Number - ncrs.fs.fed.us · PDF fileBRIAN E POWELL AND ROBI_r J.STRrOL. -2._... -_..-.- ... Recruitment, and Adult Size ... Roost Site Selection of Great Homed Owls in Relation to Black

Volume 102 N_ 2000 ._?_"_)::_-- ._-)_:,-:- Number 4

CONTENTS -::_':: :): ....

FEATURES : -A* :- - _:- ," ..... • -., . .

Nine Years After the Exxon ValderOff _ Effects on _ Bird Populationsin Prince William Sound, Alaska ;: .... _.- ..... "

DAVID B. IRONS, STEVEN _ KE/_ALL, WALLACE E ERICKSON,LYMAN L. McDONALD AND BRIAN K. LANCE . _ _ . . - ... ..... 723 :

Breeding and Post-Breeding Hal_at L_ I_,. Forest Migrant _mgbirds in theMissouri Ozarks " _.-,,:__,_:-._-_,:k.-_;i . ,&-: :. -_.,:'_ ...... - . .... "._RICHW. PAGEN, FRANKR.THOMPSONHIA-N'DDIRKE. BURHA.NS 738 --,_.... :.,--- :.

Breeding Bird Abundance in Sottemla_ _od Fore_: E_bila_ Edge, and _ ":'!-" _-.=-. ._PatchSize Effects ..-_,_-::_._._--;-_,.:-:=.• ::_:-.:-_":::..._.."_ - ..-

REX SALLABANKS, JEFFREYIL WALTERSAND SAIMEA. COLLAZO 748 -: _:.:_.. -.Breeding Bird Communities m Bereal Fen_ of Western Canada: Consequences

of Unmixing the_ _ _._ .........,............-_.....

KEITH A. HOBSON AND _ BAY]_ _(:_,::_¢_'_!_>:_--._ ., . . 759 ....!.ii.,:..::_,,._.. :_,Allozym/cGeneticStructureand NaturalHabitatFragmcn1_/_n:"DataforFive .......,_z_::_-_.

S cc_s of Amazoman Forest Birds ,:__'_"-'_+.._;_-2 :_-_::-"_,_, ":='- :!'::-_ :;)r_ p x_r n ^7_:c • - _:._. ".:'_._-_::_:_-:_:-:_-,": _<-_'"" " . _:............. •........'.............-..... 770 ._:_.:2:;,:._-_-'_?_:_':J%JZJt.Z_I J.VA. JL._.'-XZ4._O ... . . . . o :, . *y?_*_*:...o_ :* *- *-,._ .,.,. _, .-o ,. -.. . . . . . _ _._

Ambird Guildsm theLowlandCa_x_an Rainfo_stofSoutheastNicaragua -'::_,,- :: ,_

MARTIN L.CODY ..............-._+:...............,. :.-............,.. . . 784 --_. ., __!Characteristics of ForagingI-Iabit_and ChickFood Proviskmiag by Tropical :' ,-

RoseateTerns .',_-:_:_-_7,:_,-..,-:--,-:_._:....." -JAIME A. RAMOS ""'_-_:_:-_::'_':,':_:"_":":"_:":-_."- 795

Contrasting Strategies of Prov/sicaing mul Chick Growth in Two Sympatrically :.Breeding Albatrosses at Campbell Island, New Zealand -_- . ....

SUSAN M. WAUGH, HENRI WEIMERSKIRCH, YYES CHEREL AND .,- . . _ :PETER A. PRINCE .... :'_'_"_-_::':"_"_";" " -" : ' 804 -:......... :_'_c_:.7:..'-,-,_..:._.* *-" " e - * * ._ * ..... :

Reproductive Success of American Kestw2s: The Role of Prey Abundance and _;• -.. -.._.:__=.F_7:_:._:.-..... _ ....

Weather - -.,_-._'_:u._:._:_._,-:-•- "--.-: ....-..--, •-:

RUSSELL D. DAWSON AND_R. BORTOLOTTI . :-.-'_::::7",-- 814 " " " '";

NestingHabitatand Reproductive_ ofSouthwesternRiparianBirdsBRIAN E POWELL AND ROBI_r J.STRrOL. -2._... -_..-.-...... 823 -: ,.

. . .

Pair Formation and Copulation Behzv/or in Least Flycatcher ClustersSCOTT A. TAROF AND LAURENE M. RATCLIFFE 832

Artificial Nest Predation and Abundameeof Birds Along an Urbm GradientJUKKA JO_ AND ESAHUHTA....• ......... ....... 838

- atPatterns of Flock Size, Diet, and Vigilanee of Naturalized Meek Parakeets in

Hyde Park,Chicago . .-:.....".... :'_--- •JASON M. SOUTH AND STEPHEN PRUETr-_ONES 848

Do Band-tailed Pigeons Seek a Calcium Supplement at Mineral Sites?TODD A. SANDERS AND ROBERTL. JARVIS ........ ; ..... 855

A Molecular Phylogeny of the Dove Genus Zena_da: Mitochondtal and NuclearDNA Sequences J :

KEVIN P.JOHNSON AND DALE H.CLAYTON .............. 864

MitochondrialDNA and MorphologicalVm_ion ofWhite-wingedDovesinTexasCHJUSTIN L. PRUETT, scorr E. HENKE, SUSAN M. TANKSLEY,MICHAEL E SMALL, KELLY M. HOGAN AND JAY ROBERSON .. . 871

SpeciesLimitsand RecentPopuiatiomHistoryintheCurve-billedThrasherROBERT M. ZINK AND RACHELr_ C.BLACKWELL-RAGO ..... 88l

A ComparisonofPointCountsandSoundRecordingasBirdSurveyMethodsinAmazonian SoutheastPeru - . _ .- ..... .

JOHN HASELMAYER AND JAMES S. QUINN .: . .... . J. ..... 887

DefinitivePrebasicMoltofGray Catbirds at Two Sites in New EnglandCOLLEEN DWYER HEISE AND CHRISTOPHER C. RIMMER ..... 894

(Continued on inside back cover)

Volume 102, Number4 Issued 13 November 2000

:SHORT CO.MM UNIC.VFIONS

CulturalDivcrsitk-ationm theFligd_Call of the RingneckParrotin WesternAusiraliaMYRON C BAKER .............................. 905

Pacific Coast and Southwest Interior Populations of the HuRon's Vireo Differ inBasic Son,, Parameters

COLETTET._BARILAND JON C. BARLOW ............... : 911

Comparison of Co_stal Fringe and Interior Forests as Reserves for MarbledMurrelets on Vancouver Island

ALAN E. BURGER. VOLKER BAHN AND ANGELINE R. M. TILLMANNS 915

Correlates of llarlequin Duck Densities During Winter in Prince William Sound,Alaska

DANIEL ESLER. TIMOTHY D. BOWMAN, THOMAS A. DEAN,CHARLES E. O'CLAIR, STEPHEN C. JEWETT AND LYMAN L.McDONALD ................................. 920

Influence of Female Age and Body Mass on Brood and Duckling Survival,Number of Surviving Ducklings, and Brood Movements in Redheads

TINA YERKES ................................ 926

Effects of Hatching Date and Egg Size on Growth, Recruitment, and Adult Sizeof Lesser Scaup

RUSSELL D. DAWSON AND ROBERT G. CLARK ............ 930

Effects of Water Conditions on Clinch Size, Egg Volume, and Hatchling Mass ofMallards and Gadwalls in the Prairie Pothole Region

PAMELA J. PI_TZ, GARY L. KILAPU, DEBORAH A. BUHL AND DAVIDA. BRANDT .................................. 936

Reproductive Success and Survival in Relation to Experience During the FirstTwo Years in Canada Geese

DENNIS G. RAVELING, JAMES S. SEDINGER AND DEVIN S. JOHNSON 941

Impact of Brood Parasitism on Nest Survival Parameters and Seasonal Fecundityof Six Smagbird Species in Southeastern Old-field Habitat

MARIA A. WHITEHEAD, SARA H. SCHWEITZER AND WILLIAM POST 946

Roost Site Selection of Great Homed Owls in Relation to Black Fly Activity: AnAnti-parasite Behavior?

CHRISTOPH ROHNER, CHARLES J. KREBS, D. BRUCE HUNTER ANDDOUGLAS CoCURRIE ............................ 950

Observations of the Pale-eyed Blackbird in Southeastern PeruGORDON H. ORIANS AND ELIZABETH N. ORIANS .......... 956

Socialitv and Foraging Behavior of the Cerulean Warbler in Venezuelan Shade-Coffee Plantations

JASON JONES, PAOLO RAMONI PERAZZI, ERIN H. CARRUTHERSAND RALEIGH J. ROBERTSON ...................... 958

BOOK REVIEW

Hope is the Thing with Feathers. by Christopher CokinosWALTER D. KOENIG ............................. 963

NEWS AND NOTES ................................. 965

AWARD ANNOUNCEMENTS ............................ 969

REVIEWERS FOR VOLUME 102 .......................... 970

INDEX TO VOLUME 102 .............................. 972

Cover: Band-tallestPigeon (Cohtmbafasciata) landing on a final perch site beforedescending to driak mineralizedwater in the WillametteValley, Oregon. Pigeons arehypothesized to seek dietary,sodium at mineralizeddeposits, or more typically, waterlsee article by SandersandJarvis. p. 855).Photographby WorthMathewson.

Editor: WALTER D. KOENIG

Managing Editor: DAVID J. GUBERNICKEditorial Assistant: RICKEY WOLFE

Book Review Editor: BARBARA KUS

The Condor 102:738 747

© The Cooper Ornithological Society 2000

BREEDING AND POST-BREEDING HABITAT USE BY FORESTMIGRANT SONGBIRDS IN THE MISSOURI OZARKS 1

RICH W. PAGEN

Department of Fisheries and Wildlife Sciences, 302 Natural Resources Building, University of Missouri,Columbia, MO 65211

FRANK R. THOMPSON III 2 AND DIRK E. BURHANS

North Central Research Station, 202 Natural Resources Building, University of Missouri,Columbia, MO 65211, e-mail: [email protected]

Abstract. We compared habitat use by forest migrant songbirds during the breeding andpost-breeding periods in four Missouri Ozark habitats: mature upland forest, mature riparianforest, 9- to 10-year-old upland forest, and 3- to 4-year-old upland forest created by clear-cutting. Adult forest-ground species showed a decrease in abundance in all habitats duringthe post-breeding period, but hatching-year birds of one of the two forest-ground specieswere most abundant in early-successional forest during this time. Adults of the two forest-canopy species tended to increase in abundance in 3- to 4-year-old forest from breedingseason to post-breeding season. During the breeding season, some forest species were de-tected with mist-nets in the two early-successional habitats, but infrequently or not at allwith point counts in those habitats. Forest birds captured in early-successional habitats dur-ing the breeding season may have been nonbreeding floaters, or may have been foragingthere from nearby territories in mature forest. Dense shrubs or young trees in early-succes-sional forest may provide habitat for nonbreeding and post-breeding forest migrant songbirdsin the Missouri Ozarks.

Key words: early-successional, habitat use, Missouri Ozarks, post-breeding, songbirds.

INTRODUCTION adults of many species undergo a complete pre-

Neotropical migrant songbirds are the focus of basic molt (Pyle et al. 1987, Ralph et al. 1993),much attention due to declines in some species which may hinder their ability to fly (Rimmer

(Robbins et al. 1989, Askins et al. 1990). Re- 1988, Vega Rivera et al. 1998a). The availabilitysearch on the breeding, wintering, and migration of quality habitat during the post-breeding pe-ecology of some species is quite extensive (Ha- riod may be critical to survival.gan and Johnston 1992, Finch and Stangel Some recent studies of Wood Thrush (Hylo-1993). There may be other phases of this annual cichla mustelina) using radio-telemetry indicate

cycle, such as the post-breeding period, that are habitats used by hatching-year birds during theequally important to the long term stability and post-breeding period differ from habitats usedsustainability of songbird populations. We de- by adults during the breeding season. Woodfined the post-breeding period in migratory birds Thrush nest in mature upland forest, whereasas the period after nesting or fledging until the hatching-year birds often move to early- andonset of fall migration. This period may last sev- mid-successional forest after fledging (Anders eteral months for hatching-year birds produced al. 1998, Vega Rivera et al. 1998b). Adult Wood

early in the breeding season and for adults that Thrush also may move to early- to mid-succes-raise only a single brood per year (Faaborg et sional habitats or forest edges after breedingal. 1996, Anders et al. 1998). During this period, (Rappole and Ballard 1987, Vega Rivera et al.birds must build up fat reserves and, at the same 1998a).time, avoid predation (Moore et al. 1993). Both Post-breeding habitat use by other forest-

hatching-year and adult birds may be vulnerable breeding Neotropical migrant songbirds remainsduring this time because hatching-year birds largely unknown due to the difficulty in trackinghave little experience foraging on their own, and individuals from breeding territories to post-

breeding habitats, and in detecting non-singingbirds (Faaborg et al. 1996). We examined breed-

Received 13 May 1999. Accepted 11 April 2000. ing and post-breeding habitat use by forest Neo-zCorresponding author, tropical migrant songbirds. We used ANOVA

[738]

SONGBIRD POST-BREEDING HABITAT USE 739

models to determine the effects of habitat, time- were in mixed oak forests, dominated by oakof-season, age of bird, year, and the interactions (Quercus spp.) and hickory (Carya spp.). Dog-of these effects, on the relative abundance of wood (Comus florida), cherry (Prunus seroti-each species. We hypothesized that the abun- na), and sassafras (Sassafras albidum) also weredance of post-breeding adult and hatching-year present in these forests. Common shrubs includ-

forest birds would be greater in early-succes- ed black raspberry (Rubus occidentalis), poisonsional habitats than in mature forest, ivy (Toxicodendron radicans), fragrant sumac

(Rhus aromatica), and buckbrush (Symphoricar-METHODS pos orbiculatus). Riparian sites were dominated

STUDY AREA by elm (Ulmus spp.), box elder (Acer negundo),maple (Acer spp.), and sycamore (Platanus oc-

This study was conducted in 1997 and 1998 on cidentalis). Ash (Fraxinus spp.), paw paw (Asi-

the Houston-Rolla Ranger District of the Mark mina triloba), and hackberry (Celtis occidental-Twain National Forest (Pulaski, Laclede, and is) also were present, and spicebush (LinderaTexas Counties), and on Horseshoe Bend Natu- benzoin) was the most common shrub. Groundral Area, a Missouri Department of Conserva- cover was dense with forbs such as nettle (La-tion managed property owned by the Lad Foun- portea canadensis), giant ragweed (Ambrosiadation (Texas County). This area is on the Ozark trifida), and wingstem (Verbesina spp.).Plateau in south-central Missouri and is char-

acterized by rolling terrain dissected by numer- BIRD SURVEYS

ous streams. The area is heavily forested; the We surveyed bird communities by point countsthree-county area we worked in is approximately and constant-effort mist-netting. We used the

54% forested (Miles 1990, Smith 1990). The point count method because of its ability to lo-forest is primarily mature, even-aged forest that cate breeding birds (Ralph et al. 1995). We used

regenerated after extensive clearcutting during mist-netting because many Neotropical migrantthe early 1900s. Young forest stands created by songbirds are nearly silent during the post-recent timber harvests are interspersed through- breeding period (Faaborg et al. 1996), and visi-out the area. Non-forested land consists of cattle bility can be poor in early-successional habitatspastures and small towns, because of dense vegetation. Although problems

We studied bird abundance in four habitats: associated with mist-netting and point counts aremature upland forest, mature riparian forest, 9- well documented, use of both methods shouldto 10-year-old upland forest, and 3- to 4-year- reduce problems (Rappole et al. 1993, Gram andold upland forest. Three sites (replicates) of each Faaborg 1997).habitat were surveyed during 1997, and three Point counts. We conducted point counts fivedifferent sites of each habitat were surveyed in times at each site between 21 May and 17 Au-1998 for a total of 24 sites in two years. Study gust 1998, with approximately 2.5 weeks be-

sites were 2.5 to 5.2 ha, surrounded by forest, tween visits to a site. We located four evenlyand spatially dispersed throughout the study area spaced point count stations at each site. Each(Pagen 1999). point was surveyed for 10 rain, and birds were

Vegetation characteristics such as canopy cov- recorded as inside or outside a 50-m radius cir-

er, basal area, and ground cover were signifi- cle. All birds heard or seen within the study sitecantly different among the four habitats (Pagen were recorded.

1999). The mature upland and mature riparian Mist-netting. We surveyed each site with 10sites had not received any silvicultural treatment nets in 1997 and 12 nets in 1998. The distance

for at least 50 years. Both 3- to 4-year-old and between net lanes was kept constant between9- to 10-year-old forest were created by clear- years. At each site, we located net lanes approx-cutting and had abundant natural tree regenera- imately 50-m apart and at least 25-m from anytion. The 3- to 4-year-old and 9- to 10-year-old habitat edge. Net lanes were arranged as system-

forest had on average 10% and 79% canopy clo- atically as possible within each site so that theysure and a 2.5-m and 5.8-m canopy height, re- sampled comparable areas. Nets were 30-mm

spectively. Mature upland and riparian forests mesh, 12-m long, and approximately 2.6-m high.had approximately 90% canopy closure and a We surveyed birds five times between 22 May19-m canopy height (Pagen 1999). Upland sites and 21 August in 1997, and five times between

740 RICH W. PAGEN ET AL

21 May and 17 August in 1998, with approxi- ing a repeated-measures ANOVA specified as amately 2.5 weeks between visits to a site. Each nested model (SAS Institute 1990). We consid-visit entailed operating net lines for approxi- ered factors in the ANOVA model significant atmately 5.5 hr beginning 20 min before sunrise, a P-value < 0.1 to increase statistical power. WeWe banded birds with a U.S. Fish and Wildlife used the least-significant difference test

Service aluminum band and recorded the spe- (LSMEANS; SAS Institute 1990) to test for dif-

cies, age (hatching-year or adult), and sex. ferences among the levels of each factor in theBreeding condition was noted as the presence/ ANOVA and a sequential Bonferroni test (Riceabsence of a brood patch or cloacal protuber- 1989) to adjust the significance level of each testance. Banding protocol followed Ralph etal. to control for experiment-wide error rates at a(1993). P-value < 0.05.

If post-breeding adult and hatching-year for-STATISTICAL ANALYSES est birds prefer early-successional habitats, we

We selected six species for analysis that were predicted a significant habitat x time-of-seasonamong the most abundant and had small confi- interaction in the ANOVA model, and that de-dence intervals for mean detections in each hab- tections should be greater in 3- to 4-year-old for-

itat: Red-eyed Vireo (Vireo olivaceus), Northern est and 9- to 10-year-old forest than in either ofParula (Parula americana), Acadian Flycatcher the mature forest habitats during the post-breed-

(Empidonax virescens), Ovenbird (Seiurus au- ing period. However, because point counts arerocapillus), Worm-eating Warbler (Helmitheros less effective later in the season (Skirvin 1981),vermivorus), and Kentucky Warbler (Oporornis we were mainly interested in using point-count

formosus). We also analyzed two guilds of spe- data to determine abundance by habitat duringcies because confidence intervals were particu- the breeding season. Conclusions on post-breed-

larly small when data for several species were ing habitat use were based primarily on mist-pooled. These guilds were: forest-ground species netting results.(Ovenbird, Worm-eating Warbler, and Wood Mist-netting. We standardized net captures forThrush), and forest-canopy species (Red-eyed each visit to a site as captures 100-net-hr _. AsVireo, Yellow-throated Vireo [Vireo flavifrons], with point counts, visits were pooled and clas-Yellow-throated Warbler [Dendroica dominica], sifted as early or late season. We analyzed theNorthern Parula, Yellow-billed Cuckoo [Coccy- same species and species guilds used in the pointzus americanus], and Scarlet Tanager [Piranga count analysis. We screened all data for nor-

olivacea]). Kentucky Warbler, although clearly a mality and constancy of variance and log-trans-ground bird, was not included in the forest- formed data prior to analysis (Neter etal. 1996).ground guild because it is known to breed in We analyzed the effect of habitat, time-of-sea-early-successional forest (Thompson and Fritzell son, age of bird, year, and their interactions on1990). mean captures using a repeated-measures AN-

Point counts. We calculated mean detections/ OVA (SAS Institute 1990). As with point

10-rain point count for each species from the counts, we used the least-significant differencefour points surveyed during each visit to a site. test to test for differences among the levels ofWe used all detections (aural and visual) within each factor, and the sequential Bonferroni test toa 50-m radius for the calculations. Confidence control experiment-wide error rates.

intervals for mean detections point -_ for each We predicted a significant habitat X time-of-habitat decreased when individual visits were season, and habitat X time-of-season X age

pooled, so we classified visits 1 and 2 as "early class effect in the ANOVA model, and that cap-season" (which approximates the breeding sea- tures of both adults and hatching-year birdsson), visits 4 and 5 as "late season" (which ap- should be greater in 3- to 4-year-old forest and

proximates the post-breeding period), and 9- to 10-year-old forest than in either of the ma-dropped visit 3 from the analysis, ture forest habitats during the post-breeding pe-

We used a square-root transformation on riod. We also predicted that abundance of adultspoint-count data to improve normality and con- should decrease in mature forest and increase instancy of variance (Neter et al. 1996). We ana- early-successional habitats from early season tolyzed the effect of habitat, time-of-season, and late season.their interaction on mean detections point _ us- Forest-canopy species are often not detected


TABLE 1. Mean (_+ SE) abundance of forest birds detected with point counts during the breeding and post-breeding period in four habitats in Missouri, 1998. Numbers with different letters indicate significantly differentmeans among habitats within a period (P < 0.05); rows without letters are not different.

Habitat

Species Period Mature Riparian 9-year-old 3-year-old

Red-eyed Vireo Breeding 0.71 _+0.23a 0.42 _+0.04a 0.13 _+0.07b 0.00 Z 0.00bPost-breeding 0.71 _+0.00 0.29 _+0.23 0.50 - 0.26 0.00 - 0.00

Northern Parula Breeding 0.04 _+0.04 0.25 _+0.14 0.08 _+0.08 0.00 _+0.00Post-breeding 0.00 _+0.00 0.08 -+ 0.04 0.13 _+0.13 0.04 _+0.04

Acadian Flycatcher Breeding 0.04 _+0.04a 0.50 _+0.21b 0.00 - 0.00a 0.00 + 0.00aPost-breeding 0.08 + 0.08a 0.38 _+0.13b 0.00 _+0.00a 0.00 +_0.00a

Ovenbird Breeding 0.12 _+0.15 0.00 _+0.00 0.29 + 0.29 0.00 _+0.00Post-breeding 0.13 _+0.13 0.00 _+0.00 0.04 _+0.04 0.04 _+0.04

Worm-eating Warbler Breeding 0.08 _+0.08 0.04 _+0.04 0.04 + 0.04 0.04 + 0.04Post-breeding 0.04 + 0.04 0.13 _+0.00 0.13 _+0.00 0.07 _+0.17

Kentucky Warbler Breeding 0.00 _+0.00 0.08 _+0.04 0.17 _+0.08 0.08 + 0.04Post-breeding 0.00 _+0.00a 0.25 + 0.07b 0.13 + 0.07ab 0.13 _+0.07ab

Forest canopy guild Breeding 1.00 _+0.30 0.88 _+0.07 0.46 _+0.08 0.29 _+0.04Post-breeding 1.13 _+0.25 0.58 + 0.18 1.12 _+0.33 0.08 _+0.04

Forest ground guild Breeding 0.29 _+0.15 0.04 _+0.04 0.46 2 0.40 0.04 _+0.04Post-breeding 0.17 ___0.17a 0.17 _+0.04a 0.17 _+0.08a 0.21 _+0.15b

well with mist-nets in habitats with a high can- season effects (P < 0.1 and P < 0.01, respec-opy (Karr 1981, Petit et al. 1992), so a compar- tively). Three species (Northern Parula, Oven-

ison among habitats may be biased (Remsen and bird, Red-eyed Vireo) and both guilds had sig-Good 1996). Therefore, we only interpreted re- nificant habitat X time-of-season effects for

suits of the ANOVA for differences in the abun- mist-net detections (all P < 0.05), and four spe-dance of canopy species within habitats over cies and one guild had significant age-of-bird xtime and between adults and hatching year birds habitat x time-of-season effects (Kentucky War-in the same habitat, bier P < 0.01; Ovenbird P < 0.1; Wood Thrush

Red-eyed Vireo, Acadian Flycatcher, Oven- and Worm-eating Warbler P < 0.05; forest-bird, Worm-eating Warbler, and Kentucky War- ground species group P < 0.1). Non-trans-bier were detected regularly with mist-nets in at formed point count and mist-netting data for

least one of the two early successional habitats each species and guild are presented in Table 1during the early season. To determine why these and Figure 1.forest breeding birds were present in early-suc- We found four major patterns of abundance:

cessional habitat, we determined the frequency first, some species were more abundant in early-of breeding condition and sex of adults captured successional habitat than mature forest duringduring the early season in mature forest and in the late season. Significantly more hatching-yearearly-successional habitats. We also calculated Worm-eating Warblers were found in 3- to 4-the frequency of recaptured adults in mature and year-old forest and 9- to 10-year-old forest thanearly-successional forest during the breeding in mature upland forest during the late seasonseason. We calculated recapture frequencies for (Fig. 1). Hatching-year birds of the forest-visits 1-3 rather than for just visits 1 and 2 to ground guild were more abundant in 3- to 4-increase the likelihood of recapturing birds that year-old forest and 9- to 10-year-old forest than

remained in the area during the breeding season, in either of the mature habitats during the lateDue to small sample sizes and potentially low season (Fig. 1). Several other species were pre-

statistical power, we considered P-values of -< sent in early-successional habitat during the late0.1 as significant, season as well. We did not compare differences

in canopy species among habitats. Rather, weRESULTS compared how abundance in each habitatBased on point count detections, one species changed from the early season to the late season.

(Red-eyed Vireo) and one guild (forest canopy The number of adult Northern Parulas, Red-eyedspecies group)had significant habitat × time-of- Vireos, and forest-canopy birds tended to in-

742 RICH W. PAGEN ET AL.

Red-eyed Vireo Ovenbird9

A A A A A A A A 5 A A A A A A A A8A A B AB ABa Aa Bl2abCb AB A A Aa Aa Aa Aa

7 46

5 I 34 '

MU RP 9Y 3Y MU RP 9Y 3Y MU RP 9Y 3Y MU RP 9Y 3Y

Worm-eatingWarbler NorthernParula4.0 2.5

A A A A A A A A A A A A A A A A3.5 AB A B AB Aa AabAc Abe A A A A Aa Aa ABa Ba

3.0 _ T 2.0

1.5 1.0

0.5

o.o _ o.oMU RP 9Y 3Y MU RP 9Y 3Y MU RP 9Y 3Y MU RP 9Y 3Y

KentuckyWarbler AcadianFlycatcher

A A A A A A A A A

5 A A A A A B A A 2.0 A A A A Aa Aa Aa A aA B AB a A b A ab Aab4

1.5


Forestgroundguild Forestcanopyguild10 A A A A B A B A 12 ] A A A A A A A

A A B A A a A a A b A b 10 _ A A B AB ABa A a BCabq b8I

° :14 j


FIGURE 1. Mean abundance (1997, 1998) of bird species and guilds detected with mist-netting during breedingand post-breeding periods in mature upland forest (MU), mature riparian forest (RP), 9- to 10-year-old uplandforest (9Y), and 3- to 4-year-old upland forest (3Y). Bars with the same letters are not significantly different at


crease in 3- to 4-year-old forest from early to Captures of adult Worm-eating Warblers alsolate season (Fig. 1). Hatching-year birds of these were greatest in 9- to 10-year-old forest duringthree species/guilds also were present in the two the early period (Fig. 1). The forest-ground guildearly-successional habitats during the late sea- showed no habitat differences with point counts,son. but captures of adults were higher in 9- to 10-

A second pattern was that adults of some spe- year-old forest than in any other habitat during

cies/guilds decreased in abundance in all habi- the early season.tats during the late season. Numbers of adult Information on sex ratio, evidence of breedingOvenbirds and Worm-eating Warblers in 9- to condition, and frequency of recapture may help10-year-old forest tended to decrease from the explain why individuals of six species were pre-early to the late season (Fig. 1). Abundance of sent in early-successional habitat during the ear-adult Kentucky Warblers and adult forest-ground ly season (Table 2). Red-eyed Vireos and Ken-birds also tended to decline in all habitats from tucky Warblers had a more male-biased sex ratiothe early season to the late season, with a sig- in early-successional habitat than in mature for-nificant decrease of Kentucky Warblers in ma- est, whereas Ovenbirds and Worm-eating War-ture riparian forest, and a significant decrease of biers were more male-biased in mature forestforest-ground birds in mature upland forest and (Table 2). Red-eyed Vireos and Ovenbirds had9- to 10-year old forest (Fig. 1). a higher frequency of breeding condition in ma-

A third pattern was that some species were ture forest than in early-successional forest,captured in both mature forest and early-succes- whereas Kentucky Warblers had higher frequen-sional habitat during the early season, but were cy of breeding condition in early-successionalonly detected with point counts in mature forest habitat. Acadian Flycatcher, Kentucky Warbler,habitats at that time. Detections point -_ of Red- and Ovenbird had a higher frequency of recap-

eyed Vireos were greatest in mature upland for- ture in mature forest than early-successionalest during the early season, with no Red-eyed habitat, whereas Worm-eating Warblers showedVireos detected in 3- to 4-year-old forest (Table the opposite trend (Table 2). Because we could1). However, Red-eyed Vireos were frequently not reliably determine the sex of Acadian Fly-captured in the two early-successional habitats catchers, we did not report the proportion thatduring the early season. Detections point _ of were male or in breeding condition.Acadian Flycatchers were significantly higher inmature riparian forest than in all other habitats DISCUSSIONduring the early season (Table 1). No Acadian We found adult and hatching-year birds of sev-Flycatchers were detected with point counts in eral forest-nesting species in early-successionaleither of the two early-successional habitats, habitats during the post-breeding period andHowever, captures 100-net-hr -j of Acadian Fly- breeding season. Our results also highlight thecatchers were not significantly different between value of multiple survey methods. Mist-nets de-habitats in either time-of-season (Fig. 1). tected non-singing individuals of several species

Finally, some species were detected with both in early-successional habitat (Red-eyed Vireo,point counts and mist-nets in mature forest and Acadian Flycatcher), whereas point counts de-early-successional habitat during the early sea- tected several canopy species in mature forestson. Detections point _ of Ovenbirds tended to that could not be effectively netted in those hab-be highest in 9- to 10-year-old forest, followed itats (Northern Parula, Red-eyed Vireo).by mature upland forest (Table 1). Netting datashowed a similar trend (Fig. 1). Significantly POST-BREEDING SEASON HABITAT USEmore adult Ovenbirds were captured in 9- to 10- Our prediction that forest birds would becomeyear-old forest than in either 3- to 4-year-old for- more abundant in early-successional habitatsest or riparian forest during the early season, during the post-breeding period was true for sev-

,(.__

P < 0.05. The top rows of letters in each graph are comparisons of a habitat between breeding (left) and post-breeding (right) periods. The bottom rows of letters are comparisons among habitats within a period. Capitalletters compare adults (open bars); lower case letters compare hatching-year birds (hatched bars).


TABLE 2. Proportion of birds captured with mist-nets that were male (Male), in breeding condition (BC), andrecaptured (Recap) in mature forest (mature upland and mature riparian forest) and early-successional forest (3-to 4-year-old upland forest and 9- to 10-year-old upland forest) in the Missouri Ozarks, 1997 and 1998.

Mature forest Early-successional forest

Species Frequency n Frequency n

Red-eyed Vireo Male 0.50 10 0.65 34BC 0.90 10 0.77 43Recap 0.00 15 0.02 53

Acadian Flycatcher Male ....BC ....Recap 0.45 11 0.00 12

Kentucky Warbler Male 0.60 25 0.81 26BC 0.89 28 1.00 25Recap 0.32 25 0.15 41

Ovenbird Male 0.83 6 0.67 21BC 1.00 6 0.95 22Recap 0.22 9 0.14 29

Worm-eating Warbler Male 0.60 5 0.44 18BC 1.00 5 0.77 22Recap 0.00 7 0.07 41

eral species and, in particular, for hatching-year 4-year-old forest from early to late season. Thisbirds. In contrast, adults of each of the forest- increase in abundance suggests that when noground species (Ovenbird and Worm-eating longer limited by nesting requirements, theseWarbler) showed no difference in abundance be- species may move to other habitats. Morsetween habitats during the post-breeding period. (1967) reported that Northern Parulas in Loui-

We did not expect to detect many adults of these siana began foraging in the forest understory af-species with point counts because songbirds vo- ter their young had fledged the nest. Becausecalize very little during the post-breeding period. Northern Parulas reside in second growth habi-However, mist-netting also detected few adult tats during the winter (Lynch 1989), it is not

forest-ground birds in any habitat at this time. surprising that they would use early-succession-There are several explanations for low detec- al habitats during the post-breeding period as

tion rates of adult forest-ground birds during the well.

post-breeding period. First, birds may be molt- Anders et al. (1998) found juvenile Wooding, which would constrain their ability to fly Thrush used early-successional habitat during(Rimmer 1988, Vega Rivera et al. 1998a) and, the post breeding period; juveniles used earlyin turn, to encounter nets. Vega Rivera et al. successional habitat proportionally more than its(1998a) found that adult Wood Thrush hid in availability in the landscape and experienced nodense vegetation during the post-breeding period mortalities after dispersal there (Anders et al.

and were very difficult to observe or capture. 1997). The high density of vegetation in early-Second, the amount of movement and foraging successional habitats may provide a greatermay be reduced because adults no longer have abundance of food and shelter compared to ma-to provision nestlings, leading to lower capture ture forest habitats (Hollifield and Dimmickrates. Third, adults may have left the study area 1995). Birds may move into these habitats dur-

altogether. Adults sometimes leave breeding ter- ing the post-breeding period when they are noritories prior to migration (Nolan 1978, Rappole longer limited by nesting requirements that areand Ballard 1987); Nolan reported several cases found only in mature forest habitats.in which Prairie Warblers (Dendroica discolor)left territories soon after breeding to return from BREEDING SEASON HABITAT USE

unknown locations in September or October pri- In several cases, forest migrant songbirds wereor to migration, as abundant or more abundant in early-succes-

Adults of both Red-eyed Vireos and Northern sional forest than in mature forest during theParulas tended to increase in abundance in 3- to breeding season (see also Thompson et al.


1992). For example, breeding season point tad 1989), and hence would be easily missed bycounts and mist-netting indicated that Oven- point counts (Brown 1969). Nonbreeding birdsbirds, Worm-eating Warblers, and the forest- are most likely to be males (Brown 1969, Marra

ground bird guild were most abundant in 9- to and Holmes 1997), are not always in breeding10-year-old forest, with lesser numbers in ma- condition, and have a low chance of being re-ture upland forest. Other species were detected captured because they are not territorial (Welsh

with point counts almost exclusively in mature 1975). Acadian Flycatchers and Red-eyed Vir-forest during the breeding season, but were de- eos captured in early-successional habitat duringtected with mist-netting in early-successional the breeding season were likely floaters, al-forest as well. Red-eyed Vireos, for example, though some captured during May may havemay not have been detected with point counts in been late transients (Rappole et al. 1993). Both

early-successional habitat during the breeding species exhibited a low frequency of recaptureseason either because they are difficult to see in in early-successional forest, had a lower fie-

dense vegetation, or because they are silent quency of breeding condition, and Red-eyedwhen in those habitats. We detected Acadian Vireos had a sex ratio biased toward males (Ta-Flycatchers singing most often in mature ripar- ble 2).

ian forest, but did not detect them singing in Forest birds detected in early-successionaleither of the two early-successional habitats, forest during the breeding season also could be

Netting data, however, showed Acadian Fly- breeding in mature forest nearby and foraging incatchers were present in early-successional hab- early-successional forest. Mature-forest breedingitat during the early season, birds foraging in early-successional forest

Some forest birds likely bred in early-succes- should show evidence of breeding condition,sional forest. A forest bird likely bred in early- and may or may not show a male-biased sex

successional habitat if it was detected with both ratio. Because they are central place foragersmist-netting and point counts (singing) in that (Perrins and Birkhead 1983), they should havehabitat during the breeding season. Recapturing a greater chance of being recaptured at a partic-an individual bird at a particular site would also ular site than a floater (Brown 1969). Worm-eat-

suggest a bird was territorial (breeding). For ex- ing Warblers were occasionally recaptured at 3-ample, Ovenbirds and Worm-eating Warblers to 4-year-old forest sites during visits 1 to 3, and

were detected by both mist-netting and point did not exhibit a male-biased sex ratio duringcounts (singing) in 9- to 10-year-old forest, and the early season. Therefore, it is possible thatOvenbirds were recaptured at 9- to 10-year-old Worm-eating Warblers may nest in mature forest

forest sites. Kentucky Warblers were detected and occasionally forage in 3- to 4-year-old forestsinging and were recaptured in both early-suc- nearby, or may establish territories that overlapcessional habitats. We believe that Ovenbirds, with both mature forest and early-successionalWorm-eating Warblers, and Kentucky Warblers forest.at least occasionally breed in early-successionalforest, which is consistent with some other stud- CONSERVATION IMPLICATIONS

ies (Thompson and Fritzell 1990, Thompson et Maximizing the amount of mature forest habitat

al. 1992). More detailed study of pairing success in the landscape at the expense of early-succes-and nesting success among the habitats we stud- sional habitat may not always be desirable.

ied is necessary to determine the quality of these Many Neotropical migrant songbirds typicallyhabitats as breeding habitat for forest birds described as forest species were present in early-(Gibbs and Faaborg 1990, Morse and Robinson successional habitats during the breeding season,

1999). post-breeding season, or both. Maximizing ma-Some forest birds in early-successional forest ture forest habitat ignores post-breeding habitat

during the breeding season may be nonbreeding requirements, and may affect forest bird speciesfloaters (Brown 1969, Smith 1978). Early-suc- that rely on early-successional habitat during thecessional habitats may be attractive to floaters breeding season. Due to the very specific habitat

because of the lack of conspecific breeding ter- requirements of early-successional bird species,ritories, and abundant food and dense cover this strategy will obviously impact them nega-(Hollifield and Dimmick 1995). Because these tively as well. If increasing forest bird popula-birds are not territorial, they do not sing (Hogs- tions is a management goal, some mix of mature


and early-successional forest habitats may pro- Midwest: post-breeding and wintering periods, p.vide optimal habitat for most forest birds be- 189-199. In E R. Thompson III lED.], Manage-

ment of midwestern landscapes for the conserva-cause it addresses both breeding season and tion of Neotropical migratory birds. USDAForestpost-breeding season habitat needs. The appro- Serv. Gen. Tech. Rep. NC-187, St. Paul, MN.priate amount will vary depending on the extent FINCH, D. M., ANDP. W. STANGEL[EDS.]. 1993. Statusand types of other land cover in the area, and and management of Neotropical migratory birds.specific conservation objectives. For example, in USDA Forest Serv. Gen. Tech. Rep. RM-229, FortCollins, CO.a fragmented agricultural landscape, increasing GraBS, J. P., AND J. FAABORG.1990. Estimating the vi-early-successional forest at the expense of ma- ability of Ovenbird and Kentucky Warbler popu-ture forest may be detrimental because of a de- lations in forest fragments. Conserv. Biol. 4:193-crease in already limited nesting habitat and a 196.potential increase in nest predation and parasit- GRAM, W. K., ANDJ. FAABORG.1997. The distributionof Neotropical migrant birds wintering in the E1ism rates for forest birds (Donovan et al. 1995, Cielo Biosphere Reserve, Tamanlipas, Mexico.Thompson et al., in press). However, some ear- Condor 99:658-670.ly-successional habitat in a mostly forested land- HAGAN, J. M., III, ANDD. W. JOHNSTON[EDS.]. 1992.scape may be beneficial to forest birds because Ecology and conservation of Neotropical migra-

tory landbirds. Smithson. Inst. Press, Washington,of foraging and shelter benefits without a sig- DC.nificant effect on nest predation or parasitism HOGSTAD, O. 1989. The presence of non-territorialrate (Thompson et al. 1996, Donovan et al. males in Willow Warbler Phylloscopus trochillus1997). populations: a removal study. Ibis 131:263-267.

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