Northern Goshawk diet in Minnesota: An analysis using video recording systems

2005·5.8 MB·English

by  B L Smithers

,
Raptor Res. 39(3):264—273
J.
© 2005 The Raptor Research Foundation, Inc.
NORTHERN GOSHAWK DIET IN MINNESOTA:
AN ANALYSIS USING VIDEO RECORDING SYSTEMS
Brett L. Smithers^
Department ofRange, Wildlife and Fisheries Management, Texas Tech University,
Lubbock, TX 79409-2120 US.A.
Clint W. Boat
us. Geological Survey, Texas CooperativeFish and Wildlife Research Unit, Texas Tech University,
Lubbock, TX 79409-2120 U.S.A.
David E. Andersen
U.S. Geological Survey, Minnesota CooperativeFish and Wildlife Research Unit, University ofMinnesota,
MN
St. Paul, 55108-6124 U.S.A.
—
Abstract. ^We used video-recording systems to collect diet information at 13 Northern Goshawk (Ac-
cipiter gentilis) nests in Minnesota during the 2000, 2001, and 2002 breeding seasons. We collected 4871
hr of video footage, from which 652 prey deliveries were recorded. The m^ority of prey deliveries
identified were mammals (62%), whereas birds (38%) composed a smaller proportion ofdiet. Mammals
accounted for 61% of biomass delivered, and avian prey items accounted for 39% of prey biomass.
Sciurids and leporids accounted for 70% of the identified prey. Red squirrel Tamiasciurus hudsonicus
( )
eastern chipmunk Tamias striatus) and snowshoe hare (Lepus americanus) were the dominantmammals
( ,
identified in the diet, while American Crow {Corvus brachyrhynchos) and Ruffed Grouse {Bonasaumbellus)
were the dominant avian prey delivered to nests. On average, breeding goshawks delivered 2.12 prey
items/d, and each delivery averaged 275 g for a total of 551 g delivered/d. However, daily {P< 0.001)
and hourly {P — 0.01) delivery rates varied among nests. Delivery rates (P = 0.01) and biomassdelivered
{P = 0.038) increased with brood size. Diversity and equitability of prey used was similar among nests
and was low throughout the study area, most likely due to the dominance of red squirrel in the diet.
EIey Words: Northern Goshawk, Accipiter gentilis; diet, Minnesota', prey diversity, red squirrel, Tamiasciurus
hudsonicus.
DIETA DE ACCIPITER GENTILIS EN MINNESOTA: UN ANALISIS BASADO EN SISTEMAS DE GRA-
BACION EN VIDEO
—
Resumen. Empleamos sistemas de grabacion en video para recolectar informacion sobre la dieta de
Accipitergentilis en 13 nidos ubicados en Minnesota durante las temporadas reproductivas de 2000, 2001
y 2002. Obtuvimos 4871 hr de grabacion, a partir de las cuales registramos 652 entregas de presas. La
mayoria de las presas entregadas que identificamos fueron mamiferos (62%), mientras que las aves
(38%) representaron una proporcion menor de la dieta. Los mamiferos y las aves representaron el 61%
y el 39% de la biomasa entregada, respectivamente. Los sciuridos y leporidos representaron el 70% de
las presas identificadas. Los mamiferos predominantes identificados en la dieta fueron Tamiasciurus
hudsonicus, Tamias striatus y Lepus americanus, mientras que las aves llevadas a los nidos predominante-
mente fueron Corvus brachyrhynchos y Bonasa umbellus. En promedio, los individuos nidificantes entre-
garon 2.12 presas/d, y cada entrega tuvo un promedio de 275 g, para un total de 551 g entregados/d.
Sin embargo, las tasas diarias {P < 0.001) y horarias {P = 0.01) de entrega de presas variaron entre
nidos. Las tasas de entrega (P = 0.01) y la biomasa entregada (P = 0.038) incrementaron con el tamano
de la nidada. La diversidad y equitabilidad de las presas consumidas fueron similares entre nidosybajas
a traves del area de estudio, probablemente debido a la dominancia de T. hudsonicus en la dieta.
[Traduccion del equipo editorial]
^ Present address and corresponding author: P.O. Box 1363, Meeker, CO 81641 U.S.A.; email: brett^[email protected]
264
. .
September 2005 Biology 265
The Northern Goshawk {Accipiter gentilis) is a eastern Minnesota (46°50'N, 92°11'W) as described by
large, forest-dwelling raptor generally associated Boal et al. (2001) and Roberson (2001; Fig. 1). The study
area elevation ranged from ca. 200-400 m. Mean summer
with mature deciduous, coniferous, or mixed for- and winter temperatures were 18°C and —11°C, respec-
ests (e.g., Bright-Smith and Mannan 1994, Siders tively, and maximum and minimum temperature records
and Kennedy 1996, Beier and Drennan 1997, for the region were 40°C and —46°C, respectively (Daniel
Squires and Reynolds 1997). Goshawk research in and Sullivan 1981). Annual precipitation averaged 60-70
cm. The study area was dominated by pine, mixed-hard-
North America has been conducted primarily in
wood, boreal, and second-growth forests with wetland
the western halfofthe continent (Boal et al. 2003). community types interspersed among forest stands (Tes-
Consequently, there is little published literature ter 1995).
describing ecology of the species in the Western Goshawk Nests. Nests included in this study were con-
Great Lakes Region (WGLR) of North America, sidered as sampling units and were selected from all
known occupied nests in the study area (Boal et al. 2001)
where it is currently listed as a Migratory Non- With the exception ofone nest, where few data were col-
game Bird of Management Concern by the U.S. lected during 2000, diet information was not collected at
Fish and Wildlife Service (Region 3) and as a sen- any nest for more than one breeding season. Nests were
sitive species by the U.S. Forest Service (Region 9) selected randomly within the constraints of accessibility
and to include different land ownerships. Thus, our sam-
due to loss of habitat (Reynolds et al. 1992). ple is not truly random and may not be representative of
Depending on region, season, and availability, the goshawk population of our study area. However, to
goshawks capture a wide variety of prey and are examine the applicability ofour diet data to the goshawk
considered prey generalists (Squires and Reynolds population as a whole, we examined prey diversity and
1997, Squires and Kennedy 2005). Although breed- overlap among nests. High overlap and low diversity
would suggest prey use was similar among goshawk pairs
ing-season diet composition has been studied for
and that our data were representative of the population
many populations (e.g., Meng 1959, Grzybowski in general.
and Eaton 1976, Boal and Mannan 1994, Younk Video Recording. We used VHS (Model SL 800, Se-
and Bechard 1994, Lewis 2001), site-specific studies curity Labs®, Noblesville, IN U.S.A.) and 8-mm video re-
of diet are necessary for developing management cording systems (Sony® Model M-350, Fuhrman Diversi-
TX
fied, Inc., Seabrook, U.S.A.) with color or
strategies for goshawk populations at regional and black-and-white cameras (Model CCM-660W, Clover Elec-
local levels (e.g., Reynolds et al. 1992). A number tronics®, Los Alamitos, CA U.S.A.). Cameras were in-
m
of records exist of prey items collected opportu- stalled on nest trees within 0.6 of the nest or, for cam-
m
nistically at goshawk nests in the WGLR (Eng and eras with zoom lenses, on an adjacent tree up to 9
Gullion 1962, Apfelbaum and Haney 1984, Martell from the nest. Vidmeo recorders were placed in weather-
proof cases ca. 30 from the base of each camera tree.
and Dick 1996), but these reports are anecdotal Coaxial-video cables were used to convey power to and
and provide a prey list rather than a quantitative transmit images from the cameras. Recorders were pro-
assessment of food habits (Roberson et al. 2003) grammed to record from 0530-2100 H (15.5 hr of foot-
Methods used in goshawk food habits research age) at the 48-hr (1.3 frames/sec) or the 72-hr (0 8
frames/sec) setting to optimize the amount of tape used
have included indirect (i.e., identification of prey per sampling session and battery life. We replaced tapes
remains or contents of regurgitated pellets) and and batteries every 3-4 d.
direct observations of prey deliveries to nests Prey Identification. To identify prey delivered to nests,
(Meng 1959, Grzybowski and Eaton 1976, Bosa- we reviewed video footage until a prey delivery occurred,
kowski and Smith 1992, Boal and Mannan 1994). then advanced frame by frame and freeze-framed to fa-
cilitate prey identification. We identified avian and mam-
Indirect methods of assessing raptor diet can lead malian prey by morphological features and developed a
to biased results (e.g., Bielefeldt et al. 1992), list of prey species delivered by goshawks to all nests (Ta-
whereas direct methods should provide the least- ble 1). Goshawks may cache prey and retrieve cached
biased results (Collopy 1983, Marti 1987, Boal and prey items (Boal and Mannan 1994), which could bias
estimates ofdelivery rates and proportional use ofspecies
Mannan 1994). During the breeding seasons of in the diets. We attempted to identify cached prey on
2000-02, we used videography as a modified meth- basis of a successive, iterative process that included com-
od of direct observation of prey deliveries to ex- paring preyitems using flesh color, pelage orfeathercon-
amine diet of Northern Goshawks in northern dition, and time ofdelivery from review ofvideo footage,
and then remove those items thought to be cached from
Minnesota.
analysis.
Age and Biomass Estimation. We assigned avian prey
Methods
to age categories (e.g., adult,juvenile, or nestling) based
Study Area. The study area was located in the Lauren- on plumage (e.g., feathers and down) and amount of
tian Mixed-Forest Province of north-central and north- sheathing on flight feathers (Reynolds and Meslow
266 Smithers et al. VoL. 39, No. 3
Figure 1. Study area and distribution of Northern Goshawk nests in Minnesota where food habits information was
collected during the 2000-02 breeding seasons. The three-letter designations indicate individual nests. Breeding
DTK
season diet information collected at the breeding area was omitted from all analyses because of nest failure.
Breeding areas with similar prey composition are indicated with the same superscripts. Superscripts indicate cluster
number
(see Fig. 2).
1984). We categorized mammalian prey as adults or ju- mates were based on published information on mam-
veniles based on size (Bielefeldt et al. 1992). Because of malian and avian species occurring in the study area
difficulty in estimating age of small mammals, we consid- (Burt and Grossenheider 1980, Jones and Birney 1988,
ered all mammals smaller than chipmunks to be adults. Dunning 1993, Dunn and Garrett 1997, Dunn 1999, Sib-
Biomass for partial prey items was calculated using the ley 2000). We calculated mass for nestlings following
proportion of prey delivered to nests, and proportions Bielefeldt et al. (1992) using 100% of the adult mass for
were estimated qualitatively (e.g., 50% of adult size). warbler-sized species, 65% of the adult mass for robin
We
estimated biomass for prey identified to family, ge- andjay-sized species, and 55% ofthe adult mass for large
nus, or species and used the mean mass of both sexes birds such as grouse. We calculated mass of juvenile
(Reynolds and Meslow 1984, Lewis 2001). Biomass esti- red squirrel (Tamiasciurus hudsonicus), eastern chipmunk
September 2005 Biology 267
Table 1. Number, percent occurrence, and biomass of mammalian and avian prey delivered to Northern Goshawk
nests {N = 13) in Minnesota, 2000—02. Values represent pooled number of prey identified at nests during the 2000,
2001, and 2002 breeding seasons.
Biomass
Prey Category Common Name N Percent Percent
(g)
Mammals
Tamiasdurus hudsonicus red squirrel 202 31.0 38046 23.6
Tamias striatus eastern chipmunk 95 14.6 8108 5.0
Lepus americanus snowshoe hare 31 4.8 41027 25.5
Sylvilagusfloridanus eastern cottontail 7 1.1 7654 4.8
Sdurus carolinensis eastern gray squirrel 3 0.5 1679 1.0
Peromyscus spp. 2 0.3 47 0.0
Family: Muridae 1 0.2 18 0.0
Mustelafrenata long-tailed weasel 1 0.2 210 0.1
Unknown mammal (MSC1)“ 8 1.2 186 0.1
Unknown mammal (MSC2)® 9 1.4 1720 1.1
Birds
Corvus brachyrhynchos American Crow 37 5.7 14515 9.0
Bonasa umbellus Ruffed Grouse 33 5.1 18448 11.5
Aythya spp. diving duck 12 1.8 11360 7.1
Cyanodtta cristata BlueJay 8 1.2 664 0.4
Fulica americana American Coot 6 0.9 3338 2.1
Turdus migratorius American Robin 3 0.5 205 0.1
Common
Quiscalus quiscula Crackle 3 0.5 341 0.2
Family: Icteridae blackbird 3 0.5 189 0.1
Picoides spp. woodpecker 3 0.5 199 0.1
Dryocopuspileatus Pileated Woodpecker 3 0.5 861 0.5
Unknown duckling 4 0.6 400 0.2
Butorides virescens Green Heron 2 0.3 420 0.3
Perisoreus canadensis GrayJay 2 0.3 142 0.1
Agelaius phoeniceus Red-winged Blackbird 2 0.3 105 0.1
Strix varia Barred Owl 1 0.2 394 0.2
Buteo platypterus Broad-winged Hawk 1 0.2 455 0.3
Genus: Calidris 1 0.2 73 0.0
Bucephala clangula Common Goldeneye 1 0.2 900 0.6
Accipiter cooperii Cooper’s Hawk 1 0.2 439 0.3
Gallus spp. domestic chicken*’ 1 0.2
Coccothraustes vespertinus Evening Grosbeak 1 0.2 59 0.0
Pipilo erythrophthalmus Eastern Towhee 1 0.2 41 0.0
Genus: Euphagus 1 0.2 63 0.0
Acdpitergentilis Northern Goshawk 1 0.2 820 0.5
Picoides villosus Hairy Woodpecker 1 0.2 66 0.0
Charadrius vodferus Killdeer 1 0.2 97 0.1
Anas platyrhynchos Mallard 1 0.2 1082 0.7
Sitta canadensis Red-breasted Nuthatch 1 0.2 10 0.0
Sdurus aurocapillus Ovenbird 1 0.2 19 0.0
Catharusfuscescens Veery 1 0.2 31 0.0
Unknown nestling 33 5.1 1190 0.7
Unknown bird (ASCl)^ 18 2.8 173 0.1
Unknown bird (ASC2)® 23 3.5 1778 1.1
Unknown bird (ASC3)^ 6 0.9 3459 2.1
Items not identified to class
Mammalia or Aves 76 11.7
*MSCl = mouse-sized prey item; MSC2 = red squirrel-sized prey item; ASCI = warbler-sized prey item: ASC2 = robin-sized prey
Item; ASC3 = Ruffed Grouse-sized prey item.
^ Omitted from analysis.
,. , .
268 Smithers et al. VoL. 39, No. 3
{Tamias striatus), snowshoe hare {Lepus americanus), and birds delivered among nests over 5-d intervals, because
eastern cottontail (Sylvilagusfloridanus) using 95% ofthe of missing data among sampled days, with a Kruskal-Wal-
adult mass; if ages could not be determined reliably, we lis single-factor ANOVA (Zar 1999). Because observations
assigned juvenile masses to these species. within breeding areas were not independent, we exam-
To estimate biomass of unidentified prey, we pooled ined differences in provisioning rates among breeding
unidentified birds into three a priori size classes (SC) fol- areas with multivariate repeated measures ANOVA. We
lowing Storer (1966) and Kennedy andJohnson (1986) used the General Linear Model (GLM) module of STA-
that represented average mass of common species in our TISTICA (Version 6.0, StatSoft, Inc., Tulsa, OK U.S.A.)
study area: SCI = 10 g (e.g., warbler-sized), SC2 = 77 g for all statistical analyses except calculation of diet over-
(e.g., robin-sized), and SC3 = 576 g (e.g., Ruffed Grouse lap and similarity, for which we used Ecological Meth-
[Bonasa umbellus]-sized) Similarly, we pooled unidenti- odology 6.1 (Exeter Software, Setauket, NYU.S.A.). An
fied mammal prey into two apriori size classes: SCI = 23 alpha level of P = 0.05 was used for all statistical tests,
g (e.g., mouse-sized) and SC2 = 192 g (e.g., squirrel- and we present means and standard errors.
sized)
.
Prey and Biomass Delivery Rates. We calculated deliv-
Results
ery rates on the basis of number of prey delivered per
day, number of prey delivered per nestling per day, and Video Recording and Prey Identification. We in-
annudmbtehrreoefnpersetlyindgesl.ivWeeredcaplecruldaatyedatbineosmtasswsitehstoinmeat,etswoi,n stalled video monitoring systems at three, five, and
the same manner. We calculated mean deliveryrates over seven occupied goshawk nests during the 2000,
5-d intervals from hatching to 5 d post-fledging (i.e., 2001, and 2002 field seasons, respectively. We
from 0-45 d). placed cameras at nests when nestlings were ca. 8
Prey Diversity and Overlap. We calculated prey diver- d old (±1.18; range = 1-18 d). One ofthe 15 nests
sity for the study area using ungrouped prey categories
failed within 3 d of camera placement and was re-
(i.e using each prey category identified to family, genus,
or s,pecies separately) Because samples were smaller moved from analysis. Due to camera malfunctions,
.
when examining individual nests, we generalized prey we were only able to collect 16 hr offootage at one
into similar species categories (Lewis 2001) to calculate of the nests in 2000. We placed a camera at the
prey diversity for individual nests. The generalized prey
2002 nest of the same pair, but pooled data from
categories for among-nest diversity assessment were: (1)
Sciurids, (2) blackbirds and Corvids, (3) Leporids, (4) both years as one nest area for analysis. Thus, our
Ruffed Grouse, (5) diving ducks (Aythya spp.) (6) water sample of 4801 hr (x = 320 ± 42 hr/nest) ofvideo
and shore birds, (7) passerines, (8) Picidae, (9) Falcon- footage is derived from 13 nesting pairs of gos-
iforms, (10) miscellaneous mammals (e.g., long-tailed
hawks.
weaWseelca[lMcuusltaetleadfrperneaytad]i)versity using Williams (1964) and We identified 59 (8.3%) of 711 prey deliveries as
MacArthur’s (1972) modified form of the Simpson’s in- being retrievals of cached items. Of the 652 fresh
dex (Simpson 1949) and diet equitability using Smith prey deliveries, we identified 451 (69%) to the spe-
and Wilson’s index ofevenness (Smith andWilson 1996)
cies level, 20 (3%) to genus, four to family (1%),
We used prey identified to family, genus, or species to
estimate diet overlap among nests with the Simplified and four (1%) as unidentifiable ducklings (Table
Morisita’s Index of Overlap (Krebs 1999). Overlap mea- 1). Eighty (12%) birds and 17 (3%) mammals were
sures are designed to measure the degree that two spe- unidentifiable beyond class, and we were unable to
cies share a set of common resources or utilize the same identify 76 (12%) deliveries. The majority of prey
parts of the environment (Lawlor 1980). Overlap mea- =
deliveries identified to at least class {N 576) were
sures are scaled from zero to one, where zero overlap
indicates dissimilarity in resource use, and one indicates mammals (62%), whereas birds (38%) comprised
complete overlap (Krebs 1999). We also assessed similar- a smaller proportion of diet.
ity in prey use among nests with cluster analysis using When considering only those deliveries identi-
average linkage clustering (Romesburg 1984, Krebs 1999,
M(c19G8a4r)i,gawle uetsedal.th2e00u0n)-.weAisghtseudggpeasitre-dgrobuyp Rmoemtehsobduursg- fspieecdietso;fNami=ly47o6r),fitnhere rdeosmoilnutainotn p(ri.eey.,stpoecgieensuwseroer
ing arithmetic averages (UPGMA). red squirrels (41.2%), eastern chipmunks (19.8%),
Statistical Analysis. We used analysis of variance (AN- American crows (7.7%), Ruffed Grouse (6.9%),
OVA) to examine relationships between delivery rate var- and snowshoe hares (6.5%). No other individual
iables and brood size using log-transformed data (Zar
1999). Biomass of prey delivered per day per nest was species accounted for >5% of identified prey. As a
=
transformed by taking the logarithm ofbiomass delivered group, Sciurids and Leporids {N 338) accounted
per day and adding 1.0 (Zar 1999). Normality of exper- for 70% of the identified prey. Among mammals,
imental error was tested using the Shapiro-Wilk test pro- 51.8% were adults, 25.4% were Juveniles, and we
cedure, and assumptions regarding homogenousvarianc-
es were tested using Levene’s test (Zar 1999). We were unable to estimate age for 22.8%. Of the
examined differences in the number of mammals and birds, 36.7% were adults, 9.6% were Juveniles,
,
September 2005 Biology 269
27 5% were nestlings, and we could not reliably brood size. On average, daily biomass deliveredwas
.
estimate age for 26.2%. 509 g (±84 g) to nests with one nestling, 555 g
Biomass. In context of the prey species and bio- (±42 g) to broods of two, and 756 g (±107 g) to
mass proportion used by goshawks in our study, the broods of three. Despite greater amounts of bio-
delivery of one domestic chicken {Gallus spp.) was mass being provided to larger broods, this resulted
unusual and the mass would dramaticallyinfluence in nestlings in single broods receiving 509 g (±84
biomass estimates for avian prey. We therefore con- g) of biomass per day, whereas nestlings in broods
sidered it an outlier and deleted it from biomass of two each received 278 g (±3 g) of biomass per
estimates. day and nestlings in broods ofthree each receiving
We estimated the total biomass of all prey deliv- 252 g (±36 g) per day.
eries at nests as 161 kg. The mean mass for both Dietary Overlap. The diversity and equitability of
avian and mammalian prey was 281 g (±13 7 95% prey delivered to nests was low for the study area,
. ,
confidence interval = 254—308 g). Although aver- as indicated by a reciprocal of the Simpson diver-
age mass of avian prey {x = 292 g; range = 10- sity index (1/7)) of 4.28 and a Smith and Wilson
1082 g) was similar to that for mammalian prey evenness index (i^ar) of 0.30. Similarly, diversity
(275 g; range = 18-1361 g), avian prey accounted among nests was low, with a mean value of l/D =
for only 39% of biomass delivered whereas mam- 3.77 (±0.41, range = 2-09-7.35). The mean value
mals accounted for 61% of biomass delivered. of for all nests was 0.56 (±0 04 range = 0 36-
. , .
Snowshoe hare (25%), red squirrel (24%), Ruffed 0.80). Low prey diversity and evenness values may
Grouse 11%), American Crow 9%), diving ducks be attributable to goshawk diet being dominated
( (
7%), chipmunk 5%), and eastern cottontail by red squirrels and chipmunks in our study. Sim-
( (
5%) accounted for 86% of biomass used by gos- ilarly, there was high dietary overlap (>0 8 among
( . )
hawks. No other species accounted for >5% ofbio- breeding pairs of goshawks in our study (Table 2
)
mass. although one nesting area (LSP; Table 2 ap-
)
Delivery Rates. Breeding goshawks delivered peared to be measurably different from the rest.
2.12 (±0 14 prey per day (i.e., 0.14 deliveries/hr), Cluster analysis indicated there were two groups of
. )
each delivery had a mean mass of 275 g (±20 g), breeding goshawk diets that exhibited similar prey
for a total of551 g (±50 g) delivered per day. How- composition and proportion of use (Fig. 2 al-
)
ever, daily 7^ = 3 44 P < 0 001 and hourly though, again, one nest (LSP; Fig. 2 appears to
( 13,253 . , . ) )
7^ = 2 31 P = 0 01 delivery rates varied be an outlier. There was no apparent relationship
( 13,250 . , . )
among nests. between overlap measures and spatial distribution
1.3 (±0.1) prey items were delivered per nestling of nests across the study area (Fig. 1, 2).
per day, but delivery rates increased with brood
“ P = Discussion
size (7^2,271 5.23, 0.01). Daily prey delivery
rates were 1.8 (±0 1 at nests with one nestling, 2.3 Mammals were the dominant prey of breeding
. )
(±0 1 at nests with two nestlings, and 2.5 (±0 2 goshawks in Minnesota, with red squirrels and east-
. ) . )
at nests with three nestlings. Despite the increase ern chipmunks appearing to be the most impor-
in prey deliveries among nests with larger broods, tant species in terms ofboth number delivered and
there was an inverse relationship between brood biomass. These two species alone accounted for
size and the number of prey delivered per nestling 62% of all prey identified to at least family and
per day (r = —0.43, P < 0.05). Each nestling in 51% ofprey identified to at least class. Several stud-
single broods received a mean of 1.8 (±0 1 prey ies have documented red squirrels as important
. )
items per day, whereas each nestling in broods of prey for goshawks (Squires and Kennedy 2005
)
two received only 1.2 (±0 1 prey items per day, throughout their range. They may be especially im-
. )
and each nestling in broods of three received only portant during the winter when other prey may be
0.9 (±0.1) prey items per day. less available (Widen et al. 1987). Squirrels domi-
322 g (±32 g) of biomass were delivered per nated goshawk diets in Sweden in terms ofnumber
nestling. However, we observed a pattern of bio- 79%) and biomass 56%) during winters of both
( (
mass delivered to broods of different sizes that was high and low squirrel abundance (Widen et al.
similar to that of number of prey delivered to 1987). Diet information for winter goshawks in the
WGLR
broods of different sizes; biomass delivered per is not available, but the extensive use ofred
nestling per day (Eg = 5 96 P = 0 038 varied with squirrels during the summer and the patterns of
5 . , . )
1 c C
270 Smithers et al. VOL. 39, No. 3
The squirrel use during winter in other areas (Widen
CO 05 xf5 CO so t- xf5 J> rH
presented i> O] 0005 x0O5 xO 0000 0’s5h r0H5 0055 r0-5H 00 X0O0 et al. 1987) suggest this species may be of year-
o d d d d d d d d d d d
round importance to goshawks in the region. In
seasons.
terms of biomass, snowshoe hares also appear to
data
be important for goshawks in our study area, ac-
< 00 05 I-H 00 00 XO so
The breeding dcn 0d0 00d50 CdJ5 dCcOM d0005 x0dn5 00d05 c0d0O cd0O5 0d050 counting for 25% of the biomass delivered to nests.
Rabbits and hares are also used extensively by gos-
hawks throughout their range (Squires and Ken-
2002
overlap). oCM 05 I— (M o CM 05 05 ^H nedy 2005).
50 so r- 00 rH CO 00
and d00 d00 d05 d05 rd- d00 dCJ5 d05 d00 d00 Ruffed Grouse comprised 5% of prey deliveries
and 11% of biomass delivered to goshawk nests
(complete
2001, during a 3-yr period of relatively low grouse abun-
1 H xn o s0o0 IsCo so C05 00 05 C00M r- dance (Smithers 2003). There is anecdotal evi-
2000, d dCO d05 d05 dCO dC05 d05 d05 d05 dence that at least some goshawks in Minnesota
to (Z)
may rely more heavily on Ruffed Grouse than oth-
the
er prey during some time periods (Eng and Gul-
overlap) i> 0000 0050 x0O5 CO 0s5o CXOO CSOM f-H lion 1962, Apfelbaum and Haney 1984). Eng and
(no during |0*3q d ad> 0d5 0d5 xdq 0d0 Cd05 0d5 Gullion (1962) focused on Ruffed Grouse mortal-
0 ity and did not assess proportional use of grouse
in the diet ofgoshawks, and Apfelbaum and Haney
m
from Minnesota Hs IoD> o05 0050 0C05O5 0CC5OM s00o55 0C0J505 I-H (1984) reported on prey remains collected at a sin-
d d d d d d d
Cu gle nest in northern Minnesota. Because of the dif-
range in
ficulties in accurately quantifying the extent of
13)
grouse predation by goshawks (Eng and Gullion
Q 05 05 so
Values {=N U C<dyM> s0do5 0d5 J0d>5 scdoO r0d-5- 1962) and the biases associated with determining
raptor diets based on prey remains (Smithers
2003), the results of these studies need to be in-
nests
Overlap. We
tJD CM r*H terpreted cautiously. suspect that the previous
<1 r- T-^ CM
of (Z) cdn 0d0 0d5 0d5 CdD research on goshawk diet for our study area, all
hJ
Goshawk collected by indirect methods (Eng and Gullion
Index 1962, Apfelbaum and Haney 1984, Martell and
to CM CM Dick 1996), may overestimate the proportion of
CO CM 05 so
Northern (Z) dCO xdq dxO dCO birds, especially large birds such as grouse, and un-
Morisita’s
derestimate the proportion of mammals in gos-
at hawk diets.
CO CM 00 r-f Qualitative review of the data suggests the mean
nests. w 00 XO 00
Simplified collected >-n d00 0d5 0d5 delivery rate of 0.14 deliveries/hr to nests in our
study was less than that observed in Arizona (0.25
goshawk deliveries/hr; Boal and Mannan 1994), Nevada
the
data o vr> C00O I— (0.31 deliveries/hr; Younk and Bechard 1994) and
00 05
using d d two areas of southeast Alaska (0.30 and 0.23 deliv-
specific eries/hr; Lewis 2001). However, although mean
frequency
values biomass per delivery in our study (275 g) was less
X I-H
than that in Arizona (307 g/delivery) where Le-
prey Q d
designate porids and Sciurids were the dominant prey (Boal
overlap
and Mannan 1994), it was greater than the two ar-
from
w eas of Alaska (214 g and 173 g/delivery), where
codes w
Dietary p birds were the dominant prey (Lewis 2001).
generated Our study indicates that goshawks with larger
2. letter broods provision with greater delivery rates and
z biomass. Biomass per nestling was similar between
Table were three DEE DIX HAG w LSP LSA MCD PMT SHA STE WAG WFA WRI broods of two and three (16.3-18.0 g/hr), but only
.
September 2005 Biology 271
Figure 2. Cluster analysis dendrogram for food habits data collected at Northern Goshawk nests in Minnesotaduring
WAG
the 2000, 2001, and 2002 breeding seasons. Parentheses indicate cluster number (see Fig. 1). The LSP and
breeding areas exhibited the least similarity of diet composition among breeding areas.
about half as much as that received by nestlings in Based on observed frequencies of prey use, this
broods of one (33.0 g/hr). This poses an interest- would translate to the average breeding goshawk
ing question regarding energetic aspects of gos- pair capturing 29 red squirrels, 14 eastern chip-
hawk productivity; what is the minimum biomass/ munks, six American Crows, five snowshoe hares,
hr necessary to fledge young successfully? The five Ruffed Grouse, two diving ducks, one cotton-
similarity between broods of two and broods of tail, one BlueJay, and 31 miscellaneous small birds
three suggests that, at least in our study area, and and mammals. To put this level of predation in
at nests with similar prey composition, a minimum context, all of these prey captures would occur
of 16—18 g of biomass per hr may be required for within a home range of 6376 ha for a goshawk pair
successful nesting. However, a finer assessment of in the study area (Boal et al. 2003)
nestling energetics would likely require experi- Composition and richness of prey delivered to
mentation in a laboratory setting. nests was similar across the study area, and esti-
Given our prey use and delivery rate data, one mates of prey diversity and equitability were gen-
can make a generalized prediction of the relative erally low among nests. We suspect the high dietary
impact of a breeding pair of goshawks in our study overlap and similarity of prey use among breeding
area during the 45-d nestling period. With an ex- areas was most likely attributable to the dominance
pected delivery rate of 2.1 prey/d over a 45-d nest- of red squirrels and chipmunks in goshawk diets.
ling period, ca. 94 prey deliveries can be expected. However, goshawk diets were dominated by red
272 Smithers et al. VoL. 39, No. 3
squirrels and chipmunks, but snowshoe hare, idency status ofNorthern Goshawks breeding in Min-
Ruffed Grouse, and American Crow were also im- nesota. Condor 105:811-816.
portant in terms of biomass. AND R.W. Mannan. 1994. Northern Goshawk diets
As pointed out by Reynolds et al. (1992), raptor in ponderosa pine forests on the Kaibab Plateau. Stud.
Avian Biol. 16:97-102.
populations are often limited by prey availability
Bosakowski, T. AND D.G. Smith. 1992. Comparative diets
and their choice of foraging habitat is predicated
of sympatric nesting raptors in the eastern deciduous
on conditions in which prey are abundant and
forest biome. Can. Zool. 70:984-992.
J.
available. Thus, an understanding ofgoshawk prey Bright-Smith, D.J. and R.W. Mannan. 1994. Habitat use
species used and the relative importance of those by breeding male Northern Goshawks in Northern
prey species is an important step toward develop- Arizona. Stud. Avian Biol. 16:58—65.
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A
COLLOPY, M.W. 1983. comparison ofdirect observation
fosters presence of those species while incorporat-
and collections of prey remains in determining the
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