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RESEARCH Guilds of North American

RICHARD M. DE GRAAF ABSTRACT / We propose a foraging classification for USDA Forest Service North American inland, coastal, and pelagic birds. This classi- Northeastern Forest Experiment Station fication uses a three-part identification for each guild--major University of Massachusetts food, feeding substrate, and foraging technique--to classify Amherst, Massachusetts 01003, USA 672 of birds in both the breeding and nonbreeding seasons. We have attempted to group species that use similar resources in similar ways. Researchers have identified forag- NANCY G. TILGHMAN ing guilds generally by examining species distributions along USDA Forest Service one or more defined environmental axes. Such studies fre- Northeastern Forest Experiment Station quently result in species with several guild designations. While Warren, Pennsylvania 16365, USA the continuance of these studies is important, to accurately describe species' functional roles, managers need methods to STANLEY H. ANDERSON consider many species simultaneously when trying to deter- USDI and Service mine the impacts of alteration. Thus, we present an Wyoming Cooperative Wildlife Research Unit classification as a starting point for further dis- University of Wyoming cussion to aid those faced with the task of describing commu- Laramie, Wyoming 82071, USA nity effects of habitat change.

Many approaches have been taken to describe Severinghaus's guilds were not all ba~cd on habitat feeding behavior. Comparisons between different studies, requirements, to question whether the indicator concept however, have been difficult because of differences in would be effective. terminology. We propose to establish a classification Thomas and others (1979) developed lists of species scheme for North American birds by using common by life form for each habitat and successional stage in the terminology based on major food type, substrate, and Blue Mountains of . This is a use of the guild technique, using foraging guilds. concept in which assemblages of species are defined The guild concept was originally proposed by Root according to common nesting and feeding requirements. (1967) to indicate a group of species that exploit the same Another use of the guild concept was made by Short and class of environmental resources in a similar way. For Burnham (1982), who grouped species into layers of each example, Root defined a foliage-gleaning guild in habitat. For example, forest species are associated with woodlands as a group of birds whose diet consisted of subsurface, ground surface, and layer. obtained from the foliage zone of . Root Verner (1984) defines management guilds as groups pointed out that birds could be a member of several of species that respond in a similar way to a variety of guilds. Thus, seasonal changes that occur in avian feed- changes likely to affect their environment. He states that ing could be accommodated. The guild concept all bird species that depend on canopies for their food has become a useful tool in looking at , niche supply--whether fruit, buds, leaves, or --could be separation, and functional relationships within the com- in one management guild. This would give the manager munity. As a result, biologists can use the guild concept to some idea of the group of species that are affected by show how different taxa interrelate and how impacts of habitat change. Verner's use of management guilds does habitat change influence dynamics and not not fully consider the functional role originally proposed just individual species. by Root; rather, it considers responses to changes in The guild concept has been used by a number of identifiable zones of forest habitats. authors to compare or classify components of a commu- Standardization of guilds provides a means by which nity. It is possible to establish guilds on the bases of managers can compare communities of birds. different criteria. Landres (1983) reviews some efforts to The number of community components are reduced, assign species to guilds. Guilds were used by Severing- thereby facilitating the description. Likewise, biologists haus (1981) to develop the idea of indicator species; he can discuss competition and using common believed that since all members use the same resource, all terminology (Landres 1984). should respond similarly to changes in their environ- Guilds can be combined in different ways for different ment. There are enough exceptions, in part because purposes. Basically, however, we need to develop bases

Environmental Management Vol. 9, No. 6, pp. 493-536 1985 Springer-Vedag New York Inc. 494 R.M. DeGraaf and others

for classifying each species into a guild. In this article, we tion seasonally, which may require the use of different have considered food, substrate from which the food was substrates or techniques. We indicate such changes in obtained, and technique used in obtaining the food as the columns labeled as breeding period or nonbreeding peri- common ground to classify 672 species of North Ameri- od. We do not indicate gender- or age-specific differences can birds. These three criteria were selected because they in foraging guilds unless they appear vastly different and can be observed or documented for each species. The would affect the management of the species. components of major food, substrate on which food was The species in this list include all those that can be found, and techniques used to obtain food Provide realis- found in during the breeding season or tic groupings of species. The user of this information can nonbreeding season. The geographic range covered by group the species in one of the guilds presently described this list is from to Florida to to or utilize different criteria to develop the grouping. With . Birds of and the are not this classification, based on avian feeding behavior, users included unless they are regularly found north of these could compare the results of different studies for func- areas at the specified season. Accidental species are not tional differences. included. The Check-list of North American Birds, 6th Our description of guilds brings the concept as pro, edn. (American Ornithologists' Union, 1983) was used to posed by Root (1967) into use by resource managers. determine the status of these birds. Despite the different integrations and use of concept, it is now possible to have a ,dictionary" of terms characteris- tic of each species. While debates can continue about the Classification Procedures concept, our classification provides a reference point for application. For food type, we did not necessarily include all foods Birds' names are from the American Ornithologists' taken by each species--only the major food items (20% of Union (1983); scientific names are listed in the Appen- diet during a given period). dix. The major references u.'sed to develop this foraging : guild classification included A. C. Bent's Life History of Crustaceovore: North American Birds series (Smithsonian Institute, US : fruits National Museum Bulletins, , DC); Hand- Granivore: nuts book of North American Birds,' edited by Ralph S. : plants (leaves, stems, roots) Palmer (Yale University Press, New Haven, CT, 1962); : insects Waterfowl of North America, by Paul A. Johnsgard Moltuscovore: mollusks (Indiana University Press, Bloomington, 1975); , : a variety of foods including both Geese 6" of North America, by Frank C, Bellrose and plant foods (the less common food group makes (Stackpole Books, Harrisburg, PA, 1976); American up 10% of diet) Wildlife 4r Plants, by Alexander C. Martin and others : fish (Dover Publications, New York, 1951); of the : sandworms, , etc. World, by Derek Goodwin (Corneil University Press, Ithaca, NY, 1976); Audubon Guide and Substrate refers to the place where a food item is found or Audubon Land Bird Guide by Richard H. Pough (Dou- taken. bleday & Co., Garden City, NY, 1946)i Management Of air: caught in the air Migratory Shore and Upland Birds in North bark: on, in, or under bark of America, edited by Glen C. Sanderson (International coastal: waters along coast (can include brackish as Association of Fish and Wildlife Agencies, Washington, well as salt water) DC, 1977). coastal beach: beaches and/or tidal flats along coast coastal bottom: floor of continental shelf along coast coastal rock: rocks along coast Foraging Guild Designation coastal surface: surface of coastal waters Often it was difficult to limit this classification to a floral: on or in flowers single guild per species, so in several eases we have listed fresh marsh: freshwater marshes (on mud, in shallow more than one guild. An additional guild was designated water, or on marsh plants) when a second food item represented at least 20% of the freshwater: freshwater habitats (ponds, lakes, rivers, diet of a species or when the specie s spent at least 20% of streams) its time feeding on a different substrate or using a freshwater bottom: bottoms of freshwater ponds and different technique. Some species shift their food Selec- lakes Foraging Guilds of North American Birds 495

freshwater shoreline: shores of freshwater ponds, hover-gleaner: hovers in air while selecting prey (from lakes, rivers, or streams vegetation or ground) freshwater surface: surface of freshwater habitats plunger: dives from air into water to capture prey in ground: on the ground or on very low, weedy vegeta- bill or gular pouch tion prober: inserts bill into substrate (beach, mud, ground) lower /shrub: on leaves, twigs, and branches of and locates prey by touch , saplings, and lower crowns of trees sallier: perches on exposed branch or twig, waits for marsh: fresh, brackish, or saltwater marshes (on mud, to fly by, and then pursues and catches insect in shallow water, or on marsh plants) in air mud: inland on mud flats (wet fields, meadows, tun- scaler: exposes prey under bark by scaling off loose dra, or associated with freshwater habitats) bark pelagic: ocean waters away from coastlines : takes a variety of items, including refuse or pelagic surface: surface of ocean waters carrion riparian bottom: bottoms of rivers and streams screener: flies with bill open and screens prey from salt marsh: brackish or salt marshes (on mud, in air shallow water, or on marsh plants) skimmer: flies low over water and skims food from shoreline: along shoreline of both freshwater and water surface with lower mandible in water saltwater (coastal) habitats strainer: strains food items from water or mud through upper canopy: on leaves, twigs, and branches of trees lamellae along edge of bill in main canopy water: brackish, fresh- and saItwater habitats water bottom: on bottoms of fresh-, brackish, or salt- Foraging Guild Classification water habitats Nonbreeding water surface: on surface .of fresh-, brackish, or salt- Breeding period period Year round water habitats CARNIVORE: air hawker Pomarine Jaeger White-tailed Eagle Technique refers to the manner in which food is Long-tailed Jaeger Sharp-shinned obtained. Note: some techniques are associated with Glaucous particular food types and/or substrates. Cooper's Hawk ambusher: slowly stalks or waits for prey to come Merlin within reach chaser: pursues prey on ground Gyrfalcon dabbler: submerges head and neck or tips up (various Prairie Falcon water substrates) CARNIVORE: coastal beach scavenger diver: dives from surface for underwater food Laughing Gull CommonBlack- excavator: locates food in bark by drilling holes Thayer's Gull headed Gull Great Black- food pirate: steals food from other species, usually Bonaparte's Gull backed Gull Heermann's Gull other birds Mew Gull . foot plunger: catches prey by plunging from air to Ring-billed Gull water surface (or ground) and seizing prey in talons Iceland Gull forager: takes almost any food items encountered upon Yellow-footed Gull Glaucous-winged the substrate (includes all and Gull feeding on terrestrial habitats or vegetation, except grazers and grubbers) CARNIVORE: coastal food pirate Parasitic Jaeger gleaner: selects particular food items from the sub- Great strate CARNIVORE: coastal scavenger grazer: feeds on grasses, sedges, or grains in fields or Pomarine Jaeger Western Gull Iceland Gull meadows Great Skua Glaucous Gull grubber: digs up roots and tubers of either terrestrial Glaucous-winged or aquatic plants Gull hawker: flies after prey and captures it either in air or CARNIVORE: coastal surface gleaner on ground Bonaparte's Gull 496 R.M. DeGraaf and others

Foraging Guild Classification (Continued) Foraging Guild Classification (Continued) Nonbreeding Nonbreeding Breeding period period Year round Breeding period period Year round CARNIVORE:ground chaser CARNIVORE:ground scavenger (continued) Greater Roadrun- ner Crested Caraeara CARNIVORE:ground hawker CARNIVORE:pelagic food pirate American Swal- Black-shouldered Pomarine Jaeger low-tailed Kite Kite Parasitic Jaeger Gray Hawk Aplomado Falcon Sharp-shinned CARNIVORE:pelagic scavenger Pomarine Jaeger Hawk Short-tailed Alba- Northern Fulmar Long-tailed Jaeger Cooper's Hawk tross Fork-tailed Storm- Great Skua Northern Goshawk Cory's Shearwater Ring-billed Gull Common Black- Sooty Shearwater Leach's Storm- California Gull Hawk Black Storm-Petrel Petrel Ivory Gull Harris's Hawk Pomarine Jaeger Red-shouldered Great Skua Hawk South Polar Skua Broad-winged Ivory Gull Hawk CARNIVORE: Short-tailed Hawk shoreline gleaner Swainson's Hawk Mew Gull White-tailed Hawk CARNIVORE:shoreline scavenger Zone-tailed Hawk Herring Gull Red-tailed Hawk Thayer's Gull Lesser Black- Rough-legged backed Gull Hawk Golden Eagle CARNIVORE:water ambusher American Kestrel American Bittern Prairie Falcon Great Egret Common Barn- Snowy Egret Wood Eastern Screech- CRUSTACEOVORE:coastal ambusher Owl Tricolored Western Screech- Owl CRUSTACEOVORE:coastal beach gleaner Black-bellied Wilson's Plover Plover Black Northern Hawk- Snowy Plover Owl Semipalmated Northern Pygmy- Plover Owl Piping Plover Ferruginous Red Knot Pygmy-Owl QRUSTACEOVORE:coastal beach prober Spotted Owl Willet Whimbrel Great Gray Owl Marbled Long-eared Owl Sanderling Short-eared Owl Dunlin Boreal Owl CRUSTACEOVORE:coastal bottom gleaner Northern King Saw-whet Owl Steller's Eider Northern Shrike Harlequin Loggerhead Shrike Oldsquaw CARNIVORE: ground scavenger Black Parasitic Jaeger Golden Eagle Black Vulture Surf Scorer Vulture White-winged California Condor Scoter Foraging Guilds of North American Birds 497

Foraging Guild Classification (Continued) Foraging Guild Classification (Continued) Nonbreeding Nonbreeding Breeding period period Year round Breeding period period Year round CRUSTACEOVORE:coastal diver CRUSTACEOVOI~;:pelagic diver (continued) Dovekie Homed Red-faced Cormo- Black Common Murre Eared Grebe rant Xantus' Murrelet Pigeon Guillemot Ancient Murrelet Kittlitz's Murrelet Cassin's Auklet Xantus' Murrelet Parakeet Auklet Ancient Murrelet Least Auklet Cassin's Auklet Whiskered Auklet Parakeet Auklet Crested Auklet Least Auklet Whiskered Auklet Crested Auklet CRUSTACEOVORE:pelagic scavenger Rhinoceros Auklet Short-tailed Shear- water CR.USTACEOVORE:coastal rock gleaner Wilson's Storm-Pet- Black Turnstone rel Surfbird Purple CRUSTACEOVORE:pelagic surface gleaner Rock Sandpiper Bullet's Shear- Ashy Storm-Petrel water CRUSTAGEOVORE: coastal surface gleaner Least Storm-Petrel Thick-billed Thick-billed Murre Murre Razorbill CRUSTACEOVORE:freshwater bottom gleaner CRUSTACEOVORE:salt-marsh gleaner White-winged Clapper Black Rail Seaside Sparrow Scoter CRUSTACEOVORE:shoreline gleaner Hooded Merganser Black-bellied Least Sandpiper CRUSTACEOVORE:freshwaler diver Plover Pied-billed Grebe Snowy Plover Red-necked Grebe Semipalmated Plover CRUSTACEOVORE:freshwater glean er Solitary Sandpiper Piping Plover Semipalmated CRUSTACEOVORE:freshwater shoreline gleaner Sandpiper Red Knot Western Sandpiper Sanderling Rufous-necked Least Sandpiper Stint Rock Sandpiper CRUSTACEOVORE:marsh gleaner Black Rail CRUSTACEOVORE:shoreline prober CRUSTACEOVORE:marsh prober Willet Long-billed King Rail Bar-tailed Godwit Stilt Sandpiper CRUSTACEOVORE: mud gleaner Baird's Sandpiper CRUSTACEOVORE:water ambusher Tricolored Heron American Bittern CRUSTACEOVORE: mud prober Great Egret White-rumped Scarlet White Ibis Snowy Egret Sandpiper Little Blue Heron Dunlin White-faced Ibis Green-backed Heron CRUSTACEOVORE:mud strainer Black-crowned Roseate Night-Heron CRUSTACEOVORE:pelagic diver Yellow-crowned Dovekie night-heron Common Murre Wood Stork 498 R. M, DeGraaf and others

Foraging Guilds of North American Birds Foraging Guild Classification (Continued) Nonbreeding Nonbreeding Breeding period period Year round Breeding period period Year round

CRUSTACEOVORE: water bottom gleaner GRANIVORE:ground gleaner (continued) Rufous-winged Mourning Dove Hooded Merganser Sparrow Inca Dove Rufous-erowned Common Ground- CRUSTACEOVORE: water diver Sparrow Dove Pied-billed Grebe American Tree White-tipped Dove Sparrow White-collared CRUSTACEOVORE: water skimmer Chipping Sparrow Seedeater Black Skimmer Field Sparrow House Finch Black-chinned Lesser Goldfinch CRUSTACEOVORE:water surface gleaner Sparrow Lawrence's Gold- Red Lesser Yellowlegs Vesper Sparrow finch FRuGIVoRE:ground gleaner Sparrow House Sparrow House Finch Plain Chaehalaea Black-throated Sparrow FRUGIVORE:lower-canopy/shrub gleaner Sage Sparrow Band-tailed Pigeon Red-headed Wood- Ladder-backed Five-striped Gray Kingbird pecker Sparrow Rose-throated Be- Eastern Phoebe Nuttall's Wood- Savannah Sparrow card Great Crested Fly~ peeker Baird's Sparrow Eastern catcher Downy Wood- Sparrow Cassin's Kingbird pecker Song Sparrow Tree Swallow Hairy Woodpecker Lincoln's Sparrow Wrentit Phainopepla Swamp Sparrow Northern Mock- White-throated ingbird Sparrow Curve-billed Golden-crowned Thrasher Sparrow Bohemian Wax- White-crowned wing Sparrow Harris' Sparrow FRUGIVORE: upper-canopy gleaner Dark-eyed Junco Purple Finch Band-tailed Pigeon White-crowned McCown's Long- Pigeon spur Red-billed Pigeon Lapland Longspur Acorn Woodpecker Chestnut-collared Gila Woodpecker Longspur Striekland's Wood- Snow Bunting pecker McKay's Bunting Ivory-billed Wood- Red-winged Black- pecker bird Bohemian Wax- Bronzed Cowbird wing Brown-headed Cedar Waxwing Cowbird Rosy Finch GRANIVORE:freshwater dabbler Grosbeak Green-winged Teal Fulvous Wh!stling- Purple Finch No-rthern Pintail Duck Cassin's Finch Common Redpoll GlthNIVOltE:freshwater surface gleaner Hoary Redpoll Wood Duck Pine Siskin American Gold- GRANIVORE:ground gleaner finch Mallard Gray Fulvous Whistling- Eyenin_g Grosbeak California Duck Black-bellied Whistling-Duck GRANIVORE:lower-canopy/shrub gleaner Dickcissel Wood Duck American Gold- Lesser Goldfinch Cassin's Sparrow White-winged finch Lawrence's Gold- Dove finch Foraging Guilds of North American Birds 499

Foraging Guild Classification (Continued) Foraging Guild Classification (Continued) Nonbreeding Nonbreeding Breeding period period Year round Breeding period period Year round GRANIVORE:upper-canopy gleaner HERBIVORE:upper-canopy forager Purple Finch Red-cockaded Lewis' Wood- Spruce Plain Chachalaca Woodpecker pecker Blue Grouse White-headed Ruffed Grouse Woodpecker Sharp-tailed Red-breasted Nut- Grouse hatch HERBIVORE:water dabbler White-breasted Tundra American Nuthatch Trumpeter Swan Brown-headed Nuthatch Canada Pine Grosbeak Gadwall Cassin's Finch American Red Crossbill HERBIVORE:water diver White-winged Masked Duck Crossbill American Coot Pine Siskin Evening Grosbeak HERBIVORE: water grubber Snow Goose GRANIVORE:water dabbler INSEGTIVORE:air hawker Green-winged Teal American Swal- Common Black- Mississippi Kite HERBIVORE:coastal dabbler low-tailed Kite headed Gull Swainson's Hawk Whooper Swan Gray Hawk Burrowing Owl Emperor Goose American Kestrel Brant Aplomado Falcon Eurasian Wigeon Franklin's Gull Bonaparte's Gull HERBIVORE:freshwater dabbler Black Tundra Swan Black-bellied Elf Owl Trumpeter Swan Whistling-Duck Mute Swan INSECTIVORE:air sallier Canada Goose Elegant Anna's Humming- Gadwall Red-headed Wood- bird HERBIVORE:freshwater diver pecker Green Kingfisher Masked Duck Olive-sided Fly- Lewis' Wood- catcher pecker HERBIVORE: groundforager Greater Hammond's Fly- Willow Ptarmigan Ring-necked Chukar Western Wood- catcher Sage Grouse Pewee Gray Flycatcher White-tailed Ptar- Greater Prairie- Gambel's Quail Eastern Wood- Black Phoebe migan Pewee Eastern Phoebe Mountain Quail Lesser Prairie- Yellow-bellied Fly- Say's Phoebe Chicken catcher Vermillion Fly- Wild Turkey Acadian Flycatcher catcher Alder Flycatcher Ash-throated Fly- HERBIVORE:ground grazer Willow Flycatcher catcher Mute Swan Mallard Greater White- Least Flycatcher Great Crested Fly- Pink-footed Goose fronted Goose Dusky Flycatcher catcher Emperor Goose Snow Goose Western Flycatcher Great Kiskadee Brant Ross' Goose Buff-breasted Fly- Couch's Kingbird Barnacle Goose Canada Goose catcher Cassin's Kingbird American Wigeon Dusky-capped Fly- Western Kingbird catcher Scissor-tailed Fly- HERBIVORE:ground grubber Brown-crested Fly- catcher catcher Townsend's Soli- HERBIVORE:lower-canopy/shrub forager Sulphur-bellied taire Willow Ptarmigan Flycatcher Cedar Waxwing Rock.Ptarmigan Tropical Kingbird White-tailed Ptar- Thick-billed King- migan bird 500 R.M. DeGraaf and others

Foraging Guild Classification (Continued) Foraging Guild Classification (Continued) Nonbreeding Nonbreeding Breeding period period Year round Breeding period period Year round INSECTIVORE:air sallier (continued) INSECTIVORE:bark gleaner (continued) Eastern Kingbird White-breasted Gray Kingbird Nuthatch Rose-throated Pygmy Nuthatch Becard Brown-headed Bohemian Wax- Nuthatch wing Brown Creeper American Redstart Wrentit Hooded Warbler Yellow-throated Witson's Warbler Warbler Painted Redstart Pine Warbler Western Tanager Black-and-white Rufous-winged Warbler Sparrow INSECTIVORE:bark scaler Black-throated White-headed Sparrow Woodpecker Rosy Finch Three-toed Wood- INSECTIVORE:air screener pecker Lesser Common Pauraque Black-backed Common Night- Common Poorwill Woodpecker hawk Chuck-will's- ivory-billed Wood- Antillean Night- widow pecker hawk Whip-poor-will INSECTIVORE:coastal beach gleaner Buff-collared Vaux's Swift Black-bellied Wilson's Plover White-throated Plover Black Swift Swift SemipaImated Chimney Swift Tree Swallow Plover Purple Martin Violet-green Swal- Piping Plover Bank Swallow low Wandering Tattler Cliff Swallow Northern Rough- Western Sandpiper Cave Swallow winged Swallow Dunlin Barn Swallow Short-billed INSECTIVORE:bark excavator Pileated Wood- Long-billed pecker Dowiteher INSECTIVORE:coastal beach prober INSECTIVORE:bark gleaner Willet Pine Warbler Yellow-bellied Red-headed Wood- Prothonotary Sapsucker pecker INSECTIVORE:coastal rock gleaner Warbler Red-breasted Sap- Acorn Woodpecker Purple Sandpiper Painted Redstart sucker Gila Woodpecker Rock Sandpiper Williamson's Sap- Golden-fronted INSECTIVORE:fresh-marsh gleaner sucker Woodpecker Northern Jacana Red-bellied Wood- pecker INSECTIVORE:freshzoater bottom gleaner Ladder-backed Woodpecker Hooded Merganser Nuttall's Wood- INSECTIVORE:freshwater diver pecker Pied-billed Grebe Least Grebe Downy Wood- Eared Grebe pecker Red-necked Grebe Hairy Woodpecker Strickland's Wood- INSECTIVORE:fresh water gleaner pecker Wood Sandpiper Black-necked Stilt Red-cockaded Solitary Sandpiper Woodpecker INSECTIVORE:Freshwater shoreline gleaner Red-breasted Nut- Wandering Tattler Northern Water- hatch Red Knot Foraging Guilds of North American Birds 501

Foraging Guild Classification (Continued) Foraging Guild Classification (Continued) Nonbreeding Nonbreeding Breeding period period Year round Breeding period period Year round INSECTIVORE:Freshwater shoreline gleaner (continued) INSECTIVORE:ground gleaner (continued) " Sanderling Cassin's Sparrow Least Sandpiper Brewer's Sparrow Wilson's Phalarope Baird's Sparrow Water- Eastern Meadow- thrush lark Western Meadow- lark INSECTIVORE: freshwater surface gleaner Rusty Blackbird Little Gull Sabine's Gull INSECTIVORE:ground hawker Black Tern Eastern Screech- Flammulated Owl Owl Whiskered Western Screech- Screech-Owl INSECTIVORE: ground gleaner Owl Lesser Golden- Northern Pygmy- Plover Mountain Plover Owl Killdeer Greater Roadrun- Elf Owl Upland Sandpiper ner Eskimo Curlew Smooth-billed Ani INSECTIVORE:lower-canopy/shrub gleaner Surfbird Groove-billed Ani Black-billed Ash-throated Fly- Pectoral Sandpiper Rock Wren catcher Rock Sandpiper Canyon Wren Yellow-billed Carolina Wren Curlew Sandpiper Bewick's Wren Cuckoo House Wren Buff-breasted Winter Wren Mangrove Cuckoo Golden-crowned Sandpiper Sedge Wren Black-capped Kinglet Long-billed Chickadee Ruby-crowned Dowitcher Le Conte's Carolina Chick- Kinglet Bonaparte's Gull Thrasher adee Black-tailed Gnat- Mew Gull Gray-crowned Mexican Chick- catcher Ring-billed Gull Yellow-throat adee Black-capped California Gull Mountain Chick- Gnatcatcher Northern Flicker adee Hutton's Vireo Black-billed Mag- Siberian Nashville Warbler pie Boreal Chickadee Black-throated Yellow-billed Chestnut-backed Gray Warbler Chickadee Prairie Warbler Bluethroat Bridled Titmouse Common Yellow- Northern Wheat- Tufted Titmouse throat ear Verdin Bushtit White-eyed Vireo Bell's Vireo Field fare Black-capped Varied Thrush Vireo Sage Thrasher Gray Vireo Yellow Wagtail Solitary Vireo White Wagtail Blue-winged Red-throated Pipit Warbler Water Pipit Orange-crowned Sprague's Pipit Warbler Palm Warbler Virginia's Warbler Worm- Colima Warbler Warbler Lucy's Warbler Yellow Warbler Kentucky Warbler Chestnut-sided Connecticut Warbler Warbler Magnolia Warbler Mourning Warbler Black-throated Painted Redstart Blue Warbler 502 R.M. DeGraaf and others

Foraging Guild Classification (Continued) Foraging Guild Classification (Continued) Nonbreeding Nonbreeding Breeding period period Year round Breeding period period Year round INSECTIVORE: lower-canopy/shrub gleaner (continued) INSECTIVORE:upper-canopy gleaner (continued) Yellow-rumped Red-eyed Vireo Townsend's Warbler Bachman's Warbler Golden-cheeked Warbler Black-throated Warbler Golden-winged Green Warbler Kirtland's Warbler Warbler Yellow-throated American Redstart Tennessee Warbler Warbler Prothonotary Northern Parula Warbler Cape May Swainson's Warbler Warbler Blackburnian MacGillvray's Warbler Warbler Grace's Warbler Hooded Warbler Bay-breasted Wilson's Warbler Warbler Canada Warbler Blackpoll Warbler Olive Warbler Cerulean Warbler Black-throated Red-faced Warbler Sparrow Hepatic Tanager Summer Tanager INSECTIVORE:marsh gleaner Scarlet Tanager Black Rail Marsh Wren Orchard Oriole INSECTIVORE:marsh prober INSECTIVORE:water ambusher King Rail American Bittern Virginia Rail Least Bittern INSECTIVORE:mud gleaner INSECTIVORE:water bottom gleaner White-rumped Glossy Ibis Hooded Merganser Sandpiper White-faced Ibis INSECTIVORE:water diver Baird's Sandpiper Pied-billed Grebe Dunlin INSECTIVORE:water surface gleaner INSECTIVORE:riparian bottom gleaner Red-necked Phala- Solitary Sandpiper Greater Yellowlegs Harlequin Duck American Dipper rope Lesser Yellowlegs INSECTIVORE:salt-marsh gleaner : Red Phalarope Gull-billed Tern Seaside Sparrow Forster's Tern MOLLUSCOVORE:coastal beach gleaner INSECTIVORE:shoreline gleaner Black-bellied Black-bellied Killdeer Spotted Sandpiper Plover Plover Long-billed Cur- Semipalmated Semipalmated lew Plover Plover Least Sandpiper Red Knot Piping Plover Western Sandpiper Semipalmated Sandpiper MOLLUSCOVORE:coastal beach prober Western Sandpiper Whimbrel American Oyster- Rufous-necked Marbled Godwit catcher Stint Sanderling Stilt Sandpiper Dunlin MOLLUSCOVORE:coastal bottom gleaner INSECTIVORE:shoreline prober Greater Seaup Common Eider Willet King Eider Bar-tailed Godwit Marbled Godwit Steller's Eider INSECTIVORE:upper-canopy gleaner Harlequin Duck Warbler Northern Beard- Oldsquaw Yellow-throated less-Tyrannulet Black Seoter Vireo Blue-gray Gnat- Warbling Vireo catcher White-winged Philadelphia Vireo Tropical Parula Scoter Foraging Guilds of North American Birds 503

Foraging Guild Classification (Continued) Foraging Guild Classification (Continued) Nonbreeding Nonbreeding Breeding period period Year round Breeding period period Year round

MOLLUSCOVORE:coastal diver MOLLUSCOVORE:water am busher Razorbill Wood Stork MOLLUSCOVORE:coastal plunger MOLLUSCOVORE,:water bottom gleaner Red-footed Booby King Eider Tufted Duck MOLLUSCOVORE:coastal rock gleaner OMNIVORE:bark excavator Ruddy Turnstone American Black Wiltiamson's Sap- Yellow-bellied Surfbird Oystereatcher sucker Sapsucker Rock Sandpiper Black Turnstone Red-breasted Sap- sucker MOLLUSCOVORE:fresh-marsh gleaner Limpkin OMNIVORE:coastal beachforager Common Ringed MOLLUSCOVORE:fresh-marsh hover-gleaner Plover Snail Kite Ruddy Turnstone MOLLUSCOVORE:freshwater bottom gleaner OMNIVORE:coastal beach scavenger White-winged Northwestern Scoter MOLLUSCOVORE: ground gleaner OMNIVORE:coastal bottom forager Ovenbird Steller's Eider MOLLUSCOVORE:marsh prober OMNIVORE:coastal dabbler Virginia Rail Northern Pintail MOLLUSCOVORE:inud prober Dunlin OMNIVORE:floral hover-gleaner Broad-billed Hum- Ruby-throated MOLLUSCOVORE:pelagic diver mingbird Razorbill White-eared Hum- Black-chinned Pigeon Guillemot mingbird Hummingbird MOLLUSCOVORE:pelagic plunger Buff-bellied Hum- Anna's Humming- White-tailed mingbird bird Violet-crowned Costa's Humming- Red-billed Tropic- Hummingbird bird bird Blue-throated Hummingbird MOLLUSCOVORE:pelagic scavenger Magnificent Hum- Short-tailed Alba- mingbird tross Lucifer Humming- bird MOLLUSCOVORE:pelagic surface gleaner Calliope Hum- Short-tailed Alba- mingbird tross Broad-tailed Hum- Laysan mingbird Greater Shear- Rufous Humming- water bird Bullet's Shear- Allen's Humming- water bird Audubon's Shear- water OMNIVORE:fresh- marsh forager Purple Gallinule MOLLUSCOVORE:salt-marsh gleaner Sofa Clapper Rail Seaside Sparrow Whooping MOLLUSCOVORE:shoreline gleaner OMNIVORE:freshwater bottom forager Semipalmated Canvasback Plover Ring-necked Duck MOLLUSCOVORE:shoreline prober Bar-tailed Godwit Surf Scoter Marbled Godwit Common Golden- Stilt Sandpiper eye 504 R.M. DeGraaf and others

Foraging Guild Classification (Continued) Foraging Guild Classification (Continued) Nonbreeding Nonbreeding Breeding period period Year round Breeding period period Year round OMNIVORE:freshwater bottom forager (continued) OMNIVORE:ground forager (continued) Barrow's Golden- Five-striped Spar- Lark Bunting eye row Grasshopper Spar- Savannah Sparrow row Fox Sparrow Henslow's Spar- OMNIVORE:freshwater dabbler Song Sparrow row Mallard Blue-winged Teal Lincoln's Sparrow Le Conte's Spar- Cinnamon Teal Swamp Sparrow row American Coot White-throated Sharp-tailed Spar- OMNIVORE:freshwater diver Sparrow row American Coot Golden-crowned Yellow-eyed Junco Sparrow Smith's Longspur OMNIVORE:fresh water strainer White-crowned Sparrow OMNIVORE: groundforager Harris' Sparrow Gray Partridge Northern Flicker Dark-eyed Junco Ring-necked Black-billed Mag- MeCown's Long- Pheasant pie Sandhitl Crane spur Spruce Grouse Yellow-billed , Lapland Longspur Blue Grouse Magpie Rock Dove Chestnut-collared Ruffed Grouse Eastern Bluebird Golden-fronted Longspur Greater Prairie- Western Bluebird Woodpecker Snow Bunting Chicken Hermit Thrush . Red-bellied Wood- McKay's Bunting Lesser Prairie- pecker Bobolink Chicken Varied Thrush Steller's Red-winged Black- Sharp-tailed Sage Thrasher bird Grouse Water Pipit Green Jay Bronzed Cowbird Wild Turkey Sprague's Pipit Brown Jay Brown-headed Montezuma Quail Pine Warbler Scrub Jay Cowbird Long-billed Cur- Palm Warbler Gray-breasted Jay Rosy Finch lew Ovenbird Pinyon Jay Cassin's Finch Eurasian Skylark Brewer's Sparrow Clark's Common Redpoll Horned Lark Eastern Meadow- Hoar~/Redpoll lark Chihuahuan Pine Siskin Gray-checked Western Meadow- Wren American Gold- Thrush lark Townsend's Soli- finch Swainson's Thrush Rusty Blackbird taire Wood Thrush Gray Catbird OMNIVORE:ground scavenger Northern Mock- ingbird Long-billed Curve-billed Thrasher OMNIVORE:lower-canopy/shrub forager Thrasher Bendire's Thrasher Veery Northern Flicker Steller's Jay Blue Grosbeak California Gray-checked Pileated Wood- Plain Titmouse Dickcissel Thrasher Thrush pecker Cactus Wren Botteri's Sparrow Crissal Thrasher Swainson's Thrush Black-capped American Robin Rufous-crowned European Star!ing Yellow-breasted Chickadee Gray Catbird Sparrow Crested Myna Chat Carolina Chick- Brown Thrasher American Tree Northern Varied Bunting adee Long-billed Sparrow Pyrrhuloxia Rufous-winged Mexican Chick- Thrasher Chipping Sparrow Olive Sparrow Sparrow adee Bendire's Thrasher Clay-colored Spar- Green-tailed Clay-colored Spar- Mountain Chick- Crissal Thrasher row Towhee row adee Lazuli Bunting Field Sparrow Rufous-sided Sage Sparrow Siberian Tit Black-chinned Towhee Song Sparrow Boreal Chickadee Painted Bunting Sparrow Brown Towhee Hooded Oriole Chestnut-backed Audubon's Oriole Vesper Sparrow Abert's Towhee Scott's Oriole Chickadee Lark Sparrow Bachman's Spar- Common Redpolt Bridled Titmouse Sage Sparrow row Hoary Redpoll Tufted Titmouse Foraging Guilds of North American Birds 505

Foraging Guild Classification (Continued) Foraging Guild Classification (Continued) Nonbreeding Nonbreeding Breeding period period Year round Breeding period period Year round OMNIVORE:lower-canopy/shrub forager (continued) OMNIVORE:upper-canopy forager American Gold- Verdin Elegant Trogon Cape May Gray Jay finch Bushtit Black-whiskered Warbler Blue Jay Eastern Bluebird Vireo Pinyon Jay Western Bluebird Hermit Warbler Clark's Nutcracker Mountain Bluebird Western Tanager Pygmy Nuthatch Hermit Thrush Rose-breasted Altamira Oriole Sage Thrasher Grosbeak White-eyed Vireo Black-headed Solitary Vireo Grosbeak Orange-crowned Northern Oriole Warbler Pine Grosbeak Magnolia Warbler Red Crossbill Black-throated White-winged Blue Warbler Crossbill Yellow-rumped Pine Siskin Warbler Evening Grosbeak Pine Warbler OMNIVORE:water bottom forager Brewer's Sparrow Oldsquaw Canvasback OMNIVORE:marsh dabbler Black Scoter Ring-neckedDuck Mottled Duck Lesser Scaup Common Golden- OMNIVORE: marsh forager eye Yellow Rail " Barrow's Golden- Virginia Rail eye SoFa Buffiehead Whooping Crane Ruddy Duck OMNIVORE: mud forager OMNIVORE:water dabbler Spectacled Eider OMNIVORE: mud prober Short-billed OMNIVORE:water strainer Dowitcher Northern Shoveler OMNIVORE:mud strainer PISCIVORE:coastal ambusher Greater Tricolored Heron PISCIVORE:coastal diver OMNIVORE:pelagic forager Common Murre Red-throated Brandt's Cormo- British Storm- Thick-billed Arctic Loon rant Petrel Murre Pelagic OMNIVORE:pelagic scavenger Black Guillemot Yellow-billedLoon Red-faced Cormo- Black-footed Alba- Pigeon Guillemot Horned Grebe rant tross Marbled Murrelet Red-neckedGrebe Rhinoceros Auklet Red-breasted Mer- OMNIu salt-marsh forager ganser Clapper Rail OMNIVORE:salt-marsh prober Horned Puffin King Rail PISCOVORE:coastal food pirate OMNIVORE: shorelineforager Magnificent Frig- Whimbrel Fish Crow atebird Bristle-thighed Laughing Gull Curlew PISCIVORE:coastal plunger Purple Sandpiper Elegant Tern Masked Booby Northern Gannet Herring Gull Roseate Tern Blue-footed Booby Brown OMNIVORE:shoreline prober Aleutian Tern Brown Booby Royal Tern Hudsonian Godwit Sooty Tern Red-footed Booby Sandwich Tern OMNIVORE:shoreline scavenger Caspian Tern Herring Gull Fish Crow 506 R.M. DeGraaf and others

Foraging Guild Classification (Continued) Foraging Guild Classification (Continued) Nonbreeding Nonbreeding Breeding period period Year round Breeding period period Year round PISCIVORE:coastal surface gleaner PISCIVORE:water ambusher Razorbill American White Tricolored Heron Least Bittern Pelican Magnificent Frig- Little Blue Heron atebird Reddish Egret Laughing Gull Green-backed Heermann's Gull Heron Black-crowned PISCIVORE:freshwater diver Night Heron Common Loon Anhinga Greater Yellow- Yellow-billed Loon Common Mer- leggs Horned Grebe ganser Red-necked Grebe PISCIVORE:water diver Western Grebe Red-throated Loon Western Grebe Great Cormorant Hooded Merganser Arctic Loon Hooded Merganser Double-crested Red-breasted Mer- Cormorant ganser Olivaceous Cormo- rant PISCIVORE:freshwater plunger Green Kingfisher PISCIVORE:water foot-plunger Great Kiskadee Bald Eagle PISCIVORE:freshwater surface gleaner White-tailed Eagle American White Pelican PISCIVORE:water plunger Caspian Tern Forster's Tern PISCIVORE: pelagic diver Common Tern Belted Kingfisher Arctic Loon Black-legged Kitti- Arctic Tern Red-necked Grebe wake Least Tern Audubon's Shear- Red-legged Kitti- water wake PISCIVORE:water skimmer Common Murre Black Skimmer Thick-billed PISCIVORE:water strainer Murre Roseate Spoonbill Black Guillemot Pigeon Guillemot VERMIVORE:coastal beach gleaner Marbled Murrelet. Black-bellied Rhinoceros Auklet Plover Tufted Puffin Snowy Plover Atlantic Puffin Western Sandpiper Horned Puffin Dunlin VERMIVORE:coastal beach prober PISCIVORE:pelagic plunger Bar-tailed Godwit Marbled Godwit White-tailed Short-billed Tropicbird Dowitcher Red-billed Tropic- bird VERMIVORE:freshwater shoreline gleaner Least Sandpiper PISCIVORE:pelagic scavenger Pink-footed Shear- VERMIVORE:ground gleaner water American Robin Greater Shear- VERMIVORE:ground prober water White-rumped American Wood- Manx Shearwater Sandpiper cock Black-vented Dunlin Shearwater Common Snipe PlSClVORE:pelagic surface gleaner VERMIVORE:mud gleaner Brown Noddy Heerman's Gull Common Black- Semipalmated Razorbill headed Gull Plover Foraging Guilds of North American Birds 507

Foraging Guild Classification (Continued) Acknowledgments

Nonbreeding We gratefully acknowledge P. B. Landres, R. M. Breeding period period Year round Erwin, M. C. Severinghaus, W. D. Severinghaus, J. VERMIVORE:shoreline gleaner Verner, and H. L. Short for reviewing the manuscript. Black-bellied Least Sandpiper C. S. Robbins, US Fish and Wildlife Service, suggested Plover that we expand our original manuscript, limited to the VERMIVORE:shoreline prober northeastern , to include North American Stilt Sandpiper birds north of Mexico.

Appendix I. Taxonomic list of North American birds and their foraging guilds. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Red-throated Loon Piscivore: Piscivore: Gavia stellata water diver coastal diver Arctic Loon Piscivore: Piseivore: Gavia arctica water diver coastal diver pelagic diver Common Loon Piscivore: " Piscivore: Gavia immer freshwater diver coastal diver Yellow-billed Loon Piscivore: Piscivore: Gavia adamsii freshwater diver coastal diver Least Grebe Insectivore: Tachybaptus dominicus freshwater diver Pied-billed Grebe Insectivore: Insectivore: Podilym bus freshwater diver water diver Crustaceovore: Crustaceovore: freshwater diver water diver Horned Grebe Insectivore: Crustaceovore: Podiceps auritus freshwater diver coastal diver Piscivore: Piscivore: freshwater diver coastal diver Red-necked Grebe Piscivore: Piscivore: Podiceps grisegena freshwater diver coastal diver Crustaceovore: pelagic diver freshwater diver Insectivore: freshwater diver Eared Grebe Crustaceovore: Insectivore: Podiceps nigricollis coastal diver freshwater diver Western Grebe Piscivore: Piscivore: Aechmophorus occidentalis freshwater diver water diver Short-tailed Albatross Molluscovore: Diomedea albatrus pelagic surface gleaner pelagic scavenger Carnivore: pelagic scavenger Black-footed Albatross Omnivore: Diomedea nigripes pelagic scavenger Molluscovore: Diomedea immutabilis pelagic surface gleaner Northern Fulmar Carnivore: Fulmarus glacialis pelagic scavenger Cory's Shearwater Carnivore: Calonectris diomedea pelagic scavenger Pink-footed Shearwater Piscivore: Puffinus creatopus pelagic scavenger 508 R.M. DeGraaf and others

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Greater Shearwater Piscivore: Puffinus gravis pelagic scavenger Molluscovore: pelagic surface gleaner Buller's Shearwater Crustaceovore: Puffinus bulleri pelagic surface gleaner Molluscovore: pelagic surface gleaner Sooty Shearwater Carnivore: Puffinus griseus pelagic scavenger Short-tailed Shearwater Crustaceovore: Puffinus tenuirostris pelagic scavenger Manx Shearwater Piscivore: Puffinus pujfinus pelagic scavenger Black-vented Shearwater Piscivore: Pu~nus opisthomelas pelagic scavenger Audubon's Shearwater Piscivore: Pu~nus lherminieri pelagic diver MoIluscovore: pelagic surface gleaner Wilson's Storm-Petrel Crustaceovore: Oceanites oceanicus pelagic scavenger British Storm-Petrel Omnivore: Hydrobates pelagicus pelagic forager Fork-tailed Storm-Petrel Carnivore: Oceanodroma furcata pelagic scavenger Leach's Storm-Petrel Carnivore: Oceanodroma leucorhoa pelagic scavenger Ashy Storm-Petrel Crustaceovore: Oceanodrorna hornochroa pelagic surface gleaner Black Storm-Petrel Carnivore: Oceanodroma rnelania pelagic scavenger Least Storm-Petrel Crustaceovore: Oceanodroma rnicrosoma pelagic surface gleaner White-tailed Tropicbird Piscivore: Phaethon lepturus pelagic plunger Molluscovore: pelagic plunger Red-billed Tropicbird Piscivore: Phaethon aethereus pelagic plunger Molluscovore: pelagic plunger Masked Booby Piscivore: Sula dactylatra coastal plunger Blue-footed Booby Piscivore: Sula nebouxii coastal plunger Brown Booby Piscivore: Sula leucogaster coastal plunger Red-footed Booby Piscivore: Sula sula coastal plunger Molluscovore: coastal plunger Northern Gannet Piscivore: Sula bassanus coastal plunger American White Pelican Piscivore: Piscivore: Pelecanus erythrorhynchos freshwater surface gleaner coastal surface gleaner Piscivore: Pelecanus occidentalis coastal plunger Foraging Guilds of North American Birds 509

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Great Cormorant Piscivore: Phalacrocorax carbo water diver Double-crested Cormorant Piscivore: Phalacrocorax auritus water diver Olivaceous Cormorant Piscivore: Phalacrocorax olivaceus water diver Brandt's Cormorant Piscivore: Phalacrocorax penicillatus coastal diver Pelagic Cormorant Piscivore: Phalacrocorax pelagicus coastal diver Red-faced Cormorant Piseivore: Phalacrocorax urile coastal diver Crustaceovore: coastal diver Anhinga Piscivore: Anhinga anhinga freshwater diver Magnificent Frigatebird Piscivore: Fregata magnificens coastal surface gleaner coastal food pirate American Bittern Carnivore: Botaurus lentiginosus water ambusher Insectivore: water ambusher Crustaeeovore: water ambusher Least Bittern Piscivore: Isobrychus exilis water ambusher Insectivore: water ambusher Great Blue Heron Piscivore: Ardea herodias water ambusher Great Egret Carnivore: Casmerodius albus water ambusher Crustaceovore: water ambusher Snowy Egret Crustaceovore: Egretta thula water ambusher Carnivore: water ambusher Little Blue Heron Crustaceovore: Egretta caerulea water ambusher Piscivore: water ambusher Tricolored Heron Piscivore: Piscivore: Egretta tricolor coastal ambusher water ambusher Crustaceovore: Crustaceovore: coastal ambusher water ambusher Reddish Egret Piscivore: Egretta rufescens water ambusher Cattle Egret Insectivore: Bubulcus ibis ground gleaner Green-backed Heron Crustaceovore: Butorides striatus water ambusher Piscivore: water ambusher Black-crowned Night-Heron Piseivore: Nycticorax nycticorax water ambusher Crustaceovore: water ambusher 510 R M. DeGraaf and others

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Yellow-crowned Night-Heron Crustaceovore: Nycticorax violaceus water ambusher White Ibis Crustaceovore: Eudocirnus albus mud prober Scarlet Ibis Crustaceovore: Eudocimus ruber mud prober Molluscovore: mud prober Glossy Ibis Crustaceovore: falcinellus mud proher Insectivore: mud gleaner White-faced Ibis Crustaceovore: Plegadis chihi mud prober Insectivore: mud gleaner Roseate Spoonbill Piscivore: Ajaia ajaia water strainer Crustaceovore: mud strainer Wood Stork Carnivore: Mycteria americana water ambusher Crustaceovore: water ambusher Molluscovore: water ambusher Greater Flamingo Omnivore: Phoenicopterus ruber mud strainer Fulvous Whistling-Duck Granivore: Dendrocygna bicolor ground gleaner freshwater dabbler Black-bellied Whistling-Duck Herbivore: Dendrocygna autumnalis freshwater dabbler Granivore: ground gleaner Tundra Swan Herbivore: Herbivore: Cygnus columbianus freshwater dabbler water dabbler Whooper Swan Herbivore: Cygnus cygnus coastal dabbler Trumpeter Swan Herbivore: Herbivore: Cygnus buccinator freshwater dabbler water dabbler Mute Swan Herbivore: Herbivore: Cygnus olor freshwater dabbler water dabbler ground grazer Pink-footed Goose Herbivore: Anser brachyrhynchus ground grazer Greater White-fronted Goose Herbivore: Anser albifrons ground grazer Snow Goose Herbivore: Herbivore: Chen caerulescens water grubber ground grazer Ross' Goose Herbivore: Chen fossil ground grazer Emperor Goose Herbivore: Herbivore: Chen canagica ground grazer coastal dabbler Brant Herbivore: Herbivore: Branta bernicla ground grazer coastal dabbler Barnacle Goose Herbivore: Branta leucopsis ground grazer C~anada Goose Herbivore: Herbivore: Herbivore: Branta canadensis water dabbler watcr dabbler ground grazer Foraging Guilds of North American Birds 51 1

Appendix 1. Continued. Species name (Sei. name AOU list) Breeding period Nonbreeding period Year round Wood Duck Granivore: sponsa ground gleaner freshwater surface gleaner Green-winged Teal Granivore: Granivore: Anas crecca freshwater dabbler water dabbler American Black Duck Omnivore: Anas rubripes water dabbler Mottled Duck Omnivore: Anas Julvigula marsh dabbler Mallard Granivore: Herbivore: Granivore: Anas platyrhynchos ground gleaner ground grazer freshwater dabbler Northern Pintail Granivore: Omnivore: Anas acuta freshwater dabbler coastal beach forager coastal dabbler Blue-winged Teal Omnivore: Anas discors freshwater dabbler Cinnamon Teal Omnivore: Herbivore: Anas cyanoptera freshwater dabbler freshwater dabbler Northern Shoveler Omnivore: 0mnivore: Anas clypeata freshwater strainer water strainer Gadwall Herbivore: Herbivore: Anas strepera freshwater dabbler water dabbler Eurasian Wigeon Herbivore: Anas penelope coastal dabbler American Wigeon Herbivore: Herbivore: Anas americana ground grazer water dabbler Canvasback Omnivore: Omnivore: valisineria freshwater bottom forager water bottom forager Redhead Herbivore: Aythya americana water diver Ring-necked Duck Omnivore: Omnivore: Aythya collaris freshwater bottom forager water bottom forager Tufted Duck Molluscovore: Aythya fuligula water bottom gleaner Crustaceovore: water bottom gleaner Greater Scaup Omnivore: Molluscovore: Aythya marila freshwater bottom gleaner coastal bottom gleaner Lesser Scaup Crustaceovore: Omnivore: Aythya affinis freshwater bottom gleaner water bottom forager Common Eider Molluscovore: Somateria mollissima coastal bottom gleaner Crustaccovore: coastal bottom gleaner King Eider Molluscovore: Molluscovore: Somateria spectabilis water bottom gleaner coastal bottom gleaner Crustaceovore: Crustaceovore: water bottom gleaner coastal bottom gleaner Spectacled Eider Omnivorei Molluscovore: Somateria fisheri water dabbler coastal bottom gleaner Steller's Eider Omnivore: Molluscovore: Polysticta stelleri coastal bottom forager coastal bottom gleaner Crustaceovore: coastal bottom gleaner Harlequin Duck Insectivore: Crustaceovore: Histrionicus histrionicus riparian bottom gleaner coastal bottom gleaner Molluseovore: coastal bottom gleaner 512 R.M. DeGraaf and others

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Oldsquaw C)mniv~ Crustaceovore: Clangula hyemalis water bottom forager coastal bottom gleaner Molluscovore: coastal bottom gleaner Black Scoter Omnivore: Molluscovore: Melanitta nigra water bottom forager coastal bottom gleaner Crustaceovore: coastal bottom gleaner Surf Scoter Omnivore: Molluscovore: Melanitta perspicillata freshwater bottom forager coastal bottom forager Crustaceovore: coastal bottom gleaner White-winged Scoter Molluscovore: Molluscovore: Melanitta fusca freshwater bottom gleaner coastal bottom gleaner Crustaceovore: Crustaceovore: freshwater bottom gleaner coastal bottom gleaner Omnivore: Omnivore: Bucephala clangula freshwater bottom forager water bottom forager Barrow's Goldeneye Omnivore: Omnivore: Bucephala islandica freshwater bottom forager water bottom forager Bufflehead Insectivore: Omnivore: Bucephala albeola freshwater bottom gleaner water bottom gleaner Hooded Merganser Piscivore: Piscivore: Lophodytes cucullatus freshwater diver water diver Insectivore: Insectivore: freshwater bottom gleaner water bottom gleaner Crustaceovore: Crustaccovore: freshwater bottom gleaner water bottom gleaner Common Merganser Piscivore: merganser freshwater diver Red-breasted Merganser Piscivore: Piscivore: Mergus serrator freshwater diver coastal diver Ruddy Duck Omnivore: Omnivore: Oxyura jamalcensis freshwater bottom forager water bottom forager Masked Duck Herbivore: Herbivore: Oxyura dominica freslawater diver water diver Black Vulture Carnivore: Coragyps atratus ground scavenger Turkey Vulture Carnivore: Cathartes aura ground scavenger California Condor Carnivore: Gymnogyps californianus ground scavenger Osprey Piscivore: Pandion haliaetus water foot-plunger American Swallow-tailed Kite Insectivore: Elanoides forficatus air hawker Carnivore: ground hawker Black-shouldered Kite Carnivore: Elanus caeruleus ground hawker Snail Kite Molluscovorc: Rostrhamus sociabilis fresh-marsh hover-gleaner Mississippi Kite Insectivore: lctinia mississippiensis air hawker Bald Eagle Piscivorc: Haliaeetus leucocephalus water foot-plunger Carnivore: ground scavenger Foraging Guilds of North American Birds 513

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round White-tailed Eagle Piscivore: Haliaeetus albicilla water foot-plunger Carnivore: air hawker Northern Harrier Carnivore: Circus cyaneus ground hawker Sharp-shinned Hawk Carnivore: Accipiter striatus air hawker ground hawker Cooper's Hawk Carnivore: Accipiter cooperii air hawker ground hawker Northern Goshawk Carnivore: Accipiter gentilis air hawker ground hawker Common Black-Hawk Carnivore: Buteogallus anthracinus ground hawker Harris' Hawk Carnivore: Parabuteo unicinctus ground hawker Gray Hawk Carnivore: Buteo nitidus ground hawker Insectivore: air hawker Red-shouldered Hawk Carnivore: Buteo lineatus ground hawker Broad-winged Hawk Carnivore: Buteo platypterus ground hawker Short-tailed Hawk Carnivore: Buteo brachyurus ground hawker Swainson's Hawk Carnivore: Buteo swainsoni ground hawker Insectivore: air hawker White-tailed Hawk Carnivore: Buteo albicaudatus ground hawker Zone-tailed Hawk Carnivore: Buteo albonotatus ground hawker Red-tailed Hawk Carnivore: Buteo jarnaicensis ground hawker Ferruginous Hawk Carnivore: Buteo regalis ground hawker Rough-legged Hawk Carnivore: Buteo lagopus ground hawker Golden Eagle Carnivore: Carnivore: Aquila chrysaetos ground scavenger ground hawker Crested Caracara Carnivore: Polyborus plancus ground scavenger American Kestrel Insectivore: Carnivore: Falco sparverius air hawker ground hawker Merlin Carnivore: Falco columbarius air hawker Aplomado Falcon Carnivore: Falco fernoralis ground hawker Insectivore: air hawker Peregrine Falcon Carnivore: Falco peregrinus air hawker Gyrfalcon Carnivore: Falco rusticolus air hawker 514 R.M. DeGraaf and others

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Prairie Falcon Carnivore: Falco mexicanus air hawker ground hawker Plain Chachalaca Frugivore: Ortalis vetuta ground gleaner Herbivore: upper-canopy forager Gray Partridge Omnivore: Granivore: Perdix perdix ground forager ground gleaner Chukar Herbivore: Alectoris chukar ground forager Ring-necked Pheasant Omnivore: Herbivore: Phasianus colchicus ground forager ground forager Spruce Grouse 0mnivore: Herbivore: Dendragapus canadensis ground forager upper-canopy forager Blue Grouse Omnivore: Herbivore: Dendragapus obscurus ground forager upper-canopy forager Willow Ptarmigan Herbivore: Herbwore: Lagopus lag@us ground forager lower-canopy forager Rock Ptarmigan Herbivore: Herbivore: Lag@us taurus ground forager lower-canopy forager White-tailed Ptarmigan Herbivore: Herbivore: Lagopus leucurus ground forager lower-canopy forager Ruffed Grouse 0mnivore: Herbivore: Bonasa umbellus ground forager upper-canopy forager Sage Grouse Herbivore:. Centrocercus urophasianus ground forager Greater Prairie-Chicken Omnivore: Herbivore: Tympanuchus cupido ground forager ground forager Lesser Prairie-Chicken 0mnivore: Herbivore: Tympanuchus pallidicinctus ground forager ground forager Sharp-tailed Grouse Omnivore: Herbivore: Tympanuchus phasianellus ground forager upper-canopy forager Wild Turkey 0mnivore: . Herbivore: Meleagris gallopavo ground forager ground forager Montezuma Quail Omniv0re: Herbivore: Cyrtonyx montezurnae ground forager ground grubber Northern Bobwhite Omnivore: virginianus ground forager Scaled Quail 0mnivore: squamata ground fora~er Gambel's Quail Herbivore: Callipepla garnbelii ground forager California Quail Granivore: Herbivore: Callipepla californica groun d g!eaner ground forager Mountain Quail Herbivore: Granivore: Oreortyx pictus ground forager ground gleaner Yellow Rail 0mnivore: Omnivore: Coturnicops noveboracensis fresh-marsh forager marsh forager Black Rail Insectivore: Crustaceovore: Laterallus jamaicensis marsh gleaner salt-marsh gleaner Crustaceovore: marsh gleaner Clapper Rail Crustaceovore: Omnivore: longirostris salt-marsh gleaner salt-marsh forager Molluseovore: salt-marsh gleaner Foraging Guilds of North American Birds 515

Appendix I. Continued. Species name (Sci, name AOU list) Breeding period Nonbreeding period Year round King Rail Crustaceovore: Omnivore: Rallus elegans marsh prober salt-marsh prober Insectivore: marsh prober Virginia Rail Insectivore: Omnivore: Rallus limicola marsh prober marsh forager Molluscovore: marsh prober Sofa Omnivore: Omnivore: carolina fresh-marsh forager marsh forager Purple Gallinule Omnivore: Porphyrula martinica fresh-marsh forager Common Moorhen Omnivore: Gallinula, chloropus fresh-marsh forager American ,Coot Omnivore: Herbivore: Fulica americana freshwater dabbler water dabbler freshwater diver water diver Limpkin Molluscovore: Aramus guarauna fresh-marsh gleaner Sandhill Crane Omnivore: canadensis fresh-marsh forager ground forager, Whooping Crane Omnivore: Omnivore: Omnivore: Grus americana fresh-marsh forager marsh forager ground forager Black-bellied Plover Crustaceovore: Crustaceovore: Pluvialis squatarola shoreline gleaner coastal beach gleaner Insectivore: Molluscovore: shoreline gleaner coastal beach gleaner Vermivore: Insectivore: shoreline gleaner coastal beach gleaner Vermivore: coastal beach gleaner Lesser Golden-Plover Insectivore: Pluvialis dominica ground gleaner Snowy Plover Crustaceovore: Crustaceovore: Charadrius alexandrinus shoreline gleaner coastal beach gleaner Vermivore: coastal beach gleaner Wilson's Plover Insectivore: Charadrius wilsonia coastal beach gleaner Crustaceovore: coastal beach gleaner Common Ringed Plover Omnivore: Charadrius hiaticula coastal beach forager Semipalmated Plover Molluscovore: Molluscovore: Vermivore: CharadHus semipalmatus shoreline gleaner coastal beach gleaner mud gleaner Crustaceovore: Crustaceovore: shoreline gleaner coastal beach gleaner Insectivore: Insectivore: shoreline gleaner coastal beach gleaner Piping Plover Insectivore: Insectivore: Charadrius melodus shoreline gleaner coastal beach gleaner Crustaceovore: Crustaceovore: shoreline gleaner coastal beach gleaner Killdeer Insectivore: Insectivore: Charadrius vociferus ground gleaner shoreline gleaner Mountain Plover Insectivore: Charadrius montanus ground gleaner 516 R.M. DeGraaf and others

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Eurasian Dotterel Insectivore: Charadrius morinellus coastal beach gleaner American Oystercatcher Molluscovore: Haematopus palliatus coastal beach prober American Black Oystercatcher Molluscovore: Haematopus bachmani coastal rock gleaner Black-necked Stilt Insectivore: Himantopus mexicanus freshwater gleaner American Avocet Omnivore: Recurvirostra americana marsh forager Northern Jacana Insectivore: Jacana spinosa fresh-marsh gleaner Greater Yellowlegs Piscivore: melanoleuca water ambusher Insectivore: water surface gleaner Lesser Yellowlegs Insectivore: Tringa flavipes water surface gleaner Crustaceovore: water surface gleaner Wood Sandpiper Insectivore: Tringa glareola freshwater gleaner Solitary Sandpiper Insectivore: Insectivore: Crustaceovore: Tringa solitaria freshwater gleaner water surface gleaner freshwater gleaner Willet Insectivore: Insectivore: Catoptrophorus shoreline prober coastal beach prober semipalmatus Crustaceovore: Crustaceovore: shoreline prober coastal beach, prober Wandering Tattler Insectivore: Insectivore: Heteroscelus incanus freshwater shoreline gleaner coastal beach gleaner Spotted Sandpiper Insectivore: macularia shoreline gleaner Upland Sandpiper Insectivore: Bartramia longicauda ground gleaner Eskimo Curlew Insectivore: Numenius borealis ground gleaner Whimbrel Omnivore: Crustaceovore: Numenius phaeopus shoreline forager coastal beach prober Molluscovore: coastal beach prober Bristle-thighed Curlew Omnivore: Numenius tahitiensis shoreline forager Long-billed Curlew Omnivore: Insectivore: Numenius ameHcanus ground forager shoreline gleaner Crustaceovore: shoreline prober Hudsonian Godwit Omnivore: Limosa haemastica shoreline prober Bar-tailed Godwit Insectivore: Limosa lapponica shoreline prober Molluscovore: shoreline prober Crustaceovore: shoreline prober Vermivore: coastal beach prober Marbled Godwit Insectivore: Crustaceovore: Limosa fedoa _ shoreline prober coastal beach prober Foraging Guilds of North American Birds 517

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Molluscovore: Molluscovore: shoreline prober coastal beach prober Vermivore: coastal beach prober Ruddy Turnstone Omnivore: Crustaceovore: Arenaria interpres coastal beach forager coastal rock gleaner Molluscovore: coastal rock gleaner Black Turnstone Molluscovore: Arenaria melanocephala coastal rock gleaner Crustaceovore: coastal rock gleaner coastal beach gleaner Sur~ird Insectivore: IV~olluscovore: Aphriza uirgata ground gleaner coastal rock gleaner Crustaceovore: coastal rock gleaner Red Knot Crustaceovore: Molluscovore: canutus freshwater shoreline gleaner coastal beach gleaner Insectivore: Crustaceovore: freshwater shoreline gleaner coastal beach gleaner Sanderling Insectivore: Crustaceovore: Calidris alba freshwater shoreline gleaner coastal beach prober Cru'staceovore: Molluscovore: freshwater shoreline gleaner coastal beach prober Semipalmated Sandpiper Insectivore: Calidris pusilla shoreline gleaner Crustaceovore: shoreline gleaner Western Sandpiper Insectivore: Insectivore: Calidrix mauri shoreline gleaner coastal beach gleaner Crustaceovore: Molluscovore: shoreline gleaner coastal beach gleaner Vermivore: coastal beach gleaner Rufous-necked Stint Insectivore: Calidris ruficoltis shoreline gleaner Crustaceovore: shoreline gleaner Least Sandpiper Insectivore: Insectivore: Calidris minutilla freshwater shoreline gleaner shoreline gleaner Crustaceovore: Crustaceovore: freshwater shoreline gleaner shoreline gleaner Vermivore: Vermivore: freshwater shoreline gleaner shoreline gleaner White-rumped Sandpiper Insectivore: Calidris fuscicollis mud gleaner Crustaceovore: mud prober Vermivore: ground prober Baird's Sandpiper Insectivore: Calidris bairdii mud gleaner Crustaceovore: mud gleaner Pectoral Sandpiper Insectivore: Calidris melanotos ground gleaner 518 R.M. DeGraaf and others

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Purple Sandpiper Omnivore: Insectivore: Calidris maritima shoreline forager coastal rock gleaner Crustaceovore: coastal rock gleaner Rock Sandpiper Insectivore: Molluscovore: Calidris ptilocnemis ground gleaner coastal rock gleaner Crustaccovore: Crustaceovore: shoreline gleaner coastal rock gleaner Insectivore: coastal rock gleaner Dunlin Insectivore: Molluscovore: Calidris alpina mud gleaner coastal beach prober Molluscovore: Crustaceovorc: mud prober coastal beach prober Crustaceovore: Vcrmivorc: mud prober coastal beach gleaner Vermivore: Insectivore: ground prober coastal beach gleaner Curlew Sandpiper Insectivore: Calidris ferruginea grodnd gleaner Stilt Sandpiper Vermivore: Calidris himantopus shoreline prober Insectivore: shoreline gleaner Crustaceovore: shoreline prober Molluscovore: shoreline prober Buff-breasted Sandpiper Insectivore: Tryngites subruficollis ground gleaner Short-billed Dowiteher Omnivore: Insectivore: Limnodromus griseus mud prober coastal beach gleaner Vcrmivore: coastal beach prober Long-billed Dowitcher Insectivore: Insectivore: Limnodromus scolopaceus ground gleaner coastal beach gleaner Common Snipe Vermivore: Omnivore: gallinago ground prober mud forager American Vermivore: Scolopax minor ground prober Wilson's Phalarope Insectivore: Phalaropus tricolor freshwater shoreline gleaner Red-necked Phalarope Insectivore: Phalaropus lobatus water surface gleaner Red Phalarope Insectivore: Phalaropus fulicaria water surface gleaner Crustaceovore: water surface gleaner Pomarine Jaeger Carnivore: Carnivore: Stercorarius pomarinus coastal scavenger pelagic scavenger air hawker pelagic food pirate g.round hawker Parasitic Jaeger Camivore: Carnivore: Stercorarius parasiticus coastal food pirate pelagic food pirate ground scavenger Long-tailed Jaeger Carnivore: Stercorarius longicqudus air hawker ground hawker Foraging Guilds of North American Birds 519

Appendix I. Continued, Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Great Skua Carnivore: Carnivore: Catharacta skua coastal food pirate pelagic scavenger coastal scavenger ground hawker South Polar Skua Carnivore: Catharacta maccormicki pelagic scavenger Laughing Gull Carnivore: Piscivore: Larus atricilla coastal beach scavenger coastal surface gleaner coastal food pirate Franklin's Gull Insectivore; Larus pipixean air hawker Little Gull Insectivore: Larus minutus freshwater surface gleaner Common Black-headed Gull Insectivore: Larus ridbundus air hawker Carnivore: coastal beach scavenger Vermivore: mud gleaner Bonaparte's Gull Insectivore: Carnivore: Larus philadelphia air hawker coastal beach scavenger ground gleaner coastal surface gleaner Heermann's Gull Piscivore: Larus heerman n i pelagic surface gleaner coastal surface gleaner Carnivore: coastal beach scavenger Mew Gull Carnivore: Carnivore: L~2r~s CaT2~IS shoreline gleaner coastal beach scavenger Insectivore: ground gleaner Ring-billed Gull Insectivore: Carnivore: Larus delawarensis ground gleaner coastal beach scavenger Carnivore: ground hawker California Gull Insectivore: Carnivore: Larus californicus ground gleaner coastal beach scavenger Carnivore: ground hawker Herring Gull Omnivore: Carnivore: Larus argentatus shoreline scavenger shoreline scavenger Thayer's Gull Carnivore: Carnivore: Larus thayeri coastal beach scavenger shoreline scavenger Iceland Gull Carnivore: Carnivore: Larus glaucoides coastal beach scavenger coastal scavenger Lesser Black-backed Gull Carnivore: Larus fuscus shoreline scavenger Yellow-footed Gull Carnivore: Larus livens coastal beach scavenger Western Gull Carnivore: Larus occidentalis coastal beach scavenger Glaucous-winged Gull Carnivore: Larus glaucescens coastal beach scavenger Glaucous Gull Carnivore: Carnivore: Larus hyperboreus air hawker coastal scavenger Great Black-backed Gull Carnivore: Larus marinus coastal beach scavenger Black-legged Piscivore: Rissa tridactyla pelagic diver 520 R.M. DeGraaf and others

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Red-legged Kittiwake Piscivore: Rissa brevirostris pelagic diver Sabine's Gull Insectivore: Xema sabini freshwater surface gleaner Ivory Gull Carnivore: Carnivore: Pagophila eburnea ground hawker pealgic scavenger Gull-billed Tern Insectivore: Sterna nilotica water surface gleaner Caspian Tern Piscivore: Piscivore: Sterna caspia water plunger coastal plunger Royal Tern Piscivore: Sterna maxima coastal plunger Elegant Tern Piscivore: Sterna elegans coastal plunger Sandwich Tern Piscivore: . Sterna sandvicensis coastal plunger Roseate Tern Piscivore: Sterna dougallii coastal plunger Common Tern Piscivore: Piscivore: Sterna hirundo water plunger coastal plunger Arctic Tern Piscivore: Sterna paradisaea water plunger Aleutian Tern Piscivore: Sterna aleutica coastal plunger Forster's Tern Insectivore: Sterna [orsteri water surface gleaner Piscivore: water plunger Least Tern Piscivore: Sterna antillarum water plunger . Sooty Tern Piscivore: Sterna fuscata coastal plunger Black Tern Insectivore: Chlidonias niger freshwater surface gleaner air hawker Brown Noddy Piscivore: Anous stolidus pelagic surface gleaner Black Skimmer Piscivore: Rynchops niger water skimmer Crustaceovore: water skimmer Dovekie Crustaceovore: Crustaceovore: Alle alle coastal diver pelagic diver Common Murre Piscivore: Piscivore: aalge coastal diver pelagic diver Crustaceovore: Crustaceovore: coastal diver pelagic diver Thick-billed Murre Piscivore: Piscivore: Uria lomvia coastal diver pelagic diver Crustaceovore: Crustaceovore: coastal surface gleaner pelagic surface gleaner Razorbill Piscivore: Piseivore: Alca torda coastal surface gleaner pelagic surface gleaner Crustaceovore: Crustaceovore: coastal surface gleaner pelagic surface gleaner Molluscovore: Molluscovore: coastal diver pelagic diver Black Guillemot Crustaceovore: Crustaceovore: grille coastal diver pelagic diver Foraging Guilds of North American Birds 521

Appendix I. Continued., Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Piscivore: Piscivore: coastal diver pelagic diver Pigeon Guillemot Piscivore: Piscivore: Cepphus columba coastal diver pelagic diver Crustaceovore; Crustaceovore: coastal diver pelagic diver Molluscovore: pelagic diver Marbled Murrelet Piscivore: Piscivore: Brachyramphus marmoratus coastal diver pelagic diver Kittlitz's Murrelet Crustaceovore: Brachyramphus brevirostris coastal diver Xantus' Murrelet Crustaceovore: Crustaceovore: coastal diver petagic diver hypoleucus Ancient Murrelet Crustaceovore: Crustaceovore: Synthliboramphus antiquus coastal diver pelagic diver Cassin's Auklet Crustaceovore: Crustaceovore: Ptychoramphus aleuticus coastal diver pelagic diver Parakeet Auklet Crustaceovore: Crustaceovore: Cyclorrhynchus psittacula coastal diver pelagic diver Least Auklet Crustaceovore: C rustaceovore: Aethia pusilla coastal diver pelagic diver Whiskered Auklet Crustaceovore: Aethia pygmaea coastal diver pelagic diver Crested Auklet Crustaceovore: Crustaceovore: Aethia cristatella coastal diver pelagic diver Rhinocerus Auklet Crustaceovore: Crustaceovore: Cerorhinca monocerata coastal diver pelagic diver Piscivore: Piscivore: coastal diver pelagic diver Tufted Puffin Piscivore: Piscivore: Fratercula cirrhata coastal diver pelagic diver Atlantic Puffin Piscivore: Piscivore: Fratercula arctica coastal diver pelagic diver Horned Puffin Piscivore: Piscivore: Fratercula corniculata coastal diver pelagic diver Rock Dove Omnivore: Columba livia ground forager White-crowned Pigeon Frugivore: Columba leucocephata upper-canopy gleaner Red-billed Pigeon Frugivore: Columba flavirostris upper-canopy gleaner Band-tailed Pigeon Frugivore: Frugivore: Columba fasciata lower-canopy gleaner upper-canopy gleaner White-winged Dove Granivore: Zenaida asiatica ground gleaner Mourning Dove Granivore: Zenaida macroura ground gleaner Inca Dove Granivore: Columbina inca ground gleaner Common Ground-Dove Granivore: Columbina ground gleaner White-tipped Dove Granivore: LeptotiIa verreauxi ground gleaner Black-billed Cuckoo Insectivore: Coccyzus erythropthalmus lower-canopy gleaner Yellow-billed Cuckoo Insectivore: Coccyzus americanus lower-canopy gleaner 522 R, M. DeGraaf and others

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Mangrove Cuckoo Insectivore: Coccyzus minor lower-canopy gleaner Greater Roadrunner Insectivore: Geococcyx californianus ground gleaner Carnivore: ground chaser Smooth-billed Ani Insectivore: Crotophaga ani ground gleaner Groove-billed Ani Insectivore: Crotophaga sulcirostris ground gleaner Common Barn-Owl Carnivore: Tyto alba ground hawker Flammulated Owl Insectivore: Otus flammeolus ground hawker Eastern Screech-Owl Insectivore: Carnivore: Otus asio ground hawker ground hawker Western Screech-Owl Insectivore: Carnivore:, Otus kennicottii ground hawker ground hawker Whiskered Screech-Owl Insectivore: Otus trichopsis ground hawker Great Horned Owl Carnivore: Bubo virginianus ground hawker Snowy Owl Carnivore: Nyctea scandiaca ground hawker Northern Hawk-Owl Carnivore: Surnia ulula ground hawker Northern Pygmy-Owl Insectivore: Carnivore: Glaucidium gnoma ground hawker ground hawker Ferruginous Pygmy-Owl Carnivore: Glaucidium brasilianum ground hawker Elf Owl Insectivore: Micrathene whitneyi ground hawker air hawker Burrowing Owl Carnivore: Athene cunicularia ground hawker Insecuvore: air hawker Spotted Owl Carnivore: Strix occidentalis ground hawker Barred Owl Carnivore: Strix varia ground hawker Great Gray Owl Carnivore: Strix nebulosa ground hawker Long-eared Owl Carmvore: Asio otus ground hawker Short-eared Owl Carnivore: Asio flammeus ground hawker Boreal Owl Carnivore: Aegolius funereus ground hawker Northern Saw-whet Owl Carnivore: Aegolius acadicus ground hawker Lesser Nighthawk Insectivore: acutipennis air screener Insectivore: ChordeiIes minor air screener Antillean Nighthawk Insectivore: Chordeiles gundlachii air screener Common Pauraque Insectivore: Nyctidromus albicoUis air screener Foraging Guilds of North American Birds 523

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Common Poorwill Insectivore: Phalaenoptilus nuttaUii air screener Chuck-will's-widow Insectivore: Caprirnulgus carolinensis air screener Buff-collared Nightjar Insectivore: Caprimulgus ridgwayi air screener Whip-poor-will Insectivore: Caprimulgus vociferus air screener Black Swift Insectivore: Cypseloides niger air screener Chimney Swift Insectivore: Chaetura pelagica air screener Vaux's Swift Insectivore: Chaetura vauxi air screener White-throated Swift Insectivore: Aeronautes saxatalis air screener Broad-billed Hummingbird Omnivore: Cynanthus latirostris floral hover-gleaner White-eared Hummingbird Omnivore: Hylocharis leucotis floral hover-gleaner Buff-bellied Hummingbird Omnivore: Amazitia yucatanensis floral hover-gleaner Violet-crowned Hummingbird Omnivore: Amazilia violiceps floral hover-gleaner Blue-throated Hummingbird :Omnivore: Larnpornis clemenciae floral hover-gleaner Magnificent Hummingbird Omnivore: Eugenes fulgens floral hover-gleaner Lucifer Hummingbird Omnivore: Calothorax lucifer floral hover-gleaner Ruby-throated Hummingbird Omnivore: Archilochus colubris floral hover-gleaner Black-chinned Hummingbird Omnivore: Archilochus alexandri floral hover-gleaner Anna's Hummingbird Omnivore: Calypte anna floral hover-gleaner Insectivore: air sallyer Costa's Hummingbird Omnivore: Calypte costae floral hover-gleaner Calliope Hummingbird Omnivore: Stellula calliope floral hover-gleaner Broad-tailed Hummingbird Omnivore: Selasphorus platycercus floral hover-gleaner Rufous Hummingbird Omnivore: Selasphorus .rufous floral hover-gleaner Allen's Hummingbird Omnivore: Selasphorus sasin floral hover-gleaner Elegant Trogon Omnivore: Trogon elegans upper-canopy foragei Insectivore: air sallyer Belted Kingfisher Piscivore: Ceryle alcyon water plunger Green Kingfisher Piscivore: Chloroceryle americana freshwater plunger Insectivore: air sallier 524 R.M. OeGraaf and others

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Lewis' Woodpecker Insectivore: Melanerpes lewis air sallyer Granivore: upper-canopy gleaner Red-headed Woodpecker Insectivore: Frugivore: Insecuvore: Melanerpes erythrocephalus air sallier lower-canopy gleaner bark gleaner Acorn Woodpecker Frugwore: Melanerpes formicivorus upper-canopy gleaner Insectivore: bark gleaner Gila Woodpecker Insectivore: Melanerpes uropygialis bark gleaner Frugwore: upper-canopy gleaner Golden-fronted Woodpecker Insectivore: Melanerpes aurifrons bark gleaner Omnivore: ground forager Red-bellied Woodpecker Insectivore: Melanerpes carolinus bark gleaner Omnivore: ground forager Yellow-bellied Sapsucker Insectivore: Omnivore: Sphyrapicus varius bark gleaner bark excavator Red-breasted Sapsucker Insectivore: Omnivore: Sphyrapicus tuber bark gleaner bark excavator Williamson's Sapsucker Omnivore: Insectivore: Sphyrapicus thyroideus bark excavator bark gleaner Ladder-backed Woodpecker Insectivore: Picoides scalaris bark gleaner Frugivore: lower-canopy gleaner Nuttall's Woodpecker Insectivore: Picoides nuttallii bark gleaner Frugivore: lower-canopy gleaner Downy Woodpecker Insectivore: Picoides pubescens bark gleaner Frugivore: lower-canopy gleaner Hairy Woodpecker Insectivore: Picoides villosus bark gleaner Frugwore: lower-canopy gleaner Strickland's Woodpecker Insectivore: Picoides stricklandi bark gleaner Fruglvore: upper-canopy gleaner Red-cockaded Woodpecker Granivore: Insectivore: Picoides borealis upper-canopy gleaner bark gleaner White-headed Woodpecker Granivore: Insectivore: Picoides albolarvatus upper-canopy gleaner bark scaler Three-toed Woodpecker Insectwore: Picoides tridactylus bark scaler Black-backed Woodpecker Insectivore: Picoides arcticus bark scaler Northern Flicker Insectivore: Omnivore: Colaptes auratus ground gleaner ground forager lower-canopy forager Foraging Guilds of North American Birds 525

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Pileated Woodpecker Omnivore: Insectivore: Dryocopus pileatus lower-canopy forager bark excavator Ivory-billed Woodpecker Insectivore: Campephilus principalis bark scaler Frugivore: upper-canopy gleaner Northern Beardless- Insectivore: Tyrannulet upper-canopy gleaner Camptostoma imberbe Olive-sided Flycatcher Insectivore: Contopus borealis air sallier Greater Pewee Insectivore: Contopus pertinax air sallier Western Wood-Pewee Insectivore: Contopus sordidulus air sallier Eastern Wood-Pewee Insectivore: Contopus virens air sallier Yellow-bellied Flycatcher Insectivore: Empidonax flaviventris air sallier Acadian Flycatcher Insectivore: Empidonax virescens air sallier Alder Flycatcher Insectivore: Empidonax alnorum air sallier Willow Flycatcher Insectivore: Empidonax traillii air sallier Least Flycatcher Insectivore: Empidonax rninimus air sallier Hammond's Flycatcher Insectivore: Ernpidonax hammondii air sallier Dusky Flycatcher Insectivore: Empidonax oberholseri air sallier Gray Flycatcher Insectivore: Empidonax wrightii air sallier Western Flycatcher Insectivore: Empidonax difficilis air sallier Buff-breasted Flycatcher Insectivore: Empidonax fulvifrons air sallier Black Phoebe Frugivore: Insectivore: Sayornis phoebe lower-canopy gleaner air sallier Say's Phoebe Insectivore: Sayornis saya air sallier Vermilion Flycatcher Insectivore: Pryocephalus rubinus air sallier Dusky-capped Flycatcher Insectivore: Myiarchus tuberculifer air sallier Ash-throated Flycatcher Insectivore: Nlyiarchus cinerascens lower-canopy gleaner air sallier Great Crested Flycatcher Frugivore: Insectivore: Myiarchus crinitus lower-canopy gleaner air sallier Brown-crested Flycatcher Insectivore: Myiarchus tyrannulus air sallier Great Kiskadee Insectivore: Pitangus sulphuratus air sallier Piscivore: freshwater plunger Sulphur-bellied Flycatcher Insectivore: Myiodynastes luteiventris air salIier 526 R.M. DeGraaf and others

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Tropical Kingbird Insectivore: Tyrannus melancholicus air sallier Couch's Kingbird Insectivore: Tyrannus couchii air sallier Cassin's Kingbird Frugivore: Insectivore: Tyrannus vociferans lower-canopy gleaner air saIlier Thick-billed Kingbird Insectivore: Tyrannus crassirostris air sallier Western Kingbird Insectivore: Tyrannus verticalis air saltier Eastern Kingbird Insectivore: Tyrannus tyrannus air sallier Gray Kingbird Insectivore: Tyrannus dominicensis air sallier Frugivore: lower-canopy gleaner Scissor-tailed Flycatcher Insectivore: Tyrannus forficatus air sallier Rose-throated Becard Insectivore: Pachyramphus aglaiae air sallyer Frugivore: lower-canopy gleaner Eurasian Skylark Omnivore: Granivore: Alauda arvensis ground forager ground gleaner Horned Lark Omnivore: Granivore: Eremophila alpestris ground gleaner ground gleaner Purple Martin Insectivore Progne subis air screener Tree Swallow Frugivore: Insectivore: Tachycineta bicolor lower-canopy gleaner air Screener Violet-green Swallow Insectivore: Tachycineta thalassina air screener Northern Rough-winged Insectivore: Swallow air screener Stelgidopteryx serripennis Bank Swallow Insectivore: Riparia riparia air screener Cliff Swallow Insectivore: Hirundo pyrrhonota air screener Cave Swallow Insectivore: Hirundo fulva air screener Barn Swallow Insectivore: Hirundo rustica air screener Gray Jay Omnivore: canadensis upper-canopy forager Steller's Jay Omnivore: stelleri ground forager tower-canopy forager Blue Jay Omnivore: Cyanocitta cHstata ground forager upper-canopy forager Green Jay Omnivore: yncas ground forager Brown Jay Omnivorc: Cyanocorax morio ground forager Scrub Jay Omnivore: coerulescem ground forager Gray-breasted Jay Omnivore: Aphelocoma ultramarina ground forager Foraging Guilds of North American Birds 527'

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Pinyon Jay Omnivore: Gymnorhinus upper-canopy forager cyanocephalus ground forager Clark's Nutcracker Omnivore: Nuc§ columbiana upper-canopy forager ground forager Black-billed Magpie Insectivore: Omnivore: pica ground gleaner ground forager Yellow-billed Magpie Insectivore: Omnivore: Pica nuttallis ground gleaner ground forager American Crow Omnivore: brachyrhynchos ground forager Omnivore: Corvus caurinus coastal beach scavenger Fish Crow Omnivore: Corvus ossifragus shoreline scavenger Chihuahuan Raven Omnivore: Corvus cryptoleucus ground forager Common Raven Omnivore: Coruus coro.x ground scavenger Black-capped Chickadee Insectivore: Omnivore: Parus atricapillus lower-canopy gleaner lower-canopy forager Carolina Chickadee Insectivore: Omnivore: Parus carolinensis lower-canopy gleaner lower-canopy forager Mexican Chickadee Insectivore: Omnivore: Parus sclateri lower-canopy gleaner lower-canopy forager Mountain Chickadee Insectivore: Omnivore: Parus gambeli lower-canopy gleaner lower-canopy forager Siberian Tit Insectivore: Omnivore: Parus cinctus lower-canopy gleaner lower-canopy forager Boreal Chickadee Insectivore: Omnivore: Parus hudsonicus lower-canopy gleaner lower-canopy forager Chestnut-backed Chickadee Insectivore: Omnivore: Paras rufescens lower-canopy gleaner lower-canopy forager Bridled Titmouse Insectivore: Omnivore: Parus wollweberi lower-canopy gleaner lower-canopy forager Plain Titmouse Omnivore: Parus inornatus lower-canopy forager Tufted Titmouse Insectivore: Omnivore: Parus bicolor lower-canopy gleaner lower-canopy forager Verdin Insectivore: Omnivore: Auriparus flaviceps lower-canopy gleaner lower-canopy forager Bushtit Insectivore: Omnivore: Psaltriparus minimus lower-canopy gleaner lower-canopy forager Red-breasted Nuthatch Granivore: Insectivore: Sitta canadensis upper-canopy gleaner bark gleaner White-breasted Nuthatch Granivore: Insectivore: Sitta carotinensis upper-canopy gleaner bark gleaner Pygmy Nuthatch Insectivore: Sitta pygmaea bark gleaner Omnivore: upper-canopy forager Brown-headed Nuthatch Granivore: Insectivore: Sitta pusilla upper-canopy gleaner bark gleaner Brown Creeper Insectivore: Certhia americana bark gleaner Cactus Wren Omniv0re: Campylorhynchus lower-canopy forager brunneicapillus ground forager 528 R.M. DeGraaf and others

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Rock Wren Insectivore: Salpinctes obsoletus ground gleaner Canyon Wren Insectivore: Catherpes mexicanus ground gleaner Carolina Wren Insectwore: Thryothorus ludovicianus lower-canopy gleaner Bewiek's Wren Insectivore: Thryomanes bewickii ground gleaner House Wren Insectivore: Troglodytes aedon lower-canopy gleaner Winter Wren Insectivore: Troglodytes troglodytes ground gleaner Sedge Wren Insectivore: Cistothorus platensis ground gleaner Marsh Wren Insectivore: Cistothorus palustris marsh gleaner American Dipper Insectivore: Cinclus mexicanus riparian bottom gleaner Arctic Warbler Insectivore: Phylloscopus borealis upper-canopy gleaner Golden-crowned Kinglet Insectivore: Regulus satrapa lower-canopy gleaner Ruby-crowned Kinglet Insectivore: Regulus calendula lower-canopy gleaner Blue-gray Gnatcatcher Insectivore: Polioptila caerulea upper-canopy gleaner Black-tailed Gnatcatcher Insectivore: Polioptila melanura lower-canopy gleaner Black-capped Gnatcatcher Insectivore: Polioptila nigricens lower-canopy gleaner Bluethroat Insectivore: Luscinia svecica ground gleaner Northern Wheatear Insectivore: Oenanthe oenanthe ground gleaner Eastern Bluebird Insectivore: Omnivore: Sialia sialia ground gleaner grodnd forager lower-canopy forager Frugivore: lower-canopy gleaner Western Bluebird Insectivore: Omnivore: Sialia mexicana ground gleaner lower-canopy forager Mountain Bluebird Omnivore: Insectivore: Sialia currocoides lower-canopy forager ground gleaner Townsend's Solitaire Insectivore: myadestes townsendi air sallyer Omnivore: ground forager Veery Omnivore: fuscescens ground forager lower-canopy forager Gray-cheeked Thrush Omnivore: Catharus minimus ground forager lower-canopy forager Swainson's Thrush Omnivore: Catharus ustulatus ground forager lower-canopy forager Hermit Thrush Insectivore: Omnivore: Catharus guttatus ground gleaner ground forager lower-canopy forager Foraging Guilds of North American Birds 529

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Wood Thrush Omnivore: Hylocichla mustelina ground forager Field fare Insectivore: Turdus pilaris ground gleaner American Robin Vermivore: Omnivore: Omnivore: Turdus migratorius ground gleaner ground forager lower-canopy forager Varied Thrush Insectivore: Omnivore: Ixoreus naevius ground gleaner ground forager Wrentit Frugivore: Insectivore: Chamaea fasciata lower-canopy gleaner bark gleaner Gray Catbird Omnivore: Dumetella carolinensis ground forager lower-canopy forager Omnivore: Frugivore: Mimus polyglottos ground forager lower-canopy gleaner Sage Thrasher Insectivore: Omnivore: Oreoscoptes montanus ground gleaner ground forager lower-canopy forager Brown Thrasher Omnivore: Toxostoma rufum ground forager lower-canopy forager Long-billed Thrasher Omnivore: Toxostoma longirostre ground forager lower-canopy forager Bendire's Thrasher Omnivore: Toxostoma bendirei ground forager lower-canopy forager Curve-billed Thrasher Omnivore: Frugivore: Toxostoma curvirostre ground forager lower-canopy gleaner California Thrasher Omnivore: Toxostoma redivivum ground forager Crissal Thrasher Omnivore: Toxostoma dorsale ground forager lower-canopy forager Le Conte's Thrasher Insectivore: Toxostorna lecontei ground gleaner Yellow Wagtail Insectivore: MotacilIa flava ground gleaner White Wagtail Insectivore: Motacilla alba ground gleaner Red-throated Pipit Insectivore: Anthus cervinus ground gleaner Water Pipit Insectivore: Omnivore: An thus spinolet~a ground gleaner ground forager Sprague's Pipit Insectivore: Omnivore: Anthus spragueii ground gleaner .ground forager Bohemian Waxwing Insectivore: Frugivore: Frugivore: Bombycilla air sallyer lower-canopy gleaner upper-canopy gleaner Cedar Waxwing Insectivore: Bombycilla cedrorum air saUyer Frugivore: upper-canopy gleaner Phainopepla Frugivore: Phainopepla nitens lower-canopy gleaner Northern Shrike Carnivore: Lanius excubitor ground hawker Loggerhead Shrike Carnivore: Lanius ludovicianus ground hawker 530 R. M DeGraaf and others

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round European Starling Omnivore: Sturnus vulgaris ground forager Crested Myna Omnivore: Acridotheres cristatellus ground forager White-eyed Vireo Insectivore: Omnivore: Vireo griseus lower-canopy gleaner lower-canopy forager Bell's Vireo Insectivore: Vireo bellii lower-canopy gleaner Black-capped Vireo Insectivore: Vireo atricapillus lower-canopy gleaner Gray Vireo Insectivore: Vireo vicinior lower-canopy gleaner Solitary Vireo Insectxvore: Omnivore: Vireo solitarius lower-canopy gleaner tower-canopy forager Yellow-throated Vireo Insectivore: Vireo flavifrons upper-canopy gleaner Hutton's Vireo Insectivore: Vireo huttoni lower-canopy gleaner Warbling Vireo Insectivore: Vireo gilvus upper-canopy gleaner Philadelphia Vireo Insectivore: Vireo philadelphicus upper-canopy gleaner Red-eyed Vireo Insectivore: Vireo olivaceus upper-canopy gleaner Black-whiskered Vireo Omnivore: Vireo altilopuus upper-canopy forager Bachman's Warble~ Insectivore: germivora bachmanii upper-canopy gleaner Blue-winged Warbler Insectxvore: Vermivora pinus lower-canopy gleaner Golden-winged Warbler Insectivore: Vermivora chrysoptera upper-canopy gleaner Tennessee Warbler Insectivore: Verrnivora peregrina upper-canopy gleaner Orange-crowned Warbler Insectivore: Omnivore: Vermivora celata lower-canopy gleaner lower-canopy forager Nashville Warbler Insectivore: Vermivora ruficapilla lower-canopy gleaner Virginia's Warbler Insectivore: Vermivora virginiae lower-canopy gleaner Colima Warbler Insectivore: Vermivora crissalis lower-canopy gleaner Lucy's Warbler Insectivore: Verraivora luciae lower-canopy gleaner Northern Parula Insectivore: Parula americana upper-canopy gleaner Tropical Parula Insectivore: Parula pitiayumi upper-canopy gleaner Yellow Warbler Insectivore: Dendroiea petechia lower-canopy gleaner Chestnut-sided Warbler Insectivore: Dendroica pensylvanica lower-canopy gleaner Magnolia Warbler Insectivore: Omnivore: Dendroica magnolia lower-canopy gleaner tower-canopy forager Cape May Warbler Insectivore: Omnivore: Dendroica tigrina upper-canopy gleaner upper-canopy forager Black-throated Blue Warbler Insectivore: Omnivore: Dendroica caerulescens lower-canopy gleaner lower-canopy forager Foraging Guilds of North American Birds 531

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Yellow-rumped Warbler Insectivore: Omnivore: Dendroica coronata lower-canopy gleaner lower-canopy forager Black-throated Gray Warbler Insectivore: Dendroica nigrescens lower-canopy gleaner Townsend's Warbler Insectivore: Dendroica townsendi upper-canopy gleaner Hermit Warbler Omnivore: Dendroica occidentalis upper-canopy forager Black-throated Green Warbler Insectivore: Dendroica virens upper-canopy gleaner Golden-cheeked Warbler Insectivore: Dendroica chrysoparia lower-canopy gleaner Blackburnian Warbler Insectivore: Dendroica fusca upper-canopy gleaner Yellow-throated Warbler Insectivore: Dendroica dominica upper-canopy gleaner bark gleaner Grace's Warbler Insectivore: Dendroica graciae upper-canopy gleaner Pine Warbler Insectivore: Omnivore: Dendroica pinus bark gleaner lower-canopy forager ground forager Kirtland's Warbler Insectivore: Dendroica kirtlandii lower-canopy gleaner Prairie Warbler Insectivore: Dendroica discolor lower-canopy gleaner Palm Warbler Insectivore: Omnivore: Dendroica palmarum ground gleaner ground forager Bay-breasted Warbler Insectivore: Dendroica castanea upper-canopy gleaner Blackpotl Warbler Insectivore: Dendroica striata upper-canopy gleaner Cerulean Warbler Insectivore: Dendroica cerulea upper-canopy gleaner Black-and-white Warbler Insectivore: Mniotilta varia bark gleaner American Redstart Insectivore: Setophaga ruticiila lower-canopy gleaner air sallier Prothonotary Warbler Insectivore: Protonotaria citrea lower-canopy gleaner bark gleaner Worm-eating Warbler Insectivore: Helmitheros verrnivorus ground gleaner Swainson's Warbler Insectivore: Limnothlypis swalnsonii lower-canopy gleaner Ovenbird Molluscovore: Omnivore: Seiurus aurocapiUus ground gleaner ground forager Insectivore: ground gleaner Northern Waterthrush Insectivore: Seiurus noveboracensis freshwater shoreline gleaner Louisiana Waterthrush Insectivore: Seiurus rnotacilla freshwater shoreline gleaner Kentucky Warbler Insectivore: Oporornisformosus ground gleaner Connecticut Warbler Insectivore: Oporornis agilis ground gleaner 532 R.M. DeGraaf and others

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round

~ Mourning Warbler Insectivore: Oporornis philadelphia ground gleaner MacGillivray's Warbler Insectivore: Oporornis totmiei lower-canopy gleaner Common Yellowthroat Insectivore: Geothlypis trichas lower-canopy gleaner Gray-crowned Yellowthroat Insectivore: Geothlypis poliocephala ground gleaner Hooded Warbler Insectivore: Wilsonia citrina lower-canopy gleaner air sallier Wilson's Warbler Insectivore: Wilsonia pusilla lower-canopy gleaner air sallier Canada Warbler Insectivore: Wilsonia canadensis lower-canopy gleaner Red-faced Warbler Insectivore: Cardellina rubrifrons upper-canopy gleaner Painted Redstart Insectivore: Myioborus pictus air- sallier ground gleaner bark gleaner Yellow-breasted Chat Omnivore: Icteria virens lower-canopy forager Olive Warbler Insectivore: Peucedramus taeniatus lower-canopy gleaner Hepatic Tanager Insectivore: Piranga tiara upper-canopy gleaner Summer Tanager Insectivore: Piranga rubra upper-canopy gleaner Scarlet Tanager Insectivore: Piranga olivacea upper-canopy gleaner Western Tanager Omnivore: Piranga ludoviciana upper-canopy forager Insectivore: air sallier Omnivore: Cardinalis cardinalis ground forager Pyrrhuloxia Omnivore: Cardinalis sinuatus ground forager Rose-breasted Grosbeak Omnivore: Pheucticus ludovicianus upper-canopy forager Black-headed Grosbeak Omnivore: Pheucticus melanocephalus upper-canopy forager Blue Grosbeak Omnivore: Guiraca caerulea ground forager Lazuli Bunting Omnivore: Passerina amoena lower-canopy forager Indigo Bunting Omnivore: Passerina cyanea lower-canopy forager Varied Bunting Omnivore: Passerina versieolor lower-canopy forager Painted Bunting Omnivore: Passerina ciris lower-canopy forager Dickcissel Omnivore: Granivore: Spiza americana ground forager ground gleaner Olive Sparrow Omnivore: Arremonops rufivirgatus ground forager Foraging Guilds of North American Birds 533

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Green-tailed Towhee Omnivore: Pipilo chlorurus ground forager Rufous-sided Towhee Omnivore: Pipilo erythrophthalmus ground forager Brown Towhee Omnivore: Pipilo fuscus ground forager Abert's Towhee Omnivore: Pipilo aberti ground forager White-collared Seedeater Granivore: Sporophila torqueola ground gleaner Bachman's Sparrow Omnivore: Aimophila aestivalis ground forager Botteri's Sparrow Omnivore: Aimophila botterii ground forager Cassin's Sparrow Insectivore: Granivore: Aimophila cassinii ground gleaner ground gleaner Rufous-wingedSparrow Omnivore: Granivore: Aimophila carpalis lower-canopyforager ground gleaner Insectivore: air sallyer Rufous-crownedSparrow Omnivore: Granivore: Aimophila ruficeps ground forager ground gleaner Chipping Sparrow Omnivore: Granivore: Spizella passerina ground forager ground gleaner Clay-coloredSparrow Omnivore: Spizella pallida lower-canopyforager ground forager Brewer's Sparrow Insectivore: Omnivore: Spizella breweri ground gleaner lower-canopyforager ground forager Field Sparrow Omnivore: Granivore: Spizella pusilla ground forager ground gleaner Black-chinnedSparrow Omnivore: Granivore: Spizella atrogularis ground forager ground gleaner Vesper Sparrow Omnivore: Granivore: Pooecetes gramineus ground forager ground gleaner Lark Sparrow Omnivore: Granivore: Chondestes grammacus ground forager ground gleaner Black-throated Sparrow Insectivore: Granivore: Amphispiza bilineata lower-canopygleaner ground gleaner ground forager Five-striped Sparrow Omnivore: Granivore: Amphispiza quinquestriata ground forager ground gleaner Lark Bunting Omnivore: Calamospiza melanocorys ground forager Savannah Sparrow Omnivore: Granivore: Passerculus sandwichensis grou_nd forager ground gleaner Baird's Sparrow Insectivore: Granivore: Ammodramus bairdii ground gleaner ground gleaner Grasshopper Sparrow Omnivore: Ammodramus savannarum ground forager Henslow's Sparrow Omnivore: Ammodramus henslowii ground forager Le Conte's Sparrow Omnivore: Ammodramus leconteii ground forager Sharp-tailed Sparrow Omnivore: Ammodramus caudacutus ground forager 534 R.M. DeGraaf and others

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Seaside Sparrow Crustaceovore: Ammodramus maritimus salt-marsh gleaner MolIuscovore: salt-marsh gleaner Insectivore: salt-marsh gleaner Fox Sparrow Omnivore: Granivore Passserella iliaca ground forager ground gleaner Song Sparrow Omnivore: Granivore: Melospiza melodia lower-canopy forager ground gleaner ground forager Lincoln's Sparrow Omnwore: Granivore: Melospiza lincolnii ground forager ground gleaner Swamp Sparrow Omnivore: Granivore: Melospiza georgiana ground forager - ground gleaner White-throated Sparrow Omnivore: Granivore: Zonotrichia albicollis ground forager ground gleaner Golden-crowned Sparrow Omnivore: Granivore: Zonotrichia atricapilla ground forager ground gleaner White-crowned Sparrow Omnivore: Gramvore: Zonotrichia leucophrys ground forager ground gleaner Harris' Sparrow Omnivore: Granlvore: Zonotrichia querula ground forager ground gleaner Dark-eyed Junco Omnivore: Granivore: Junco hyernalis ground forager ground gleaner Yellow-eyedJunco Omnivore: Junco phaeonotus ground forager McCown's Longspur Omnivore: Granivore: Calcarius mccownii ground forager ground gleaner Lapland Longspur Omnivore: Granivore: Calcarius lapponicus ground forager ground gleaner Smith's Longspur Omnivore: Calcarius pictus ground forager Chestnut-collared Longspur Omnivore: Granivore: Calcarius ornatus ground forager ground gleaner Snow Bunting Omnivore: Granivore: Plectrophenax nival# ground forager ground gleaner McKay's Bunting Omnivore: Granivore: Plectrophenax hyperboreus ground forager ground gleaner Bobolink Omnivore: Dolichonyx oryzivorus ground forager Red-winged Blackbird Omnivore: Granivore: Agelaius phoeniceus ground forager ground gleaner Tricolored Blackbird Omnivore: Agelaius tricolar ground forager Eastern Insectivore: Omnivore: Sturnella rnagna ground gleaner ground forager Western Meadowlark Insectivore: Omnivore: Sturnella neglecta ground gleaner ground forager Yellow-headed Blackbird Omnivore: Xanthocephalus ground forager xanthocephalus Rusty Blackbird Insectivore: Omnivore: Euphagus carolinus ground gleaner ground forager Brewer's Blackbird Omnivore: Euphagus cyanocephalus ground forager Great-tailed Grackle Omnivore: Quiscalus mexicanus ground forager Foraging Guilds of North American Birds 535

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Boat-tailed Grackle Omnivore: Quiscalus major ground forager Omnivore: Quiscalus quiscula ground forager Bronzed Cowbird Omnivore: Granivore: Molothrus aeneus ground forager ground gleaner Brown-headed Cowbird Omnivore: Granivore: Molothrus ater ground forager ground gleaner Orchard Oriole Insectivore: Icterus spurius upper-canopy gleaner Hooded Oriole Omnivore: Icterus cucullatus lower-canopy forager Altamira Oriole Omnivore: Icterus gularis upper-canopy forager Audubon's Oriole Omnivore: Icterus graduacauda lower-canopy forager Northern Oriole Omnivore: Icterus galbula upper-canopy forager Scott's Oriole Omnivore: Icterus parisorum lower-canopy forager Rosy Finch Omnivore: Granivore: Leucosticte arctoa ground forager ground gleaner Insectivore: air sallyer Pine Grosbeak Omnivore: Granivore: Pinicola enucleator upper-canopy forager upper-canopy gleaner ground gleaner Purple Finch Granivore: Granivore: Carpodacus purpureus upper-canopy gleaner ground gleaner Frugivore: upper-canopy gleaner Cassin's Finch Omnivore: Granivore: Carpodacus cassinii ground forager upper-canopy gleaner ground gleaner House Finch Frugivore: Granivore: Carpodacus mexicanus ground gleaner ground gleaner Red Crossbill Omnivore: Granivore; Loxia curvirostra upper-canopy forager upper-canopy gleaner White-winged Crossbill Omnivore: Granivore: Loxia leucoptera upper-canopy forager upper-canopy gleaner Common Redpoll Omnivore: Granivore: Carduelis flammea ground forager ground gleaner lower-canopy forager Hoary Redpolt Omnivore: Granivore: Carduelis hornemanni ground forager ground gleaner lower-canopy forager Pine Siskin Omnivore: Granivore: Carduelis pinus upper-canopy forager upper-canopy gleaner ground forager ground gleaner Lesser Goldfinch Granivore: Carduelis psaltria ground gleaner lower-canopy gleaner Lawrence's Goldfinch Granivore: Carduelis laurrencei ground gleaner lower-canopy gleaner American Goldfinch Omnivore: Granivore: Carduelis tristis lower-canopy forager lower-canopy gleaner ground forager ground gleaner 536 R.M. OeGraaf and others

Appendix I. Continued. Species name (Sci. name AOU list) Breeding period Nonbreeding period Year round Evening Grosbeak Omnivore: Granivore: Coccothraugtes vespertinus upper-canopy forager upper-canopy gleaner ground gleaner House Sparrow Granivore: Passer domesticus ground gleaner Eurasian Tree Sparrow Granivore: Passer montanus ground gleaner

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