BIOLOGICAL

CONSERVATION

Biological Conservation 118 (2004) 377–387

www.elsevier.com/locate/biocon

Area-sensitive forest birds move extensively among forest patches

Gail S. Fraser *, Bridget J.M. Stutchbury

Biology Department, York University, Toronto, Ont., Canada M3J 1P3

Received 1 November 2002; received in revised form 2 May 2003; accepted 19 June 2003

Abstract

Habitat fragmentation is thought to create a barrier to individual movements particularly for area-sensitive species which, by

definition, prefer to breed in large tracts of forest. For two breeding seasons, we radio-tracked an area-sensitive species, the scarlet

tanager Piranga olivacea, in a fragmented landscape in northeastern PA. We found that scarlet tanagers made extensive and frequent

movements among fragments. Paired males were less likely to leave their capture fragment, and traveled shorter distances. Unpaired

males in fragments had two distinct tactics we labeled: ‘‘Sedentary’’ and ‘‘Mobile’’. Sedentary males stayed at one fragment and sang

at high rates, while mobile males spent 58% of the total time tracked off their capture fragment and traveled over a kilometer away

from the capture site over open fields and through forests. Mobile males were not floaters per se because they were territorial (i.e.,

singing) in multiple sites. Habitat structure of a fragment did not correlate with the percent time a male spent off the capture

fragment. Males in fragments experienced lower pairing success and were more likely to be first time breeders compared to males in

continuous forest. Our results suggest that movements by scarlet tanagers in fragmented landscapes are not restricted during the

breeding season, and that these movements are related directly to pairing status and indirectly to population density.

� 2003 Elsevier Ltd. All rights reserved.

Keywords: Forest fragmentation; Neotropical migrant; Pairing success; Corridors; Radiotelemetry; Space-use; Scarlet tanager; Piranga olivacea

1. Introduction

Many North American songbird populations are

declining as result of habitat fragmentation on their

breeding grounds (e.g., Robinson et al., 1995). Many

studies suggest that Neotropical migrant species that

prefer larger tracts of forests (‘‘area-sensitive’’ birds; see

Villard, 1998) are especially sensitive to forest frag-mentation (e.g., Askins et al., 1987; Freemark and

Collins, 1992; Lee et al., 2002). Area-sensitive birds are

expected to (1) experience lower reproductive and pair-

ing success with a decrease in forest area (e.g., Gibbs

and Faaborg, 1990; Porneluzi et al., 1993; Villard et al.,

1993; Robinson et al., 1995; Roberts and Norment,

1999); (2) frequently be absent in small forest fragments

(e.g., Askins et al., 1987; see also Brown, 1984); and (3)prefer to use corridors and frequently avoid crossing

open, non-forested areas (Sieving et al., 1996; Desro-

chers and Hannon, 1997; B�elisle et al., 2001; B�elisle and

* Corresponding author.

E-mail address: fraserg@tamug.tamu.edu (G.S. Fraser).

0006-3207/$ - see front matter � 2003 Elsevier Ltd. All rights reserved.

doi:10.1016/j.biocon.2003.06.006

St. Clair, 2001). While the effects of fragmentation on

the distributions of Neotropical migrant species (e.g.,

Lynch and Whigham, 1984; Blake and Karr, 1987;

Freemark and Collins, 1992), and factors influencing

reproductive success (e.g., Robinson et al., 1995; Friesen

et al., 1999) have been extensively studied, little work

has been done on how individuals move within and

among fragments.Community-level, temperate breeding studies have

driven concepts on how populations are affected by

forest fragmentation in Neotropical migrant research

(Hunter, 1992), but an important gap in our under-

standing concerns how individuals of different species

move through a mosaic environment (Wiens, 1994).

Individual movements and habitat selection are key

components in understanding the effects of forest frag-mentation on population dynamics (Wiens, 1994). For

example, edge avoidance, or a reluctance to cross open

areas, are considered typical traits of area-sensitive and/

or forest interior Neotropical migrants breeding in the

temperate eastern forests of North America, yet for

most species we have little or no empirical data to

378 G.S. Fraser, B.J.M. Stutchbury / Biological Conservation 118 (2004) 377–387

support these ideas (Villard, 1998). Although edge

avoidance or a reluctance to cross open areas appear

uncharacteristic for migratory bird species, some evi-

dence suggests that migratory birds, upon arriving at the

breeding grounds, do not disperse or colonize new areasas rapidly as resident bird species (Bensch, 1999), and

many of these species are particularly sensitive to habi-

tat fragmentation (e.g., Freemark and Collins, 1992).

Low breeding dispersal rates or colonization into new

areas may be linked to behaviors such as site fidelity

(Bensch, 1999) or higher predation risks when crossing

open areas (Wiens, 1994; Grubb and Doherty, 1999).

Recent radio-tracking studies on two area-sensitivebird species demonstrate that movements between

fragments are not only possible, but can occur regularly

(Bayne and Hobson, 2001a; Norris and Stutchbury,

2001). Norris and Stutchbury (2001) found that male

hooded warblers Wilsonia citrina leave their small forest

fragments on a daily basis through the incubation and

nestling stages. They crossed gaps up to 460 m in search

of extra-pair copulations, despite being paired andhaving an active nest. Bayne and Hobson (2001a) ob-

served no differences in the amount of movement made

by male ovenbirds Seiurus aurocapillus in continuous

forest compared to fragmented habitat in the post-

fledging period. However, males with fledglings moved

less compared to those without young and rarely crossed

open areas and consequently those on forest fragments

were restricted in their movements compared to males incontinuous forest. These studies suggest that moderate

fragmentation and isolation of forest patches on the

breeding grounds may not inhibit movement among

patches by birds during certain periods of the breeding

season. Clearly, more data on habitat use are required to

understand how area-sensitive species behave in frag-

mented landscapes and to facilitate better strategies for

their conservation.We chose to study scarlet tanager Piranga olivacea

behavior in forest fragments because this species is

negatively influenced by forest fragmentation (Rosen-

berg et al., 1999), and scarlet tanagers are canopy birds

that prefer mature forests (Mowbray, 1999). There are a

number of studies that describe a positive relationship

between forest area and the presence of tanagers (e.g.,

Robbins et al., 1989; Askins et al., 1987; Freemark andCollins, 1992). Rosenberg et al. (1999, Project Tanager)

found a positive relationship between forest cover at the

landscape scale and the occurrence of tanagers, although

the response of scarlet tanagers to forest fragmentation

varied among regions. Roberts and Norment (1999)

recorded a complete absence of scarlet tanagers in

fragments less than 10 ha in a highly forested area

(northwestern NY). Although tanagers can occur insmall woodlots (e.g., 3 ha, Galli et al., 1976; 6.5 ha,

Blake and Karr, 1984), it is unclear whether these birds

were breeders or vagrants (see Mowbray, 1999) because

none of these studies involved radio-tracking of indi-

viduals. Here, we present the results of a radio-tracking

study on scarlet tanagers in northwestern PA and de-

scribe the movements and pairing success of individuals

in a fragmented landscape.

2. Methods

2.1. Study area

We studied scarlet tanagers at the Hemlock Hill Bi-

ological Research site (41�460N, 79�560W) and sur-rounding region (approximately 740 km2) in Crawford

County, northwestern PA in 2000 and 2001. Male

scarlet tanagers were located on 10 small isolated

woodlots (<10 ha, mean¼ 5.3� 2.5 ha, range 3.2–9.6

ha), 2 mid-sized fragments (44 and 75 ha), and 2 rela-

tively large forest plots (150, 165 ha), hereafter referred

to as continuous forest. All fragmentation in this area

was caused by agricultural activities and eight fragmentswere patches completely isolated by fields or roads,

while four woodlots were connected to other forest

habitat by small (<40 m wide) corridors. Mean distance

to nearest forests for the 12 fragments was 138� 21.9 m

(range 10–325 m).

We calculated the area and perimeter–area ratio of

forest woodlots with a planimeter (Planix 7) from 1993,

1:40,000 aerial photographs. Perimeter–area ratio wascalculated as the ratio of edge (m) to area (ha) for each

fragment. Percent forest cover within 1 km of each

woodlot was also estimated from aerial photographs.

Based on USGS topographical maps, total forest cover

for the region (740 km2) was approximately 39%. Al-

though forest cover in some areas has increased slightly

(<1%) since the production of these maps (personal

observation), all of the fragments used in the study ap-peared to be the same size as shown on the 1993 aerial

photographs. Furthermore, no major logging has oc-

curred in any of the fragments except for one that was

selectively logged (i.e., the fragment is still the same size)

in the past ten years. We concluded that there has been

little change in forest cover in the region since the early

nineties.

The four most predominant tree groups in the forestsfragments were maple Acer spp., hemlock Tsuga

canadensis, American beech Fagus grandifolia, and

hickory Carya spp. In the continuous forest plots the

mixture was similar except elm Ulmus spp. was more

common than American beech.

2.2. Radio-telemetry

We captured tanagers with song playbacks, a tanager

decoy (taxidermic mount), and one 12 m long mist-net

placed at standard height. Each bird was banded with a

Table 1

A summary of telemetry on scarlet tanagers in Northwestern PA

2000 2001

Continuousa Fragment Continuousb Fragment

Total tracking (h) 110.8 114.2 30.1 104.1

Mean tracking time (h� SD) 11.1� 3.6 12.7� 4.5 10.0� 3.5 10.4� 4.1

Range of tracking time (all males) 6–14.7 6–19.5 6–12.1 6–18

Number of males tagged 10 9c 3c 9

aMales tracked on 150 ha plot only.bMales tracked on 150 and 165 ha plots.cOne male each year was captured in a fragment and then settled in continuous forest.

G.S. Fraser, B.J.M. Stutchbury / Biological Conservation 118 (2004) 377–387 379

US Fish and Wildlife aluminum band, and an individ-

ually distinct color band combination. We measured

wing chord, tarsus length, and body mass (using a 50 g

Pesola scale) of each adult. We aged tanagers based on

plumage coloration; second-year (SY) males have brown

on their primaries, whereas after second-year males

(ASY) have all black primaries (Mowbray, 1999). Males(n ¼ 31; Table 1) were fitted with a small (1.4 g), BD-2G

radio transmitter (Holohil Systems Ltd, Carp, Ont.,

Canada) attached with a figure-8 cotton embroidery

thread harness (see Rappole and Tipton, 1991). The

transmitter and harness together weighed about 5% of

adult body mass (average �30 g) and the harness was

designed to fall off by the end of the field season.

Transmitters lasted up to 8 weeks and the range wasapproximately 1.5 km.

While we attempted multiple recaptures of all tana-

gers at the end of each season to remove the transmitter,

we had a low success rate (n ¼ 3). Nine transmitters (5 in

2000; 4 in 2001) fell off before the end of the study. In

the continuous forest, males with transmitters left on at

the end of the 2000 season experienced a similar return

rate (67%, n ¼ 6) to males with transmitters removed(57%, n ¼ 7). Surprisingly, 3 males with transmitters

attached in 2000 returned with transmitters still in place

upon arrival on the breeding grounds in 2001 (contin-

uous forest plot, all ASY males in 2000). These three

males were the first to return in 2001, attracted females

within a few days, and lost the transmitters within a

month of arriving. Also, all males that were recaptured

at the end of each study year had a mass that was within1 g of initial capture mass (mean initial capture

mass¼ 27.8� 1.0; mean recapture mass¼ 27.8� 0.29,

n ¼ 3). From these data, we concluded that scarlet

tanagers were not negatively affected by wearing a radio

transmitter (see also Powell et al., 2000).

We radio-tracked tanagers in 2 h bouts between 06:00

and 14:00 h, usually on foot 20–30 m away from the

focal bird, however on a few occasions (2.6% of totaltracking time) tanagers were tracked by car. Observa-

tions were not made during wet weather. We had a total

of 359 tracking hours and each male used in our anal-

yses had a minimum of 6 hours of tracking time (Table

1). For both years, birds were tracked between 13 May

and 30 June.

Because many studies have described scarlet tanagers

as forest interior or area-sensitive birds, we were par-

ticularly interested in documenting whether males

moved off the isolated woodlots. We report these results,

for paired and unpaired males, in percent time off cap-ture fragment (minutes off fragment divided by total

minutes tracked), and the rate off capture fragment

(number of trips off fragment per hour tracked). Percent

time off fragment for paired males was not divided into

different breeding stages because only two of 11 pairs

fledged young during our study and nests were difficult

to locate, so stage of nesting was most often unknown.

We also calculated percent time off territories for malesin continuous forest and compared them to paired males

in forest fragments with a t test. The distance traveled

was calculated from topographical maps (fragments) or

from grid marks (continuous forest; see below) and re-

ported as average values. When relevant, we also re-

ported corridor use as a percent of time tracked. All

averaged values are reported with �1 SE.

We determined whether a male was paired based onrepeated sightings of a female with the focal male, or

seeing her carry nest material, on a nest, or carrying

food for young. We used a v2 test to compare age dis-

tribution and pairing success between fragments and

continuous forest. We measured song rates (number of

song bouts during a tracking period) during the radio-

telemetry observations from males on forest fragments

and compared paired and unpaired male rates with a ttest.

We had two different methods for recording the lo-

cation of a male during a tracking session and deter-

mining subsequent territory size, depending on which

plot a male was tracked in. The 150 ha forest had con-

spicuous grid marks every 50 m; the location of a male

tracked on this plot was recorded by noting the closest

grid mark whenever the bird moved. We measured ter-ritory size with a planimeter for each tracking session

and reported the average territory size for each male.

The location of males in forest fragments or in the 165

ha plot were recorded by noting their position relative to

380 G.S. Fraser, B.J.M. Stutchbury / Biological Conservation 118 (2004) 377–387

obvious landmark features. At the end of each season,

we mapped all movements for males, measured the ex-

tent of these movements with a 50 m tape and used a

planimeter to determine territory size (i.e., area defended

with song). Movements off territories for continuousforest males were assumed when males moved greater

than twice the distance away from the perimeter of their

territory (i.e., usually 100 m or more) and were in other

males� territories. Movements off fragments for males in

forest fragments were considered time off territory and

these movements were not included in the area estimate.

Four males used and defended an entire woodlot,

therefore we considered the whole fragment to be theirterritory.

We found that some males in fragments not only

gained a partner mid-season, but that they also switched

territories mid-season (i.e., from a fragment to contin-

uous forest); therefore, we report the final status (e.g.,

paired or not, and in fragment or continuous forest) of

male tanagers as they were by the end of the telemetry

work (30 June for both years, or sooner if the trans-mitter fell off).

2.3. Habitat structure and movements off territories

Habitat structure, which can influence prey abun-

dance (Smith and Shugart, 1987), may also determine

how often males leave their fragments or territories. We

examined (1) whether there was a relationship betweenamount of movement off a forest fragment and the

fragment�s habitat features, (2) whether movements in

continuous forest were related to the habitat features of

territories, and (3) whether paired males� habitat differedbetween continuous forest and forest fragments. We

measured habitat characteristics with the James and

Shugart (1970) method. For each territory we used 5–

0.02 ha plots. The initial plot was positioned at thecenter of the territory, while the other 4 plots were po-

sitioned 50 m from the center, in each cardinal direction.

For each territory we determined tree density (number

of trees per ha), average diameter of tree at breast height

(cm), number of tree species, total log basal area, %

canopy cover, canopy height (m), and shrub density

(number of shrub stems per ha). Many of our variables

were correlated, therefore we log transformed all thevariables and used a Principle Components Analysis

(PCA; correlations matrix) to collapse our dataset

(McGarigal et al., 2000). We used principle components

(PC) I and II, % forest cover within 1 km, and perime-

ter–area ratio, in a backwards step-wise regression with

percent time off fragment (arcsin transformed; Zar,

1996) as the dependent variable for males on 12 frag-

ments (Mobile males not included, see Section 3). If afragment had more than one male in both study years

(n ¼ 3), we averaged the percent time off territory for all

males in the fragment. In addition, if a fragment had

more than one vegetation plot (n ¼ 1), we averaged the

values for this analysis.

We repeated the PCA and backwards step-wise re-

gression for 13 territories in continuous forest, excluding

perimeter–area ratio as a variable with the dependentvariable as percent time off territory. Percent forest

cover was included in the regression because some males

were on the edge of the forest (n ¼ 5). We included edge

and interior territories in one step-wise regression be-

cause we found no difference in percent time off territory

between males in the two areas (p ¼ 0:8) and because of

our limited sample sizes. Finally, we compared habitat

features (canopy cover and height, log total basal area,and shrub and tree density) between continuous forest

(n ¼ 12) and fragment plots (n ¼ 10) for paired males in

a series of t tests.

2.4. Occupancy of forest fragments

We used a playback system to survey 13 woodlots in

May 2000 (mean area� SD¼ 10.1� 19.6, range 3.0–75ha) and 15 woodlots in May 2001 (mean area¼11.9� 19.6, range 3.0–75 ha) to estimate percent occu-

pancy of isolated forest fragments. Fragments with

tanagers (either banded or unbanded) observed more

than once were considered occupied. In all but two cases

each year, the males with transmitters were the only

color-banded individuals present in the fragments.

3. Results

3.1. Radio-telemetry

3.1.1. Movements of males in forest fragments: paired

versus unpaired behavior

Paired males were less likely to leave their capturefragment than were unpaired males (t test, n ¼ 11; 8; t ¼�2:8; p ¼ 0:01) and traveled shorter distances than un-

paired males (t test, n ¼ 6; 8; t ¼ �3:15:0; p ¼ 0:01; Ta-ble 2). Paired males also had significantly lower song

rates than unpaired males (paired: mean¼ 0.4� 0.1

songs/min; unpaired: mean¼ 2.5� 0.3 songs/min; (t test,

n ¼ 11; 6; t ¼ �7:4; p < 0:001). We found no differences

in territory size between paired and unpaired males(Mobile males not included-see below) in fragments

(t test, n ¼ 11; 4; t ¼ �2:0; p ¼ 0:06; Table 2).

Observations of paired males off territories (i.e., off

fragment) can provide insights into the reasons for these

movements. Only six of 11 paired males moved off their

forest fragments and we observed considerable variation

in the number of times these movements occurred. Three

paired males only left once out of the total time tracked(24.2 h; mean per male¼ 8.1 h): two males (with no

young) left for unknown reasons (no visual observations

of them) and the third (nesting status unknown) was

Table 2

Movements of radio-tracked male scarlet tanagers in fragments and continuous forest

Continuous Fragment

Paired Unpaired Paired Unpaired

Stay and sing Move and

sing

Total

No. of males 12 1 11 4 4 8

Percent time off fragment or territory 4.7� 4.3n:s: 1.1 5.7� 2.2 6.7� 2.6 57.8� 8.5 32.3� 10.5

No. times off fragment or territory per hour 0.2� 0.2 0.17 0.2� 0.07 0.2� 0.09 N.A.c 0.2� 0.09

(n ¼ 4)

Distance traveled (m)a 106� 16.5��

(n ¼ 10)

25 173� 27

(n ¼ 6)

467.0� 144.8 1120� 204 793.4� 169.1

Territory size (ha)b 0.9� 0.2� 0.6 1.6� 0.3 3.2� 0.9 N.A.c 3.2� 0.9

(n ¼ 4)

Values are mean� SD.aCorridors were not used in movements. Distances are calculated as one-way trips. Sample sizes differ from number of males tagged because mean

distance traveled is only from males that left their territory.b Territory size calculated from tagged males only.c Because Move and Sing males were off their capture fragment more than 50% of the time tracked and did not make discrete movements related to

their capture fragment we did not calculate the number of times off fragment per hour or territory size. t Tests between paired males in fragments

versus paired males in continuous forest.* p ¼ 0:05.** p ¼ 0:04.

G.S. Fraser, B.J.M. Stutchbury / Biological Conservation 118 (2004) 377–387 381

observed foraging off territory. One paired male (with

no young) left twice out of total time tracked (12.1 h),

and on one of these occasions he was engaged in a male-

male chase. The last two paired males left multiple times

(>3) over the course of the total time tracked (33.7 h;

mean per male¼ 16.9 h). One male spent 17% of his time

in a neighboring fragment (200 m away) and was also

observed there with a female. This pair also spent 52%of their time in an adjacent corridor (40� 10 m) where

on one occasion, they were observed foraging with

fledglings on ripe elderberry berries (Sambucus nigra).

The second male increased his time off the capture

fragment from 2% during incubation stage to 28%

during the fledgling stage. He was observed singing

during the incubation stage in a neighboring fragment

and foraging during the fledgling stage in this samefragment and once more in another nearby forest.

We found that tanager pairs (4/11) on forest frag-

ments sometimes moved territories mid-season. Two

males disappeared immediately after nest predation

events, and they were not subsequently observed (either

visually through playbacks or by radio transmitter sig-

nals) in their fragment or in nearby forests for the re-

mainder of the field season. Two other pairs movedthree times to different territories within a season. Final

territories were 300 and 400 m away from the original

territories and were entirely off the capture fragments;

one pair moved to a neighboring fragment (approxi-

mately twice as large as the capture fragment), the other

pair moved to continuous forest.

We found that unpaired males (all SY; n ¼ 8) adop-

ted one of two distinct tactics that we describe as: (1)Sedentary and (2) Mobile. Sedentary males (n ¼ 4) spent

the majority of their time in their capture fragment

singing at high rates (Table 2). Mobile males (n ¼ 4)

moved over open fields and through forests and were off

their capture fragment more than 50% of the time

tracked (e.g., Fig. 1(a); Table 2). Unlike secretive float-

ers (e.g., Zack and Stutchbury, 1992), Mobile males

advertised by singing on the original capture fragment

(territorial behavior that allowed us to initially capturethem). Mobile males were difficult to follow because we

did not know where they would be from one day to the

next and they were often out of transmitter range (ap-

proximately 1.5 km) at the time of a scheduled tracking

session. For example, one male was out of transmitter

range 11 out of 18 days we attempted tracking and he

was located in the capture fragment on only three out of

the remaining seven days. Yet when he was on thecapture fragment he would respond to playbacks (i.e.,

was territorial). A conservative estimate of the total area

utilized by Mobile males (n ¼ 3) averaged 122� 99 ha.

The fourth Mobile male likely had a similar range as the

other males, but we only located him once off the cap-

ture fragment and consequently could not estimate the

total area traveled.

The mating status of unpaired males sometimeschanged comparatively late in the breeding season.

Three males gained females mid-season (two Sedentary

– 14 June and 7 July; one Mobile – 18 June). The two

Sedentary males (both the sole occupants of each

fragment) were counted as unpaired in the data anal-

ysis because one male�s transmitter fell off before

pairing occurred and the other male gained a female

after the end of our study. The one Mobile male wewere able to track after he was paired behaved

Ferris unpaired

+*53.8%

+9.5%

+12%

24.6%+

1 km

+100%+

Ferris paired(b)

(a)

Fig. 1. An example of (a) an SY, unpaired male who adopted the Move

and Sing strategy, tracked from 13 May to 13 Jun 2001 for 14 h. Song

rate was 1.2 songs/min. The boxes represent available forest. The star

represents the capture fragment (3.6 ha) and the crosses are the dif-

ferent areas in which this male was tracked. The percent time is the

time spent in each area out of the total time tracked (14 h). The dashed

line encompasses the total area of movements by male. (b) The same

male later in the breeding season after establishing a territory in con-

tinuous forest and attracting a female. Tracked from 18 to 26 Jun 2001

for six hours. Song rate was 0.008 songs/min. Percent time is spent in

territory. Note the marked reduction movements and the decline in

song rate after pairing.

Table 3

Comparisons of scarlet tanager pairing success, age structure, and

territory size, between continuous forest and forest fragments

Continuous forest Forest fragments

No. malesa 30 17

Age ratio (SY:ASY) 3:27 14:3

No. paired males

(SY, ASY)

27 (2; 25) 10 (7; 3)

% Pairing success 90% 59%

One SY unpaired male was captured in a fragment, but settled with

a female in continuous forest (see Fig. 1). Another ASY paired male

was initially in a fragment for two weeks (20 May–3 June) then moved

to continuous forest with, presumably, the same partner. Both of these

males were placed in the continuous forest category.a Includes all birds intensively followed from both study years, in-

cluding individuals that were not banded. For radio-tagged birds lo-

cation and pairing status of males is that at end of study (30 June).

382 G.S. Fraser, B.J.M. Stutchbury / Biological Conservation 118 (2004) 377–387

similarly to paired males observed at the beginning of

the season (Fig. 1).

3.1.2. Age structure and pairing success: fragments versus

continuous forest

SY males were more likely to be in forest fragments(14/17), whereas ASY males were more likely to be

found in the continuous forest (27/30; v21 ¼ 24:6; p <0:0001; Table 3). SY males (9/17) also experienced lower

pairing success than ASY males (28/30; v21 ¼10:6; p ¼0:001; Table 3). All three ASY males in forest fragments

were paired, while only one SY male in continuous

forest was unpaired.

3.2. Habitat structure, movements off territories, and

corridor use

Principle component scores I and II explained 64% of

the variance in the data matrix for forest fragmenthabitat. We interpreted PCI to be representative of

canopy height and cover (magnitudes of loading 0.52,

0.49, respectively; all other variables<�0.42) and PCII

to be representative of tree density (loading 0.77; all

other variables<�0.34) for forest fragment habitat.

Canopy height and cover (PCI), tree density (PCII),

edge density, or the amount of forest cover within 1 km

did not influence the probability of a male leaving hisfragment (backward stepwise regression, no variables

remained in model, R2 ¼ 0:00; n ¼ 12; does not include

Mobile males).

Principle component scores I and II explained 65% of

the variance in the data matrix for continuous forest

territories. We interpreted PCI primarily as diameter at

breast height (DBH) (loading 0.61; all other vari-

ables<�0.42) and PCII was most influenced by canopyheight and the number of tree species (loading 0.51,

0.49, respectively; all other variables<�0.47). DBH

(PC I), the number of tree species and canopy height

(PC II), or the amount of forest cover within 1 kin did

not influence the probability of a male leaving his ter-

ritory (backward stepwise regression, no variables re-

mained in model, R2 ¼ 0:00, n ¼ 13).

Paired males in continuous forest resided in forestswith significantly taller canopy and increased total basal

area compared to paired males in fragments (Table 4). In

the two years of our study, we tracked scarlet tanagers in

12 fragments. In four fragments we captured both paired

and unpaired males in both study years. Because both

paired and unpaired males were captured on these frag-

ments,we suggest that there are no special habitat features

related to pairing success. In six fragments, we capturedonly single paired males, and in the last two fragments we

captured only single unpaired males (Table 4).

Table 4

Mean (�SD) habitat characteristics for fragments and continuous forest plots

Continuous forest Forest fragments

Paired (n ¼ 12)a Unpaired (n ¼ 1) Paired (n ¼ 10) Unpaired Sedentary (n ¼ 2)b

Canopy cover (%) 56� 12n:s: 60 50.1� 26.3 64� 31

Canopy height (m) 25.6� 5.2� 35.4 17.7� 7.7 28.4� 6.5

Tree density (trees per ha) 436.3� 100.3 525 454.5� 124.1 377.5� 17.7

Shrub density (shrub stems per ha) 29,343� 22,878n:s: 37,910 46,428� 36,324 28,098� 16,399

Log basal area 1.4� 0.08� 1.6 1.3� 0.1 1.4� 0.17

Edge density (m of edge per ha) NA NA 35.3� 10.4 58.7� 19.2

Fragment size (ha) NA NA 15.3� 24.5 7.5� 3.0

% Forest cover within 1 km 47.6� 3.8 45.5 28.3� 6.8 36.9� 14.2

a t Tests between paired males in fragments and continuous forest.bCharacteristics of capture fragments; n ¼ 3 (instead of n ¼ 4) because on two fragments we captured two males (one Sedentary and one paired) in

opposing study years (see Section 3 for further information).* p < 0:01.

G.S. Fraser, B.J.M. Stutchbury / Biological Conservation 118 (2004) 377–387 383

Four fragments had forest corridors available for

tanagers to use in their movements. Six males (two un-

paired, four paired) were tracked on these fragments

(Table 5). Two males were observed using (or flying inthe direction of) a corridor when leaving the same

fragment (different years). The first male just went into

the corridor and no further. The second male not only

used a corridor to go to an adjacent fragment, but he

also spent 52% of the time tracked in the corridor and

we considered this area part of his territory (Section

3.1.1). All other movements, on both the remaining

fragments with corridors and the eight fragments with-out corridors, during tracking sessions, were across

fields and roads (Table 5).

3.3. Occupancy of fragments

In May of 2000 and 2001, 77% (10/13) and 73% (11/

15) of the fragments surveyed were occupied. Eighty-one

Table 5

Number of males radio-tracked on each fragment and fragment characterist

Fragment Size Closest forest

(m)

No. males radio tagged

(yr 1, yr 2)

Corridor

(no.)a

ST 3.2 110 ð1; 1Þ No

RP 3.2 150 ð1; 0Þ No

FR 3.6 20 ð1; 1Þ No

PC 3.7 200 ð1; 1Þ Yes (1)

AR 3.7 125 ð0; 1Þ No

MY 3.8 130 ð1; 0Þ Yes (1)

CE 4.8 100 ð0; 1Þ No

FL 5.4 100 ð0; 1Þ Yes (1)

LT 8.0 110 ð2; 1Þ No

DC 9.6 325 ð0; 1Þ No

FG 44 100 ð1; 1Þ Yes (1)

GF 75 200 ð0; 1Þ No

aNumber of corridors attached to fragment.bMale went in direction of corridor or was observed using corridor.cMobile males, see text for more information.d This male spent 52% of his time tracked in the corridor.

percent (9/11) of fragments of <10 ha were occupied in

2000 and 83% (10/12) were occupied in 2001.

4. Discussion

Current theory predicts that area-sensitive, or forest-

interior birds, occur more frequently in larger habitat

patches, experience lower pairing success with a decrease

in forest area, and avoid crossing open gaps. Yet scarlet

tanagers in our study made extensive and frequent

movements among fragments without using corridorsand had a high occupancy rate in small forest fragments

<10 ha, observations not supportive of current predic-

tions for area-sensitive species. On the other hand, some

of the patterns we observed were consistent with the

negative effects of fragmentation: the proportion of SY

tanagers was greater in forest fragments than in con-

tinuous forest and males in fragments experienced lower

ics

? Total no. times

tracked off

fragment per male

No. times corridor

in or throughb

No. males in

fragment per

season

2,3 N/A 1

12 N/A 1

1,Mc N/A 1

2,8 l,7d 2,2

0 N/A 1

0 0 1

3 N/A 1+

3 0 1

M,0 N/A 2,1

M N/A 1

0,1 0 3,3

0 N/A 2+

384 G.S. Fraser, B.J.M. Stutchbury / Biological Conservation 118 (2004) 377–387

pairing success. Tanagers in large forests had higher

pairing success and were more likely to be ASY, but SY

males on forest fragments were not restricted in their

movements by the fragmented landscape.

4.1. Movements off fragments

Tanagers� movements were not restricted to their

capture fragment. Movements off fragments seemed to

have a variety of causes including foraging, male-male

chases, and mate searching. Paired males were less likely

to leave their capture fragment than were unpaired

males, which suggests that a primary motivation forunpaired male movements off fragments was to find a

mate. The extreme of this behavior was the Mobile male

strategy. Because there was no difference in the amount

of movement off territories for paired males in fragments

and continuous forest, we conclude that paired males

react similarly to the surrounding landscape indepen-

dent of whether they are in fragments or continuous

forest.Comparisons to other radio-tracking studies are

limited because very few exist for Neotropical migrants

on the temperate breeding grounds (Norris and Stut-

chbury, 2001; Norris et al., 2000 and Bayne and Hob-

son, 2001a). Compared to hooded warblers, paired

tanager males left their fragments less frequently. Norris

and Stutchbury (2001) found that both paired and un-

paired male hooded warblers left their fragments andreturned, as much as once every two hours, and crossed

gaps in search of extra-pair matings through the incu-

bation and nestling stage.

Some tanager movements off fragments were perma-

nent changes in territory location after nest failure. In

the two cases where we were able to follow the birds,

both moved to areas where other tanagers were present

and one moved into continuous forest. Changing terri-tory sites may be an attempt to increase the likelihood of

success of the second nesting attempt, and this behavior

has been noted in other species. For example, prairie

warblers (Dendroica discolor) frequently changed mates

and territories within a breeding season in association

with nest failure (Nolan, 1978).

We discovered a previously undescribed behavior

used by forest birds to search for mates, Mobile males.We found that Mobile males moved and advertised their

position among both fragments and continuous forest in

search of a mate, a form of continuous dispersal

throughout the breeding season. Both mate finding

tactics (i.e., Sedentary and Mobile) sometimes were

successful at attracting females mid-season and this

success was likely related to re-nesting attempts, al-

though only three of the nine unpaired males foundmates. The success of Sedentary males in attracting a

female suggests that females were also moving among

forest fragments. Males that pair later in the breeding

season might still have an opportunity to rear chicks

because we have preliminary evidence that at least some

scarlet tanagers in this area will double brood and re-

nest (unpublished data). Thus, scarlet tanager females

may be reproductively available for longer periods oftime than in single-brooded species. With some level of

double brooding and re-nesting, scarlet tanagers may be

able to maintain viable populations in some fragmented

habitats, as Friesen et al. (1999) concluded for wood

thrush (Hylocichla mustelina) breeding in small forest

fragments in southern Ontario. More work is needed to

understand the role that female tanagers have in nest site

selection in a fragmented landscape, and in mate choiceand retention within and between seasons.

Our study demonstrated variation in male scarlet

tanager behavior was mostly related to pairing status.

Unpaired males were much more likely to leave their

capture fragment and sing in multiple sites, which may

mean that during annual breeding bird surveys (Sauer et

al., 2002) tanager densities may be overestimated.

However, paired males had significantly lower songrates than unpaired males, so they may go undetected

more often during breeding censuses. These two effects

may partially cancel each other out.

4.2. Age structure and pairing success

Pairing success can be influenced by factors such as

age (e.g., Sherry and Holmes, 1989; Bayne and Hobson,2001b) and location of breeding territory (i.e., fragment

or continuous forest). In our study, 83% of scarlet tan-

ager males in forest fragments were SY birds that ex-

perienced lower pairing success compared to males in

the continuous forest plot. Although most males in the

fragmented habitat were unpaired SY males, ASY males

with territories in forest fragments (n ¼ 3) experienced

100% pairing success, suggesting that age is also animportant factor in female mate choice in scarlet tana-

gers. Roberts and Norment (1999) observed lower

pairing success for tanagers in smaller forest fragments

(areas <1000 ha), though pairing success was higher in

their forest plots compared to ours (see below). Differ-

ences in pairing success due to area effects has been

documented in ovenbirds (Villard et al., 1993; Van Horn

et al., 1995), but Villard et al. (1993) suggest that lowpairing success for males in forest fragments may also be

related to higher local population density.

4.3. Habitat structure, movements off capture fragments,

and corridor use

In their extensive study on tanagers, Rosenberg et al.

(1999) found that regional forest cover affects the sen-sitivity of scarlet tanagers to forest fragmentation. In

areas with higher amounts of regional forest cover, such

as the US northeast, tanagers may occur in smaller

G.S. Fraser, B.J.M. Stutchbury / Biological Conservation 118 (2004) 377–387 385

forest fragments, whereas in areas with less regional

forest cover, tanagers may occur only in larger forest

plots (Rosenberg et al., 1999). Our study area has a

moderate amount of hardwood forest cover, and is in

the center of this species� breeding range, a combinationthat may result in high population density and higher

occupation of available habitat (Brown, 1984). In our

study area, not only did we find high occupancy rates of

small forest fragments, but we also tracked tanagers

moving among fragments. Thus, our findings appear to

support Project Tanager�s (Rosenberg et al., 1999)

conclusions on the importance of regional forest cover,

and help explain why scarlet tanager area-sensitivityvaries across regions.

As population density increases SY males may have

less opportunity to establish a territory on preferred

larger forest patches through competitive interactions

with ASY males (see Sherry and Holmes, 1989) and

consequently experience lower pairing success. It follows

that as density decreases the occupation of smaller forest

fragments should also decrease, but the proportion ofpaired SY males should increase (Weins, 1976; Brown,

1984; Villard et al., 1993). The differences we observed in

scarlet tanager pairing success, patch size occupancy,

and territory size between our study and one conducted

less than 150 km away in upstate NY (Roberts and

Norment, 1999) implicates regional population density

as a major factor in determining these differences.

Roberts and Norment (1999) found that scarlet tanagerswere not present in forest fragments of less than 10 ha in

upstate NY, whereas we frequently found scarlet tana-

gers on forest fragments of <10 ha. Roberts and Nor-

ment (1999) also measured higher rates of pairing

success (>75%) on their forest fragments (10–150 ha)

compared to the 59% pairing success we observed for

tanagers in smaller forest patches (3.2–75 ha). Finally,

Roberts and Norment (1999) also reported larger terri-tory sizes compared to those estimated in our study;

however, differences in estimated territory size may also

be due to differences in methodologies.

Male scarlet tanagers did not require the presence of

a corridor to travel among forest fragments (Table 5).

Scarlet tanagers are strong fliers and can cross 100 m

gaps in seconds (personal observation). Therefore, we

conclude that absence of corridors did not alter themovements of adult tanagers, although tanagers may

have moved more frequently if more corridors were

available. While it appears that corridors are not nec-

essary for tanager movement among forest fragments,

this study was not rigorously designed to test corridor

use (see Nicholls and Margules, 1991), nor is it known

whether corridors may be important for other aspects of

tanager reproductive success, such as juvenile dispersal(see Lens and Dhondt, 1994). This study supports the

view of Hunter (1992) and Wiens (1994), who argued

that a long-distance migrant like the scarlet tanager

should not be restricted in their movements in a frag-

mented landscape despite their preference for large

forests.

4.4. Conclusions

Our results show that a species exhibiting an ‘‘area-

sensitive’’ pattern of fragment occupancy (e.g., Roberts

and Norment, 1999) cannot be assumed to be limited to

forest-interior habitat. Based on Project Tanager results

(Rosenberg et al., 1999), and the fact that scarlet tana-

gers are canopy birds that prefer mature forests (Mow-

bray, 1999) we would never have predicted that scarlettanagers would move around extensively once they es-

tablished a territory in a fragment. This kind of empir-

ical work challenges long-held assumptions about how

birds behave in fragmented landscapes, how far they

move, and to what extent gaps represent a barrier to

daily movements. To date, three of three ‘‘area-sensi-

tive’’ birds studied (hooded warbler, ovenbird, and

scarlet tanager; Norris and Stutchbury, 2001; Bayne andHobson, 2001a, this study, respectively) are actually not

area-sensitive in terms of movements. Nonetheless,

species may still be considered area-sensitive if their

probability of occurrence increases with habitat patch

size. However, area-sensitive species should not be as-

sumed to avoid edge habitat, rely on corridors, or avoid

crossing open fields (Villard, 1998).

Many researchers emphasize the importance of coreforest area, or forest interior, when assessing how bird

populations and nesting success are affected by frag-

mentation. We suggest that small fragments are more

important to these forest birds than previously appre-

ciated. In our study area, smaller forest patches located

amongst larger forested areas appeared to allow SY

unpaired males to readily move around and prospect for

breeding sites or mates in different locations and theymay be particularly important in regions with higher

population densities. If regional density can be esti-

mated, managers may be able to make informed deci-

sions about how important smaller fragments are for

tanagers and potentially use tanagers as a model for

other Neotropical migrants (Wiens, 1994). In our study

area, all but one of the fragments was privately owned,

which emphasizes the importance of private land in re-serve networks (Wiens, 1994). Landowners educated

about the importance of these woodlots could choose to

make informed decisions on tree harvesting. For ex-

ample, one landowner in the area harvested two large

trees off 3.7 ha fragment, but removed them in a rela-

tively non-destructive manner (using a horse) leaving the

canopy fairly intact.

Rosenberg et al. (1999) stress caution in extrapolatingresults from one study to the whole species because of

the variation in tanager response to fragmentation

among different regions. Observed variation in breeding

386 G.S. Fraser, B.J.M. Stutchbury / Biological Conservation 118 (2004) 377–387

behavior just within one region (US northeast; Roberts

and Norment, 1999, this study) is further support for

caution in suggesting broad recommendations on habi-

tat management for scarlet tanagers. From this study,

we concluded that many of the reasons for movementsin a fragmented habitat are related to breeding adult

dispersal. However, our study also demonstrates the

need to understand: (1) the array of behavioral re-

sponses to fragmentation in various regions with dif-

ferent levels of forest cover and population densities; (2)

whether smaller forest fragments are reproductive

‘‘sinks’’ rather than ‘‘sources’’ (see Pulliam, 1988); and

(3) how fledgling dispersal may be affected by frag-mented habitat.

Acknowledgements

We thank Hemlock Hill Biological Research Area

and all of the land owners for permission to conduct

research. This project was funded by awards to BJMSfrom the Molson Foundation, the Premier�s Research

Excellence Award, and the Natural Sciences and Engi-

neering Research Council of Canada. Bianca Capuano,

Kevin J. Cooper, Ryan Norris, and Mari Ortwerth all

provided excellent field assistance. A special thanks to

M. Ortwerth and K. Cooper for their hard work.

Thanks to Steve Woolfenden for his help on calculating

territory and fragment sizes, and to Eugene Morton andBonnie Woolfenden for lively tanager discussions. Bev

Brown, Laurence Packer, Scott Robinson, Scott Rush,

and two anonymous reviewers contributed useful com-

ments on an earlier draft.

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