History of Grassland Birds in Eastern North America Robert A

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9-1999 History of grassland birds in eastern North America Robert A. Askins Connecticut College, [email protected]

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Recommended Citation Kurosawa, R. and R. A. Askins. 1999. Differences in bird communities on the forest edge and in the forest interior: Are there forest- interior specialists in Japan? Journal of the Yamashina Institute for Ornithology 31:63-79.

This Article is brought to you for free and open access by the Biology Department at Digital Commons @ Connecticut College. It has been accepted for inclusion in Biology Faculty Publications by an authorized administrator of Digital Commons @ Connecticut College. For more information, please contact [email protected]. The views expressed in this paper are solely those of the author. Studies in Avian Biology No. 19:60-71, 1999.

HISTORY OF GRASSLAND BIRDS IN EASTERN NORTH AMERICA

ROBERT A. ASKINS

AhSWUCt. Until recently the severe decline in the populations of many species of grassland birds in eastern North America has aroused relatively little concern or conservation action. This response appears to be rooted in the perception that grassland birds invaded the East Coast from western grasslands after European settlers cleared the forest. Detailed historical accounts and analysis of pollen deposits, however, show that open grasslands existed on the East Coast of North America at the time of European settlement. Extensive grasslands resulted from burning and agricultural clearing by Native Americans. Natural disturbances, such as wildfire and beaver (Castor canadensis) activity, produced grasslands even before Native Americans cleared the forest. The presence of specialized grassland birds in Pleistocene deposits and in the earliest ornithological collections from eastern North America, and the existence of distinctive eastern populations of the Greater Prairie-Chicken (5mpanuchus cupido), Henslows’ Sparrow (Ammodramushenslowii), and Savannah Sparrow (Passerculussand- wichensis),indicate that grassland birds are an ancient component of biological diversity on the heavily forested East Coast of North America.

LA HISTORIA DE LAS AVES DE PASTIZAL EN EL ESTE DE NORTEAMERICA Sinopsis. Hasta hate poco tiempo, la declinacidn severa de las poblaciones de muchas especies de aves de pastizal en el este de America de1 Norte ha causado poco inter& o accidn de conservation. Esta respuesta se basa en la perception de que las aves de pastizal invadieron la costa este de America de1 Norte de 10s pastizales occidentales despues de la tala de1 bosque por 10s colonos europeos. Sin embargo, relatos historicos detallados y analisis de depositos de polen indican que pastizales abiertos existian en la costa este de America de1 Norte en la Cpoca de la colonizacidn europea. Quemas y desbrozos agricolos de 10s indigenas norteamericanos se tradujeron en pastizales extensivos. Disturbios naturales, coma 10s hechos por incendios y castores (Castor canadensis), produjeron pastizales antes de que 10s indigenas talaran el bosque. La presencia de aves de pastizal especialistas en 10s depdsitos pleistocenos y en las colecciones ornitoldgicas m&s antiguas de1 este de America de1 Norte, y la existencia de poblaciones orientales distintivas de Tympanuchus cupido, el Gorrion de Henslow (Am- modramushenslowii), y el Gorridn Sabanero (Passerculus.sandwichensis), indican que las aves de pastizal forman una parte antigua de la diversidad biologica en la arbolada costa este de America de1 Norte. Key Words: beaver; bird populations; disturbance; eastern grasslands; grassland birds.

Many grassland bird species were common or routes scattered throughout the United States even abundant along the East Coast through and southern Canada where birds are counted most of the nineteenth century, but their num- each year (Peterjohn 1994). The results for all bers diminished noticeably between the late of the survey routes east of the Mississippi River nineteenth and mid-twentieth centuries. Griscom indicate that since 1966, when the surveys be- (1949) described the Upland Sandpiper (Bartru- gan, the abundance of 14 of the 19 species of mia longicauda), Bobolink (Dolichonyx oryzi- grassland and savanna birds in eastern North vorus), Eastern Meadowlark (Sturnella magna), America has declined significantly (Table 1). and Grasshopper Sparrow (Ammodramus savan- Some have shown rapid population changes. Be- narum) as formerly common but declining in tween 1966 and 1994, for example, Grasshopper Massachusetts. Forbush (1925:449) mourned the Sparrows decreased at a rate of 6% per year, virtual disappearance of the Upland Sandpiper whereas the annual rates of decline were 3% for from New England: “our childrens’ children Vesper Sparrows (Pooecetes gramineus), 9% for may never see an Upland Plover in the sky or Henslows’ Sparrows (Ammodramus henslowii), hear its rich notes on the summer air. Its cries and 3% for Eastern Meadowlarks. In contrast, are among the most pleasing and remarkable only 2 of 40 species of forest-dwelling migra- sounds of rural life.” tory birds-a group that has received consider- Although the decline of grassland birds was able attention from conservationists-decreased obvious to any careful observer, it is only in re- at a rate of more than 2% per year during ap- cent decades that we have been able to calculate proximately the same period (Askins 1993). the precise rate and extent of these population Another indication that grassland birds in the changes. The best evidence for this comes from eastern United States are in trouble comes from the Breeding Bird Survey, a system of roadside state lists of endangered and threatened species

60 HISTORY OF EASTERN GRASSLAND BIRDS-A&ins 61

TABLE 1. POPULATIONTRENDS OF GRASSLANDAND SAVANNA SPECIALISTSIN NORTH AMERICA EAST OF THE MISSISSIPPIRIVER BETWEEN1966 AND 1994

Grassland speciesb Northern Harrier (Circus cyaneus) +2.1 * 227 Ring-necked Pheasant (Phasianus colchicus) -2.2 _ 522 Northern Bobwhite (Colinus virginianus) -3.3 *** 880 Upland Sandpiper (Barrramia Zongicauda) il.2 201 Horned Lark (Eremophila alpestris) 0.0 _ 642 Dickcissel (S&a americana) -4.3 *** 269 Vesper Sparrow (Pooecetes gramineus) -3.3 *** 650 Lark Sparrow (Chondestes grammucus) -6.9 * 39 Savannah Sparrow (Passerculus sandwichensis) -1.7 *** 727 Grasshopper Sparrow (Ammodramus savannarum) -5.9 *** 730 Henslows’ Sparrow (A. henslowii) -9.3 *** 137 Bobolink (Dolichonyx oryzivnrus) -1.4 *** 745 Eastern Meadowlark (Sturnella magna) -3.4 *** 1,330 Western Meadowlark (S. neglecta) ~7.2 *** 174 Savanna speciesC Common Ground-Dove (Columbine passerina) -3.4 *** 127 Red-headed Woodpecker (Melanerpes etythrocephalus) -2.1 *** 667 Eastern Bluebird (Sialia sialis) +2.2 *** 1,185 Loggerhead Shrike (Lanius ludoviciunus) -4.3 *** 379 American Goldfinch (Carduelis tristis) -1.1 *** 1,349

Note: Data are from the Breeding Bird Survey datahaw, U.S. Fish and Wildlife Serwce (Sauer et al 1995). Habitat clars~fications are based on DeGraaf and Rudis 1986, DeGraaf et al. 1991, and Askms 1993. a * P < 0.05, *** P < 0.001. h Species dependent on open habitats dominated by grass and forhs, with 11ttIe woody vegetation. C Species found primarily in open grassland with scattered trees or shrub\

(Vickery 1992). Of the 40 species listed as en- many of these species,” which are now declin- dangered, threatened, or of special concern in ing. The implication is that this is a return to three or more northeastern states, 13 are grass- ecosystems more similar to those before Euro- land or savanna specialists and only 3 are forest pean settlement and therefore should not be a specialists. For example, Upland Sandpiper, cause for concern. According to Whitcomb, Northern Harrier (Circus cyaneus), Loggerhead these species could survive only with active Shrike (L.anius ludovicianus), and Grasshopper, management to preserve grassland “in a region Henslows,’ and Vesper sparrows are listed in all where [grassland] is inappropriate as an equilib- or most of the New England states (Vickery rium community” (Whitcomb 1987: 165). 1992). The populations of many of these species Many historians and botanists have depicted have declined in other parts of the eastern Unit- the landscape of the ancient East Coast of North ed States, both in heavily forested areas along America as carpeted with forest, a forest so con- the East Coast and in the more agricultural Mid- tinuous that a squirrel could travel from the At- west (Herkert 1991, Bollinger and Gavin 1992). lantic Ocean to the Mississippi River without What Mayfield (1988) called the “quiet de- touching the ground (Day 1953). Clearly forests cline” of grassland birds has attracted surpris- in eastern North America were extensive, and in ingly little attention or concern from most gov- some areas they were essentially unbroken (Sic- ernment wildlife agencies and conservation or- ganizations. This response is rooted in the wide- cama 1971, Lorimer 1977, Bormann and Likens ly held view that before Europeans cleared the 1979, Runkle 1990, Seischab and Orwig 1991). land, unbroken forest stretched from the Atlantic Since the early 19OOs, however, some botanists to the Great Plains, leading to the general im- have argued that the forest was not always con- pression that grassland species invaded the east- tinuous; in coastal areas and even in some inland em states from western savannas and prairies af- areas, it was interrupted by scrubland, barrens, ter the clearing of the forest for agriculture. For glades, and even, in places, prairielike grass- example, Whitcomb (1987) argued that this in- lands (Day 1953). If this is true, then grassland vasion of the eastern “neosavanna” created by birds would have had a place in the landscape agriculture has been a “failed experiment for before European settlement. 62 STUDIES IN AVIAN BIOLOGY NO. 19

WERE THERE GRASSLANDS ON THE area was developed. Blueberry barrens, which EAST COAST BEFORE EUROPEAN are open expanses covered with lowbush blue- SETTLEMENT? berry (Vaccinium ungustifolium) shrubs and grasses, still cover large areas in eastern Maine, When Conrad (1935) visited Long Islands’ where they are maintained by burning for blue- Hempstead Plains in the 1930s much of the area berry production. Some of the largest East Coast was little bluestem (Schizachyrium scoparius) populations of Upland Sandpipers, Vesper Spar- prairie, a yellow-green grassland dotted with the rows, and other species of grassland birds breed small, bright green hemispheres of wild indigo on these barrens (Vickery et al. 1994). (Baptista tirzcroria). In May the prairie was blue Many of these grasslands may have resulted with the blossoms of birdfoot violets (Viola pe- from the activities of Native Americans before data). As Conrad pointed out, this grassland on European settlement. Early explorers and colo- the New York coast was remarkably similar to nists frequently encountered open landscapes the tallgrass prairies of Iowa and Nebraska. created by firewood harvesting, agricultural Moreover, the Hempstead Plains had a rich com- clearing, and burning to enhance hunting. For munity of grassland birds: Upland Sandpipers, example, Giovanni da Verrazano described the Bobolinks, and Vesper and Grasshopper spar- area around Narragansett Bay (Rhode Island) in rows were all common there in the 1920s (Bull 1524 as open plains, without forests or trees, for 1974). many leagues inland (Day 1953). Samuel de European travelers described the Hempstead Champlain and John Smith reported extensive Plains as treeless in the 1600s (Harper 1911), so areas of cleared land along the New England this grassland was not a product of European coast before Europeans colonized the area agriculture. The Hempstead Plains were char- (Whitney 1994). Moreover, an early settler in acterized by thin soil resting on a porous foun- Salem, Massachusetts, described “open plains, dation of quartz and granite pebbles (Conrad in some places five hundred acres. . .not much 1935, Cain et al. 1937), features that, in com- troublesome for to cleere for the plough to goe bination with periodic fires, appeared to favor in” (Day 1953:331). These clearings were not the growth of grasses and herbs rather than trees restricted to coastal areas; accounts of early set- and shrubs. tlers indicate that river valleys had been cleared The Plains once covered more than 20,000 ha, by Native Americans for farming and hunting and for many years the area was used primarily (Patterson and Sassaman 1988). for grazing sheep and racing horses (Svenson Early assessments that Native American ag- 1936, Stalter and Lamont 1987). Large areas of riculture had relatively little effect on the land- grassland remained in the 1930s but after World scape were based on population estimates after War II these open areas were subdivided for European settlement, but population densities housing or plowed for truck farms. Today only were much higher before contact with Europe- a few acres of this prairie survive: an 8-ha parcel ans triggered massive epidemics that killed a belonging to Nassau County Community Col- large proportion of the people in most tribes lege and managed by The Nature Conservancy, (Crosby 1972, Cronon 1983, Denevan 1992, and a 19-ha parcel managed as a nature preserve Whitney 1994). As Kulikoff (1986:29) wrote re- by Nassau County (Antenen et al. 1994). The garding the Chesapeake Bay area, “though En- smaller preserve has been maintained with con- glish settlers did not find a wilderness, they did trolled burning. create one” ; extensive agricultural clearings re- Although the Hempstead Plains may have verted to forest as Native American populations been one of the largest and most distinctive declined. Pilgrims traveling through the area grasslands on the East Coast, it was not the only near Warren, Massachusetts, in 1621 “saw the one. Another grassland, the Montauk Downs, remains of so many once occupied villages and covered approximately 2,400 ha of eastern Long such extensive formerly cultivated fields that Island (Taylor 1923), and several large grass- they concluded thousands of people must have lands, called “glades,” characterized a plateau lived there before the plague” (Russell 1980: in the Allegheny Mountains of western Penn- 24). Maps, drawings, and written accounts of the sylvania (Whitney 1994). Also, in the 1600s a landscape around Native American settlements savanna where occasional large oaks (Quercus) in the southeastern United States before Euro- broke an expanse of tall, wiry grass (probably pean settlement provide evidence of extensive bluestem) stretched for 24 km along the Quin- clearings created by farming and of “parklands” nipiac River north of New Haven, Connecticut maintained by controlled burning (Hammett (Olmsted 1937). After decades of overgrazing, 1992). In New York and southern New England, this area became the almost desertlike North Ha- relatively high population densities combined ven Sand Plains, and subsequently most of the with slash-and-bum agriculture (Whitney 1994) HISTORY OF EASTERN GRASSLAND BIRDS-Askins 63

would have resulted in extensive areas of trolled burning. These support a diversity of cleared land in the form of both active and aban- breeding grassland birds (Vickery et al. 1994). doned fields. This would have produced a “mo- The extent of forest clearing by Native Amer- saic of forests and fields in varying stages of icans in the northeastern United States probably succession” (Patterson and Sassaman 1988: paled in comparison with the extensive agricul- 115). Another view is that Eastern tribes used tural fields created by the Moundbuilders, who large permanent agricultural fields from which lived along the Mississippi River and its tribu- tree stumps had been removed rather than tem- taries in much of what is now the southeastern porary fields cut out of the forest for slash-and- and midwestem United States. Moundbuilding bum agriculture (Doolittle 1992). This perma- cultures existed in the lower Mississippi River nent farmland would have had to be rested oc- Valley as early as 1500 B.C. The early Mound- casionally, however, producing “weed-covered” building cultures probably depended on a mix- fallow fields of the sort seen by Champlain near ture of hunting, gathering of nuts, fishing, and the site of Boston in 1605 (Doolittle 1992). Re- small-scale farming based on native plants such gardless of whether Native Americans used as sunflower (Helianthus) and marsh elder (Zva slash-and-bum or permanent-field agriculture, frutescens; Shaffer 1992). Later, during the Mis- their activities would have produced open hab- sissippian Period, which lasted from A.D. 700 itats (abandoned or fallow fields) that could have until the early 17OOs,large-scale agriculture sup- been used by grassland birds. ported a dense population living in closely There also is good evidence that the Native spaced villages. Corn (Zea mays) and beans (Le- Americans of the East Coast burned large areas guminosae) from Mexico replaced indigenous to create open woodlands and grassland for crops, and large areas were cleared for farming hunting. For example, Roger Williams wrote in (Shaffer 1992). The largest population center the 1640s that Native Americans in New En- was Cahokia, located on the Mississippi River gland “burnt up all the underwoods in the Coun- near its confluence with the Missouri River trey, once or twice a yeare and therefore as No- (Shaffer 1992). This center covered 800 ha, with ble men in England possessed great Parkes . . 160 ha enclosed in a wooden palisade. The site was dotted with as many as 120 earthen mounds, onely for their game” (Williams 1963:47). In the largest of which rose to 30 m and covered 1818, B. Trumbull reported that the Native more than 6 ha. The mounds supported wooden Americans of Connecticut “so often burned the buildings, and the area below the mounds was country, to take deer and other wild game, that densely packed with rectangular thatched-roof in many of the plain dry parts of it, there was houses where an estimated 15,000-38,000 peo- but little small timber. Where the lands were ple lived. Cahokia and the many villages and thus burned there grew bent grass, or as some towns around it were supported by farming the called it, thatch, two, three and four feet high” American Bottom, a 324~km2 strip of rich allu- (Olmsted 1937:266). Native Americans in New vial soil in the floodplain along the eastern bank York not only burned the woods each autumn to of the Mississippi River. In A.D. 1000 there were create a more open understory but also burned 50 villages and 8 other large or medium-sized plains and meadows to improve hunting (Whit- centers within 40 km of Cahokia. ney 1994). There are no historical accounts of Cahokia Although fires were probably infrequent in because it was abandoned after A.D. 1200 (Shaf- most forests that were remote from Native fer 1992). Early Spanish visitors visited similar American settlements (Russell 1983), fire and sites that were still occupied in the 15OOs,how- other disturbances near settlements provided ex- ever. A chronicler of Hemando de Sotos’ expe- tensive habitat for early successional species, in- dition (1539-1542) described a Moundbuilder cluding potential habitat for grassland birds. An town along the Mississippi River as being “in analysis of charcoal deposits in the sediments of an open field, that for a quarter of a league over 11 lakes in New England demonstrated that be- was all inhabited; and at the distance of from fore European settlement, fires were frequent in half a league to a league off were many other densely populated coastal areas but infrequent in large towns, in which was a good quantity of inland and northern areas (Patterson and Sassa- maize, beans, walnuts [Juglans], and dried man 1988). Moreover, Winnes’ (1988) analysis Ameims [persimmons]” (Boume 1904: 149). of pollen and charcoal in lake sediments showed The Moundbuilding centers were abandoned that the area around Pineo Pond in eastern long before Europeans settled the southeastern Maine has been characterized by frequent, mod- United States or Mississippi River Valley. This erate fires and scrubby, fire-adapted vegetation culture may have been destroyed by Old World for at least 900 yr. Today this area is dominated diseases that swept inland from European out- by blueberry barrens that are maintained by con- posts on the Florida and Gulf Coasts (Crosby 64 STUDIES IN AVIAN BIOLOGY NO. 19

1986). Before the collapse of this agricultural their origin in the work of the beaver” (Whitney society, however, there were extensive areas of 1994:304). Coles and Orme (1983:99) argued open fields in many parts of the Southeast, es- that ancient forests in England must have been pecially the lower Mississippi River Valley. “moth-holed with clearings wherever beaver were present.” These “grassy meadows of relict GRASSLANDS BEFORE NATIVE pools” were also an important feature of the pre- AMERICAN AGRICULTURE settlement landscape of eastern North America. The patterns of Native American land use ob- Although beaver meadows are generally too served in the 1600s began to emerge about small to accommodate many species of special- 2,000 yr ago (Smith 1989, 1995). Many species ized grassland birds, some other species (e.g., of grassland birds may have colonized cultivated Eastern Meadowlark and Savannah Sparrow) areas after the initiation of Native American, occur in patches of grassland of similar size (5- rather than European, agriculture, and their cur- 10 ha; Herkert 1994, Vickery et al. 1994). rent decline represents a return to conditions be- Many regions of the East were subject to dis- fore humans began to substantially modify the turbances that created large openings in the for- vegetation of eastern North America. est (Runkle 1990). Grassland, savanna, and Undoubtedly, many apparently “natural” grassy scrub were probably created by large open grasslands of eastern North America were fires, particularly fires that burned following the product of human activities. For example, hurricanes or tornadoes that periodically leveled historical accounts and analysis of the pollen re- forests. These catastrophic disturbances were cord indicate that the extensive heathlands and probably most frequent in low-lying sandy areas sandplain grasslands on Nantucket Island, Mas- on the coastal plain. In many regions farther in- sachusetts, resulted from the clearing of oak for- land, fires, windstorms, and other disturbances est and grazing of sheep after Europeans settled were infrequent, and consequently the forest that island (Dunwiddie 1989). canopy was almost continuous, with few large Some of the open habitats on the East Coast openings (Lorimer 1977). For example, the may predate disturbance by Native Americans or northern hardwood forest of western New York Europeans, however. Smaller shrubby and and of the White Mountains of New Hampshire grassy openings in the eastern forest result from probably formed an almost unbroken canopy dam-building by beavers (Castor canadensis). (Bormann and Likens 1979, Seischab and Orwig After beavers exhaust the food supply around a 1991). Large grassy openings may have oc- pond, they move to another area. When the curred in some of the river valleys in the interior, abandoned pond drains, the pond bed often be- however. John Winthrop (Hosmer 1959:85) de- comes a “beaver meadow,” a patch of shrubby scribed how one of the first European expedi- vegetation or grassland. This meadow eventually tions to the White Mountains passed through is overgrown with young forest, and after lo- “many thousands of acres of rich meadow” as 30 yr beavers may recolonize the site and initiate it paddled birch-bark canoes up the Saco River another cycle (Remillard et al. 1987). in what is now Maine. Although beaver meadows are largely restricted to flood plains, their total area was probably A LONGER VIEW PLEISTOCENE STEPPE extensive before beavers were extirpated in most ANDSAVANNA parts of eastern North America (Naiman et al. When continental glaciers covered much of 1988). In Ontarios’ Algonquin Provincial Park, Canada and the northern United States, the re- where beavers are protected, there is a high den- gions immediately south of the glaciers were sity of beaver ponds and meadows (Coles and dominated by a spruce (Picea) parkland, a Orme 1983). After beavers became reestablished grassy savanna with scattered spruce trees at Quabbin Reservoir in Massachusetts in 1952, (Webb 1988). Samples of pollen from lake sed- the population grew rapidly until the density iments deposited 18,000-12,000 yr ago show reached 0.8 colony per kilometer of stream that this savanna stretched westward from the (Howard and Larson 1985). The impact of a Atlantic Coast to the Great Plains. Most species dense beaver population can be considerable. In of deciduous trees, and presumably the closed the Adirondack Mountains of New York, beaver forests where these species grow today, were re- dams created patches of disturbance that covered stricted to the extreme southeastern United an average area of 7 ha, with a maximum area States (Webb 1988). Thus, in eastern North of 12 ha (Remillard et al. 1987). Bela Hubbard, America there was a gradient from savanna in who surveyed land in Michigan before European the north to dense forest in the south. settlement, reported that one-fifth of the area Vegetation zones shifted and changed as the within 19 km of what is now Detroit was cov- glaciers retreated northward beginning about ered with “marshy tracts or prairies which had 12,000 yr ago. Spruce parkland largely disap- HISTORY OF EASTERN GRASSLAND BIRDS-As&s 6.5

peared, and a new gradient, from eastern forest [Pinicola enucleator]) lived alongside grassland to western prairie, gradually formed (Webb species such as Sharp-tailed Grouse, Greater 1988). Before this transition occurred, however, Prairie-Chicken (Zympanuchus’ cupido), Horned the spruce parkland was occupied by a diversity Lark (Eremophila alpestris), meadowlark (Stur- of large open-country mammals; caribou (Ran- nella sp.), pocket gopher (Geomys spp.), and gifer tarandus), mastodons (Mammut american- thirteen-lined ground squirrel (Parmalee and urn), and long-nosed peccaries (Mylohyus nasu- Klippel 1982). - A tus) are frequently found in fossil deposits from A key question is whether grassland birds the time of the spruce parkland (Km-ten and An- could have survived the northward spread of derson 1980). closed-canopy forest over eastern North Ameri- One of the best samples of spruce parkland ca after the continental glaciers melted. This is animals comes from a site called New Paris No. an issue not only for the current warm intergla- 4 in Pennsylvania (Guilday et al. 1964). Ap- cial period but also for previous interglacial pe- proximately 11,000 yr ago, a deep sinkhole act- riods. In all previous interglacial periods and at ed like a pitfall trap, collecting the skeletons of the beginning of the current postglacial period, more than 2,700 animals that fell into the crev- large browsers such as mastodons and giant ice and died. The mixture of small mammals and ground sloths (Megalonyx jeffersonii and other skeletal remains of Sharp-tailed Grouse (Tym- species) may have created and maintained open- panuchus phasianellus) at this site suggests that ings in the forest in much the same way as Af- there was extensive grassland habitat in this re- rican elephants (Zaxodonta africana) maintain gion. open savannas in East Africa today (Dublin et A better picture of the birdlife of the postgla- al. 1990). European ecologists have recognized cial period comes from another site, the caves that giant herbivores, particularly the extinct rel- of Natural Chimneys in Virginia, where skele- atives of elephants, probably opened the forest, tons were deposited at about the same time as creating glades, parklike woods, or even savan- at the New Paris No. 4 site. The skeletons at nas in areas that would otherwise be dominated Natural Chimneys were deposited in owl pellets, by dense forest (Andersson and Appelquist so both small mammals and birds were well rep- 1990, Puchkov 1992). Such openings would resented (Guilday 1962). Although remains of have supported a variety of animal and plant these animals may have accumulated over a long species that depend on grassy habitats. Through period while the vegetation was changing, they most of the past 1 million yr, as forests retreated provide a glimpse of the bird community of the and advanced in response to the shrinking and spruce parkland. The bones of Sharp-tailed growing of glacial ice sheets, woodland habitats Grouse, Northern Bobwhite (Colinus virgini- may have been modified and opened by giant anus), Upland Sandpiper, Red-headed Wood- herbivores. Only in the present interglacial did pecker (Melanerpes erythrocephalus), Black- mastodons, ground sloths, and other giants dis- billed Magpie (Pica pica), and Brown-headed appear from North America, perhaps as a result Cowbird (Molothrus a&r), along with other of the invasion of the continent by people who grassland vertebrates such as thirteen-lined had already developed efficient tools and strat- ground squirrel (Spermophilus tridecemlinea- egies for hunting large animals (Martin and tus), point to a landscape with large amounts of Klein 1984). Human activities such as burning open savanna or grassland. The remains of and agricultural clearing subsequently may have woodland species such as Red-bellied Wood- substituted for giant herbivores in creating a mo- pecker (Melanerpes carolinus), Eastern Wood- saic of forest and openings (Andersson and Ap- Pewee (Contopus virens), and Red-breasted Nut- pelquist 1990), permitting open-country species hatch (Sitta canadensis) in the same deposits to persist in eastern woodlands. suggest either that woodland and savanna were found in the area at the same time or that wood- THE ORIGIN OF EASTERN GRASSLAND land invaded and replaced the savanna while the BIRDS bones accumulated at Natural Chimneys. In ei- The common impression that many species of ther case, it is clear that grassland birds occurred grassland birds spread eastward from the prairies in eastern North America before the spruce of the Midwest to the newly cleared farmland of parkland receded and disappeared. the East Coast is substantiated by several well- A site farther south, at Duck River, Tennessee, documented examples of range expansion. For revealed that in the late Pleistocene typical co- example, the prairie subspecies of Homed Lark niferous forest species (Northern Hawk-Owl (Eremophila alpestris praticola) spread east- [Surnia ulula], Boreal Owl [Aegolius funereus], ward from Illinois and Wisconsin, reaching Northern Saw-whet Owl [A. acadicus], Gray Jay Michigan and Ontario in the 1870s New York [Perisoreus canadensis], and Pine Grosbeak in the 1880s New England by 1891, and Penn- 66 STUDIES IN AVIAN BIOLOGY NO. 19

Sylvania and Maryland by 1910 (Forbush 1927, dramus henslowii susurrans) has a breeding Thomas 1951, Hurley and Franks 1976). The range restricted to central New York and south- Dickcissel (Spizu americana) spread eastward em New England south to Virginia, eastern West from the tallgrass prairies in the early 1800s but Virginia, and North Carolina (Smith 1968). It is its range contracted after 1850, and it eventually darker than the western subspecies, with a deep- disappeared as a regular breeding bird along the er bill and more buff on the underparts and more East Coast (Hurley and Franks 1976). Western yellow in the wing (Smith 1968). Meadowlarks (Stumella neglecta) expanded east The “Ipswich” Sparrow (Passerculus sand- into Wisconsin and Michigan after 1900 (Lan- wichensis princeps), a subspecies of the Savan- yon 1956), and Lark Sparrows (Chondestes nah Sparrow, is also restricted to the East Coast grammacus) spread eastward from the prairies (Wheelwright and Rising 1993). This population into agricultural areas in the Ohio Valley, West is so distinctive that it was considered a separate Virginia, and western Maryland (Brooks 1938). species until 1973. After reclassifying it as a Although these eastward range expansions subspecies of Savannah Sparrow, the American were well documented, there is no similar evi- Ornithologists ’ Union (1957) recommended that dence for invasion of the East by the species that it be designated by its vernacular name in quotes are most abundant and widespread in eastern (“Ipswich” Sparrow). Vernacular names are grasslands. Upland Sandpipers, Grasshopper only officially recognized for particularly dis- Sparrows, Bobolinks, Eastern Meadowlarks, and tinctive subspecies (Stobo and McLaren 1975). other common grassland birds were reported by The “Ipswich” Sparrow was not described the earliest ornithologists who systematically until 1868, when C. J. Maynard collected one in documented the distribution of birds on the east- the coastal dunes at Ipswich, Massachusetts (El- em coast of North America. Alexander Wilsons’ liott 1968). Its breeding range on Sable Island, American Ornithology (originally published be- a 32-km-long island about 135 km off the coast tween 1808 and 1814; Brewer 1839) and John of Nova Scotia, was not discovered until 1884. James Audubons’ Ornithological Biography “Ipswich” Sparrows spend both summers and (Audubon 1831-1849) were published more winters in extremely open habitats. During the than 100 yr after most of the eastern seaboard breeding season they are virtually restricted to had been cleared, so it is possible that grassland low shrubby vegetation and stands of marram birds colonized the meadows and pastures cre- grass (Ammophila breviligulata) on Sable Island ated by Europeans long before their occurrence (Stobo and McLaren 1975). In winter they occur was initially documented. Some seventeenth- primarily in a narrow zone of dunes near Atlan- century European observers, such as John Jos- tic beaches from Nova Scotia to Florida, with selyn (Lindholt 1988) and William Wood the highest densities on relatively undeveloped (Vaughan 1977), described gamebirds and the barrier islands and sandy peninsulas between more conspicuous songbirds, but only a few spe- New Jersey and Virginia (Stobo and McLaren cies are recognizable because descriptions are 1971). sketchy and the names of British birds were fre- The “Ipswich” Sparrow is adapted to living quently used for North American species. Mark in the dunes and sandy scrub adjacent to the Catesbys’ Natural History of Carolina, Florida, ocean. It is paler gray than other Savannah Spar- and the Bahama Islands, completed in 1747, in- row subspecies, so it tends to be well camou- cludes descriptions and paintings of many spe- flaged in the light-colored dune and beach areas cies of eastern birds, including two species of where it lives (Stobo and McLaren 1975). Also, grassland songbirds, Eastern Meadowlark and it averages 9% larger than other eastern subspe- Bobolink (Feduccia 1985). It is not surprising, cies of Savannah Sparrow, which might be an however, that there are relatively few descrip- adaptation to feeding on the exceptionally large tions of grassland songbirds from this period. seeds of dune grasses such as marram grass and Many grassland birds are small, inconspicuous, sea oats (Uniola paniculata; Stobo and McLaren and dull colored, so they could have been over- 1975). Finally, unlike other Savannah Sparrow looked by early observers. subspecies, the “Ipswich” Sparrow has a short Significantly, the East Coast populations of tail, making it more similar to Grasshopper three species of grassland birds were distinctive Sparrow, Seaside Sparrow (Ammodramus mari- enough from western populations to be consid- timus), and other sparrows that live in open hab- ered separate subspecies. This suggests that itats with few tall shrubs or trees. These specific these populations have existed in isolation in the adaptations indicate that grassy habitats must East for many thousands of years, perhaps since have existed along the outer beaches of the East unbroken grasslands reached from the Great Coast for a long time. Plains to the Atlantic during the last glacial pe- The now-extinct Heath Hen (Tympanuchus riod. The eastern Henslows’ Sparrow (Ammo- cupido cupido) was the eastern subspecies of the HISTORY OF EASTERN GRASSLAND BIRDS-Ask& 67

Greater Prairie-Chicken. During the early years level of abundance characteristic of the preset- of European settlement, the Heath Hen was tlement landscape. Many of these species are common or even abundant in open grasslands now in danger of regional extinction, and they and scrublands on Long Island and around Bos- deserve the same attention from conservationists ton, and it ranged along the coast from southern as birds associated with eastern forests, marshes, Maine as far south as Virginia (Gross 1932). Be- and lakes. cause it was an important game species, it was described by many early settlers. In the 1600s CONSERVATION OF GRASSLAND BIRDS William Wood, Thomas Morton, and other ob- Many of the original grasslands, such as bea- servers wrote that the Heath Hen was common ver meadows and recently burned areas, were in eastern Massachusetts (Forbush 1927, Gross ephemeral. Other areas may have been disturbed 1932, Vaughan 1977). Heath Hens inhabited frequently enough to create stable grasslands; sandy scrub-oak plains, pine (Pinus) barrens, the Hempstead Plains in New York and some of blueberry barrens, and other open habitats (For- the barrens in eastern Maine are obvious can- bush 1927, Johnsgard 1983). In the nineteenth didates. Temporary grasslands are created much century they were common in the open grass- less frequently today because beavers are less land of the Hempstead Plains on Long Island abundant and fires are controlled, and most of (Bull 1974). the areas that may have been stable grasslands The abundance of Heath Hens at the time of have been developed for agriculture or housing. European settlement, and the recognition of the The blueberry barrens of eastern Maine are an Heath Hen and eastern populations of two other exception; these open habitats have been main- species of grassland birds as distinct subspecies, tained in a seminatural state by controlled bum- suggest that grassland birds inhabited the East ing to sustain commercial blueberry production Coast long before Europeans arrived or even be- (Vickery et al. 1994). fore Native Americans started clearing the land With the exception of Maines’ blueberry bar- for farming. This is consistent with the evidence rens and a few other areas remaining as semi- on grassland plants. Several plant species are re- natural open habitat for centuries, present-day stricted to eastern grasslands (Mehrhoff 1997; I? habitats used by grassland birds along the East Dunwiddie, pers. comm.), suggesting that they Coast are highly artificial. Populations of grass- evolved in isolation from the grasslands of the land species have diminished primarily because Great Plains. Bushy rockrose (Helianthemum much of the farmland in the Northeast and parts dumosum) is found from Massachusetts to Long of the Southeast has been abandoned and has Island; sandplain agalinis (Agalinis acuta) from reverted to forest, and because the remaining Massachusetts to Maryland; and sickle-leaved farmland is now managed more intensively for golden aster (Pityopsis [Chrysopsis] falcata) agricultural production (Hart 1968, Askins from Massachusetts to New Jersey (Gleason and 1993). For example, hayfields have become less Cronquist 1991). In addition, a subspecies of suitable as nesting habitat for Eastern Mead- northern blazing star (Liatris scariosa var. no- owlarks, Bobolinks, and some other grassland vae-angliae) is found only in eastern grasslands. species because they are mowed earlier in the Because of the paucity of historical records of summer, before the end of the nesting season, small birds in the 1600s and 1700s it is likely and because they are rotated more frequently that only carefully dated skeletal remains could (Bollinger and Gavin 1992). In southern New provide definitive evidence of the occurrence of England, most of the remaining populations of most species of grassland birds before European Grasshopper Sparrows and Upland Sandpipers settlement. There is strong evidence, however, are found in extensive mowed areas at airports that extensive grasslands and savannas occurred and military airfields (Veit and Petersen 1993, in eastern North America at the time of Euro- Bevier 1994, Melvin 1994). The farmland once pean settlement. We also know that some grass- used by these species has either disappeared or land species were found in the spruce parkland become unsuitable for nesting. of postglacial times, about 11,000 yr ago, and Regional populations of grassland birds can that distinctive eastern subspecies evolved in be maintained with proper management of arti- three grassland species. Therefore, it is reason- ficial grasslands such as fallow farmland and the able to conclude that many open-country species mowed areas near airport runways. The Conser- are native to the region, not recent invaders from vation Reserve Program (CRP), which pays the western prairies. During the eighteenth and farmers to take land out of production in order nineteenth centuries these species probably be- to manage it for conservation of soil and wildlife came much more abundant than they had been (Dunn et al. 1993), could potentially benefit before Europeans cleared the land, but subse- grassland birds in the East as it already has in quently they may have declined far below the some western prairie regions (Johnson and 68 STUDIES IN AVIAN BIOLOGY NO. 19

Schwartz 1993, Johnson and Igl 1995). How- only destroyed natural grasslands directly, but ever, most of the CRP land is concentrated in they also have interrupted or suppressed many the northcentral United States (Rodenhouse et al. of the natural processes of disturbance, such as 1995), and abandoned farmlands in the East fires and beaver activity, that once created the quickly become wooded, so a better approach early successional habitats that grassland species might be to compensate farmers who use less need. In the near term, artificial grasslands rep- intensive farming methods to create traditional resent our best hope for maintaining grassland hay meadows and other types of farmland that species. These species are an important, and once sustained grassland birds. This approach probably ancient, component of biological di- has been successful in preserving open-country versity along the East Coast of North America. species in the Netherlands (Beintema 1988). Relatively simple changes in airport manage- ACKNOWLEDGMENTS ment (e.g., removing woody vegetation and I have gained insights about the questions addressed changing mowing schedules to avoid the nesting in this paper in discussions with F? D. Vickery, W. A. season) have sustained or improved habitat for Niering, G. D. Dreyer, and R. M. DeGraaf. I? D. Vick- grassland birds at Westover Air Reserve Base in ery, J. R. Herkert, and an anonymous reviewer made helpful suggestions for improving the manuscript. My Massachusetts (Melvin 1994), Bradley Intema- research was supported by funds from the USDA For- tional Airport in Connecticut (Crossman 1989), est Service, Northeastern Forest Experiment Station. and Floyd Bennett Field, a former naval air base An earlier version of this paper was published in the on Long Island, New York (Lent and Litwin book Grasslands of Northeastern North America: 1989). Habitat management at Westover resulted Ecology and Conservation of Native and Agricultural in substantial increases in the abundance of Landscapes (I? D. Vickery and I? W. Dunwiddie [ed- Grasshopper Sparrows and Upland Sandpipers itors]. 1997. Massachusetts Audubon Society, Lincoln, between 1987 and 1994 (Melvin 1994). MA). This paper is also being revised for a-chapter in Even when grassland birds are absent from an a book by the author that will be published by Yale University Press. area, it should be possible to create habitat that will attract them. Probably because eastern LITERATURE CITED grassland birds have always depended on patch- AMERICAN ORNITHOLOGISTS ’ UNION. 1957. Check-list es of ephemeral habitat, they have a remarkable of North American birds. 5th ed. American Omi- ability to find and colonize remote sites, even thologists ’ Union, Washington, D.C. sites far from other bird populations of the same ANDERSSON,L., AND T. APPELQUIST.1990. The influ- species. When a field in Lincoln, Massachusetts, ence of the Pleistocene megafauna on the nemoral was managed to maintain tall grass for nesting and the boreonemoral ecosystem: a hypothesis with Bobolinks, in 1995 it attracted a breeding pair implications for nature conservation strategy. of Henslows’ Sparrow, a species that has almost Svensk Botanisk Tidskrift 84:355-368. disappeared from Massachusetts, with no known ANTENEN, S., M. JORDAN, K. MOTIVANS, J. B. WASHA, AND R. ZAREMBA. 1994. Hempstead Plains fire man- breeding records in the preceding 20 yr (Ells agement plan, Nassau County, Long Island, New 1995). Similarly, when abandoned strip mines in York. Report to Long Island Chapter of The Nature heavily forested areas of West Virginia were re- Conservancy, Cold Spring Harbor, NY. stored and seeded with grass, they were colo- ASKINS, R. A. 1993. Population trends in grassland, nized by Homed Larks, Eastern Meadowlarks, shrubland, and forest birds in eastern North Ameri- and Savannah, Vesper, and Grasshopper spar- ca. Pp. l-34 in D. M. Power (editor). Current or- rows (Whitmore and Hall 1978). These new nithology, vol. 11. Plenum Publishing, New York, grasslands were extremely isolated from other NY. grasslands supporting grassland birds, but they AUDUBON, J. J. 183 l-1849. Ornithological biography, or an account of the birds of the United States of still attracted breeding populations of several America. Vol. 1, J. Dodson, Philadelphia, PA; ~01s. species. 2-5, A. and C. Black, Edinburgh, Scotland. Expending scarce resources to maintain BEINTEMA, A. J. 1988. Conservation of grassland bird meadows, fallow fields, and airfields may seem communities in the Netherlands. Pp. 105-111 in I? unwise to many conservationists who are accus- D. Goriup (editor). 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