j. RaptorRes.

34(3):187-195
¸ 2000 The Raptor ResearchFoundation,Inc.

POPULATION FLUCTUATIONS OF THE HARRIS' HAWK

(PARABUTEO UNICINCTUS) AND ITS REAPPEARANCE
IN CALIFORNIA

MICHAEL A. PATTEN
Department
of Biology,University
of California,Riverside,
CA 92521 U.S.A.

RICHARD A. ERICKSON
LSA Associates,One Park Plaza, Suite 500, Irvine, CA 92714 U.S.A.

ABSTRACT.--TheHarris' Hawk (Parabuteo unicinctus)was consideredextirpated from California in the

mid-1960s.Most sightingsin the past 30 yearswere, therefore, consideredto be escapedor released
birds.The specieshasrecentlystagedan incursioninto southernCalifornia and northern BajaCalifornia
in the 1990s,involvingnearly 50 individualsand local breeding.This incursionwasapparentlyanother
in a long-term seriesof populationfluctuationsof the Harris' Hawk, each bringing large numbersto
the north and west of its establishedrange in Arizona and Baja California. Although first recorded at
the state border in the 1850s, the Harris' Hawk was not recorded as a breeder until an incursion in the
late 1910s and 1920sbrought hundreds to the state, including the first known breeders.Numbers
declinedagain in the 1940s,built up again in the 1950s,and thereafterdrasticallydeclinedto the point
of their absenceby the mid-1960s.Therefore, the recentincursionwasnot anomalousbut rather follows
historicalpatternsof occurrence,indicatingthat Californiais on the fringe of the natural range of the
Harris' Hawk,with emigrationbringingbirdsinto the stateand subsequent populationdecreases leading
again to "extirpation."
KEYWORDS: Harris'Hawk;Parabuteounicinctus;Baja California;California;
population
fluctuations.

Fluctuacionespoblacionalesde Parabuteo
unicinctus
y su reaparicitn en California
RESUMEN.--E1 gavilfinde harris (Parabuteounicinctus)fue consideradocomo extirpado de California a
mediadosde 1960. La mayoria de los avistamientosde los filtimos 30 aftosfueron consideradoscomo
avesescapadas o liberadas.La especieha incursionadoen el surde Californiay norte de BajaCalifornia
en los aftos90, incluyendounos 50 individuosy algunoseventosde reproduccitn locales.Estaincursitn
es aparentementeuna mils de las ocurridasa largo plazo por estaespecie.Cada una trayendograndes
nfimeros de individuos al norte y oeste de su rango establecidoen Arizona y Baja California. Aunque
por primera vez fue registradoen el borde del estadoen 1850, el gavilfinde harris no fue reportado
en reproduccitn hastasu incursitn en 1910 y 1920 con cientosde individuosincluyendolos primeros
registrosde reproduccitn. Los nfimerosde individuosdeclinaron otra vez en 1940, aumentaron en
1950, y declinaron drfisticamentehastaconsiderarlosausentesen 1960. Por lo tanto, la reciente incur-
sitn no es antmala, al contrario, siguelos patronesde ocurrencia indicando que California estfien el
limite del rango natural del gavilfinde harris, con su emigraci0n trayendo avesdentro del estadoy la
subsecuentedeclinacitn la cual conllevaa su extirpacitn.
[Traducci6n de Ctsar Mfirquez]

The Harris' Hawk (Parabuteounidnctus) ranges
from the southwestern United States
through Central America to central Chile and cen-
tral Argentina, with a geographicallydisjunctpop-
ulation on the Baja California peninsula. In the
United States, it occurs from southern Arizona,
southeastern New Mexico, and central Texas south-
ward (Fig. 1; American Ornithologists' Union
1998). Its range in Arizona, New Mexico, and Tex-
as has been expandingnorthwardin recent years
southward (Bednarz et al. 1988, Bednarz 1995, Dawson 1998).
In California, the Harris' Hawk wasfound formerly
throughout the lower Colorado River Valley and in
the Imperial Valleysouthof the Salton Sea (Grin-
nell and Miller 1944). By the mid-1950s,it wasex-
tirpated from California as a breeder (Remsen
1978, Walton et al. 1988), with the last definite wild
bird recorded north of Blythe on 28 November

187
188 PATTENAND ERICKSON VOL. 34, NO. 3

LakeTahoe

Colorad/e•'River

California

Arizona
Salton Sea

Figure1. The northwestern

portionof the currentrange(theshadedarea)of the Harris'Hawk(Parabuteoun2-
cznctus),
modifiedfromBednarz(1995)andDawson(1998).The dashedline signifies
the formerwesterly
limitsof
its rangein southwestern
Arizonaand southeastern
California.
SEPTEMBER2000 HArems' HAwI• IN CALIFOm'4IA 189

1964 (Garrett and Dunn 1981, Rosenberg et al. mate of the magnitude of the incursion and to examine
1991). its geographicextent.
A reintroduction project for the Harris' Hawk THE 1994 INCURSION
wasinitiated in California in 1979 by variousstate,
federal, and private groups (Stewart1979, 1982, Despitean annual "background"escape/release
Walton et al. 1988). Eight birds were releasedthat rate throughout California of >3 Harris' Hawks
year and severalmore were releasedeachyear un- (Bloom pers. comm., Walton pers. comm.), a dif-
til 1989, for a total of 222 releases (Linthicum ferent phenomenon began 15 April 1994, whenJ.
1989, Linthicum pers. comm.). The first pair nest- Rudley,P. Jorgensen,and M. Jorgensenobserved
ed successfullyin 1983, three pairs bred success- three adults together in Borrego Valley. Between
fully in 1986 (Walton et al. 1988, Rosenberget al. 1994-96, at least 34 individuals had been found in
1991), and five nested in 1989 (Linthicum 1989, southern California (Table 1; McCaskie 1995). The
Bednarz 1995). However, it is unlikely that this largest groups of birds consistedof at least eight
population is viable, as birds are now infrequently individuals in the Borrego Springs region of the
noted (Rosenberget al. 1991, Patten pers. obs.). Anza-Borrego Desert and up to five individuals
Sincethe mid-1960s,virtuallyall recent records both at the former George Air Force Base near
of the Harris' Hawk in California are of birds con- Victorville and at Boulevard (Table 1). During this
sidered to have escapedfrom falconers(Garrett apparentlynatural incursion (Bednarz 1995, Mas-
and Dunn 1981, Unitt 1984). In some cases,birds sey1997,Walton pers.comm.), Harris' Hawkswere
have been observedwith jessesand clearly came found north of their historical range as far asVic-
from this source. In other cases,there appearsto torville in the Mojave Desert, with scatteredindi-
be some tendency for natural occurrence such as vidualsreported around the Salton Sea and else-
sightingsof immatures along the Colorado River where (Fig. 2). Additional birds in cismontane
near Blythe in September (Roberson 1980) and valleysat Riversideand in centralSanDiego Coun-
December1978 (Rosenberget al. 1991) and at the ty may or may not have been naturally occurring,
south end of the Salton Sea on 25 June 1989 with individualsfar westin coastalOrange County
(McCaskie 1989). Nevertheless,records of individ- and in the Antelope Valley being particularlysus-
ual birds are perhapsalwayssuspectgiven that the pect given the apparent geographicextent of the
speciesremains popular with falconersand reha- influx (Fig. 2). Indeed, the Orange County bird
bilitated birds are occasionallyreleased,as were a showed signsof being in captivity (Bloom pers.
few around the Salton Sea in the 1970s and 1980s comm., Danielspers. comm.).
This influx into southern California was con-
(Walton et al. 1988). Herein, we document a major
comitant with at least 22 individuals well north of
increasein sightingsbeginning in April 1994 that
wasapparentlya natural influx involvingnearly50 the species'normal range in northern Baja Cali-
individuals throughout southern California and fornia (Table 2; Radamakerpers. comm., Wurster
northern Baja California. Further, we hypothesize pers.comm.) and in adjacentnorthwesternSonora
that such incursions are the rule rather than the (Russell and Monson 1998). During this period,
exceptionfor the occurrenceof this speciesin Cal- Harris' Hawks bred in California at Borrego
ifornia. Springs (Massey1997), Boulevard (Unitt pers.
comm.) and Laguna Dam (McCaskie1996, Massey
METHODS 1997), and in northern BajaCaliforniaat Valle San
For the recent incursion,we gatheredrecordsand doc- Telmo (Bloom pers. comm.). Small numbershave
umentation from various field observers (see Acknowl- persistedin BorregoValleyas recentlyas 7 March
edgments)and from files of the California Bird Records 1999 (Jorgensenpers. comm.) and in Valle San
Committee. All specificdata gathered are on file at the Telmo on 31 January 1999 (Patten pers. obs.).
Western Foundation of Vertebrate Zoology (WFVZ),
Camarillo, California U.S.A. Recent and historical data HISTORICAL TRENDS AND CURRENT STATUS
were gathered from seasonalreports for the Southern
Pacific CoastRegion publishedin FieldNotes(now North The historical distribution of the Harris' Hawk
AmericanBirds),ChristmasBird Countsand specimensat in California is not clear. The specieswas first re-
the San Diego Natural History Museum (SDNHM), San
Diego, California U.S.A., National Museum of Natural corded along the Colorado River on the Arizona
History (USNM), Washington, D.C.U.S.A. and WFVZ. side in February 1854 (Kennerly 1859, Swarth
We tabulatedand mapped thesedata to obtain an esti- 1914), but Elliot Couesnever recorded the species
190 PATTEN AND ERICKSON VOL. 34, NO. 3

Table 1. California records of the Harris' Hawk (Parabuteounicinctus)from spring 1994 through winter 1996-97
(Fig. 2), arranged chronologically.Birds nestedat Boulevard(1994) and Laguna Dam (1996), and exhibited nesting
behavior (copulations,carrying sticksand food) at Borrego Valley (1994-95), with immatures observedin 1995. Data
are on file with the CBRC.

DATE (S) LOCATION MAXIMUM

15 April 1994-January1999+ San Diego County;Borrego Valley

1 June 1994-31 October 1995 San Diego County; Boulevard
26 November1994-29January 1995 San Diego County; Santee
? December 1994 RiversideCounty;Blythe
7-18 December 1994 Imperial County; Westmorland
10-12 December 1994 Orange County; Irvine
31 December 1994-29 January 1995 RiversideCounty; n. end Salton Sea
2-21 January 1995 San Bernardino County;Victorville
27 June-23 July 1995 RiversideCounty; Riverside
6-10 July 1995 San Diego County;Carrizo Canyon
3 April 1996 San Diego County;Escondido
5 April 1996 San Bernardino County;Vidal Wash
25 March-30 December 1996 Los AngelesCounty;Antelope Valley
29 March-April 1996 Imperial County; Laguna Dam
31 March 1996 San Diego County;Spring Valley

during his extensivesurveysof the lower Colorado first for the lower ColoradoRiverValley,asCooper
River Valleyin the 1870sand 1880s.It apparently (1870) made no specificmentionof encountering
wasnot recorded at the Colorado River again until this species.By the mid-1940s,it was "locally com-
August (Stephens1903) and December 1902 (Wil- mon" in California (Grinnell and Miller 1944).
der 1916), when individuals were noted on both There were occasionalrecords of large numbers,
sidesof the river. Two specimenscollected in the although reports of 400-500 between Calexicoand
Rio Colorado delta of northeasternBaja California Heber, Imperial County, on 22 October 1920
bracket these records,a male along the Rio Alamo (Chambers 1921) and 250 near Calexico on 28 Au-
southwestof Pilot Knob on 7 April 1894 (USNM gust 1923 (Chambers 1924) are perhapsbest con-
133726)and a subadultalongthe Rio Hardy on 16 sidered tentative given that the speciesdoes not
April 1905 (USNM 197921).Thus,a fewbirdswere travel in large flocks (Bednarz pers. comm.).
in the area from the mid-1890s until the early The few recordsof the Harris' Hawk prior to the
1900s; however,following the few noted in 1902, late 1910s may have involved occasionalstraysto
the species again went unrecorded in California the westof its establishedrange given its apparent
for a decade. For example, Joseph Grinnell and spread into western Arizona. This species"has a
party did not find the Harris' Hawk during their reputation for being somewhat nomadic" (Bed-
exhaustivesurveyof the length of the lower Colo- narz et al. 1988), with straysbeing recorded north
rado River Valley 14 February-15 May 1910 (Grin- to Ohio (Earl 1918) and Kansas(Bunker 1919,Sny-
nell 1914). Given the paucity of records through der 1919), east to Louisiana (Coombs 1892), and
this period, Grinnell (1915) consideredthe species west to Utah and Nevada (Palmer 1988). The spe-
to be only a "summer visitant" to the Colorado cies has bred opportunisticallyeven at the fringes
River, basedsolelyon Stephens' (1903) records. of its range, including occasionalnestingin Kansas
The Harris' Hawk was not documented as a (Parmaleeand Stephens1964) and Louisiana(Bai-
breeder in southeastern California until the late ley and Wright 1931). On a smaller scale,groups
1910s,with the first evidencefound on 25July 1916 of Harris' Hawks have been documented to invade
(Wiley 1917, Bancroft1920); the first breedingev- and subsequentlynest in severalregionsin south-
idence for northeastern Baja California was con- ern Arizona east of its normal range (Bednarz
comitant (WFVZ 83655). Thus, it is probable that 1995). Furthermore, there are two historical re-
the specieshad only recently expanded its range cords of the Harris' Hawk for coastalSan Diego
into the area. Thesebreeding recordswere alsothe County,California: one collectedat MissionValley
SEPTEMBER 2000 H2xmus' HAWK IN CALIFORNIA 191

Colorado
River

Los

San
Dieg
( . ß

ß 1 individual
Gulf of
ß 2-3 individuals
California
ß 4+ individuals
Approximate northern
•' limit
ofrange
(Wilbur
1987)

Figure 2. Recordsof the Harris' Hawk (Parabuteounicinctus)in southern California and northern Baja California
since April 1994. The double line identifies the apparent geographic limits of the 1994 incursion. Recordswithin
this line are best considered naturally occurring, records falling on the line are debatable, and records well to the
west or east are problematic (see text). Shading representsurban/agricultural areas.

on 17 November 1912 (Grey 1913, SDNHM 1842; 3 December 1902 (Wilder 1916). These numbers
the second record for California) and one ob- followed many decadesof no recordsfor the lower
served at Oceanside from 1-6 November 1942 ColoradoRiverValley.Elliot Couesnever recorded
(Kent 1944). the speciesduring manyyearsof work at Yumaand
Although there are no long-term population the specieswas apparently absent again by 1910
censusdata, availabledata suggestthat the Harris' (Grinnell 1914). This dearth was followed by an
Hawk has undergone four influxes into California influx in the late 1910s and early 1920s that was
during the 20th century (Fig. 3). The first major apparently an order of magnitude larger than the
northwesterlyexpansionwas around the turn of incursionsof 1902 or 1994, asevidencedby reports
the century when "10-20 [were] in the air at a of large numbers in the Imperial Valley (Chambers
time" along the lower Colorado River between 1- 1921, 1924), a new westerlyoutpostfor the species.
192 P^TT}•NANDEPdCICSON VOI•. 34, No. 3

Table2. BajaCaliforniarecordsof the Harris' Hawk (Parabuteo unicinctus)

north of itsnormalrangefromfall 1993
throughfall 1995 (Fig. 2). Recordsare arrangedchronologically.Breedinghasbeen documentedin the ValleSan
Telmo,where at leastone wasstill presentas of 31 January1999 (Pattenpers.obs.);in addition,an adultwasstill
presentat Leyesde Reformaon 7 November1998(Erickson pers.obs.).Forthesakeof completenesswealsoinclude
one record from this timeframefor the Rio Coloradoin extremenorthwesternSonora(Russelland Monson1998).
See Methods for data sources (now on file at WFVZ).

DaTE (S) LOCATION MAXIMUM

5 September 1993-fall 1995+ Valle San Telmo 2

20 February 1994 20 km s. of San Quintin 3
9 April 1994 E1 Doctor (Sonora) 1
10 April 1994 Valle las Palmas 1
23 April 1994 Laguna Hanson 1
21 May 1994 Campo Christiano 3
21 May 1994-5 March 1995 Valle San Matias 5
30 May 1994 Ejido Sinaloa 1
13 November 1994-fall 1995+ Leyesde Reforma 2
13 November 1994 Valle Trinidad 1
26 March 1995 La Rumorosa 1
10 November 1995 H6roes de la Independencia 1

200

__ 100s in ImperialValley and

alongColoradoRiverby early1920s

flocks of 10-20 and

first California record in 1902 numbersbuildup to
•oo flocksof 30 by 1950

first record for

Colorado River
(Arizon• in 1854 over 30 individuals
unrecordedby
Grinnell in 1910 stage influxin
50
unrecordedby
Coues in late 1800s extirpated
'1964

0
1840 1860 1880 1900 1920 1940 1960 1980 2000
YEAR
Ftgure 3. Timeline of the fluctuatingoccurrenceof the Harris' Hawk (Parabuteo
unicinctus)in Californiaand the
lower Colorado River Valley.
SEPTEMBER 2000 HARMS' HAWK IN CALIFORNIA 193

There was apparently another influx of Harris' nia and northern Baja California, it explains nei-
Hawks into California during the late 1940s and ther the magnitude nor the rapidity of the 1994
early 1950s (Bednarz 1995). During this period event. Instead, our investigation supports the hy-
numbers again built to double digits (e.g., 30 at pothesisthat the speciesundergoesperiodic pop-
HavasuNational Wildlife Refugeon 27 December ulation fluctuationsthat result in rapid range ex-
1950) following two decadesof only a few individ- pansionsfollowed by adventitiousbreeding and,
uals being regularly recorded (Rosenberg et al. typically, slow range contraction (Millsap 1981,
1991). Bednarzet al. 1988). Each expansion-retraction cy-
Two major factorshavebeen implicatedin caus- cle differs in magnitude and may bring individuals
ing the extirpation of this speciesin California. into areaswhere they had not been recorded pre-
First, the sport of falconry had an upsurgein pop- viously,such as the Mojave Desert. The breeding
ularity in the 1950s,and the Harris' Hawk wasand biologyof the Harris' Hawk promotesrapid expan-
remains a favored bird (White 1988, 1989). Rem- sion in numbers when conditions allow. While
sen (1978) suggestedthat nestlingswere taken in most nest in spring (March-June), it is able to
California until the population wascompletelyde- breed year-roundin temperate-climatedesert hab-
pleted, but there is no evidencethat falconersever itats in North America and may produce second
harvested Harris' Hawks in California (Walton and third clutches (Bednarz 1987, 1995). We be-
pers. comm.). Instead, birds were harvestedin Ar- lieve that most birds observed during the 1994
izona and Texas and most flown now are from cap- event originated in northern Baja California,
tive breeding. Shootingwasundoubtedlycommon where rainfall totals at Ensenada were 140% of av-
and may alsohave contributedsomewhatto its de- erage during winter 1991-92 and 168% in 1992-
cline. 93, but were 90% averagein 1993-94 (Mellink
Second, habitat loss along the Colorado River pers. comm.). Perhapstwo yearsof favorable con-
from agricultural clearing and water diversionpro- ditions allowed for an increase in numbers suffi-
jectswasextensivebetween1930-60 (Whaley1986) cient to send birds far afield during lessfavorable
and similarly occurred during this time in the Im- conditions in 1993-94, although causesare likely
perial Valley (Steere 1952). In addition to direct more complex.
clearing, erratic water levelsled to periodic flood- In summary,the availableevidencesuggeststhat
ing and desiccation,killing mostsuitablenesttrees. the Harris' Hawk has alwaysbeen on the fringe of
Throughout its range, the Harris' Hawk inhabits its natural range in California, with occasionalir-
savannah-typehabitatsin arid and semiarid areas, ruptive occurrencesinto the state every few de-
including open woodland, open scrub, mesquite cades.Many stayto breed or linger for significant
(Prosopis spp.) woodland, and riparian woodland periods, but eventually numbers decline as ap-
bordering open spaces.Trees, especiallycotton- peared to be happening alreadyin the wake of the
woods (Populusspp.), or large cacti such as the sa- influx of 1994. As noted above for Louisiana and
guaro (Carnegieagigantea),are used for nesting; Kansas, this species is capable of adventitious
however,this specieswill useutility polesand other breeding following a lengthy dispersal.Further-
artificial structures.In recentyears,thisspecieshas more, the more general pattern of range expan-
been steadilyincreasingits range in southeastern sion and contraction has been documented re-
Arizona (Bednarz 1995, Dawson 1998) where it has peatedly in Arizona and New Mexico (Bednarz
become more tolerant of human settlements. In-
1995, Dawson1998), althoughit is perhapsmore
deed, Dawson (1998) noted that "the willingness dramatic in California given that the speciesmay
of Harris' Hawk to nest in urban areas offers some
not occur in the statefor yearsat a time.
hope of mitigating habitat loss to development,"
although post-fledgingsurvivalof suchbirdsis low ACKNOWLEDGMENTS
(Bednarz pers. comm.). We are indebted to P.D. Jorgensen, K.A. Radamaker,
Perhapssuch tolerancefor urban settingis a re- R. Theriault, and T.E. Wurster for graciously providing
cent advent, for if it were alwaysthe casethen the extensive summaries of field observations of the Harris'
Harris' Hawk may have persistedin many areas Hawk in southern California and northern Baja Califor-
nia. P.H. Bloom, B.E. Daniels, G. McCaskie, E. Mellink,
where it formerly occurred.However,we feel that J. Linthicum, P. Unitt, and B.J. Walton supplied unpub-
whereasthis urban tolerancemay play somesmall lished information and shared their views of this incur-
role in the recent incursion into southern Califor- sion. California Bird Records Committee files (housed at
194 P^TTE•q AND EP,ICKSO•q VOL. 34, NO. 3

WFVZ) were an invaluable resource for information ical engineers, in 1853 and 1854: No. 3, report on
about recent occurrences. R. Corado and J. Fisher birds collected on the route. Pac. RailroadRep.10:19-
(WFVZ), J.P. Dean and C. Ludwig (USNM), and P. Unitt 35.
(SDNHM) provideddata about and/or allowedaccessto KENT, W.A. 1944. Rare birds seen in southern California.
specimensin their care. We thank J.C. Bednarz, P.H. Condor 46:129-130.
Bloom, BJ. Walton, and S.R. Wilbur for their careful re-
viewsof the manuscript. LINTHICUM,J. [ED.]. 1989. Peregrine Falcon monitoring,
nestmanagment,hack site,and cross-fostering efforts,
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Received30 October 1999; accepted14 March 2000