats (especially Larids, Alcids, jae~ers, ground squirrels, and ptarmigan). In general, the prey taken by a nesiing Gyrfalcon pair on the Seward Pcnin- sula-reflects the habitat characteristics of the hunting ranse of t11at pair such that pairs nesting in the vicinity of a particular species !!concentration!! ''rill prey substan- tially on that species. For examp~e, pairs (eyrie number~ one and four) nesting on the coast will take a substantial nu~ber of jaegers, Alcids, and Larids compared with a pair nesting in the uplands where ptar~igan and ground squirrels 11ill _predominate. Little i8-kno1·m about pair or incli vidual prefer- ence. It _is note-worthy that the nesting pair at the eyrie des_ignated._::,_': 2" in the tables took large numbers of Lonr:;- -- tailed ~~~~ers during a time when ~ock Ptarmigan were very 119 plentiful in the immediate vicinity of their nest (about 40 territorial male Rock Ptarmigan were in view of their nest- ing cliff). In 1969 this same pair (as judged by the color of the members of the pair) accounted for at least six Long- tailed Jaegers at a time when jaegers were much scarcer and when ~1 oth~r nesting pairs of Gyrfalcons accounted for only ten Long-tailed Jaegers (based on the prey remains). Tables 12~,b ~o 27a,b describe a continuum of the variability of Gyrfalcon prey over a large sample of eyries in differing habitats and situations. It is possible to fit ~ll pr~vious Gyrfalcon food studies reoorted in the literature into this broad picture. A broad continuum of situations exists ranging fron insular seabird eater~ to interior ptarmigan/grotlnd squirrel eaters. This general continuum will be affected,. for example, hy the nearness of waterfowl habitat (Bengtson, 1971). Each general ar·ea of G~~falcon nesting habitat can be defined by geographical criteria {the existence of marshes, coast lines, rivers and uplands, for example) and each of these areas will tend to have its own general falcon/prey relationship. E8ch Gyrfalcon pair, dependent upon their location in relation to prey species, will produce a unifor~ dietary prey list, but that.list will be strongly related to those p~cy spe- cies T6~nd in the vicinity of the Gyrfalcons' nest. -su~h 120 a dietary list from one pair may or may not closely resem- ble the dietary lists from other Gyrfalcon pairs depending upon whether or not all the pairs are nesting in similar habitat Situations. SUMMARY AND CONCLUSIONS 1. Breedin~ Gyrfalcon populations on the Seward Peninsula during the summers of 1968, 1969, and 1970 were st~ble on a region-wide basis at a high level. Of 131 nestings observed, 311 occurred in 1968, lJ8 in 1969, and l.J9 in 1970. The low figure for 1968 reflects poor survey -coverage in tha~ year. 2. Local shifting of populations within smaller units of the peninsula occurred which were correlated witl1 prey abundance. 3~ Nesting-cliff tenacity 0as low. Over the three summers only four per cent of the nest1ng sites were oc~upied all three years. The interrelationships between Rough-legg~d Hawks, Golden Eagles, and Ravens, all of ~hitih also utilize cliffs as nesting sites~ is complex anct as yet p6brly ~nderstood. 4. Distribution on the peninsula is not uniform, but controlled by the availability of nesting cliffs. Over all three years 66 per cent of the observed nestings occurred in only about 2,200 sauare miles of the peninsula or approximately 13 per_cent. 5. Prey remains were collected from 37 nestings· and 1,483 kills were identified, representing 32 species bf birds and eight species of mammals. This is a considerably 121 . i : 122 more diverse list of Gyrfalcon prey than has previously been renorted. 6. Four species vJere of overriding importance: Rock Ptarmigan, Willow Ptarmigan, the Arctic Ground Squirrel, and the~Long-tailed Jaeger. These four made up 81 per cent by number and 92 per cent by weight of the sample. 1. Prey utilization varie~ with availability both temporally and spatially. ~~ile the four species mentioned-above continued to be important during all summers, it is apparent that Gyrfalcons readily tions to members of a wide spectrum of other prey species as availability dictates. LITERATURE CI'l'F.D Bengt~o~, Sveh~Axel. 1971. · Hunting methods and choice of prey of Gyrfalcons Falco rusticolus at Myvatn in northeast Iceland. Ibis, 113:~68-476. Bond, R. M. 1936. Eating habits of falcons with special reference to pellet analysis. Condor, 38:72-76. Brown, Leslie, and Dean Amadon. falcons of the world. 1968. Eagles, hawks, and McGraw Hill, N.Y. 945 pp. Brull, H. 1937. Das Leben de~tscher Greifvogel. Jena: Gustav Fischer. Not seen. Cade, T. J. 1960~ Ecology of the peregrine and Gyrfalcon populations in Alaska. Univ. Calif. Publ. Zool., 63:151-290. Dementiev, G. P., and N. N. Gortchakovskaya. biology pf the Norwegian Gyrfalcon. 559-565. 1945. On the Ibis, 87: Erring-ton, P ~ I.!; 1930. The pellet analysis Method of raptor food habits study. Condor, 32:292-296. 1932. Technique of ~aptor fbod habits study. Condor, 34:75-86. Gudmundsson, Fir.nur. 1970, The predator-prey relationship of the Gyrfalc6ri (Falco rusticolus) and the Rock Pta~migan (Lagopus mutus) in Iceland. Excerpts from the Abs~racts of the XV International Orni- thological Congress. In Raptor Research News, 4: 178. Hagen, Yngvar. l9S2. Tile g,y 1•..,-falcon irr Dovre, Norway. Skrifter Norske Videnskaps-Akademi. No. 4. (Falco rusticolus L.) Utigitt av Det -- I. Mat-Naturv. Klasse, Murie, A .. 1946. Observations on the birds of Mount ·.McKinley Nation~l Park, Alaska. · Coridor, 48: 253-261. 123 ~ ~ I ~ t I I I J I I i I l 12lJ Roseneau ,-D. n. 1910. -Numbers and productivity of Gyr- falcons on the Seward Peninsula, Alaska~ In A~aska Departr:Jent of . ..,:ish and Game Bird ~Ze1-~r:1en~ !}eport Vol. XI. Ratcliffe, D. A.· 1962. Breeding density of the peregrine (Falco peregrinus) and Raven (Corvus corax). -Ibis ,-lOLl: 13-39. Salomonsen, F. 1951. Gr¢nlands Fugle, Vol. 3. Copenhagen. Voous, I<.-H. 1960·. Atlas of European birds. Ne"I'T Yor1<:. Nelson. 284 pp. Weir, D. N. 1967. A possible source of error in raptor food analysis. Bird Study, 14:194. · 'dhite, ____ ., C_. J.Y., and H. K. Springer.. 1965 ~ Notes on the Gyr- falcon in western coastal Alaska. Aul<, 82:104- 105. and~-R. B. ~~eden. 1966. falcons and behavior of Condor~ 68:517-519. Hunting methods of Gyr- their prey (ptarmigan).