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Home , California condor, Condor

Chancellor,R. D. & B.-U.Meyburg eds. 2000 Raptorsat Risk WWGBP/Hancock House

The CaliforniaCondor Recovery T\rrogtamme

Lloyd Kiff

ABSTRACT Relativelywidespread over through the latePleistocene, the Catifomia hasbeen a rare throughoutits recordedhistory. Despitemuch habitat preservation,research, education,and enforcement measures, it steadilydeclined in numbersfor nearly200 ycars, reaching a low point of 2l-22 in 1983.Losses were due almostentirely to direci humanpersecution, primarily shootingand indirectanthropogenic causes, including lead and possibly other forms of poisoning. "hands-on" In 1980a managementprogftrmme was initiated uy ttreuniteo StatesFish andWildlife Serviceand the NationalAudubon Society. Wild birds weretquippedwith radios!o determine causesof mortality and to identify critical habitat,breeding behaviour was studied intensively, andseveral , young, and adults were taken to zoosto "stablisha captivebreeding programme' Despiteefforts to sustainthe wild population,the speciessuffered a disastrousdecline in the winter of 1984-85to only oneknown breedingpair, and authorities decided to trap the last free-flying individuals for their own securityano to aad them to the captive breedingprogramme. The last free-flyingindividual was broughtinto captivityon 19 April 1987.The first successful captive breedingwas accomplishedin 1988,and pioduciiviryof the captiveflock has inqeased annually sincethen at breedingfacilities at the SanDiego Wild park, Los AngelesZoo, and World CenterforBirds of hey. Thefirst releasesof captiveprogeny occumdin 1992.Subsequently, 16releases have been conducted at four sitesin Califomiaandtwo in .By theena of tSSi there were 147condors in existence,including 42 n the wild (22 in Arizonaand 20 in ) and 105in captivity.As a resultof theaggressive management programme, the total condorpopulation has increasedby seventimes since the early 1980s,and th; is increasinghope that this unusual speciescan be savedfrom extinction.

INTRODUCTION The California CondotGyrwtogyps californianushas always been rare in modemtimes. Judging from fossilevidence, it wasmore common and widespread in thepleistocene, ranging east*.ra al the way to Florida(Brodkorb 1964) nd New York (Steadman& Miller t987), buiby historical timesit occurredonly alongthe Pacific Coast from BritishColumbia south to northernBaja Califomia (Koford Norte, 1953).Following the arival of Europeanman on the pacific Coastabout 200 ago,condor numbers and range declined even more precipitously, almost entirely because

3U of anthropogenicfactors. As recentlyas the 1970s,many writersbegan predicting the Catifornia Condor'scertain and imminent demise. However, an aggressivemanagement programme begun in 1980may yet save it. Knowledgegained from both the positive and negative aspects of theprogramme mightbe usefulto personsatrempting to saveother problem species-.

BACKGROUND

Early history of condor rrsearrchand management The first detailedsnrdy of the CaliforniaCondor of any kind was conductedby the wildlife photographenWilliam Finleyand Herman Bohlman at EatonCanyon, Los Angeles Counry, Califomia in the springof 1906.The four-panaccount of their exploitsat a condorneit andthe behaviourof the nestling,which they took into captivity,still makesfascinating reading (Finley 1906,l90ga 1908b,19l0). Therewas linle formalattention paid to thespecies until threedecades later when Cyril S. Robinson, a British-bomemployee of the U.S. ForestService, made a pioneeringfield studyof the roosting andfeeding habits of condorsin theLos PadresNational Forest, mostiy in SantaBarbara Counti Califomiabetween 193640 (Robinson 1940). The fint actualgovemmental land management action on behalfof the condorwas the establishmentof the 1,198-acreSisquoc Condor Sanctuary by the U.S.Forest Service in 1937at theurging of localrancher-conservationist Robert E. Eastonand the NationalAudubon Society (Koford 1953).The sanctuaryarea included an importantcondor roost, nestsites, and bathing pool. In a ssnse,the condor management era began with thisevent, since the sanctuarywas henceforth closed to public entrywithout permit. At aboutthe same time, a prominentamateur omithologist and wildlife film pasaden4 "Bill" makerfrcm Califomia,J.R' Pemberton,began effons to film condorsin theSespe area of VenturaCounty, California"He intercstedJohn Baker, president of theNational Society, and Joscph Grinneit, a professorat the Universityof Califomiaat Berkeleyand the leadingomithologist in rhestate, in theidea ofsupporting a graduatestudent to conducta comprehensivestudy ofthe CaliforniaCondor underGrinnell's direction. With financialsupport from Pemberton and the National Audubon Society, thanksto Baker'sefforts, Carl Koford wasassigned by Grinnellto rheproject, and he conducteda landmarkstudy of thespecies and its breedingbehaviour between 1939-46 (Koford 1953).Koford workedmainly in the backcountry of VenturaCounty, and his findingsled to theestablishment of the35,00G'acre , which was enlarged to 53,000acres in l95l (Wilbur I97g). In his I 953monograph, Koford (op cit.) estimatedthat the total condor popularion consisted of only 60 individuals,although he privately admittedthat this was a conservativefigure and that the populationmight contain as many as 100birds (Koford in liu.). Condornumbers continued to decline,however, and the National Audubon Sociery commissioned a short-termassessment of thestatus of thepopulation in 1963-64by two Califomiaconservationist ranchers,Ian and Eben McMillan. Their findings, which were published with thecollaboration of Alden H. Miller of the Universityof California Berkeley(Miller er al. 1964),suggesred that the condorpopulation had declined to 40 birds. In 1965'Fred C. Sibleybecame the first U.S. Fish and Wildlife Service biologist charged with studyingthe condor population with thespecific assignmenr of predictingthe effects of a proposed damon Sespecreek, which flows throughthe sespecondor Sancruary,on rhespecies. Also in 1965,the National Audubon Society hired John Borneman as a "condorwarden," armed only with a strongsense of publicrelations and a frnelydeveloped sense of humour.The U.S. Forest Servicesoon had its owncondor biologist, Dean Carrier, and his assignmentwas to preparea condor managementplan for thevast Los PadresNational Forest, which, by then,comprised a largeportion of thecondor's range (Canier 197 I ). Canier'sposition was mainrained between l96g to 1973. Sibley'sassignment lasted for four years,and his conclusionthat thedam would likely havea disastrousimpact on thecondor population (Sibley 1969) coincided with thedefeat of theproject by the localelectorare. During his tenure,Sibley, Robert D. Mallette(california Departmentof Fish

308 andGame), and Borneman, initiated an annualsurvey of thecondor population, which washeld for two dayseach October from 1966to 1980and involved the participation of 50-100 trained observers (Mallette& Bomeman1966). The combined results of severalsurveys indicated that the total condor populationwas between50-60 individualsbetween 1966-70 (Wilbur 1978)and confirmedwhat manyhad suspected,i.e., that the populationestimate of the McMillans (Miller et at. 1964)was probablytoo conservative. SanfordR. Wilbur followedSibley as the U.S. Fish and Wildlife Servicecondor biologist in late 1969,and he held that position until 1980.During this entire period, he wasthe only U.S. Fish and "prograrnme." Wildlife Serviceemployee assigned to the condor Wilbur produceda monographic accounton the species,including an exhaustivebibliography (Wilbur 1978),and madedetailed studieson condorfood resources(IVilbur 1972), (Wilbur 1975),and populationstatus (Wifbur et al. 1'972,1976). He alsocollaborated in a studyof DDE-inducedeggshell thinning in (Kiff er al. 1979),conducted supplemental feeding experiments near the SespeCondor Sanctuary(Wilbur et al. 1974),and madeseveral expeditions to the SierraSan PedroMarth Baja CaliforniaNorte, Mexico in a fruitlessattempt to confirm the existenceof condorsin that range (Wilbur & Kitr 1980).Finally, he organizedthe First InternationalSymposium on theVultures, co- sponsoredby the WestemFoundation of VertebrateZoology andheld at the SantaBarbara Museum of NaturalHistory in 1979.The proceedingsappeared as a major book on Old and New World vulturebiology and conservation (Wilbur & Jackson1983), and the symposiumgrearly facilitated collaborationbetween researchers on severalcontinents. As a partof theFederal mandate to recovercondor populations, a "CalifomiaCondor Recovery Team" was establishedin 1972.Consisting of five members,representing the four govemment agenciescooperating in the programme,plus the NationalAudubon Society,the Tlnm preparcda rrecoveryplan for thespecies (U.S. Fish andWldlife Service1974).ltwas formalized in 1975,the first of its kind everapproved by theU.S. Fish and Wildtife Service.The plan washeavily oriented towardprotecting condor habitat as a meansof savingthe species. Habitatprotection, nest protection, and enforcement had been the themes of condorrecovery for manyyears, yet the Fendsin theannual survey results and the completedisappearance of condors from areaswhere they hadoccuned regularly up until very recentyears indicated that the species wascontinuing to decline.By thelate 1970s, Wilbur (1978) estimated thar the total condor population consistedof no morethan 50, andpossibly as few as40, individuals.By now,even some biologists werepredicting the inevitableextinction of theCalifornia Condor.

CALIFORNIA CONDORRECOVERY PROGRAMME

Modern manrgement era Reactingto the increasinglyhopeless situation, Richard Plunken, then vice-presidentof the NationalAudubon Society, proposed enlisting the help of theAmerican Ornithologists' Union to convenea panelof expertsfrom varioussubdisciplines of biology to considerthe situationand suggeststeps that might be takento savethe condor.Following severalmeetings, the panel,headed by RobertRicklefs, soonrecommended that a vigorouscondor management programme be set into motionimmediately. The principalingedients of thesuch a programmewere to includeradio tracking to obtain informalion on causesof mortality and critical habitat,detailed behavioural observations of birdsin the wild, especiallyat thenest, and, if necessary, @icklefs 1978). This "Califomia amountedto an endorsementof a draft CondorContingency Plan," preparedby the CondorRecovery Team in 1976under Sanford Wilbur's directionand approvedby the U.S. Fish andWildlife Servicein 1977,that had alsorecommended captive breeding and other"hands-on" actions(U.S. Fish andWildlife Service1996). The appearanceof thesereports gave momentumto the establishmentof the Condor Research Center,situated in Ventura,Califomia, by the U.S. Fish and Wildlife Service(USFWS) and the NationalAudubon Society in 1980.Noel F.R.Snyder was chosen to serveas the USFWScondor

309 biologist,replacing Wilbur, and John Ogden was placed in chargeof theNational Audubon Society portion of the programme.The California Departmentof Fish and Gamealso maintaineda condor biologistposition between 1982 to 1989. A divisionof labourwas determined early in theprogramme, and Ogden's crew focused on the identificationof criticalhabitat, while Snyder'sgroup located nests and made observations of birds throughouttheir breeding cycle. In May 1980Federal and State permits authorizing the caphrre of a condorfor captivebreeding and equippingl0 wild condorswith radiotelemetrydevices were approved,but they wererescinded in the following monthwhen a condorchick died while being examinedby a field teamfrom the CondorResearch Center. After nearlytwo yearsof hearingsand difficult negotiations,the California StateFish and GameCommission granted permission for researchersto placeradios on condors,and nine individualswere eventually tilgged by the NAS field crew.Paflly as a deviceto capturebirds, the NAS group engagedin supplementalfeeding of condorson large rancheswhere they were known to forage.Captured birds were weighed,blood samplestaken, and equipped with patagially-mountedradio transmitterswith bold numbersspecific to eachindividual. Up until this time, the perennialproblems in condor managementincluded (l) obtaining an accurateestimate of the numberof birds still in existence,(2) undcrstandingthe actualrange of the speciesand identifying critical habitat,and (3) identifying causesof monality. Noel Snyderand Eric Johnstondeveloped a novel techniqueto answerthe first question(Snyder & Johnston1985). They censused the population by comparingthe distinctive wing patterns,produced by differential patternsof featherloss and replacement,of individual condorsfnom photographs. Basedon analysesof hundredsof photographsfrom all partsof the condor'srange, they concluded thatthe wild populationconsisted of only 22 birdsin 1982(Fig. l). This wasthe lowestnumber of California Condorsin the recordedhistory of the species.It was also the first time that the entire condorpopulation had ever beenaccurately counted down to the last individual, a feat that would not havebeen possible with conventionalobservational techniques.

Figure 1. Condor Populations since 1982 160 J' 1rtO WILD CAPTIVE 129 100 80 60 40 zCI 0 r*l l-i 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99

Building on the years of informationabout condor movementscompiled by Koford, the McMillans,Sibley, Wilbur, and the annualOctober surveys, the NAS crewobtained a muchbetter understandingofthe currentrange ofthe condorby followingradioed birds with fixed-wingaircraft andon the ground.In addition,simply havingmore condor observers in the field led to increased

310 understandingof thewhereabouts of thebirds. It wasobvious that the effective range of thespecies haddeclined greatly since the early 1970swith a particularlyconspicuous decline in thenumber of birdsrecorded in thecoastal mountains north of SantaBarbara County. Meretsky and Snyder (1992) integrateddata from theradiotelemetry studies and the photographic survey, reporting that individual condorsoccasionally travelled more than 200 km, visitingall portionsof thestill vastcondor range in a singleday. Despitethe size of thebirds and the prominence of therecovery programme, surprisingly little is knownabout the causes of condormonality, even in recenttimes. Shooting was probably by far the mostfrequent cause of condorlosses historically, but suehincidents were also more likely to come to the attentionof condorchroniclers, especially since many birds were savedas museum specimens(Wilbur 1974,1978). Other known causes of condorlosses historically included kiuing condorsfor Indianceremonial use, capturing birds for sportor forpets, collecting, and accidental deaths,including striking man-made objects (Koford 1953,Wilbur 1978).Poisoning from various agentsmay havebeen a major problemal certaintimes and placesin the condor'shistory but convincingdocumentation of suchepisodes is surprisinglylacking. Severe DDE-induced eggshell thinningoccurred in thecondor population during the 1960s,and probably earlier (Kiff et al. lg79), but after the banningof DDT use in the United Statesin 1972,the incidenceof thin-shelledeggs declinedalmost immediately. Inexplicably, two badlythinned eggs were laid by a singlefemale in 1986(Kiff 1989),but by thenthis phenomenonwas no longera problemfor thecondor population asa whole. Partly as a result of the radiotelemetryprogamme, the causeof deathof threewild condorsin the 1980swas shownto be from a previouslyundocumented cause - (Snyder & Snyder1989). The birds had apparently ingested lead bullet fragments or leadshot from shotdeer or other carcasseswithin their rangein sufficientamounts to causetheir deaths.This finding had implicationsfor thepast, present, and frrture ofthe species.As for thepast, this could have been the causeof deathof severalcondors reportedly found sick andunable to fly (Koford 1953).Based on anadmittedly small sample size, it wasthe major cause of condorlosses during the 1980s,when the mostintensive field snrdieswere made of the species.In the westemUnited States, where hunting andcarrying guns arejealously guarded privileges, the incidence ofshooting on theopen range does not seemlikely to declinesoon, so leadpoisoning could still figureprominently in thefuture of the condor.Unforrunaely, only oneof thesix birdslost from thewild flock in thewinter of 1984-85was radioed,and the otherbirds were never found, so the actual causes oftheir deathsrcmains unknown. Snyderand his associatesmade detailed observations at condornests, adding greatly to the informationprovided earlier by Koford (Snyder1983, Snyder e t al.1986, Snyder& Snyder1989). Followingother difficult negotiationswith stateofficials, rhe team was finally allowedin 1983to begintaking wild-laid eggsfor hatchingin captivity in an effort to graduallybuild up a captive breeding flock. It had already been confirmed that Califomia Condors,like the related Andean condors,would lay replacementclutches (Harrison & Kiff 1980,Snyder & Hamberl9g5), and biologiststook advantageof thisbehaviour to double-and sometimes triple-clutch wild pairs.From 1983to 198515 eggs were taken from the wild andincubated at the SanDiego Wild Animal Park with hatchabilityof 86.7%and a survivalrate of 92.3%(Kuehler & Witman 1988,Kuehler er a/. l99l). Theoverallcondorpopulationbegan to increase forthe firsttimeinrecorded history following thelow pointin 1981-82(Fig. 1). Even thoughthe numberof birds in the wild continuedto decline(Ftg. 1), the situationwas lookingmore hopeful by the fall of 1984,when it seemedpossible that therecould be as manyas five breedingcondor pairs in thecoming breeding season (Toone & Wallace1994). Unexpectedly, however,six out of theremaining 15 wild birds(40% of theentirc wild population)were lost in the winterof 1984-85,and the numberof known breedingpairs was reducedto one.This prompted USFWSofficials to recommendin the fall of 1985that the lastrcmaining individuals be captured for their own securityand to bolsterthe captiveflock. Sincethe principal problemfor the condorhas alwaysbeen excessive adult mortality, this wasone way of reducingit, i.e.,bringing the birds into captivity where they were safer.The plan was strongly opposedby the NAS, who obtained an injunctionto block captureof the birds, but the usFws position,which was supportedby the majorityofcondor biologists,prevailed in court in June1986. The lastfree-flying condor, a male "AC-9" bearingthe tag (= adultcondor no. 9), wascaptured on EasterSunday, 19 April 1987. Captivebreeding: WhenAC-9 wasbrought into captivity,the captive flock toralled27 birds,including 10 captured asfree-flying individuals between 1967-87, 13 from eggslaid in thewild andhatched in captivity between1983-86, and 4 removedfrom thewild asnestlings between 1982-86. Fortuitously, the sex ratio wasabout even among the 27 birds.The flock wasabout evenly split betweentwo facilities, oneat theSan Diego Wild Animal Parkat SanPasqual, Califomia and the other at theLos Angeles Zoo.A third majorbreeding facility wasestablished in 1993at theWorld Center for Birdsof Preyin Boise,Idaho. Captivebreeding was first accomplishedin 1988,when a pairof wild-caughtcondors at theSan Diego Wild Animal Park produceda chick christened"Molloko" (Toone& Risser1988). Four youngwere produced in the following ,and chick productionof the captiveflock hasincreased annually(Fig. 2), albeitmore slowly thananticipated.

Figure 2. Captive breeding production 20 18 16 U 14 ,T2 f0 8 z 6 ii 4 2 0 _l__ 1988 1989 1990 1991 1992 1993 19941995 1996 1997 1998 ESFfim-wcBp I -L.L.zaa

DNA fingerprintinganalyses revealed that the captive condor flock included14 founders from "clans" three in whichthe between-group relatedness seemed to besignificantly less than the within- grouprelatedness (Geyer et al. 1993).The implicationfor condormanagers is thatthe founder gene contributionsof the subgroupsare moreimportant than the contributionsof individualfounders. Theseconsiderations weigh heavily in determiningpairs of thecaptive birds, which arecomputer- modelgenerated to maximizeheterozygosity in theprogeny. There is alreadyknown to be a lethal genewithin thepopulation that causes embryonic skeletal dysplasia (M. Wallacein litt.).Since the homozygousgene responsible is possessedby bothmembers of oneof themost productive pairs, it led to greatlyreduced egg success in thispair beforethe cause was determined. From 1988to 1998226 eggswere laid by the captiveflock. Of theseeggs, 167 Q3.9%) were fertile,and 141(84.4%) hatched. A high rateof infertilityamong first-time breeders and the effects ofthe aforementioneddeleterious gene account for mostofthe non-hatching.Ofsome interesthas beenthe high frequencyof egg laying in January,far earlierthan any recordedhistorically in the wild (Snyder& Hamber1985, Harvey er al. 1996),and breeding attempts by five-yearold birds

312 (onesuccessful) and even one four-year old bird. Previorisly,the age of first breedingin thespecies wasthought to be six years,when the full adultplumage is attained,or later(Koford 1953).

Reintroductionto the wilcl Followingthe capture of the last free-flyingCalifomia Condor in 1987,the CondorRecovery Teamrecommended the initiation of anexperimental release of thesimilarAndean Condor in southem Californiato refinecondor release techniques, test the criteria being used to selectrelease sites, field test condorrearing methods, identify potentialproblems peculiar to the Californiaenvironment, evaluateradiotelemetry packages and other equipment, and train a field crewfor anticipatedCalifornia Condorreleases (U.S. Fish and Wildlife Service 1996). Between 1988 to 1991,13 yearling female AndeanCondors were obtained from severalU.S. zoos and released from threedifferent sites in the vicinity of theSespe Condor Sanctuary area (Wallace l99l). Onlyone of the 13birds, an individual thatstruck a powerline, was lost during the experiment. Following their recapture, the female Andean Condorswere sent to Colombiaand Venezuela to be re-releasedin areaswhere males of the same agehad been released earlier (Lieberman et al. l99l). The sunogaterelease experiment wasjudged to be highly successful,and it helpedthe condor recovery progftrmme and habitat protection efforts to maintainmomentum during the period when Califomia Condors were not in the wild. In October1986 the Condor Recovery Team established several criteria to be satisfiedbefore a releaseof captive-bredCalifomia Condors could take place. These included having three actively breedingpairs of condors,three chicks behaviourally suitable for release,and retaining at leastfive offspringfrom eachbreeding pair contributing to therelease (U.S. Fish and Wildlife Service1996). In June1989, a provisionwas added to retaina minimumof sevenprogeny in captivityfor founders thatwere not reproductivelyactive. By 1991,several years earlier than originally predicted, two ofthe captive-producedchicks met thesecriteria. Although the aforementioned criteria had called for a minimumof threeeligible birds, it wasdecided to go aheadand seize the earliest opportunity to conducta release,using two young of the sameage to createa larger social group. The first releaseof captive-reared CaliforniaCondors occurred on 14January 1992in theSespe Condor Sanctuary, when "Xewe" (a "Chocuyens" female)and (a male)were setfree. The AndeanCondors were returned to captivity in September1992, officially marking the end of the experimentwith the speciesin California. Chocuyenswas found dead on 8 October1992, having died from ingestingethylene glycol, a principal componentof anti-freeze,which the bird probablyfound in a campgroundparking lot. Otherreleases soon followed, however, with six morecondors being released on I December 1992at thesame Sespe Condor Sanctuary site. By theend of I 998, I 6 separatereleases of captive- producedCalifomia Condors had been conducted at six localitiesin two states(Fig. 3, Tables 1,2). The Arizonareleases were finally initiatedin 1996by The PeregrineFund, eight yearsafter the recoveryteam had recommended such an action and following years ofcontentious public hearings, lawsuits,and negotiations between a myriadof govemmentalagencies and Native American tribes. TheCondor Recovery Team established a protocol for theselection ofrelease sites, and it includes suchcriteria as rcmoteness (but with sufficientaccess for thefield team),protection from mammalian predators,availability of suitableroost sitesand water,historical condor use, and properwind conditionsfor soaring.Not least,areas have been selected where local communiry support for releases exceedslocal opposition, although all of therclease sites are on landowned by theFederal govemment. Partof theoverall sratcgy in addingnew release sites has been to find localitiesthat are suffrciently disjunctfrom earliersites to allow thenew birdsto developtheir foragingand flight skills without interferencefrom olderbirds, yet closeenough to expectthat birds from separaterelease sites will eventuallymeet and, over time,incorporate the interstitialareas into their effectiveranges. Althoughtherc has been quite a bit of variabiliryin releaseprocedures, depending as much(or more)on politicalfactors as biologicalones, the main ingredientshave remained the same:Birds selectedfor releaseare usually raised together in a speciallydesignated pen at oneof the breeding facilitiesfor four-fivemonths (equivalent to their normalnestling period) or less,then helicoptered to therelease site, where they are held in anopen pen covered with nettingfor a periodranging from

3r.3 only two wecksto threemonths, during which the birds becomeacclimatised to their surroundings and further socialize with eachother. During this period of confinement,they are provided with unlimitedfood, using techniques to minimizethe associationof food with humans.The achralrelease is initiated by the removalof the nening from the enclosureduring nighttime, so that the birds are free to leavethe pen when they choose.Most birds havebeen released at aboutthe age when they would haveleft a nestsite normally, but membersof onecohort were not releaseduntil well into theirsecond year.

Figure 3. Condor ReleaseSites

Releasedbirds are provided with food (stillborndairy calves) at designatedfeeding sites, which are movedfrom time to time, as long as they continueto utilize suchsources. Although released birdshave tended to beginfeeding to someextent on itemsthey have found on theirown afterabout two yearsin thewild, all still dependmostly on thefood provided to themby biologists.This assures them of a contaminant-freefood source,and at leastin the early monthsfollowing release,it encouragesdlem to remainin the moresecure area near the releasesi!e. It is probablethat this "wlture specieswill alwaysutilize such restaurants"to someextent in thefuture, depending on the "narural" seasonalavailabiliry of food sources(including livestock). Thercleascd birds in Califomiahave not proved to beparticularly more mobile than the historical condorpopulation, but, as might be expected,they tend to wandermore widely as they grow older. As a pan of therccovery process, the original Condor Recovery Team designated critical habitat in variousponions of the vast(>2 million hectares)recent historical range of the species,and it has beengratifying to seethe released birds gradually begin to reoccupymany ofthese areas. Certain of theindividuals released at theVermilion Cliffs in Arizonahave routinely moved widely throughout theregion from southemUtah to thecentral portion of theGrand Canyon. One bird evenwandered

314 as far north as FlamingGorge, Wyoming, located on the ColoradoRiver, before returning to its releasesite, a distanceof 560km (S. Farrypers. comm.). Three others also moved north along the channelof the ColoradoRiver to GrandJunction, Colorado, where they spentseveral days in the vicinity of a parkvisitor's cente. They retumed to therelease site, nearly 400 km SW in lessthan a day (S. Farryop cit.).

Table 1. History of California Condor releases'1992'1998. Male Female Unknown SespeCondor Sanctuary, Ventura Co., California: 14 luwary 1992 I 1 2 December1992 2 4 Lion Canyon,Santa Barbara Co., California: a 8 December1993 J 8 February1995 I 29August 1995 I 19 November1996 2 ) 14 November1997 I 5 CastleCrags, San Luis ObispoCo., California l3 February1996 2 2 VentanaWilderness Area, Monterey County, Califomia: 19January 1997 J I 12December 1997 2 VermilionCliffs, CoconinoCo., Arizona: 12December 1996 2 4 l4May 1997 z 2 27 May 1997 3 2 20 November1997 4 4 23 November1998 HurricaneRidge, Coconino Co., Arizona: 18November 1998 3

Tabte 2. Surnmary of California condor releasercsults' January l92'December 1998 (compiled by Robert Mesta, U.S. Fish and Wildlife Service) Totalreleases t6 Totalcondors released 75 Totaldied 19 Lossesin wild: Collisionswith Powerlines ) Drowned 2 Causeunknown* 2 Poisoned Electrocutedon Powerline Killedby GoldenEagle * Onepossibly killed bYcoYote Disappearedand presumed dead: Recaptured& died in caPtivity: Malnutrition Cancer Gunshotwound Total recapturedfor behaviouralreasons 14 Totalsurviving in wild 42 Percentof releasedbirds still in wild 56

315 Early in the releaseprogramme, the birds were attractedto powerpoles as perchsites, and this led to severalfatal collisionswith powerlines. To addressthis problern MichaelWallace, I-osAngeles Zoo Curatorand also the CondorRecovery Team Leader, placed fake powerpoles in the penswhere the birdswere held prior to release,wiring themso thatbirds landing on thesestructures received mild electricalshocks. Judging from the fact that only one bird hasbeen lost subsequentlyfrom strikinga powerlineand that none have been recorded attempting to perchon powerpoles since this techniquewas initiated, the method seems to haveworked. The principalproblem for the releaseprogramme, and one not yet solved,is the tendencyfor manyof thebirds to be drawnto humanstructures and activities. This hasresulted in manybizarre episodesinvolving released condors and humans, much to the concernand frustration of the field crewsfollowing thebirds. In Californi4 somebirds have tended to landon theroofs of housesand other structures,where they have tom out insulation and causedother property damage,even to vehicles.In Arizon4 the birdshave been attracted to the campsof fishermenalong the Colorado River,where they have torn up tents,sleeping bags, and other gear. All of thesebehaviours seem to derivemore from the curiosityofthe condorsand their naturaltendency to play,rather than from hunger.While someepisodes are initially entertaining,they eventuallyannoy the generalpublic, especiallythe victims of condorvandalism, and they erode support for therecovery programme. In aneffort to discouragesuch behaviour and to makecondors warier, some of theearlier release "aversive cohortswere subjected to variousforms of conditioning,"or systematicharassment, by theLos Angeles Zoo staff.Some biologists have also strongly favoured parent rearing of thebirds to be releasedover puppet rearing, arguing that such individuals remain warier of humansin thewild (seediscussion in Wallace1994). Empirically, however, there seems to havebeen little conelation betweenthese earlier experiences in thezoo setting,and the eventual behaviour of birdsonce they arereleased. The pressureto releaseas many birds as possible, regardless of theirprovenance, in an effort to establishthe speciesin thewild rlssoon as possible has precluded rigorous field testingof rearingmethods. Regrettably, the tendencyof someindividuals to be constantlyattracted to anthropogenicsituations has forced condor biologists to recapture14 of them and removethem from thewild permanently(Table 2). The resultsof the releasesto theend of 1998are summarized in Table2. It is encouragingthat thereis no primarycause of mortality,especially now that the incidenceof collisionwith power lineshas been minimized, and it is alsoencouraging that only oneof thebirds has been shot. Only about.25vo(19 of75) ofthe releasedbirds have actually died, and including the birds recaptured and returnedto captivityfor behaviouralreasons as losses,survival of the releasedcondors has been 56%.Thisis remarkablygood for theearly, still experimentalyears of sucha programme,and, if this rateof successcan be maintainedor improved,it is likely that the condorcan be successfullyre- establishedin severalregions. Oneominous undercunent of therelease programme is thatseveral birds have been recaptured andbrought back into captivitybecause of elevatedlead levels in theirblood. They weresubjected to chelationtreatment to reducethe leadlevels, apparently recovered, and werere-released to the wild. In the opinionof theattending veterinarian (C. Stringfieldpers. comm.) the birds would not havesurvived without such treatment. One bird hassuffered two suchepisodes, yet is now backin thewild. In oneinstance, several condors were actually observed feeding on a deergutpile that was suspectedto containlead bullet fragments,and they weresoon trapped by the field team.Blood analysesconfirmed that the birds had alarmingly high lead levels. There is somehope that there will be a generalconversion to safernon-lead alloys for bulletsin the nearfuture, aided by a pushfrom theU.S. military establishment. Shon of theabandonment of lead as the bullet of choiceby hunrers andother shooters in opencountry, this factorremains a seriousthreat to wild condors. Additionalrelease sites being credibly considered or suggested(in descendingorder ofprobabiliry) arein BajaCalifomia Norte, Mexico (Siena San Pedro Martir), (Ladder Ranch), various ponionsof northernCalifornia, and San Diego County, Catifornia (Frg. 3). The CondorRecovery Teamhas formally recommendedthat releases be conducted at thefirst two sites.Other areas, including theChannel Islands off southernCalifomi4 theColumbia River Gorge of Oregon,and the Big Bend areaof WestTexas have also been suggested.

316 Presentsituation and future directions The condorrecovery programme is unusualamong rescue efforts of this in that both the captiveand wild populationsare beingbuilt up in parallel,and, so far, this strategyseems to be working.By theend of 1998there were 147condors, including 105 in captiviryand 42 in thewild (Fig. 1), a seven-foldincrease in thepopulation since the low pointof 1982.From this standpointthe recoveryprogramme has been remarkably successful. Some of theolder birds (now five yearsold) releasedin Califomiaane beginning to form pairbondsand engage in adolescentversions ofcourtship displays(R. Mestapers. comm.). Successful breeding by condorsin the wild is the lastremaining proximatemilestone, short of establishinga completely self-sustaining population, always the ultimate goalof therecovery programme. Accordingto the mostrecent revision of the CaliforniaCondor Recovery Plan (U.S. Fish and 'the Wildlife Service1996) minimumcriterion for reclassificationto threatenedis themaintenance of at leasttwo non-captivepopulations and one captive population. These populations (1) musteach numberat least150 individuals,(2) musteach contain at least15 breedingpairs and (3) be reproductivelyself-sustaining and have a positiverate of populationgrowth. In addition,the non- captivepopulations (4) mustbe spatially disjunct and non-interacting, and (5) mustcontain individuals descendedfrom eachofthe 14 founders." Themost serious remaining obstacles to achievingthese objectives, as I seeit, are:(l) Increasing therate ofcaptive breeding, which has never approached the level theoretically possible, (2) solving theproblem of condorattraction to humansand areas of humanactivities in thewild, and(3) solving theproblem of leaduse in therange of thecondor. Of probablefuture concem are (1) thepossibility of additionaldeleterious genes appearing in thepopulation, (2) habitatloss in someregions, and (3) politicalopposition to thecondor recovery programme and programmes in general, resultingin thewithdrawal of essentialgovemment support. Fundingthe programme has always been a problem.It coss The PeregrineFund about $2,000/ yearto maintaineach individual condor (W. Burnhampers. comm.), so merely maintaining a captive populationof at least150 individuals would cost$300,000 annually at thatrate. Indeed, some (e.g., Tom Cade)are questioningthe wisdomof keeping150 or morecondors in captivityindefinitely. Perhapseventually the captive flock couldbe dispersedto manyzoo facilitiesso thatthe financial burdencould be sharedby moreorganizations and more educational oppornrnities could be created, The field effortsare also quite expensive, since the programme is still at thelevel wherebiologists attemptto monitorthe whereaboutsof eachindividual. With time andmore birds in the wild, this intensivelevel of managementcan probably be reduced without harm to theoverall wild population. ln the meantime,the condorrecovery programme can only survivethrough the partnershipof privatesector organizations likeThe PeregrineFund and the Zoological Society ofSan Diegoand governmentalentities, especially the andthe U.S. Fish and Wildlife Service.Above all, it is the continuedsuccess of the programmethat maintainsbroad public interestand support, withoutwhich this specieswould havelittle hopeof survival.

ACKNOWLEDGEMENTS TheCalifomia Condor Recovery Programme is succeedingbecause of theefforts of hundredsof dedicatedgovernment officials, non-govemment organizations, biologists, and volunteers,and I owe mostof thema personaldebt. In specificregard to this update,the assistance of RobenMesta who faithfully maintainsthe scorecardof condorgains and losses, is especiallyappreciated.

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317 CARRIER' W.D. 1971.Habitat management plan for theCalifornia Condor. U.S. Forest Service. 5l pp. FINLEY' w.L. 1906.Life hisroryof thecalifomia condor.part I. Condorg:r3s-142. FINLEI W.L. lX)Ea. Life historyof theCalifomia Condor. pan Il. Condor10:5_10. FINLEI w.L. 1908b.Life historyof thecalifomia condor.partllJ. condor 10:59-65. FINLEY, W.L. 1910.Life hisroryof theCalifomia Condor. pan IV. Condorl2:5-l l. CEYER' CJ.' O.A.RYDER, L.G. CIIEMNICK & E.A. THOMPSON. l.993.Analysisof relatednessin the CaliforniaCondors from DNA fingerprints.Molecular Biol.Evot.10:571-5g9. HARRISON' E.N. & L.F. KIFF r9E0. Apparentreplacement clutch laid by wild CaliforniaConbr. Condor 82:351-352. HARVEY' N.C.' K.L. PRESTON & AJ' LEEIE 1996.Reproductive behaviour of captiveCalifomia Condors ( Gyttnogy ps califurnianus ). fuo Biol. I 5 : I I 5- I 25. KIFF'L.F. 1989.DDEandtheCaliforniaCondor (Gynvagyps catifomianus):theendof astory?pp.477480in B.-u. Meyburgand R.D. chancellor(cds.), Raptors in themdern world.wwGBp, Berlin. KIFF' LJ'.' D.3. PEAKALL & S.R. WILBUR 1979.Recent changes in Califomia Condoreggshells. Condor 81:166-172. KOFORD' C. 1953.The Califomia Condor.Nationat Audubon Society Research Repon no. J, New york. I 54 pp. KUEHLE& c.M., DJ. STERNER, D.s. JoNEs, R.L. USNIK & s. KASIELKE 1991.Report on "uptin" hatchesof califomia condorsGymnogyps californianus: 1983-1990. zno Biot.l0:65-6g KUEHLE& C.M. & PN.WIIITMAN lgSs.Artificial incubationofCalifomiaCordror califomiarus eggsremoved from thewild. bo Biol.7:123-132. LIEBERMAN' A.' J. WILEY' J.V. RODRIQT EZ & JJ0'/'.PAF;Zl9gl. First experimentalreinroduction of captiverearedAndean Condors Vulnr gryphusinroColombia, S.A. Pp. 129-l3l inAmericanAssoc. Zoological Parksand Aquariums Conf. Proc. l99l , AAZPA, Wheeling,West Virginia. MALLETTE' R.D. & J.C. BORNEMAN f966. First cooperativesunrey of the Califomia Condor.Calif. Fish Game52:185-203. MERETSKY'vJ. & N.F.R.SNYDER 1992. Rangeuseandmovements of CalifomiaCondors.Condorg4:313- 335. RICKLEFS' R.E. (Ed.) 1978.Report of the advisorypanel on the CalifomiaCondor. Narl. Audubon Soc. Consen'.Rept. no. 6. 27 pp. ROBINSON, C.S. 1940.Notes on the Califomia Condor,collected on l"os PadresNational Forest,Califomia. U.S.Forest Service, Santa Barbara, Califomia. 2l pp. SIBLEY' F.C. 1969.Effeca of 0reSespe Creek Project on theCalifomia Condor.U.S. Fish andWildlife Service, L,aurel,Maryland. l9 pp. SNYDE& N.F.R.1983. califomia condorrcproducrion, past and prcsent . Bird cowem. l:67-g6. SNYDER' N.tr'.R.& J.A. HAMBER 1985.Replacement-clutching and annual nesting of CaliforniaCondon. Condor87:374-378. SNYDER' N.F.R. & E.v. JOHNSON 1985.Photographic censusing of the 1982-1983California Condor population.Condor 8'l :l -13. SNYDER' N.F.R" R.R. RAMEY & F.C. SIBLEY 19E6.Nest-site biology of the CalifomiaCondor. Condor 88:228-241. SNYDER' N.nR. & H. SNYDER 1989.Biology andconservation of the CalifomiaCondor. Pp. 175-267in D.M. Power(ed.),Currcnt omithology.Yol.6, Plenum Press, New york. STEADMAN, D.W. & N.G. MILLER 19E7.Calilomia Condorassociated with spruce-pinewoodland in the [,atePleistocene of New York. QuatemaryResearch 28:415426. TOONE, W.D. & A.C. RISSER 198E.Captive managcment of theCalifomia CondorGymnogyps catifumianus. Int. bo Yearbo ok 27:50-58. TOONE, W.D. & M-P. WALLACE 1994.The extinctionin thewild andreinuoducrion of theCalifomia Condor (Gytrnogypscalifornianus). Pp. 411-419in P.J.s.olney, G.M. Mace, and A.T.c. Feismer(eds.), cteative conservation:interactive management of wild and captiveanimals. chapman & Hall, London. U.S.FISH AND WILDLIFE SERVICE 1996.California Condor recovery plan, third rcvision.USFWS, Portland, Oregon. WALLACE, M.P. 199f. Methodsand strategies for releasingCalifomia Condon to the wild. Pp. l2l-128 in AmericanAssoc.ZoologicalParksandAquariumsAnnualConf.Proc.lggl,AAZPA,Wheeling,WestVirginia. WALLACE' M.P. f 994. Contol of behaviouraldevelopment in the conrcxtof reintroductionprograms for birds.7no Biol. 13:49l -499. WILBUR' S.R. 1972.The food resourcesof the CalifomiaCondor. U.S. Fish andWildlife Servicc.Patuxent Wildlife ResearchCenter, Laurel, Maryland. l8 pp. WILBUR, S.R. 1974.Califomia Condor specimens in collections. Witson BulI.86:jl-72.

318 WILBUR, S.R. 1975.California Condor and molt asfield studyuds. Catif. FishGame 6l:144-148. WILBUR, S.R. 1976.Stanrs of theCalifomia Condor, 1972-1975. Am Bids 30:789-790. WILBUR, S.R.197E. The Califomia Condor, 1966-'16: a lookat its pastand futur€. Nonh American Faunano. 72. t36 pp. WILBUR, S.R.,W.D. CARRIER, J.C. BORNEMAN & R.D. MALLETTE 1972.Distribution and numbers of theCalifomia Condor, I 966-I 97l. Am.B irds26: 8 I 9-823. WILBUR, S.R., W.D. CARRJER & J.C. BORNEMAN 1974.Supplemental feeding program for Califomia Condors.J. Wildl. Manage.38:343-346. WILBUR, S.R,& J.A. JACKSON, JR. 19E3.Vulore biologyand management. Univ. Calif. Prcss,Berteley. 550pp. WILBUR. S.R.& L.n KIFF 1980.The CalifomiaCondor in BaiaCalifomia. Am Bids 34:856-859.

Lloyd Kiff The PeregrineFund 566West Flying Hawk Lane Boise,ID83709, U.S.A.

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