United States Department of Agriculture

Research Note

PNW-RN-570

Composition of Diets Selected by 

Sitka Black-Tailed Deer on Channel 

Island, Central Southeast Alaska

Thomas A. Hanley, Michael P. Gillingham, and Katherine L. Parker1

Pacifi c Northwest 

Research Station September 2014

Forest

Service

DEPAR TMENT OF AGRICULT URE

1 Th omas A. Hanley is a research wildlife biologist, U.S. Department of Agriculture, Forest Service, Pacifi c 

Northwest Research Station, Forestry Sciences Laboratory, 11175 Auke Lake Way, Juneau, AK 99801. Michael P. 

Gillingham and Katherine L. Parker are professors, Natural Resources and Environmental Studies Institute, 

University of Northern British Columbia, 3333 University Way, Prince George, BC, Canada V2N 429.

Abstract 

Composition of diets selected by tame but free-ranging deer in a natural forest 

environment was studied throughout a 24-month period and summarized by mean 

monthly percentage composition on the basis of dry-matter intake. A modifi cation of the standard bite-count method of diet determination was used. All forages 

were identifi ed by species and plant part (leaf, twig). Major forages comprising 

the diet were identifi ed seasonally and considered in relation to their role in the 

nutritional ecology of the deer. Overall, a simple but consistent pattern was evident 

in the data. When green, leafy forages (herbs and shrub leaves) were available, they 

were consistently targeted by the foraging deer, but as they became less available 

in winter, and especially with their burial by snow, diet composition shift ed to less 

digestible woody forages (shrub twigs and conifer foliage) supplemented with highly 

digestible arboreal lichens. Although a wide array of potential forages are available 

during the growing season, dietary options shrink to a narrow range of important 

evergreen forbs, a few ferns, shrub twigs, and lichens in winter.

Keywords: Odocoileus hemionus sitkensis, plant unit, habitat, food, browsing, 

grazing, dietary, dry-matter intake, understory vegetation, Tsuga heterophylla, Picea 

sitchensis, Vaccinium, Alectoria sarmentosa, Lysichiton americanus, Dryopteris 

expansa, Cornus canadensis, Rubus pedatus, Coptis aspleniifolia, Tiarella trifoliata.
RESEARCH NOTE PNW-RN-570

2

Introduction

Knowledge of diet composition is essential to understanding animal-habitat interactions because diet composition directly determines the nutritional quality of the 

animal’s food intake, and it also determines the selective pressures an animal exerts 

on its potential food base. For herbivores, knowledge of diet composition can be 

used to identify specifi c plant species and parts from within the entire vegetation 

community that are most important to the animal’s nutritional ecology and are most 

directly affected by herbivory. Diet composition for a given species of herbivore 

differs greatly, however, with differences in relative availabilities, innate palatabilities, and nutritional qualities of forages, all of which differ geographically, seasonally, and phenologically. Diet composition, therefore, tends to be most relevant at a 

local and time-specifi c scale. Nevertheless, it is only through many studies of diet 

composition that a broad, general picture of how animals relate to their environment through nutritional interactions can emerge for a given animal species within 

a given ecological region.

Diet composition for Sitka black-tailed deer (Odocoileus hemionus sitkensis) in 

the western hemlock–Sitka spruce (Tsuga heterophylla–Picea sitchensis 2

) forests of 

southeast Alaska has been studied by numerous investigators since the late 1950s 

(reviewed by Hanley 1984: table 2), although most of the early work consisted 

simply of lists of major forages identifi ed in the rumen contents of harvested deer. 

More quantitative study began in the 1980s with studies of both rumen contents and 

fecal composition (determined microscopically by identifi cation of plant fragments 

on the basis of their cell wall structure and other unique identifying features) 

expressed on a percentage ovendry-weight basis (Hanley et al. 1985, Hanley and 

McKendrick 1985, Lewis 1994, Pierce 1981). Although the dry-matter composition 

of rumen contents and feces can be determined more or less accurately for forages 

comprising the bulk of the sample collected, composition of neither rumen contents 

nor feces is assured to be directly related to composition of the diet that produced 

them. The principal problems are that plant species (and even tissues within a species) differ greatly in their recognizability after having been chewed and digested, 

and they differ greatly in their retention time in the rumen, their rate of digestion, 

and their overall dry-matter digestibility. The consequence is that easily identifi able 

and poorly digested species tend to be overestimated while poorly identifi able and 

highly digested species tend to be underestimated relative to the true composition 

of the diet. These problems have been well known for many years (Dearden et al. 

2

 See appendix for full scientifi c names, sources, and common names for all plants in 

this report.
Composition of Diets Selected by Sitka Black-Tailed Deer on Channel Island, Central Southeast Alaska

3

1975, Gaare et al. 1977, Havstad and Donart 1978, Slater and Jones 1971), and 

although partial solutions, including adjusting the results for dry-matter digestibility, have been suggested and used, the true composition of the actual diet can 

never really be determined by those techniques. For example, chewed and digested, 

young, highly digestible leaf tissue with few or no unique visually identifi able 

features will go unidentifi ed particularly in the feces, no matter what its “correction 

factor” might be, yet that tissue might be some of the most nutritiously important 

in the diet.

In 1989 and 1990 (October 1988 through September 1990), a unique, intensive 

study of the nutritional ecology of Sitka black-tailed deer was conducted on Channel Island, near Wrangell, Alaska (Parker et al. 1999). Its primary purpose was to 

fi eld test theory and hypotheses derived from independent studies of nutritional 

processes determining protein and energy balance of black-tailed deer in forest 

environments. The study involved closely monitoring the year-around activity, 

dietary intake, energy expenditure, and changes in body weight and composition of 

tame but free-ranging deer living without supplemental food on the island. Accurate 

M.P. Gillingham recording bites taken by a deer as it feeds on small forbs in a tidally infl uenced beach fringe habitat on 

Channel Island. 

Thomas A. Hanley
RESEARCH NOTE PNW-RN-570

4

determination of the true dry-matter composition of the diets selected by those deer 

was essential for the success of the study. Indeed, it was one of the principal reasons 

that tame deer were needed. The technique that was used involved identifying, 

counting, and tracking virtually all bites taken by animals under close observation by an observer while the animals were going about their usual daily activities 

(Parker et al. 1993a). 

The data on diet composition from the Channel Island study are unusual and 

especially valuable in several respects: (1) they provide accurate estimates of actual 

dietary intake (not rumen or fecal composition); (2) observers were able to see 

plants and plant parts eaten that would never have been observed or identifi ed in 

any other way; and (3) the data were obtained for every month over a full 2-year 

cycle. Although the results were used in many analyses and published reports from 

the study (e.g., Gillingham et al. 1997, 2001; Parker et al. 1993b, 1996, 1999), they 

were never published at the species level of diet composition, except in two composite fi gures identifying only a few of the most dominant species and major forage 

groups (fi gs. 5 and 8 in Parker et al. 1999). More recently, the need for more specifi c 

data on diet composition has become evident (e.g., in some aspects of habitat modeling—Hanley et al. 2012), and the value of those unique, highly accurate results 

from the Channel Island study has increased. Therefore, our purpose in this report 

is to provide a summary of those data at the species-specifi c and plant-part (leaves, 

twigs) levels of detail.

Methods

Study Area and Animals

The study was conducted on Channel Island (56° 22’ N latitude, 132° 10’ W longitude), a completely forested island of about 65 ha, approximately 20 km southeast of 

the town of Wrangell in central-southeast Alaska. Elevation ranged from 0 to about 

125 m above mean sea level. Forest overstory was a mixture of old-growth western 

hemlock–Sitka spruce and variously disturbed younger stands resulting from 

windthrow and some old hand-logged sites. Understory included virtually all the 

major understory species typical of the region with the exception of a notable few 

such as salal (Gaultheria shallon), willows (Salix spp.), and cedars (Thuja plicata, 

Callitropsis nootkatensis). The vegetation occurred within three major associations 

or community types, named after their dominant species: blueberry-spreading 

woodfern (Vaccinium ovalifolium–Dryopteris expansa); skunkcabbage (Lysichiton 

americanus); and devilsclub (Oplopanax horridus). Additionally, there was an open 

“beach” habitat that rimmed the southeasterly edges of the island and was subject to 

occasional inundation by seasonally high tides; tidal fl ood was suffi cient to exclude 
Composition of Diets Selected by Sitka Black-Tailed Deer on Channel Island, Central Southeast Alaska

5

trees but seldom enough to allow a species-rich wetland fl ora to persist (see Parker 

et al. 1999 for more details of the vegetation, including a vegetation map and measures of seasonal availability). No resident deer were believed to inhabit Channel 

Island at the time the study began, and the study animals were the only deer known 

to inhabit the island during the study, although an occasional transient wild deer did 

come and go. Therefore, the vegetation of the island was virtually ungrazed before 

the study, and it remained lightly to moderately grazed during the study.

The study animals were nine Sitka black-tailed deer (two males and two 

females born in 1987, and two males and three females born in 1988). All were 

either born in a captive animal facility in Juneau, Alaska, or captured in the wild, 

separated from their mothers shortly after birth and transported to a small island 

next to Channel Island where they were bottle-raised and then weaned onto the natural vegetation of Channel Island (Parker et al. 1999). This hand-rearing technique 

imprinted the deer on their human handlers so they accepted the handlers without 

fear or concern throughout the study. The handlers were always able to work at very 

close proximity to the deer (e.g., within 0.5 m if desirable) without disturbing them. 

The technique had the disadvantage of the deer not learning from their mothers, 

but all nine deer quickly adjusted to the Channel Island vegetation and were able 

to select diets that adequately met their nutritional needs for maintenance, growth, 

and even reproduction (four singletons and two sets of twins were produced by the 

fi ve females during the 2 years of data collection). Other studies have shown that 

hand-reared animals usually consume the same foods as maternally raised ones 

(Olsen-Rutz and Urness 1987, Spalinger et al. 1997). All deer utilized the entire 

island throughout the study. They all had seemingly unlimited access to all of the

major forage species. 

Diet Composition

Data were collected for the full 2-year period of October 1988 through September 

1990. A unique technique for estimating food intake was developed and fi eld-tested 

specifi cally for this study (Parker et al. 1993a). It was a modifi cation of the conventional bite-count method of studying dietary intake of tame deer (Wallmo and Neff 

1970) and consisted of defi ning a “plant unit” for each and every plant species and 

plant part (leaf, twig), where the plant unit was defi ned as an average-size piece of 

tissue commonly eaten by the deer, and intake was recorded in plant units instead 

of simply bite counts. In the case of small to medium-size leaves, the plant unit 

was often average leaf size; for large leaves (e.g., skunkcabbage), it was a square of 

specifi ed size, about average mouth width of the deer. A two-dimensional template 

of the plant unit was made for each forage species or forage species of comparable 

size, and template sizes were memorized by the observers. Forage composition 

and intake rate were determined while observing foraging deer by counting and 
RESEARCH NOTE PNW-RN-570

6

immediately recording (by plant species code in a portable data logger) the number 

of plant units of each forage consumed as it occurred. Representative plant units of 

all available forages were routinely collected, weighed, oven dried and reweighed 

throughout the study, thereby providing a basis for converting units consumed into 

both fresh and ovendry (100 °C) weights. This plant-unit technique worked well 

in the Channel Island communities of easily identifi able forbs and shrubs. Tests of 

the technique, comparing estimated weights of hand-harvested samples of plant 

material with actual weights, yielded an average correlation coeffi cient (r) of 0.985 

(range of 0.970 to 1.00, with N ranging from 7 to 284) for 21 forages of widely 

varying size and form (Parker et al. 1993a). Throughout the study and this report, 

we differentiate between leaf and twig plant parts of the same species because they 

differ greatly in their nutritional value as forages for deer.

Individual animals were randomly selected for detailed study of their activity 

(including foraging) during 2- to 8-hour periods every several days throughout the 

24 months of study. Although some activity trials occurred at night, only results 

from daytime trials are included in this report. Average diet composition and intake 

were determined for each of the animals on a monthly basis (see Parker et al. 1999 

for details). Here we are concerned only with diet composition on an ovendryweight basis, averaged across all animals each month.

Results and Discussion

Major Species and Plant Parts

Rather than presenting the raw monthly data for all plant species consumed in each 

of 24 months, we have summarized the results in terms of seasonal ranges in mean 

monthly diet composition for the four seasons of spring (April and May), summer 

(June, July, and August), fall (September, October, and November), and winter 

(December, January, February, and March) (table 1). Specifi c values of monthly 

means are not so informative, as they mainly refl ect very local circumstances. But 

ranges of values, by major season, are informative because the range provides a 

more general description relative to seasonal variation, and the seasonal grouping of months provides a meaningful pooling of major environmental differences 

affecting both availability and nutritional quality of the forage resources. Note that 

the seasons we defi ned are not of equal length. Spring is the period of new greenup 

and early, rapid growth of previously dormant vegetation; summer is the period of 

maximum biomass and phenological maturation of most forages; fall is the period 

of onset of seasonal senescence, withdrawal of reserves from leaves to storage 

organs, leaf-drop and winter senescence, but mostly frost- and snow-free conditions 
Composition of Diets Selected by Sitka Black-Tailed Deer on Channel Island, Central Southeast Alaska

7

under forest canopies at sea level; and winter is the period of leafl ess deciduous 

plants, full winter dormancy, and frequently frozen or snow-covered ground.

Over the course of the study, the deer ate at least a small amount of virtually 

all plant species occurring on the island, including even a few that were available 

only after washing ashore (Parker et al. 1999). Some forages were eaten only sparingly, perhaps as a taste or simply because of a desire for dietary diversity. Such 

rarely eaten species may not have been encountered during the diet-composition 

observations.

Overall, diets in spring, summer, and fall were highly diverse, with few species comprising large proportions of the diet (table 1). Forages that were eaten 

most were bunchberry (Cornus canadensis), false lily of the valley (Maianthemum 

dilatatum), skunkcabbage, Pacifi c silverweed (Argentina egedii), spreading woodfern, deer fern (Blechnum spicant), sedge (Carex spp.), rusty menziesia (Menziesia 

ferruginea) leaves, devilsclub leaves, blueberry leaves and twigs, western hemlock 

current annual growth (leaves and twigs together), Sitka alder (Alnus viridis) leaves, 

crabapple (Malus fusca) leaves, mushrooms, and witch’s hair lichen (Alectoria 

sarmentosa), each of which constituted more than 10 percent of the diet in at least 

one month. Skunkcabbage, spreading woodfern, deer fern, menziesia leaves, and 

blueberry leaves and twigs were the dietary dominants (each >25 percent at some 

time), with skunkcabbage reaching as high as 49 percent of the diet in June of 1989.

Diets were much less diverse in winter, when forage availability was much 

reduced by both phenology (deciduousness) and burial by snow. Most of the summer forages in southeast Alaska are deciduous so are simply not present in winter, 

and Sitka black-tailed deer do not dig through snow for forage, so any food buried 

by snow is not available to them. Forages that constituted major proportions of the 

winter diet were bunchberry, spreading woodfern rhizomes, dead leaves of sedge, 

blueberry twigs, red huckleberry (Vaccinium parvifolium) twigs, western hemlock 

current annual growth, rockweed alga (Fucus furcatus), and witch’s hair lichen, all 

of which comprised at least 10 percent of the diet in at least one month (table 1). 

Blueberry and witch’s hair lichen were the most dominant forages then (each reaching 30 to 32 percent, respectively, at some time).

However, data on diet composition alone do not provide measures of relative 

importance of the various forages to deer because they are strongly affected by relative availabilities and do not include nutritional value. The most highly nutritious 

forages are often relatively rare on the landscape and so cannot be eaten in great 

quantities, whereas forages of marginal quality are often very abundant and may 

be eaten as simply a matter of last recourse. For example, in winter, the evergreen 

forbs bunchberry, fi ve-leaved bramble, fern-leaf goldthread (Coptis aspleniifolia), 
RESEARCH NOTE PNW-RN-570

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Table 1—Seasonal diet composition (percentage dry-matter intake) of black-tailed deer 

diets during the 24 consecutive months of observation

 Spring Summer Fall Winter

Forage class and species (April–May) (June–Aug.) (Sept.–Nov.) (Dec.–March)

 Percentage dry-matter intake 

Forbs: 

 Achillea millefolium t t–1 0–1 0

 Argentina egedii t–1 1–10 0–3 0

 Coptis aspleniifolia 0–1 t t–8 t–4

 Cornus canadensis t–2 t–2 3–17 t–11

 Equisetum arvense 0–5 t–1 0–t 0

 Glaux maritima 0–t t–2 0–t 0

 Lysichiton americanus 6–33 14–49 3–22 0–8

 Maianthemum dilatatum 3–21 1–10 t–1 0–t

 Plantago macrocarpa 0–t 2–4 0–t 0

 Plantago maritima 0–6 t 0–t 0–t

 Rubus pedatus t–1 t t–8 t–5

 Streptopus amplexifolius t–4 t–3 0–t 0

 Streptopus streptopoides t–7 t–2 0–t 0–t

 Tiarella trifoliata 0–t t t–3 0–2

 Total forbs 31–52a 38–74 14–51 t–24

Ferns:

 Athyrium fi lix-femina 3–7 t–4 0–t 0–t

 Blechnum spicant 0–1 t 0–28 0–8

 Dryopteris expansa t–26 t–2 1–31 t–37

 Polypodium glycyrrhiza 0–t 0–t 0–3 0–2

 Total ferns 4–31 2–5 2–35 1–43

Graminoids:

 Carex spp. 1–18 t–3 0–t 0–10

 Deschampsia cespitosa t–3 t–1 0–t 0–3

 Elymus arenarius t–2 0–t 0–t 0–6

 Total graminoids 5–21 1–4 0–t 0–10

Shrub leaves:

 Menziesia ferruginea 0–t 1–23 0–29 0–t

 Oplopanax horridus 0–4 6–14 t–10 0–t

 Rubus spectabilis t–1 t t–3 0–t

 Vaccinium ovalifoliumb

 t–26 t–8 t–3 0–t

 Vaccinium parvifolium 0–t t t–3 0–2

 Vaccinium speciesc 0–t t t–3 t–2

 Total shrub leaves t–33 16–40 4–45 t–3

Shrub twigs:

 Vaccinium ovalifoliumb 0–27 0 0–t 2–30

 Vaccinium parvifolium 0–t 0 0–t 2–23

 Total shrub twigs 0–27 0 0–t 4–52

Trees:

 Alnus viridis leaves t–2 2–10 0–4 0–4

 Malus fusca leaves 0–3 t–2 t–21 0–t

 Tsuga heterophylla CAGd 0–3 0–t 0–10 5–16

 Total trees 1–5 2–11 3–23 8–18
Composition of Diets Selected by Sitka Black-Tailed Deer on Channel Island, Central Southeast Alaska

9

and to a lesser degree, foamfl ower (Tiarella trifoliata) are among the most nutritious (Parker et al. 1999) and palatable forages, but they are very low growing and 

are among the fi rst forages to be buried by snow. Their relatively low composition 

in the diet during winter (table 1) does not indicate a low importance; it indicates a 

low availability. In fact, those forbs are so important that their availability during 

snow-free conditions plays a major role in meeting the overall winter nutritional 

requirements for survival of deer in these forests (Parker et al. 1999). On the other 

hand, blueberry twigs are a marginally satisfactory winter forage in terms of their 

nutritional quality (Parker et al. 1999), but they are often the only nutritionally 

adequate forage rising through a snowpack. In their case, their high proportion of 

the diet does indicate relatively high importance, but only because of the relative 

Table 1—Seasonal diet composition (percentage dry-matter intake) of black-tailed deer 

diets during the 24 consecutive months of observation (continued)

 Spring Summer Fall Winter

Forage class and species (April–May) (June–Aug.) (Sept.–Nov.) (Dec.–March)

 Percentage dry-matter intake 

Other:e

 Alectoria sarmentosa f t–10 t–1 t–15 8–32

 Fucus furcatus 0–t 0–t 0–t t–15

 Lobaria spp. 0–t 0–t 0–1 t–6

 Mushrooms 0 0–12 4–23 0–3

 Peltigera spp. 0–t 0–t 0–t 0–1

Note: Values are ranges of monthly means, summarized by four seasons (spring, summer, fall, winter). Only forages that 

constituted at least 1 percent of mean dietary dry-matter intake during at least one month are included in the table. t = trace

= >0 but <1 percent. See appendix for full scientifi c nomenclature and common names. a “Total” values are the range of monthly totals; they are not a sum of data in the column above them.

b Includes Vaccinium alaskaense, which hybridizes with V. ovalifolium. c Evergreen, decumbent, sexually immature form of V. alaskaense, V. ovalifolium, and V. parvifolium. d CAG = current annual growth (current year’s leaves and twigs together).

e Additional forages that were observed to be consumed but never constituted at least 1 percentage of the mean monthly 

dry-matter intake in at least one month included the following (with season in parentheses—spring, Sp; summer, Su; fall, 

F; winter, W): 

Forbs—Angelica lucida (Sp, Su, F, W), Aster species (Su, F), Castilleja hyperborea (Sp, Su, F), Clintonia unifl ora (Sp, Su, 

F, W), Cochlearia groenlandica (Su, F), Conioselinum gmelinii (Sp, Su, F), Dodecatheon pulchellum (Sp, Su), Epilobium

species (Su), Fragaria chiloensis (Sp, Su, F), Fritillaria camschatcensis (Sp, Su), Galium trifi dum (Su, F), Heracleum 

maximum (Su), Lathyrus japonicus (Sp, Su, F), Listera cordata (Sp, Su, F, W), Lupinus nootkatensis (Sp, Su, F), Moneses 

unifl ora (Sp, Su, F, W), Osmorhiza species (Sp), Prenanthes alba (Sp, Su, F, W), Ranunculus occidentalis (Sp, Su, F), 

Rumex species (Su), Trifolium species (Sp), Veratrum album (Su), Viola glabella (Sp, Su, F); 

Ferns—Gymnocarpium dryopteris (Sp, Su, F, W); 

Graminoids—Hordeum brachyantherum (Sp, Su, F); 

Shrub leaves—Cornus sericea (Su, F), Ribes laxifl orum (Sp, Su, F), Ribes species (Sp, Su, F), Rubus parvifl orus (Sp, Su, 

F, W), Sambucus racemosa (Sp, Su, F, W), Viburnum edule (Su); 

Shrub twigs—Menziesia ferruginea (F, W); 

Trees—Callitropsis nootkatensis (W), Picea sitchensis (Sp, Su, F, W), Sorbus aucuparia (Su);

Other—Lycopodium species (F,W), Marchantiophyta (Sp), moss species (Su, F).

f Includes Usnea species that looks very similar to the more common A. sarmentosa.
RESEARCH NOTE PNW-RN-570

10

unavailability of better forages. An understanding of the relative importance of 

any given forage, therefore, requires consideration of both its availability and its 

nutritional quality in relation to those of all other forages in the habitat at the same 

time. It requires either detailed study (e.g., like that of Parker et al. 1999) or analysis 

within a nutritionally based model (e.g., like that of Hanley et al. 2012).

Some of the forages identifi ed as major dietary components (above) were 

eaten only at certain times or under certain circumstances, and their apparent 

high use within a season is an artifact of relatively high use at one specifi c time. 

For example, the use of alder and crabapple leaves was high only at their time of 

early senescence (alder in late August) and leaf-fall (crabapple in September). At 

other times, these two species were eaten only occasionally while green during the 

growing season or as dead leaves in winter. Similarly, sedges were a large dietary 

component in spring when they were young, fast-growing, succulent, and highly 

digestible (reaching a peak at 18 percent in May 1989), but they were virtually 

ignored at other times, except for heavy use (10 percent) in February 1990 during 

the deepest snow of the study when they provided one of the few alternatives to 

blueberry twigs and western hemlock foliage. The sedge leaves were dead then, but 

they occurred in the beach habitat where the snow was melted by tidal fl ood, and 

dead sedge leaves were the only “leafy” material available. That was also the same 

time that rockweed alga (another beach species of very low nutritional value–

Hanley and McKendrick 1983, Parker et al. 1999) reached its high of 15-percent 

diet composition. At times and circumstances such as that, almost any available 

forage might be eaten just for variety; the spike in diet composition says virtually 

nothing about value or importance.

Many of the forbs in table 1 occurred in the beach habitat and are not typical 

of upland forest habitats. They often seemed to be eaten as simply an addition 

for dietary diversity, although we cannot know that for certain. In any case, their 

dietary roles were mainly an artifact of habitat and vegetation circumstances of 

Channel Island; they do not refl ect broader patterns of diet composition of deer in 

forest habitats of the region, nor are they necessary foods for deer of the region. 

Other, common forest forbs could easily take their place.

Several new, important insights were gained in this study, and we believe they 

are likely to be generally true throughout the region. They involve specifi cally 

targeted plant tissues that came as a surprise to us or plant species or amounts that 

exceeded our expectations. Perhaps most surprising was the role that spreading 

woodfern played in the seasonal diets. Spreading woodfern is a common species 

widespread through many forest habitats of the region, and it was known from previous studies to be a fairly common dietary component for black-tailed deer, having 
Composition of Diets Selected by Sitka Black-Tailed Deer on Channel Island, Central Southeast Alaska

11

been found frequently in both rumen contents and feces. However, only by direct 

observation was it possible to identify that different parts of the plant are eaten at 

different times of the year and that the plant is specifi cally targeted by deer in both 

the shoulder seasons of spring and fall. In early spring as the snow retreats and deer 

are in their most depleted body condition (Parker et al. 1999), spreading woodfern 

becomes available before most other species leaf out, and deer target the fern’s 

fi ddlehead still buried in the moss or duff of the forest fl oor. Timing of spring is an 

especially critical time for deer, both for surviving winter and for the nutritional 

demands of a gestating fawn; and fi ddleheads of spreading woodfern are highly 

nutritious then (Parker et al. 1999). As summer progresses, deer consume the fronds 

of spreading woodfern, but only occasionally (table 1) and not importantly. However, in late fall after leaf fall of blueberry and other deciduous forage but before the 

forest fl oor freezes, deer actively target the rhizomes of the fern, buried in the forest 

fl oor but clearly evident because of the dead fronds on the ground (Gillingham et al. 

2000); that activity carries into winter, too, but only when the ground surface is not 

frozen or snow covered. Spreading woodfern, therefore, again becomes an important forage because the rhizomes are a large bundle providing energy and protein at 

a time when other highly digestible forages have become mostly unavailable. Before 

the Channel Island study, the targeting and relative importance of spreading woodfern fi ddleheads and rhizomes had never been noticed, or at least reported.

Similarly, skunkcabbage is another critically important forage in early spring, 

especially before green-up of other forages. It is usually the fi rst plant to break 

forth from the ground surface, sometimes even through still-frozen ground (e.g., 

in March, table 1), and deer actively seek the bright yellow buds. Skunkcabbage 

is always very highly digestible and very protein-rich (Hanley and McKendrick 

1983, Parker et al. 1999), but especially so in early spring at the very time that other 

forage resources are at their worst (Hanley and McKendrick 1985, Parker et al. 

1999). Although skunkcabbage has long been known to be very important to deer 

then and also as a signifi cant diet component throughout summer, it was surprising 

to see exactly how strong a diet component it could be during summer—reaching as 

high as 49 percent of dry-matter intake. Diet composition that high was surprising 

because skunkcabbage is such an extremely wet forage (ovendry weights are usually less than 10 percent of wet weights). Also, because of its very high dry-matter 

digestibility (e.g., 87 percent in spring, 76 percent in summer—Hanley and McKendrick 1983), it has been diffi cult to accurately estimate the true diet composition of 

skunkcabbage from rumen or fecal samples, even with adjustments for dry-matter 

digestibility. The very large leaf sizes, however, enable deer to have very high 
RESEARCH NOTE PNW-RN-570

12

intake rates while eating it. Skunkcabbage was targeted by deer throughout the 

growing season on Channel Island and was the only species that noticeably suffered an obvious decline in its availability because of deer use (although it always 

remained widely available throughout the island).

False lily of the valley is a species that was known to be eaten by deer in spring 

and summer, but was never suspected of playing such a substantial role as 22 percent diet composition (in May 1990; 16 percent in May 1989). It is one of the early 

species to leaf-out in spring, and its leaves are especially succulent and digestible 

then (especially while still curled before unfurling in early May). We suspect that 

the leaf tissue is relatively lacking in uniquely identifi able morphological features 

then, which is the reason for its apparent underreporting in previous studies of 

rumen and fecal contents. On the other hand, we suspect that mushrooms, heavily 

targeted by the deer in the fall (and peaking at 23 percent of the diet in September 

1990), have been underestimated in rumen and fecal studies because of their high 

digestibility, as their spores should be identifi able microscopically.

Another species that was eaten more than we expected was rusty menziesia. 

In both years of the study, consumption of leaves (but virtually never twigs) of 

this shrub peaked in late August to September (24 and 29 percent, 1989 and 1990, 

respectively), possibly refl ecting their high availability combined with an elevated 

drive in the deer to maximize food intake for replenishing body reserves before 

winter. Daily dry-matter intake, especially among females recovering from the 

demands of fawn-rearing, peaked then, reaching as high as during peak lactation 

(fi g. 8 in Parker et al. 1999). September is a month when both nutritional quality 

and available biomass of deciduous forages are declining rapidly with the onset of 

fall. Menziesia leaves retain their color and nutritional quality then (Hanley and 

McKendrick 1983); but perhaps more importantly, their growth in clumped whorls 

instead of isolated individuals makes them especially profi table food items in 

terms of dry-matter intake rate (Shipley and Spalinger 1992, Spalinger et al. 1988). 

Regardless of the reason, such high use of menziesia was surprising, and we suspect 

its use elsewhere might often be underestimated because of very low use of its 

twigs. Whereas browsed twigs are very obvious, missing leaves are not so obvious, 

especially in fall.

Similarly, but in an opposite pattern, we were surprised at the relatively low use 

of blueberry and red huckleberry leaves during summer. Red huckleberry leaves 

were seldom eaten year-around, but that might have largely refl ected their relatively 

low availability on Channel Island. Oval-leaf blueberry, on the other hand, was the 

dominant shrub in most of the vegetation of the island, yet its leaves were eaten 
Composition of Diets Selected by Sitka Black-Tailed Deer on Channel Island, Central Southeast Alaska

13

primarily in spring (up to 26 percent of the diet in May 1990) with less use later. 

Its twigs, too, were eaten then (up to 22 percent of the diet in April 1990) as they 

were recently emerged, rapidly growing, and at their peak nutritional value. In fact, 

dry-matter digestibility and digestible energy concentration not only peak in spring 

for blueberry (as for most other species, too), but leaves and twigs of blueberry 

have similar values then (Hanley and McKendrick 1983, Parker et al. 1999). By late 

August, though, digestible energy concentration of blueberry leaves was only 76 to 

79 percent of that for menziesia leaves, and use of blueberry dropped to less than 

that of menziesia. The same pattern of high use of blueberry in early summer, shifting to highly digestible forbs and devilsclub leaves in midsummer, and greater use 

of menziesia than blueberry in late summer or fall has been observed elsewhere in 

the region, too (Hanley and McKendrick 1985). However, when shrub twigs became 

such an important dietary component in winter, then both blueberry and red huckleberry were the staple foods. Despite its much lower availability, huckleberry was 

avidly sought and eaten in almost similar proportion as blueberry then (up to 23 and 

30 percent of the diet, respectively).

Several other species also were used less than we had expected. The four 

winter-evergreen forbs—bunchberry, fern-leaf goldthread, fi ve-leaved bramble, and 

foamfl ower—comprised very small percentages of the diet in spring and summer 

(0 to 2 percent). It wasn’t until other, more succulent and digestible forbs became 

deciduous in the fall that the intake of these four species increased. Fall and snowfree conditions of early winter were the peak times of their use; they also were the 

times of their highest relative nutritional quality—relative to other forages. 

Forage Classes and Temporal Pattern

Graphs of monthly changes in diet composition throughout the study (e.g., fi gs. 5 

and 8 of Parker et al. 1999 or a plotting of totals for the major forage classes in table 

1) are complex. However, if we view the forages in their most simple, nutritionally 

relevant classes of leafy material (herbs and shrub leaves), woody material (shrub 

twigs and conifer foliage), and highly digestible arboreal lichen (witch’s hair and 

beard lichens), then a simple, meaningful, and easily remembered pattern emerges 

(fi g. 1). Furthermore, we are confi dent that this pattern is a consistent, highly 

predictable pattern for black-tailed deer throughout southeast Alaska because it is 

clearly related to relative availabilities and nutritional values of forages. Whenever 

green leafy material (highly digestible and nutritious) is available, it is avidly eaten; 

but when it is not available in winter, the diet shifts to woody browse supplemented 

with highly digestible lichen. 
RESEARCH NOTE PNW-RN-570

14

The correlation (r) matrix for the data on diet composition during the 24 months 

of study (shown in fi g. 1) is -0.950 for “Herbs and Shrub Leaves” with “Shrub 

Twigs and Trees”; -0.854 for “Herbs and Shrub Leaves” with “Alectoria”; and 

0.713 for “Shrub Twigs and Trees” with “Alectoria.” But more important than the 

r values, per se, is that the pattern and correlations make sense nutritionally.

A species-diverse diet of green leafy material provides a balanced diet that is 

rich in digestible energy, digestible protein, and other essential nutrients. Diet diversity is important in balancing both the nutritional needs of deer and the deleterious 

effects of tannins, nontannin phenolics, toxins, and other secondary compounds 

in plants (McArthur et al. 1993). However, during winter, especially with snow 

on the ground, green leafy material is not available, and shrub twigs (Vaccinium

spp.) become the next best forage; yet alone, they are marginal at best for meeting the deer’s requirement for digestible energy and might even be inadequate for 

sustained, over-winter survival. Therefore, the very highly digestible, energy-rich 

arboreal lichens are especially important then as a supplement to the energy-poor 

diet of woody twigs. For example, whereas the concentration of digestible energy 

Figure 1—Mean monthly diet composition (percentage dry-matter intake) of black-tailed deer diets 

throughout the 24 consecutive months of observation, grouped by major forage classes of deciduous 

leaf material (herbs and shrub leaves), woody twigs and conifer foliage (shrub twigs and trees), and 

wind-blown arboreal lichen (Alectoria). “Herbs” are forbs, ferns, and graminoids.
Composition of Diets Selected by Sitka Black-Tailed Deer on Channel Island, Central Southeast Alaska

15

was about 9.94 to 10.23 kJ/g for blueberry and red huckleberry twigs, respectively, 

in winter, that of witch’s beard lichen was about 13.90 kJ/g (Parker et al. 1999). That 

36 to 40 percent difference in digestible energy concentration is an enormous and 

very important difference to the winter energy budget of deer (Parker et al. 1999), 

especially if the deer are bulk-limited in their food intake while on the strongly 

woody diet. Although witch’s hair and beard lichens contain such low concentrations of nitrogen that their digestible protein concentrations are actually negative 

numbers (-1.2 to -2.2 percent—Parker et al. 1999, Robbins 1987), protein is not 

a limiting factor for deer in winter because protein requirements are so low then 

and because the shrub component of the diet provides suffi cient digestible protein 

(Parker et al. 1999). The importance of Vaccinium shrubs and witch’s hair and beard 

lichens in winter diets of both Sitka and Columbian (O.h. columbianus) black-tailed 

deer in coastal Alaska and British Columbia, Canada, has been well known for 

many years (Hanley et al. 1989, Klein 1965, Rochelle 1980). On Channel Island, 

the deer commonly searched for and foraged on those wind-blown arboreal lichens 

during and after windstorms. Therefore, while the detailed, monthly data from the 

Channel Island study refl ect many items that were very specifi c to only that study 

area at that specifi c time (e.g., the many wetland forbs in the beach habitat, the 

timing and amounts of snow, the heavy use of dead sedge leaves during the deepest snow 3 ), the basic pattern of dietary change in relation to seasonal changes in 

forage availability (fi g. 1) and many of the insights gained about key forage species 

and plant parts are very likely true throughout the range of Sitka black-tailed deer 

in southeast Alaska. Forage species will differ with habitat, especially in summer 

(e.g., use of alpine habitats by seasonally migratory deer), but the overall pattern 

of a diverse array of forbs dominating the summer diet, winter-evergreen forbs, 

Vaccinium shrubs, and wind-blown arboreal lichens being critically important in 

winter, and certain species such as skunkcabbage and spreading woodfern being 

highly sought in early spring captures much of the essential linkages among deer, 

their habitat, diet, and nutritional requirements.

Acknowledgments

We thank Jeffrey Barnard, Mary Friberg, and Jason Dungan for their helpful 

reviews of an earlier draft of the manuscript.

3

 Note that because sedge is a graminoid, and graminoids are included with “herbs” 

in fi gure 1, the sedge 10 percent diet composition in February, 1990, elevates the total 

herbs and shrub leaves disproportionately for that month. Herbs and shrub leaves are a 

high-quality forage class, but dead sedge leaves are very low quality.
RESEARCH NOTE PNW-RN-570

16

English Equivalents

When you know: Multiply by: To get:

Centimeters (cm) 0.394 Inches

Meters (m) 3.28 Feet

Hectares (ha) 2.47 Acres

Square meters (m2) 10.76 Square feet

Grams (g) 0.0352 Ounces

Kilograms (kg) 2.205 Pounds

Kilojoules (kJ) 0.2388 Kilocalories (kcal)

Kilojoules (kJ) 0.948 British thermal units (BTU)

Kilojoules per gram (kJ/g) 26.932 BTU/ounce 

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Composition of Diets Selected by Sitka Black-Tailed Deer on Channel Island, Central Southeast Alaska

19

Appendix: Scientifi c and Common Names of Plant Species 

in This Reporta

Species Common name

Achillea millefolium L. Common yarrow

Alectoria sarmentosa (Ach.) Ach. Witch’s hair lichen

Alnus viridis (Chaix) DC. ssp. sinuata (Regel) Á. Löve & D. Löve Sitka alder

Angelica lucida L. Seacoast angelica

Argentina egedii (Wormsk.) Rydb. ssp. egedii Pacifi c silverweed

Aster L. spp. Aster

Athyrium fi lix-femina (L.) Roth Common ladyfern

Blechnum spicant (L.) Sm. Deer fern

Callitropsis nootkatensis (D. Don) Oerst. ex D.P. Little Alaska cedar

Carex L. spp. Sedge

Castilleja hyperborea Pennell Northern Indian paintbrush

Clintonia unifl ora (Menzies ex Schult. & Schult. f.) Kunth Bridge’s bonnet

Cochlearia groenlandica L. Danish scurvygrass

Conioselinum gmelinii (Cham. & Schltdl.) Steud. Pacifi c hemlockparsley

Coptis aspleniifolia Salisb. Fernleaf goldthread

Cornus canadensis L. Bunchberry dogwood

Cornus sericea L. ssp. sericea Redosier dogwood 

Deschampsia cespitosa (L.) P. Beauv. Tufted hairgrass

Dodecatheon pulchellum (Raf.) Merr. Darkthroat shootingstar

Dryopteris expansa (C. Presl) Fraser-Jenkins & Jermy Spreading woodfern

Elymus arenarius L. ssp. mollis (Trin.) Hultén American dunegrass

Epilobium L. spp. Willowherb

Equisetum arvense L. Field horsetail

Fragaria chiloensis (L.) Mill. Beach strawberry

Fritillaria camschatcensis (L.) Ker Gawl. Kamchatka fritillary

Fucus furcatus Esper Rockweed

Galium trifi dum L. Threepedal bedstraw

Gaultheria shallon Pursh Salal

Glaux maritima L. Sea milkwort

Gymnocarpium dryopteris (L.) Newman Western oakfern

Heracleum maximum W. Bartram Common cowparsnip

Hordeum brachyantherum Nevski Meadow barley

Lathyrus japonicus Willd. var. maritimus (L.) Kartesz & Gandhi Beach pea

Listera cordata (L.) R. Br. Heartleaf twayblade

Lobaria (Schreb). species Lung lichen
RESEARCH NOTE PNW-RN-570

20

Appendix: Scientifi c and Common Names of Plant Species 

in This Reporta (continued)

Species Common name

Lupinus nootkatensis Donn ex Sims Nootka lupine

Lycopodium L. spp. Clubmoss

Lysichiton americanus Hultén & H. St. John American skunkcabbage

Maianthemum dilatatum (Alph. Wood) A. Nelson & J.F. Macbr. False lily of the valley

Malus fusca (Raf.) C.K. Schneid. Oregon crabapple

Marchantiophyta, a division of bryophytes Liverworts

Menziesia ferruginea Sm. Rusty menziesia

Moneses unifl ora (L.) A. Gray Single delight

Oplopanax horridus (Sm.) Miq. Devilsclub

Osmorhiza Raf. spp. Sweetroot

Peltigera Willd. Felt lichen

Picea sitchensis (Bong.) Carrière Sitka spruce

Plantago macrocarpa Cham. & Schltdl. Seashore plantain

Plantago maritima L. Goose tongue

Polypodium glycyrrhiza D.C. Eaton Licorice fern

Prenanthes alba L. White rattlesnakeroot

Ranunculus occidentalis Nutt. Western buttercup

Ribes laxifl orum Pursh Trailing blackcurrent

Ribes L. spp. Current

Rubus parvifl orus Nutt. Thimbleberry

Rubus pedatus Sm. Five-leaved bramble

Rubus spectabilis Pursh Salmonberry

Rumex L. spp. Dock

Salix L. spp. Willow

Sambucus racemosa L. Red elderberry

Sorbus aucuparia L. European mountain ash

Streptopus amplexifolius (L.) DC. Claspleaf twistedstalk

Streptopus streptopoides (Ledeb.) Frye & Rigg Small twistedstalk

Thuja plicata Donn ex D. Don Western redcedar

Tiarella trifoliata L. Threeleaf foamfl ower

Trifolium L. spp. Clover

Tsuga heterophylla (Raf.) Sarg. Western hemlock

Usnea Dill. ex Adans. spp. Beard lichen

Vaccinium L. spp. Blueberry

Vaccinium alaskaense Howell Alaska blueberry
Composition of Diets Selected by Sitka Black-Tailed Deer on Channel Island, Central Southeast Alaska

21

Appendix: Scientifi c and Common Names of Plant Species 

in This Reporta (continued)

Species Common name

Vaccinium ovalifolium Sm. Oval-leaf blueberry

Vaccinium parvifolium Sm. Red huckleberry

Veratrum album L. White false hellebore

Viburnum edule (Michx.) Raf. Squashberry

Viola glabella Nutt. Pioneer violet

a

 Source of nomenclature is PLANTS database, http://plants.usda.gov/java/.
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