United States 

Department of 

Agriculture 

Forest Service 

Southeastern Forest 

Experiment Station 

Research Paper 

S E-259 

Forest Stands Selected 

by Foraging 

Red-Cockaded Woodpeckers 

Robert G. Hooper and Richard F. Harlow 
Forest Stands Selected by Foraging 

Red=Cockaded Woodpeckers 

Robert G. Hooper, Wildlife Biologist 

Charleston, South Carolina 

Richard F. Harlow, Wildlife Biologist1 

Clemson, South Carolina 

ABSTRACT 

Selection of forest stands by 18 clans of foraglng 

red-cockaded woodpedtsrs (Plcoldes boreal is) was studfed in their year-round home ranges on the Francfs 

Marfon National Forest in South Carolina. The foragfng use of 276 stands relative to thelr avallablllty 

wfthln the home ranges was compared to several stand 

characteristics. Selection among stands wfth slml lar 

characteristics was highly variable and red-cockaded 

woodpeckers foraged In stands wfth a greaC varlety of 

condftions, suggesting the birds were not very selective. Although stand selection Increased sfgniflcantly as the density of pines 224 cm d.b.h. increased 

and as the basal area of hardwoods 212 cm d.b.h. dw 

creased, these relationshfps were weak. Beyond 30 

years, stand age was not found to be a slgnlffcant 

factor in stand selectlon. 

Keywords: Endangered specles, foraging habltat, habitat selection, loblolly pine, longfeaf pfne, Picoides 

borea l is. 

Foraging red-cockaded woodpeckers 

these various ways (Hooper and others 

1982). Within the limits set by soil 

and other natural site factors, conditions of forest stands used by redcockaded woodpeckers are determined to a 

great extent by forest management. A 

k now1 edge of stand conditions sel ected 

by the bird for foraging enhances assessment of management options for 

this endangered species. This study 

compared selection of forest stands by 

foraging red-cockaded woodpeckers to 

stand age, density and size of pine 

stems, and basal area of hardwoods. 

Met hods 

The study was conducted on the 

Francis Marion National Forest in South 

Carolina, With more than 400 clans, the 

area has one of the largest popul ations 

of red-cockaded woodpeckers (Lennartz 

and others 1983). The size and apparent 

stabil ity of the population suggest 

relatively good habitat conditions for 

the species on the study area. 

clearly prefer pines over other trees Stands of loblolly (Pinus taeda) and tend to select trees that are larg- and longleaf (P. pal ustriS)pinescur er than the average available ones on the higher Xd better drained sites, (Detotelle and others 1983; Hooper and interspersed with hardwood and cypress Lennartz 1981; Porter and Labisky 1986; (Taxodium spp.) stands in drainages and Skorupa 1979). Pines grow in a variety l ow areas. The most common hard~gods of conditions from stands of nearly pure are tupelo (Nyssa spp.), oaks (Quercus pines to those with a few pines scattered among hardwoods. Even stands that spp.) , red mF(Acer rubrum), sweetgum 

are dominated by pines differ in the ( Li quidambar styraciflu-dbay 

(Persea borbonia) , magnol ias (Magnol ia amount of hardwood understory and mid- s'mand holl ies (Ilex spp.). Stands story, and they vary in the density, are second growth and essentially even- size, and age of the pines. Year-round aged; however, some have scattered older home ranqes of red-cockaded woodpeckers 

a re large and are typical ly composed of 

a mosaic of forest stands that differ in INOW retired. 
trees in the overstory or an understory 

of younger trees. Since 1944, pine 

stands have been prescribed burned on 

1- to 4-year intervals. Even-age timber 

management has been practiced on the 

National Forest since 1950, 

Eighteen different clans (six/year) 

of col or-banded red-cockaded woodpeckers 

were studied between May 1976 and April 

1979, Clans were family groups of two 

to nine birds consisting of the breeding 

pair, juveniles, and adult male helpers. 

Clans were followed 2 to 14 hours/day on 

454 days for a total of 2,350 hours. 

Birds were out of their cavities from 9 

to 14 hours, depending on time of year, 

and the average time spent following a 

clan was 5.2 hours/day. Thus, sampl ing 

was skewed toward the morning hours, and 

a bout 80 percent of the observations 

were collected in the first 5 hours 

after the birds left their roosts, 

At 5-minute intervals, the coordinates of each clan were determined 

from 1 :24,000 scale aerial photographs 

gridded to a scale of 15,2 m with each 

10th 1 ine shown on the photographs. 

Major grid points (152 x 152 m) were 

marked in the forest when sufficient 

natural features for determining location of the clans were lacking. After 

del ineation of the home ranges (Hooper 

and others 1982), maps showing forest 

stands were prepared for each home 

range. Stands were defined as areas 

with aggregations of trees and other 

vegetation of sufficient uniformity in 

composition, age, and density to be 

distinguished from the vegdation and 

conditions of adjoining stands, Coordinates for the foraying locations of 

clans were matched with stand coordinates to determine the use of each 

s land, 

Although red-cockaded woodpeckers 

are territorial, we were unable to de1 ineate enti re territorial boundaries 

(Hooper and others 1982); however, most 

clans did share some mutual boundary 

with adjacent clans. If an adjacent 

clan was not apparently aware of it, a 

clan would sometimes violate the boundary and travel into the adjacent territory. Although foraging occurred on 

these extraterritorial trips, areas 

across territorial boundaries were not 

considered avail able foraging habitat, 

Stands ranged in sire from 0.5 to 

35,9 ha, By chance alone, larger stands 

would be expected to receive more foraging use than smaller stands, Thus, to 

Gompare foraging use among stands, stand 

size was taken into account by calculation of a uselavai labil ity ratio (UAR) 

for each stand: 

where: 

UARai = the UAR for stand i in home 

range a 

Ra i = the number of Foraging observations in stand i of home range a 

A,! = the area of stand i in home range 

a 

Because on most days we did not 

fol low clans a1 1 day, results could be 

biased in favor of forest stands close 

to the roost trees and against stands 

near the home range boundary. To test 

for bias, we compared the length of 

dai ly observation period to the proportion of observations recorded in stands 

various distances from the colony, In 

10 home ranges we followed clans a1 l day 

Or) 48 days, These observations were 

free of bias that could have resulted 

from fol lowing birds for only part of a 

day, Thus, the proportion of observations in a stand, based on the sum of 

observations for whole-day monitoring 

within a home range, was considered an 

unbiased estimate far the comparison, 

Whole days were divided into I-hour 

periods, and the number of observations 

for each of the stands was summed wl'thii? 

hourly periods, Beginning each time 

with the first hourly period, these sums 

were aecumul ated across hour1 y periods 

to create partial -day observation periods of 1 hour, 2 ho~~s, 3 hours, and so 
on, Next, the proportion of nbservaLions in each stand for the observation 

periods was determined. The deviation 

of the estimate for the partial-day 

observation From the who1 @-day estimate 

for each stand was determined by subtracting the whole-day value from each 

partial-day estimate, Thus, for @ach of 

140 stands in these 10 home ranges, a 

deviation from the unbiased who1 e-day 

estimate was obtained for each length of 

observation, The mean of the distances 

from the breeding male's roost tree to 

the proximal and dish1 boundary of each 

0% t * * 

I i Hour 

041 : * 

* re * * * **' 1 -"I **** * "*' * 

I * -" 

-03 1' 

0 200 400 600 BOG IOOG I200 140C 

stand was determined. The deviations 

for individual stands were plotted by 

mean stand distance for each of the 

created partial -day observation periods, 

Figure 1 shows results for periods 

ending in odd-numbered hours, 

Consistently higher or 1 ower esti - 

mates by partial -day observation periods 

when compared with who1 e-day estimates 

woul d suggest that parti a1 -day estimates 

are biased, Such appeared to be the 

case if birds were fol lowed for only 1 

hour (fig, 1). Stands within 200 m of 

0 5 

o 4 7 Hours 

9 Hours 

a;? L. i 01 1 * * 

**I* 

- i -02 /. 

-03 li 1 

0 ZOO 400 609 800 1000 t2CO 1400 

MEAN STAND D 

Figure I*--8evlatlon of partial-day frm whole-day estfma$es of Phe proportion sf foraging time spent 

by red-cockaded weo+ecksrs In 140 forest stands a9 varlsus distances frm the breedfng males' roost 

cav:tles. 4 pasitfve devlatian indicates an inflated partial-day estimate, and a negative dsvlaPIon 

indicates an underestimate. Data were calculated from 48 whole days of observation in 18 h 

an Phe Francis Marion Naf-fonal Forest In South Carolina* 
the breeding ma1 e' s roost tree appeared 

to have inflated estimates of foraging 

time, whereas stands at more than a 

600-m distance appeared to have underestimated values. If birds were followed 5 hours, this trend was no longer 

apparent. Fol lowing birds longer than 5 

hours reduced the variation in observed 

values, but we used 5-hour days as the 

minimum to include in the analysis. By 

using 5-hour and longer days, the sampling intensity was twice that of 7-hour 

and 1 onger days : 283 days and 1,910 

hours vs. 109 days and 971 hours, respectively. Data collected on f 71 days 

(442 hours) were not used in the analysis because they were col lected on days 

when we followed clans less than 5 

hours. 

Characteristics of the vegetation 

were measured in the 276 stands contained within the home ranges of the 18 

clans. Depending on stand size, 3 to 20 

sampl ing points were located systematical ly from a random starLing point, 

Each sample point served as the center 

of a I-m2/ha factor wedge prism plot and 

a fixed radius plot of 0.02 ha. The 

prism plot sampled trees >12.0 cm d.b.h. 

and the 0.02-ha radius plot sampled 

stems 3.0 to 71.9 cm d.b.h. All stems 

were recorded by species and d.b.h. to 

allow calculation of basal area and density of stems for hardwood, pines, and 

cypress. Stand age was determined by 

increment boring a dominant tree on each 

plot and averaging plot values for each 

stand. 

For analysis, forest stands were 

the observational unit and UAR was the 

dependent vari abl e. Regressi ons were 

run by using various measures of stand 

conditions (table 1) as independent 

variables. 

Results 

The density, size, and age of 

pines and the basal area of hardwoods 

varied greatly among forest stands, providi ng red-cockaded woodpeckers a wide 

choice of conditions from which to select foraging areas (table 1). Given 

Table 7.--Summary of characteristics of 276 forest 

stands used by 18 clans of red-cockade4 woodpeckers 

as year-round home ranges on Francis Marion National 

Forest, South Carol i na 

- --- 

Variable &an Range SD 

Stand age (years) 

Qumber pine treesiha 

(by d,b.h, in cm) 

Basal area pines 

(m2fhaf 

Ba sa 1 area hardwoods 

Im2/ha) 

>12,0 crn d,b.h, 

the range of conditions available to the 

birds, perhaps the most notable result 

of the study was the variability in the 

selection of stands with similar conditions (figs. 2,3,4,). These results 

suggest that the red-cockaded woodpeckers were not very selective of forest stands in which they foraged. nespite the variation in stand selection, 

statistical ly significant re1 ationships 

were found between stand selection and 

the major stand characteristics that we 

weasured. 

Several weak but statistically 

signi f icant trends were found between 

stand selection and the density and size 

of pines, Regressions that used the 

density of pines 224 cm, 236 cm, and >48 

cm d.b.h. were significant, but the percentage of the variation in stand selection that was accounted for was very 

small (fig. 2). Regressions that included pines (24 cm d.6.h. were not 

significant (P<0.05). - 

Although small, the amount of variation in stand selection that was accounted for by the regression models 

increased as the minimum dobehe of pines 

used in the independant variable became 

larger (fig. 2). Specifically, the 

density of pines >36 crn deheh, had a 

stronger re? ationshi p to stand selection 

than pines 224 cm d.b,h,, and pines 248 

cm d,b,h, had a stronger relationship to 

stand selection than pines 236 cm d,h.h, 

These results appear to suggest that 

stands with pines >48 crn d,h,h, were 

preferred over those with smaller trees, 

Other results, however, do not support 

such a conclusion, Me found no significant difference in selection of stands 

with and without pines 248 cm d,b.h, 

(mean UAR=1,3 and 1.2, n=141 and 93, 

respectively, P>OeS1 based on t-test ; 

a1 1 stands use'd" in this comparTson had 

210 pines/ha that were 224 cm debah.). 

In addition, when the area and associated foraging observation within 60 m 

of cavity trees were removed from the 

analysis, the relationship of stand 

selection to density of pines >48 cm 

d.b.h. was not significant (fig. 2,D). 

In contrast, without the effect of cavity trees the rela%ionskip between stand 

selection and the density of pines 224 

cm d.b,h, was sonewhat stronger than 

when a1 l observations were used 

(R2=0.07, P<0.000?, n=271 and R2=0.03, 

- P<0.002, ny276, respectively). The 

re1 ationshi p between stand selection and 

the density of pines 237 cn d,b.h, was 

the same whether or not the area around 

cavity trees was included, Removing the 

60-m area and observa%ions around cavity 

trees From the analysis made little 

change in the availabitity of stands 

witk pines 248 cm d,b.h, Only 5 of the 

28 stands with cavity trees were dropped 

Prom the analysis, leaving 145 stands 

witk pines 348 crn d,b,h, In total, th~ 

775 ha of stands with pines a48 em 

d.k,h, was reduced by only 7 percent. 

Thus, any apparent preference For stands 

with pines 248 cm d,h,h, seems to be due 

primarily to the attraction of caviLy 

trees. 

Preference for Foraging in stands 

tended to decrease as the basal area of 

hardwoods 212 cm d,h,h, increased (fig, 

3). Although statistically significant, 

RSQUARE = 0.02 

P < 0.02 

N = 276 

(SQUARE METERS PER HECTARE) 

BASAL AREA OF HARaWOBD TREES >= 12 CM DBH 

Figure 3.--Use!avallabi lity ratlo cwnpared -to Phe basat area of hardwood trees 212 cm d.blh* 

ln forest stands selected by 18 clans of fsragfng red-cockaded woodpe&ers, DaPa are from 

year-round horn ranges on the Francis Marian National Forest in SouPh Carolina. 
the regression accounted for virtually 

none of the variation in stand selection, suggesting that the density of 

hardwoods was not an important factor in 

stand selection, Exduding the area and 

f oragi ng observati sns around cavity 

trees did not change the regression of 

stand selection and hardwood basal area 

(R2=0.02, - P<O.O2, n=27l). 

Stand selection was signi ficantly 

related to stand age, but tke amount of 

variation accounted for by the regression was very small (fig. 4.A). Two 

factors appeared to he responsible for 

the relationship beLween stand selection 

and stand age, The first factor was 

that the seleaion for stands (30 years 

oad was radical fy di fferent from selection for older stands in that all the 

UAR values were low (fig. 4,A). The 

second facLor was that most cavity trees 

occurred in older stands and the attraction of cavity trees tended to inflate 

the UAR values, When stands (30 years 

old and the area and foraging observation within 60 m of cavities trees 

were removed from the analysis, the regression was no longer significant 

(fig. 4,B). Using 30 years as the point 

to delete young stands was arbitrary, 

but the same result occurred when stands 

(25 years old or (20 years old were used 

(R2=0.003, P>0.35, n=256 ; and R2=0.005, 

- PXl.25, n=259, respecti vely) . Discounting the area around cavity trees had 

little effect on the availability of 

stands ~80years old. Of the 133 ha in 

stands 280 years old, only 23 ha were 

exc 1 uded , 

Red-cockaded woodpeckers on the 

Francis Marion National Forest had a 

wide rawge sf tree densities, tree 

sizes, hardwood basal area, and stand 

ages from which ta select stands for 

foraging. The resul ts suggested that 

all these measures of stand conditions 

had some bearing on stand seleetisn, 

However, the birds were highly variable 

in their selection, and the observed relationships between stand conditions and 

stand sel eeti on were consequently weak, 

Several reasons for the variation in 

stand selection can be suggested. It 

was possible that clans in some, but not 

all, home ranges had more foraging 

resources than they needed (Hooper and 

others 1982; Wood and others 1985). If 

true, this might explain why some stands 

of agparently preferred foraging conditions received little or no use during 

the study. On the other hand, some home 

ranges may have been deficient in preferred foraging stands. If so, birds 

could have quickly exhausted stands with 

preferred conditions and have been 

forced to use less desirable stands at a 

higher rate than the more desirable 

ones, Another possible source of variation was related to stand size, For 

small stands, a few foraging ohservations could result in a stand having a 

high uselavailabil ity ratio (UAR), A1 so 

large stands could receive a considerable amount of use and still have a 

relatively low UAR. The UAR accounted 

for the inequiGies of stand size to a 

l arge exdent, as evidenced by the correlation between stand size and use 

(r=O,62) and between stand size and 

uselavailability (r=0.07). Nonetheless, 

stand size possibly contributed to the 

large variation in UAR's for stands of 

similar conditions, 

The importance of the density of 

pines as a factor in stand selection 

was corroborated to some extent by other 

studies, but direct comparison of resul ts are confounded. In one home 

range, Wood (1983) found the most intensively foraged area had the highest tree 

density, the moderately foraged area had 

an intermediate density, and the area of 

1 imited use had the least density of 

trees. In central Florida, DeLoteSle 

(pers. commun.) found a positive relationship between density of pines >6 em 

d,b,h, and stand selection. That relationship of smaller pines to stand selection might be explained by the lack sf 

pines 230 cm d.b.h. (DeLotelle and 

others 1983). Foraging areas in one 

home range in North Carol ina had fewer 

pine trees/ha than the home range as a 

whole, whereas the foraging areas in one 

other home range had a higher density 

than that home range as a whole (Repasky 

1984). Repasky included pines 22.5 cm 
A 

RSQUARE = 0.05 

P < 0.0003 

N = 276 

0 25 50 75 100 125 150 

STAND AGE (YEARS) 

B 

RSQUARE 0.002 

P > 0.47 

N = 252 

STAND AGE (YEARS) 

Figure 4.--A, Uselavailabllfty ratio compared to the age of forest stands selected by 18 clans 

of foraging red-cockaded woodpeckers; B, the same cornpartson excepP the area and associated 

foraging observations within 60 m of cavity trees in stands (30 years old have been mftted. 

Data are from year-round home ranges on the Francis Marion National Forest In South Carolina. 
d.b.h.; thus his findings were consistent with the lack of relationship we 

found between density of pines 23 cm 

d,b.h, and stand selection, But, in 

four home ranges in north Florida, 

Porter and Labisky (1986) found an inverse relationship between three levels 

of selection and the density of trees. 

Because they apparently incl uded smal 1 

pines, their results may not be inconsistent with ours, given that we found a 

1 ack of re1 ationshi p between stand 

selection and pines 23 cm d,b.h. 

Over and above the density of pines 

224 em d,b.h,, there was no evidence 

that the density or occurrence of even 

1 arger pines enhanced stand selection. 

Thus, pines 24.0 to 35.6 cm d.b.h. appeared to be equivalent to those 236.0 

cm d.beh. It has been suggested that 

larger trees could be foraged more efficiently than smaller trees because of 

presumably more total arthropod biomass 

and perhaps more biomass per unit area 

(Hooper and Lennartz 1981; Jackson 1979; 

fkorupa 19799, Indeed, several studies 

found that red-cockaded woodpeckers se1 ected individual trees for foraging 

that were on the average larger in dimeter than the mean d.b.h. of available 

trees (DeLotel le and others 1983; Hooper 

and Lennartz 1981; Porter and Labisky 

1986; Skorupa 1979). However, the results of these studies did not indicate 

that the rate of selection of individual 

trees continued to increase with the 

continued increase in diameter. Specifically, Hooper and Lennartz (1981) found 

strong selection for pines 224 cm d.b.h., 

but there was little difference in selection for pines 24.8 to 35.9, 36.0 to 

47.9, and 248.0 cm d.b.h. They also 

found that while the use of pines 12.0 

to 23.9 cm d,b.h. only equaled their 

availabil ity, pines of that size were 

foraged upon as often as pines a36 cm 

d.b.h. (27.6 and 26.8 percent, respectively). Porter and Labisky (1986) presented convincing evidence of selection 

for pines larger than those available 

on average, but the mean diameter of 

selected trees converged rapidly with 

the mean diameter of available trees as 

the latter increased. DeLotel le and 

others (1983) found a preference for 

trees 227 cm d.b.h. for foraging, but 

smal ler trees accounted for 83 percent 

of trees selected. Skorupa (1979) 

reported selection for larger than 

average pines but did not present data 

that allowed comparison of tree setection by diameter. 

Most of the statistical relationship between stand age and stand sef ection resulted from stands t30 years of 

age receiving little use as foraging 

areas and from the attraction of cavity 

trees in the oldest stands. Whether or 

not to discount the 60-m area around 

cavity trees is problematical , There 

is no doubt woodpeckers met some of 

t hei r nutritional requi rements when 

foraging in the immediate vicinity of 

cavity trees. On the other hand, it is 

questionable if the foraging time around 

cavity trees was determined primarily by 

stand conditions. Specifically, the 

birds began and ended each day at their 

cavities. They spent additional time in 

the vicinity of cavity trees engaged in 

cavity excavation, maintenance, and defense. Although during these times some 

cl an members typical ly foraged cont i nually and most clan members foraged at 

least intermittently, the primary attraction appeared to be the proximity of the 

cavity trees. Other areas of the home 

range did not have this same attraction 

to the birds. Most important, discounting the area around cavity trees had 

1 ittle effect on the availability of 

stands a80 years old. Of the 133 ha in 

stands 280 years old, only 23 ha were 

excluded. Thus, the birds had considerable opportunity remaining to select the 

oldest stands preferentially but did 

not do so. 

The negative effects of hardwoods 

on stand selection were consistent with 

the 1 iterature. Repasky (1984) found 

theoretical basis for hardwoods substantially increasing the cost in energy of 

foraging. A1 so, Hooper and tennartz 

(1981) and Repasky (1984) found that the 

proportion of hardwoods sel ected as 

foraging substrate was much less than 

that available. 
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H-er, Robert GI 

Forest stands selwted by fwaglng red-cockaded woodpeckers. Reso Pap. 

SE-259. Ashevl lle, NC: U.S. DsparMnt of Agriculture, Forest Service, 

SoutheasMn Forest Experimnt Statlon; 1986. '(opp. 

Selection of forest stands by 18 clans of foregfng red-cockaded woodpeckers was studfsd In thelr ysar-rwnd h ranges. The foraging use of 

276 stands relative to their avallabt 11ty within the home ranges was compared to several stand charactwlstfcs. Selectlm among stands with simftar characterlstlcs was RfghOy variable and red-cockadeds foraged In 

stands wlth a great varlesty of condltlons. Stand selection increased sfgnlffcantly as hnslty of plnes 3 24 crn Qeb.h. Increased and as basal area 

of hardwoods 2 12 ern debehe decreased, but theses relationships were weak. 

Bsyond 30 years, s-tand age was no* s slgnlffcant factor In selection. 

S: Endangered spwlss, fwaglng habitat, habitat selection, loblolly pine, longleaf plns, Plcoldes bwsalls* 

Hooper, Robsrt 6. 

Forest stands selected by foraging re&-c-aded w txkers. Res. Pap. 

SE-259. Ashevllle, NC: Urnso Depar-nt of Agriculture, Forest Service, 

Southeastern Forest Experiment Statlon; 1986. 1Qpp. 

Selection of forest stands by 18 clans of foraging red-cdaded 

peers was studled ln thelr year-round ham ranges* The foraglng use of 

276 stands relative to their avallabllfty within the hm ranges was campared to several stand characterfstfcs. Setecflan amng stands with slmilar characterfstfcs was highly variable and red-cockadeds forapd ln 

stands wlth a great variety of conditions. Stand selectton increased significantly as density of plnes 2 24 em d.b.h. Inweased and as basal area 

of har&occ;ds 2 12 crn drbrh. decreassd, but these relatlonshtps were weak. 

Beyond 30 years, stand age was not a sfgnlffcant factor In si~lectlon. 

KEYWRDS: Endangsred specfes, foraging habitat, habitat selection, lablofly pine, longleaf pine, Plcoides borealfs.