AMERICAN WOODCOCK FALL MIGRATION ECOLOGY IN THE … · The American woodcock, Scolopax minor, is a...
Transcript of AMERICAN WOODCOCK FALL MIGRATION ECOLOGY IN THE … · The American woodcock, Scolopax minor, is a...
FALL MIGRATION ECOLOGY OF AMERICAN WOODCOCK IN THE CENTRAL REGION OF THE UNITED STATES
FALL MIGRATION ECOLOGY OF AMERICAN WOODCOCK IN THE CENTRAL REGION OF THE UNITED STATES
A thesis submitted in partial fulfillment of the requirements for the degree of
Master of Science
By
NICHOLAS ANTHONY MYATT, B.S. Northland College, 2002
August 2004 University of Arkansas
This thesis is approved for Recommendation to the Graduate Council Thesis Director: ______________________ David G. Krementz Thesis Committee: ______________________ W. Fredrick Limp ______________________ Kimberly G. Smith
THESIS DUPLICATION RELEASE
I hereby authorize the University of Arkansas Libraries to duplicate this thesis when needed for research and/or scholarship.
Agreed___________________________
Refused__________________________
ACKNOWLEDGMENTS
I would like to thank everyone who made my research possible. Dr. David
Krementz, my major advisor, provided advice and support throughout my project, as
well as provided me with an excellent start to a career in wildlife management and
ecology. Dr. Fred Limp and Dr. Kimberly Smith provided valuable help while
serving on my thesis committee. This project was made possible through funding
from the U.S. Fish and Wildlife Service – Region IV and the Biological Resources
Division of the U.S. Geological Survey.
I especially thank Dr. David Andersen, Dr. Scott Lutz, Dr. John Bruggink,
Kevin Doherty, Jed Meunier, Eileen Oppelt and their field crews who captured and
radio-marked woodcock, and provided information on departure dates.
Housing was provided by Swan Lake National Wildlife Refuge (NWR), Two
Rivers NWR, Marais de Cygnes NWR, Mingo NWR, Lower Hatchie NWR, and St.
Catherine Creek NWR. I would also like to thank my pilot Jimmy Goad for his
incredible abilities to withstand endless hours of aerial radio telemetry and for his
flexibility to my constantly changing schedule.
This project would not have been as enjoyable without the help and support
from my fellow graduate students at the Arkansas Cooperative Fish and Wildlife
Research Unit. I would also like to thank Alicia Korpach who conducted a pilot
season before I began at the Co-op Unit. I would not have been able to complete the
GIS portions of this project without the technical support of John Wilson. Lastly, I
would like to thank my fiancé Jill Babski who has always provided an endless supply
of support and put up with my constant absence during many field seasons.
iv
TABLE OF CONTENTS
Page
CHAPTER 1: Introduction--------------------------------------------------------------------- 1
Literature Cited-------------------------------------------------------------------------- 9
Figures------------------------------------------------------------------------------------ 14
CHAPTER 2: Fall migration of American woodcock using Central --------------------- 16 Region band recovery and wing receipt data
Abstract----------------------------------------------------------------------------------- 17
Introduction ------------------------------------------------------------------------------ 17
Methods ---------------------------------------------------------------------------------- 18
Migration Progression -------------------------------------------------------------- 19
Migration Direction and Destination --------------------------------------------- 19
Results ------------------------------------------------------------------------------------ 20
Migration Progression -------------------------------------------------------------- 20
Migration Direction and Destination --------------------------------------------- 21
Discussion---------------------------------------------------------------------------- 21
Management Implications---------------------------------------------------------- 24
Literature Cited-------------------------------------------------------------------------- 25
Figures------------------------------------------------------------------------------------ 27
CHAPTER 3: Fall migration rates, routes, and habitat use of American --------------- 67 woodcock in the Central Region
v
vi
Abstract ------------------------------------------------------------------------------------ 68
Introduction ------------------------------------------------------------------------------- 68
Study Area--------------------------------------------------------------------------------- 72
Methods------------------------------------------------------------------------------------ 73
Capture --------------------------------------------------------------------------------- 73
Telemetry ------------------------------------------------------------------------------ 74
Habitat---------------------------------------------------------------------------------- 76
Results ------------------------------------------------------------------------------------- 78
Sample Size---------------------------------------------------------------------------- 78
Telemetry ------------------------------------------------------------------------------ 79
Migration Distance ------------------------------------------------------------------- 80
Stopover Duration -------------------------------------------------------------------- 80
Migration Habitat --------------------------------------------------------------------- 80
Winter Habitat------------------------------------------------------------------------- 82
Fall Migration Routes ---------------------------------------------------------------- 82
Potential Habitat Map ---------------------------------------------------------------- 83
Discussion --------------------------------------------------------------------------------- 83
Management Implications --------------------------------------------------------------- 90
Literature Cited --------------------------------------------------------------------------- 92
Figures ------------------------------------------------------------------------------------- 97
Tables -------------------------------------------------------------------------------------- 115
CHAPTER 1
INTRODUCTION
The American woodcock, Scolopax minor, is a popular, migratory game bird
throughout the eastern half of the United States, annually providing an estimated 3.4
million days of recreational hunting (U.S. Department of Interior 1988). Woodcock
population management is separated into two management units, the Eastern and
Central Regions (Coon et al. 1977) (Fig. 1). The U.S. Fish and Wildlife Service
monitors woodcock populations in each region using a series of singing ground
surveys that exploit the courtship display of male woodcock on breeding grounds.
Each randomly selected survey route is sampled annually during peak seasonal
courtship activities. Population indices are calculated using average number of
singing woodcock per route, weighted for land area (Tautin et al. 1983). Since annual
surveys began in 1968, population indices have annually declined 2.3% in the Eastern
Region and 1.8% in the Central Region (Kelley 2003). Breeding population indices
were the lowest in 1997 in the Central Region and 1995 in the Eastern Region (Kelley
2003).
Widespread habitat alteration and loss caused by human development and
forest succession are thought to be the primary causes of woodcock population
declines. In the northeastern United States, hardwood seedling-sapling stand area has
decreased by 26% over the last 20 years (Desseckar and Pursglove 2000).
Furthermore, the total area of aspen (Populus spp.) forest in Minnesota, Michigan and
1
Wisconsin has decreased by 21% since the mid-1960s (Desseckar and Pursglove
2000).
In the U. S., the largest concentration of bottomland hardwood forest, primary
woodcock wintering habitat, occurs in the Lower Mississippi Alluvial Valley (MAV).
Until the 1930s, the MAV has remained largely untouched and undeveloped due to
seasonal flooding. Extensive reduction and degradation has taken place since that
time, largely due to water control, which was followed by land clearing (Newling
1990). From the 1950s to 1970s, bottomland hardwood forests were lost at a rate
exceeding 120,000 ha per year (MacDonald et al. 1979). Of the original 10 million
hectares of bottomland hardwood forest in the MAV, 7.2 million hectares have been
cleared for agriculture (King and Keeland 1999).
Although hunting is not considered to be a major cause of woodcock
population declines, the U.S. Fish and Wildlife Service restricted hunting in the
Eastern Region in 1985, with additional restrictions in the Eastern and Central
Regions in 1998 (Woehr 1999). These restrictions included a reduction from a 65-
day hunting season and a 5-bird limit to 45 days in the Central Region and 30 days in
the Eastern Region, each with a 3-bird limit (Kelley 2003). Woodcock population
indices continue to decline despite reductions in season length and daily bag limit.
Little research has been conducted on the fall migration ecology of this
species. Although the timing of departure from the breeding grounds and arrival on
the wintering grounds has been documented, little is known about what happens
during the migration period. To effectively manage woodcock, managers need to
consider fall migration habitat use, timing, and routes.
2
The woodcock range is restricted to the eastern half of North America south
of the Taiga (Keppie and Whitting 1994). In the Central Region, woodcock primarily
breed in the states and provinces surrounding the Great Lakes, but woodcock have
been documented breeding in low densities as far south as Louisiana. Woodcock
primarily over-winter in the Gulf Coastal states, with the highest densities thought to
occur in southern Louisiana (Glasgow 1958). The northern extent of the wintering
range varies due to winter severity, with birds spreading throughout Louisiana during
mild, wet winters and concentrating in south central Louisiana during cool, dry
winters (Glasgow 1958). Straw et al. (1994) estimated wintering densities using 5
years of Christmas Bird Count (CBC) data collected within a 2-week period around
25 December. They concluded that woodcock winter farther north and west than
previously thought, with low densities of wintering birds reaching to southern
Missouri and the western extent reaching across the eastern half of Texas.
Woodcock habitat use has been studied extensively on the northern breeding
grounds and to a lesser extent on the wintering grounds. Understory structure is more
important than species composition; however, species composition can affect food
supply (e.g., earthworm density) and habitat structure (Straw et al. 1994). Woodcock
use a wide range of structural habitat types on the breeding and wintering grounds,
with very dense or very open habitats used least (Cade 1985). Dense shrub or tree
cover may inhibit flight and sparse cover may not provide protection from avian
predators. Vegetative cover may be more important than prey availability in
determining diurnal habitat use on the wintering grounds (Johnson and Causey 1982,
Boggus and Whiting 1982).
3
Diurnal woodcock cover on the breeding grounds primarily consists of early
successional, second growth hardwood forest, especially those containing aspen or
alder (Godfrey 1974, Gregg 1984, Straw et al. 1994) with moist soils. Conifer stands
are seldom used, except during drought conditions (Sepik et al. 1983).
Woodcock use a greater variety of habitats on the wintering grounds than on
the breeding grounds (Keppie and Whiting 1994). Bottomland hardwood forest are
historically considered to provide the best habitat, but pinelands and associated
drainages also offer wintering habitat throughout the southeastern U.S. (Reid and
Goodrum 1953, Glasgow 1958, Pursglove 1975, Kroll and Whiting 1977, Pace and
Wood 1979, Johnson and Causey 1982). Glasgow (1958) found wintering woodcock
in Louisiana typically in bottomland hardwood forest with scattered cane
(Arundinaria gigantean) thickets and blackberry (Rubus spp.). Wintering woodcock
in eastern Texas were found in 2-year-old pine clearcuts, as well as pole and mature
sized mixed pine/hardwood stands (Kroll and Whiting 1977).
Pine forests used vary from clear-cuts of loblolly (Pinus taeda) and shortleaf
(Pinus echinata) pine (Kroll and Whiting 1977, Boggus and Whiting 1982) to mature
stands of longleaf (Pinus palustris) pine (Johnson and Causey 1982). When moisture
is limited, mixed pine-hardwood stands and hardwood drainages provide more
suitable habitat than predominantly pine areas (Boggus and Whiting 1982).
Bourgeois (1977) hypothesized that woodcock were selecting diurnal cover based on
structural characteristics rather than species composition. He found that woodcock
hens primarily used early successional forests characterized by high densities of
4
seedling-saplings. Britt (1971) determined that canopy closure (structure) was more
important than overstory species composition.
Earthworms are the primary food of woodcock, comprising 79% and 75% of
their diet on the breeding and wintering grounds, respectively (Sperry 1940). The
distribution and density of earthworms in different soils is highly variable due to
factors such as soil moisture, pH, temperature, texture, type and land use history. On
the breeding grounds, Aporrectodea tuberculata and Dendrobaena octaedra prefer
soil moistures of 15-80%, high pH, and temperatures of 10-18ºC (Reynolds et al.
1977). If soil moisture becomes low, earthworms aestivate in a mucus cocoon or
move deeper into the soil (Edwards and Lofty 1977) below the probing depth of
woodcock. Earthworms are found in a wide variety of soils, ranging from gravelly
sand to clay, but the greatest densities occur in light loam soils (Guild 1948,
Nicholson and Owen 1982, Owen and Galbraith 1989).
Most woodcock leave the breeding grounds between October and December
(Godfrey 1974, Coon et al. 1976, Gregg 1984, Sepik and Derleth 1993, Keppie and
Whitting 1994). In Maine and Wisconsin, woodcock do not begin lipogenisis until
early-October so they were not physiologically ready to migrate until mid-October
(Owen and Krohn 1973, Gregg 1984). Woodcock initiate migration from
Pennsylvania between late-November and early-December (Coon et al. 1976),
October and November in Wisconsin (Gregg 1984), and early November in
Minnesota (Godfrey 1974).
Information on the timing of migration flights in mid-latitude states comes
largely from anecdotes and harvest data. In Missouri, mean woodcock harvest date,
5
determined using 1972-1979 wing-receipt data, was 6 November (Murphy 1983).
Peak migration in Kentucky and Tennessee begins after the first week of November
with the largest number of birds present in the second and third week of November
(Russell 1958, Roberts 1978). Woodcock begin arriving in Oklahoma in mid-
October with peak sightings occurring between 11 November and 10 December
(Barclay and Smith 1977). Woodcock occupy the wintering grounds in Louisiana
from November to February (Martin et al. 1969) with maximum numbers arriving in
mid-December (Glasgow 1958, Martin et al. 1969, Keppie and Whitting 1994).
Abundance and distribution of woodcock in Louisiana varies with winter severity
(Williams 1969) and precipitation.
Woodcock migration studies have used band recovery data, flush counts,
anecdotal evidence, and small samples of radio-marked woodcock to determine
migration timing (Glasgow 1958, Krohn et al. 1977, Roberts 1978, Sepik and Derleth
1993). A woodcock, radio-marked by Sepik and Derleth (1993) was shot in New
York on 9 November, two days after it was last located 680 km away in Maine. Coon
et al. (1976) initiated a study in Pennsylvania using radio telemetry to gather data on
migration timing. This is the only published study to date that radio-tracked
woodcock after fall migration began. The majority of their radio-marked woodcock
departed between 2.5 hours after sunset and midnight. All woodcock departures
during the study occurred during an 11-day period preceding a full moon (Coon et al
1976). They tracked two hatch-year females during portions of their migratory
flights. In 1973, one female traveled 56 km on her first night of migration. The
following evening she resumed migration and was followed for 56 km until weather
6
conditions became unsuitable for the tracking aircraft. In 1974, a female flew 53 km
on her first night of migration. The following evening she resumed migration and
was followed for nearly 148 km until the signal was lost (Coon et al. 1977). The
direction that both woodcock flew coincided with the direction of a river valley,
which led Coon et al. (1976) to suggest that woodcock might follow land features
during migration.
Commencement and progression of fall migration is influenced by weather
with the heaviest flights coinciding with cold fronts (Sepik and Derleth 1993). Gregg
(1984) found that severe cold snaps forced nearly all woodcock out of northern
Wisconsin by 1 November in some years, but mild weather in other years enabled
birds to remain into early winter. Godfrey (1974) reported that woodcock in
Minnesota departed between the retreat of a low-pressure system and the entrance of
a high-pressure center, when winds were most favorable for southerly flight.
Heaviest flights of woodcock have been reported at Cape May, New Jersey, after cold
nights with northwesterly winds (Sheldon 1967). Woodcock initiated migration in
Pennsylvania when high-pressure cells approached from the north and west or when
low-pressure cells retreated to the north and east (Coon et al. 1976). Krementz et al.
(1994) reported an association between woodcock spring migration departure dates
and both moon phase and the passage of weather fronts.
Glasgow (1958) investigated woodcock fall migration routes by plotting 175
band recoveries. Based on these band recoveries, he determined two possible fall
migration routes in the central U.S. (Fig. 2). He described birds migrating along a
western route originating in Minnesota, Wisconsin, Upper Peninsula of Michigan,
7
and western Ontario. These birds funneled into the Mississippi River Valley, which
they followed south to central Missouri. The woodcock then spread out across
southern Missouri and Arkansas before reaching Louisiana and east Texas. The
second route described is the central route, which is used by woodcock migration
from Lower Peninsula Michigan, Indiana, Ohio, and eastern Ontario. This route
passed through west Kentucky and west Tennessee, and then followed the eastern
edge of the lower Mississippi Alluvial Valley before reaching southern Mississippi
and Louisiana. Sheldon (1967) plotted 225 additional band recoveries and
hypothesized that birds from Minnesota and Wisconsin are migrating south along the
Mississippi River through the MAV (Fig. 2).
Migration stopovers may be crucial refueling sites for migrants traveling from
the breeding to the wintering grounds. Making a series of short flights is
energetically cheaper than one long flight (Piersma 1987). When there is an
abundance of suitable stopover locations, birds divide their migration into many short
flights, or "hops" (Alerstam and Lindstrom 1990). Hops are frequent, short flights
with many brief refueling periods (Piersma 1987). Habitat availability determines the
flight strategy used by birds. As the distance between suitable stopover sites
increases, migratory flights shift from "hops" to "jumps", where jumps are longer
flight with few but longer stopovers.
Stopover duration is the length of time an individual bird remains at a
stopover site during fall migration. Little information is known about the stopover
duration of woodcock. Other than the observations of two stopover durations by Coon
et al. (1976), there is only anecdotal evidence provided by woodcock hunters.
8
Hunters report that birds can be abundant in coverts one day and gone the next.
These reports have led biologist to believe that stopover duration is short and that
woodcock generally resume migration when weather conditions become favorable.
Accurate information on the distribution and amount of woodcock habitat
within the U.S. remains largely unknown (Cushwa et al. 1977, Duncan 2000).
Managers need better information on the distribution and changes in habitat at the
national level. Forest inventory data have been used to identify trends and
distribution of woodcock habitat (Woehr 1999). Using these data, direct comparisons
between years and states are not always possible due to differing data collection and
reporting methodologies. The advancement of Geographic Information System (GIS)
technologies has potential applications in mapping national woodcock habitat
availability (Duncan 2000). Woodcock select habitat based on structure and age
instead of species composition (Burgeois 1977, Straw et al. 1994), but spatial land
cover data are classified by cover type, generally disregarding structure or
successional stage. Although current data are insufficient to map present distribution
of suitable woodcock habitat in the U.S., those areas that provide no woodcock
habitat can be mapped to provide clues into regional habitat availability, as well as
identify large areas of no potential woodcock habitat.
In 2001 I initiated a project to better understand woodcock fall migration in
the Central Region. The objectives of the project were to use radio telemetry, band
return data, and wing receipt data to: 1) determine the timing and progression of fall
migration 2) document fall migration routes in the Central Region 3) determine
stopover duration of woodcock during fall migration and 4) investigate woodcock
9
habitat use during fall migration. As part of this project, I mapped the distribution of
potential woodcock habitat and identified priority areas for potential future woodcock
management in the Central Region.
LITERATURE CITED
Alerstam, T., and A. Lindstrom. 1990. Optimal bird migration: The relative importance of time, energy, and safety. Pages 331 - 351 in E. Gwinner, editor. Bird migration: physiology and ecophysiology. Springer-Verlag, Berlin, Germany.
Barclay, J. S., and R. W. Smith. 1977. The status and distribution of woodcock in
Oklahoma. Proceedings of the Woodcock Symposium 6:39-50. Boggus, T. G., and R. M. Whiting, Jr. 1982. Effects of habitat variables on foraging
of American woodcock wintering in East Texas. U.S. Fish and Wildlife Service Wildlife Research Report 14:148-153
Bourgeois, A. 1977. Quantitative analysis of American woodcock nest and brood habitat. Proceedings of the Woodcock Symposium 6:108-118. Britt, T. L. 1971. Studies of woodcock on the Louisiana wintering ground. Thesis,
Louisiana State University, Baton Rouge, Louisiana, USA. Cade, B. S. 1985. Habitat suitability index models: American woodcock (wintering).
Biological Report 82. U. S. Fish and Wildlife Service, Washington, D.C., USA.
Coon, R. A., P. D. Caldwell, and G. L. Storm. 1976. Some characteristics of fall migration of female woodcock. Journal of Wildlife Management 40:91-95. _____. 1977. Nesting, habitat, fall migration, and harvest characteristics of the
American woodcock in Pennsylvania. Thesis, Pennsylvania State University, College Junction, Pennsylvania, USA.
_____, T. J. Dwyer, and J. W. Artmann. 1977. Identification of potential harvest
units in the United States for the American woodcock. Proceedings of the Woodcock Symposium 6:147-153. Cushwa, C. T., J. E. Barnard, and R. B. Barnes. 1977. Trends in woodcock habitat in
the United States. Proceedings of the Woodcock Symposium 6:31-37.
10
Dessecker, D. R., and S. R. Pursglove, Jr. 2000. Current population status and likely future trends for American woodcock. Pages 3-8 in D. G. McAuley, J. G. Bruggink, and G. F. Sepik, editors. Proceedings of the Ninth American Woodcock Symposium. U.S. Geological Survey, Biological Resources Division Information and Technology Report USGS/BRD/ITR-2000-0009, Patuxent Wildlife Research Center, Laurel, Maryland, USA.
Duncan, P. S. 2000. American woodcock management, past, present, and future.
Pages 1-2 in D. G. McAuley, J. G. Bruggink, and G. F. Sepik, editors. Proceedings of the Ninth American Woodcock Symposium. U.S. Geological Survey, Biological Resources Division Information and Technology Report USGS/BRD/ITR-2000-0009, Patuxent Wildlife Research Center, Laurel, Maryland, USA.
Edwards, C. A., and J. R. Lofty. 1977. Biology of earthworms. John Wiley and Sons, New York, New York, USA.
Glasgow, L. L. 1958. Contributions to the knowledge of the ecology of the
American Woodcock, Philohela minor (Gmelin), on the wintering range in Louisiana. Dissertation, Texas A & M College, College Station, Texas, USA.
Godfrey, G. A. 1974. Behavior and ecology of American woodcock on the breeding
range in Minnesota. Dissertation, University of Minnesota, Minneapolis, Minnesota, USA.
Guild, W. F. M. L. 1948. The effects of soil type on the structure of earthworm populations. Annals of Applied Biology 35:181-192. Gregg, L. 1984. Population ecology of woodcock in Wisconsin. Wisconsin Department of Natural Resources. Technical Bulletin 144. Johnson, R. C., and M. K. Causey. 1982. Use of longleaf pine stands by woodcock
in southern Alabama following prescribed burning. U.S. Fish and Wildlife Service, Wildlife Research Report 14:120-125.
Kelley, J. R., Jr. 2003. American woodcock population status, 2003. U.S. Fish and
Wildlife Service, Laurel, Maryland, USA. Keppie, D. M. and R. M. Whiting, Jr. 1994. American woodcock (Scolopax minor).
The birds of North America, no. 100. The American Ornithologists’ Union, Washington, D.C., USA, and The Academy of Natural Sciences, Philadelphia, Pennsylvania, USA.
King, S. L., and B. D. Keeland. 1999. Evaluation of reforestation in the Mississippi
River Alluvial Valley. Restoration Ecology 7:348-359.
11
Krementz, D. G., J. T. Seginak, and G. W. Pendleton. 1994. Winter movements and spring migration of American woodcock along the Atlantic Coast. Wilson Bulletin 106:482-493.
Krohn, W. B., J. C. Rieffenberger, and F. Ferrigno. 1977. Fall migration of
woodcock at Cape May, New Jersey. Journal of Wildlife Management 41:104-111.
Kroll, J. C., and R. M. Whitting. 1977. Discriminate function analysis of woodcock
winter habitat in east Texas. Proceedings of the Woodcock Symposium 6:63-71.
MacDonald, P. O., W. E. Frayer, and J. K. Clauser. 1979. Documentation,
chronology, and future projections of bottomland hardwood habitat loss in the lower Mississippi Alluvial Plain. Volume 1, basic report. HRB-Singer, Inc., State College, Pennsylvania, USA.
Martin, F. W., S. O. Williams, J. D. Newsom, and L. L. Glasgow. 1969. Analysis of
records of Louisiana banded woodcock. Proceedings of the Southeastern Association of Game and Fish Commissioners 23:85-96.
Murphy, D. W. 1983. Ecology of American woodcock in central Missouri. Thesis,
University of Missouri, Columbia, Missouri, USA. Newling, C. J. 1990. Restoration of bottomland hardwood forests in the Lower
Mississippi Valley. Restoration and Management Notes 8:23-28. Nicholson, C. P., and R. B. Owen, Jr. 1982. Earthworm abundance in selected forest habitats in Maine. Megadrilogiea 41:78-80. Owen, R. B., and W. B. Krohn. 1973. Molt patterns and weight changes of the
American woodcock. Wilson Bulletin 85:31-41. _____, and W. J. Galbraith. 1989. Earthworm biomass in relation to forest types,
soil and land use: Implications for woodcock management. Wildlife Society Bulletin 17:130-136.
Pace, R. M., III, and G. W. Wood. 1979. Observations of woodcock wintering in
coastal South Carolina. Proceedings of the Southeastern Association of Game and Fish Commissions 33:72-80.
Piersma, T. 1987. Hop, skip, or jump? Constraints on migration of arctic waders by
feeding, fattening, and flight speed. Limosa 60:185-194. Pursglove, S. R. 1975. Observations on wintering woodcock in northeast Georgia.
12
Proceedings of the Southeastern Association of Game and Fish Commissions 29:630-639.
Reid, V., and P. Goodrum. 1953. Wintering woodcock populations in west central
Louisiana, 1952-53. U.S. Fish and Wildlife Service, Special Scientific Report Wildlife 24.
Reynolds, J. W., W. B. Krohn, and G. A. Jordan. 1977. Earthworm populations as
related to woodcock habitat usage in central Maine. Proceedings of the Woodcock Symposium 6:135-146.
Roberts, T. H. 1978. Migration, distribution, and breeding of American woodcock. Tennessee Wildlife Resources Agency, Technical Report 79.
Russell, D. M. 1958. Woodcock and Wilson’s snipe studies in Kentucky. Kentucky
Department of Fish and Wildlife Resources. Project Report W-31-R. Sepik, G. F., and E. L. Derleth. 1993. Premigratory dispersal and fall migration of American woodcock in Maine. Biological Report 16:36-40. ____, _____, and T. J. Dwyer. 1983. The effect of drought on a local woodcock
population. Transactions of Northeast Fish and Wildlife Conference 40:1-8. Sheldon, W. G. 1967. The book of the American woodcock. University of Massachusetts Press, Amherst, Massachusetts, USA. Sperry, C. C. 1940. Food habits of a group of shorebirds: Woodcock, snipe, knot,
and dowitcher. U.S. Biological Survey, Wildlife Resources Bulletin 1. Straw, J. A., D. G. Krementz, M. W. Olinde, and G. F. Sepik. 1994. American
woodcock. Pages 97-114 in T.C. Tacha and C.E. Braun, editors. Migratory shore and upland game bird management in North America. International Association of Fish and Wildlife Agencies, Washington, D.C., USA.
Tautin, J., P. H. Geissler, R. E. Munro, and R. S. Pospahala. 1983. Monitoring the
population status of American woodcock. Transactions of the North American Wildlife Conference 48:376-388.
U.S. Department of Interior. 1988. 1985 National survey of fishing, hunting and
wildlife associated recreation. U.S. Government Printing Office, Washington, D.C., USA.
Williams, S. O., III. 1969. Population dynamics of woodcock wintering in
Louisiana. Thesis, Louisiana State University, Baton Rouge, Louisiana, USA. Woehr, J. R. 1999. Declines in American woodcock populations: Probable cause
13
and management recommendations. The report of the Woodcock Task Force to the Migratory Shore and Upland Game Bird Subcommittee, Migratory Wildlife Committee, International Association of Fish and Wildlife Agencies.
14
Figure 1. Woodcock management regions determined by Coon et al. (1977) using band recovery data.
15
Figure 2. Possible American woodcock fall migration route proposed by Glasgow (1958) and Sheldon (1967) using band return data.
16
Chapter 2
FALL MIGRATION OF AMERICAN WOODCOCK USING CENTRAL REGION BAND RECOVERY AND WING RECEIPT DATA¹
_________________ ¹Myatt, N. A., and D. G. Krementz. To be submitted to The Journal of Wildlife Management.
17
Abstract: Band recovery and wing receipt data have potential to provide information
on fall migration ecology of American woodcock in the Central Region, yet these
extensive data sets have not been recently analyzed. I examined all direct recoveries
of woodcock banded in Michigan, Minnesota and Wisconsin, as well as wing receipt
data, to determine the progression of fall migration and the migration direction and
final destination of woodcock migrating from these states. Migration initiation was
not observed until late October and early November, with most migration occurring
during November. Wing receipt data showed a similar trend, with most change in
mean receipt latitude occurring from 1 November – 5 December. During November,
wing receipts were spread through the entire Central Region. Band recovery data
demonstrated that during 1-15 December, birds were spread throughout the southern
U.S. During 15-31 December, 92.3% (n=26) of band recoveries were on the
wintering grounds (south of 33º N latitude). Most banded woodcock from Michigan,
Minnesota, and Wisconsin wintered in Louisiana, but some Michigan banded
woodcock were recovered as far east as Georgia and South Carolina. Hunting
seasons on the breeding grounds start in late-September before migration initiation.
Managers need to consider the effects of harvesting locally produced birds on
woodcock populations.
INTRODUCTION
Little information has been published on the fall migration ecology of
American woodcock in the Central Region (Keppie and Whiting 1994), which
includes the woodcock’s range west of the Appalachian Mountains (Coon et al.
18
1977). The only available information on woodcock fall migration timing and routes
were based on anecdotal evidence, reports of migration initiation in small numbers of
radio-marked woodcock (Coon et al. 1976, Gregg 1984), and analysis of band return
data from the 1950s to the early-1980s (Glasgow 1958, Sheldon 1967, Krohn et al.
1977, Gregg 1984). Since then, many woodcock bands have been recovered and
computer technologies have advanced to aid in mapping and analysis of band
recovery data.
The U.S. Fish and Wildlife Service (USFWS) collects data on annual
woodcock reproductive rates using a parts collection survey. Survey participants
were asked to submit one wing from each woodcock harvested (Kelley 2003).
Murphy (1983) used woodcock wing receipt data to estimate peak fall migration of
woodcock in Missouri. While wing receipt densities are influenced by hunter
densities, hunting season dates, and participation in the program, these data can
provide valuable information on woodcock migration.
I analyzed Central Region woodcock banding and wing receipt data to: 1)
determine the timing and progression of fall migration, and 2) to determine the
direction and final destination of woodcock migrating from Minnesota, Wisconsin,
and Michigan.
METHODS
I obtained woodcock banding data from the Bird Banding Lab for 1929-2001.
Band returns from Central Region banded woodcock with known day, month, year,
and 10-minute block were used in my analysis. The USFWS also provided historic
19
wing receipt data from 1963-2002 (J.R. Kelley, Jr. US FWS, personal
communication). I used ESRI Geographic Information System (GIS) software to
query, analyze, and map band return and wing receipt densities.
Migration Progression
In my analysis of migration progression, I used all direct recoveries of birds
banded in the northern half of the U.S. portion of the Central Region and recovered
between 1 September and 31 December, 1929-2001. I queried these recoveries by 2-
week periods from 1 September-31 December. Banding and recovery locations were
mapped for each period with lines connecting the two locations to facilitate
interpretation. I determined mean band recovery latitude by computing the average
latitude of all band returns during each 2-week period.
In my analysis of migration progression, I used all woodcock wing receipts
from woodcock shot in the Central Region between 1 September and 31 December
1963-2002. I queried the data by 5-day periods from 1 September-31 December.
During each period, wing receipts for each county were tallied and receipt densities
mapped. I determined the mean wing receipt latitude by computing the average
latitude of all wing receipts during a 5-day period. The center point of each county
was used to determine the latitude of each wing receipt.
Migration Direction and Destination
I investigated the final destination of migrating woodcock using all direct
recoveries of woodcock banded in Minnesota, Wisconsin, and Michigan and
recovered in a different state from 1 September-31 December. I mapped these
locations and drew a line from the banding location to the recovery location. I used
20
an ArcView script to measure the distance and angle between banding and recovery
locations. This script created an Azmuthal map projection centered on the individual
banding location and then measured the distance and angle to the recovery location.
The Azmuthal projection was chosen because it most accurately estimated distances
and angular data from its center point (J. Wilson, Center for Advanced Spatial
Technology, personal communication).
I used program ORIANA for Windows (Provalis Research) to estimate mean
migration angle and associated circular statistics of the migration angles (angle
between banding and recovery location) of all band recoveries outside of the banding
state. Using these data, I created roses were to visually display migration angles for
each state.
RESULTS
Migration Progression
Mean woodcock band recovery latitude remained relatively constant
throughout the early fall, until a slight shift south was observed during 16-31 October
(Fig. 1). I observed the largest change in mean latitude between 1-15 November.
After this period, mean latitude gradually decreased during the final two periods (Fig.
1). Looking at the migration progression maps created from banding data (Fig. 2 - 9),
no migration was evident until 1-15 November when woodcock were being
recovered throughout the northern half of the Central Region (Fig. 6). During 16-30
November, woodcock were being recovered throughout the southern half of the
Central Region (Fig. 7).
21
Mean woodcock wing receipt latitude followed a pattern similar to band
recovery data. No substantial change in latitude was observed until 21-25 October
(Fig. 10). Mean latitude steadily decreased throughout November until 1-5 December
when mean latitude leveled off (Fig. 10). Wing receipt density maps (Fig. 11 - 34)
showed that throughout November, woodcock wings were returned throughout the
Central Region. Densities of wing returns in Louisiana gradually increased during
the later half of November and reach high densities by 11-15 December (Fig. 31).
Migration Direction and Destination
The map of Minnesota (Fig. 35) out-of-state direct recoveries (n=7) indicated
that woodcock migrated straight south to the winter grounds in Louisiana. Wisconsin
birds (n=37) were recovered during migration as far east as Kentucky and as far west
as Oklahoma, with the majority of recoveries on the winter grounds in Louisiana (Fig.
36). Michigan recoveries (n=37) were more widely scattered than recoveries from
the other two states (Fig. 37). Michigan banded birds were recovered on the winter
grounds in east Texas, Louisiana, Mississippi, Alabama, and Georgia. Two Michigan
birds were recovered after traveling east of Michigan into Pennsylvania and
Massachusetts (Fig. 37).
Minnesota direct recoveries (n=7) had a mean migration angle of 168º (95%
CI: 188° - 206°) (Fig. 38). This angle was skewed east by two recoveries in
Wisconsin. Wisconsin direct recoveries (n=37) were further west with a mean
migration angle of 182º (95% CI: 179° - 185°) (Fig. 38). Michigan direct recoveries
(n=37) had a migration angle further west than the other two states, with a mean
migration angle of 197º (95% CI: 188° to 206°) (Fig. 38).
22
DISCUSSION
Woodcock were harvested in the Great Lakes States throughout November. This
wide range of harvest dates showed that migration initiation was drawn out over a
period of a few weeks. The dates of peak migration vary among years further adding
to this range. Migration occurred over a relatively short time period, with nearly all
evidence of migration between 1 November and 15 December. The variances in
mean wing receipt and band recovery latitude were greatest during 15-30 November,
at which point migrating woodcock were spread throughout the entire Central Region.
During migration, shorebird species are thought to concentrate along defined
routes and rely on a few historic stopover areas where they can replenish fat reserves
(Myers et al. 1987, Skagen 1997). Band recovery and wing receipt data did not
support the idea of exact migration routes or historic stopover areas for woodcock;
however, the direction of general migration routes was evident. Woodcock banded in
Minnesota and Wisconsin appeared to migrate straight south to the winter grounds.
The majority of Michigan banded woodcock appeared to have migrated south through
eastern Indiana and Illinois, and then south to the wintering grounds through east
Kentucky, east Tennessee, and Mississippi.
Several wing receipts were suspect. These locations were woodcock reported
harvested in the far north in late-December or in the Gulf Coastal states in mid-
September. I realize there are several possible explanations for this, but regardless of
the cause of these locations, they are so few that they did not significantly affect my
results. Another factor that could have possibly affected my results was a difference
23
in hunting pressure. Woodcock are hunted heavily in the Great Lakes states and
Louisiana, so the majority of wing receipts and band recoveries were from those
locations.
Compared to other states, few wing receipts were received from Iowa or
Arkansas. Why these two states had noticeably fewer wing receipts is not clear, but
may include: 1) fewer woodcock being harvested in either state, 2) fewer hunters
participated in the Parts Collection Survey, or 3) woodcock migration stopover
duration was shorter or absent in those states. Maps of all Central Region wing
receipts (Fig. 39) showed a possible trend of densities decreasing from the Great
Lakes states into Iowa and northern Illinois, then increasing in Missouri and southern
Illinois, decreasing again in Arkansas, northern Mississippi, and northern Alabama,
and finally increasing again in Louisiana. This trend provides evidence that
woodcock were possibly migrating over the areas of low abundance and stopping
over in areas of high abundance (i.e. the mid-latitude states). Woodcock were
harvested in all states in the Central Region during fall migration, so I know that birds
were not migrating in one long flight from the breeding to wintering grounds.
Making a series of short flights is always energetically cheaper than one large
flight due to the cost of transporting extra fat (Piersma 1987). Limited availability of
high quality foraging sites is thought to be the reason for shorebirds making long
flights (Piersma 1987). Further research into the availability and quality of woodcock
habitat in the central U.S. might provide information on woodcock migration routes
and flight duration.
24
Most woodcock band recoveries from Minnesota and Wisconsin appeared to
follow the same direction, but there was a large variance in the Michigan recoveries.
Two woodcock banded in Michigan were recovered nearly straight east of their
banding location in Pennsylvania and Massachusetts. Recognize though that such
recoveries could be reporting errors in the Bird Banding Lab files. Woodcock banded
in Minnesota and Wisconsin appeared to overwinter in Louisiana, east Texas, and
west Mississippi. Michigan banded woodcock were recovered on the wintering
grounds throughout the southeastern U.S., however all band recoveries of Michigan
banded woodcock in Alabama, Georgia, and South Carolina were recovered >25
years ago.
There have been a variety of woodcock publications describing their wintering
range. Sheldon (1967) reported that woodcock mainly winter in southeastern
Arkansas, Louisiana, and south-western Mississippi. Straw et al. (1994) analyzed
Christmas Bird Count data and reported that wintering woodcock are common to
abundant in southern Louisiana and east Texas but their wintering range in the central
region extended north to central Missouri. Woodcock winter further north in some
years than others (Williams 1969, Britt 1971, Roberts 1993). Despite these few birds
that overwinter north of traditional areas, I found that most woodcock winter south of
33º N latitude and arrive at their wintering location by 15 December. Of the bands
recovered from 16-31 December, 92.3% were south of 33º N latitude, which
supported Glasgow's (1958) findings that the majority of woodcock are on the
wintering grounds by 15 December. This is further supported by the wing receipt
data, which indicated that harvest leveled off in Louisiana by 15 December.
25
MANAGEMENT IMPLICATIONS Band recovery and wing receipt data both showed little sign of woodcock migration
until 1 November. Woodcock hunting seasons in the Great Lakes states open in late-
September, over a month before migration occurs. Managers need to consider the
effects of harvesting locally produced birds on woodcock populations in their own
states/provinces.
Further investigation is needed to determine the cause of the lack of wing
receipts in Iowa and Arkansas. Outreach efforts might be needed to increase the
amount of public participation in the USFWS Parts Collection Survey in these areas.
Woodcock banding efforts in the Great Lakes states need to be continued to obtain
more information on woodcock fall migration ecology through increased sample
sizes.
LITERATURE CITED Britt, T. L. 1971. Studies of woodcock on the Louisiana wintering ground. Thesis,
Louisiana State University, Baton Rouge, Louisiana, USA. Coon, R. A., P. D. Caldwell, and G. L. Storm. 1976. Some characteristics of fall
migration of female woodcock. Journal of Wildlife Management 40:91-95. _____, T. J. Dwyer, and J. W. Artmann. 1977. Identification of potential harvest
units in the United States for the American woodcock. Proceedings of the Woodcock Symposium 6:147-153. Gregg, L. 1984. Population ecology of woodcock in Wisconsin. Wisconsin Department of Natural Resources. Technical Bulletin 144. Kelley, J.R., Jr. 2003. American woodcock population status, 2003. U.S. Fish and
Wildlife Service, Laurel, Maryland, USA.
26
Keppie, D. M. and R. M. Whiting, Jr. 1994. American woodcock (Scolopax minor). The birds of North America, no. 100. The American Ornithologists’ Union, Washington, D.C., USA, and The Academy of Natural Sciences, Philadelphia, Pennsylvania, USA.
Krohn, W. B., J. C. Rieffenberger, and F. Ferrigno. 1977. Fall migration of
woodcock at Cape May, New Jersey. Journal of Wildlife Management 41:104-111.
Murphy, D.W. 1983. Ecology of American woodcock in central Missouri. Thesis, University of Missouri, Columbia, Missouri, USA. Meyers, J.P., R. I. G. Morrison, P. Z. Antas, B. A. Harrington, T. E. Lovejoy, M.
Sallaberry, S. E. Senner, and A. Tarak. 1987. Conservation strategy for migratory species. American Science 75:18-26.
Piersma, T. 1987. Hop, skip, or jump? Constraints on migration of arctic waders by feeding, fattening, and flight speed. Limosa 60:185-194. Roberts, T. H. 1993. The ecology and management of wintering woodcocks.
Biological Report 16:87-97 Sheldon, W. G. 1967. The book of the American woodcock. University of Massachusetts Press, Amherst, Massachusetts, USA. Skagen, S. K. 1997. Stopover ecology of transitory populations: The case of
migrant shorebirds. Ecological Studies 125:244-269. Williams, S. O. III. 1969. Population dynamics of woodcock wintering in Louisiana. Thesis, Louisiana State University, Baton Rouge, Louisiana, USA.
27
Band Recoveries
28
30
32
34
36
38
40
42
44
46
48
1 - 15 16 - 30 1 - 15 16 - 31 1 - 15 16 - 30 1 - 15 16 - 31
Latit
ude
September October November December Figure 1. Mean (+/- SD) latitude of all woodcock banded in Minnesota, Wisconsin, and Michigan and directly recovered during 15-day periods between 1 September and 31 December 1929-2001.
28
Figure 2. Woodcock banding and direct recovery locations for individuals banded in Minnesota, Wisconsin, and Michigan and recovered between 1-15 September, 1929-2001. Banding and associated recovery locations are connected with a line.
29
Figure 3. Woodcock banding and direct recovery locations for individuals banded in Minnesota, Wisconsin, and Michigan and recovered between 16-30 September, 1929-2001. Banding and associated recovery locations are connected with a line.
30
Figure 4. Woodcock banding and direct recovery locations for individuals banded in Minnesota, Wisconsin, and Michigan and recovered between 1-15 October, 1929-2001. Banding and associated recovery locations are connected with a line.
31
Figure 5. Woodcock banding and direct recovery locations for individuals banded in Minnesota, Wisconsin, and Michigan and recovered between 16-31 October, 1929-2001. Banding and associated recovery locations are connected with a line.
32
Figure 6. Woodcock banding and direct recovery locations for individuals banded in Minnesota, Wisconsin, and Michigan and recovered between 1-15 November, 1929-2001. Banding and associated recovery locations are connected with a line.
33
Figure 7. Woodcock banding and direct recovery locations for individuals banded in Minnesota, Wisconsin, and Michigan and recovered between 16-30 November, 1929- 2001. Banding and associated recovery locations are connected with a line.
34
Figure 8. Woodcock banding and direct recovery locations for individuals banded in Minnesota, Wisconsin, and Michigan and recovered between 1-15 December, 1929-2001. Banding and associated recovery locations are connected with a line.
35
Figure 9. Woodcock banding and direct recovery locations for individuals banded in Minnesota, Wisconsin, and Michigan and recovered between 16-31 December, 1929-2001. Banding and associated recovery locations are connected with a line.
36
Wing Receipts
28
33
38
43
48
53
1 - 5 11 - 15 21 - 25 1 - 5 11 - 15 21 - 25 1 - 5 11 - 15 21 - 25 1 - 5 11 - 15 21 - 25
Latit
ude
September October November December Figure 10. Mean (+/- SD) latitude for all woodcock wings sent to the U.S. Fish and Wildlife Service parts collection survey from birds shot during 5-day periods between 1 September and 31 December 1963-2003. The center point of each county was used as the harvest latitude for each wing receipt.
37
Figure 11. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 1 - 5 September, 1963 - 2002.
38
Figure 12. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 6 - 10 September, 1963 - 2002.
39
Figure 13. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 11 - 15 September, 1963 - 2002.
40
Figure 14. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested from 16 - 20 September, 1963 - 2002.
41
Figure 15. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 21 - 25 September, 1963 - 2002.
42
Figure 16. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 26 - 30 September, 1963 - 2002.
43
Figure 17. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 1 - 5 October, 1963 - 2002.
44
Figure 18. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 6 - 10 October, 1963 - 2002.
45
Figure 19. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 11 - 15 October, 1963 - 2002.
46
Figure 20. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 16 - 20 October, 1963 - 2002.
47
Figure 21. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 21 - 25 October, 1963 - 2002.
48
Figure 22. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 26 - 31 October, 1963 - 2002.
49
Figure 23. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 1 - 5 November, 1963 - 2002.
50
Figure 24. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 6 - 10 November, 1963 - 2002.
51
Figure 25. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 11 - 15 November, 1963 - 2002.
52
Figure 26. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 16 - 20 November, 1963 - 2002.
53
Figure 27. Density of woodcock wings received by the U.S. Fish and Wildlife Service parts collection survey from woodcock harvested between 21 - 25 November, 1963 - 2002.
54
Figure 28. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 26 - 30 November, 1963 - 2002.
55
Figure 29. Density of woodcock wings received by the U.S. Fish and Wildlife Service parts collection survey from woodcock harvested between 1 - 5 December, 1963 - 2002.
56
Figure 30. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 6 - 10 December, 1963 - 2002.
57
Figure 31. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 11 - 15 December, 1963 - 2002.
58
Figure 32. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 16 - 20 December, 1963 - 2002.
59
Figure 33. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 21 - 25 December, 1963 - 2002.
60
Figure 34. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested between 26 - 31 December, 1963 - 2002.
61
Figure 35. Woodcock direct recoveries (n=7) from birds banded in Minnesota and recovered in a different state between 1 September to 31 December, 1929 - 2001. Each banding and associated recovery location is connected with a line.
62
Figure 36. Woodcock direct recoveries (n=37) from individual birds banded in Wisconsin and recovered in a different state from 1 September to 31 December, 1929 - 2001. Banding and associated recovery locations are connected with a line
63
Figure 37. Woodcock direct recoveries (n=37) from individual birds banded in Michigan and recovered in a different state from 1 September to 31 December, 1929 - 2001. Banding and associated recovery locations are connected with a line
64
Minnesota n = 7
Minnesota n = 7
Wisconsin n = 37
Michigan n = 37
Figure 38. Mean migration angle and 95% confidence intervals of woodcock banded in Minnesota, Wisconsin, and Michigan and directly recovered in a different state between 1 September to 31 December, 1929 - 2001.
65
Figure 39. Density of woodcock wings received by the U.S. Fish and Wildlife Service Parts Collection Survey from woodcock harvested from 1 September – 31 December, 1963 - 2002.
66
Figure 40. Woodcock band recovery locations from 1 September to 31 December, 1929-2001.
67
Chapter 3
FALL MIGRATION RATES, ROUTES, AND HABITAT USE OF AMERICAN
WOODCOCK IN THE CENTRAL REGION¹ __________________ ¹Myatt, N. A., and D. G. Krementz. To be submitted to The Journal of Wildlife Management.
68
Abstract: American woodcock (Scolopax minor) ecology has been extensively
studied on the breeding grounds and to a lesser extent on the wintering grounds, but
little research has been conducted on the migration ecology of this declining species.
In Fall 2001 I began a 3-year study to document woodcock fall migration routes,
rates, and habitat use in the Central Region of the U.S. From 2001-2003, 582 radio-
marked woodcock initiated migration from 3 study sites in Minnesota, Wisconsin,
and Michigan. Aerial searches were conducted from fixed-wing aircraft during each
fall migration period in the Central Region. During 224 hours of aerial telemetry, I
located 42 radio-marked woodcock in 6 states. Radio-marked birds were located in
upland habitats more frequently than bottomland habitats (80% vs. 20%,
respectively). Migrating woodcock used a higher proportion of mature forest than
expected. Stopover duration often exceeded 4 days, with some birds stopping longer
than a week. Using locations of radio-marked birds, I speculated woodcock
migration routes in the central U.S. GIS was used to map potential woodcock habitat
in the Central Region. Based on my results, I have identified priority areas for future
woodcock management in the Central Region.
INTRODUCTION
American woodcock ecology has been studied extensively on the northern
breeding grounds and to a lesser extent on the wintering grounds, but little research
has been conducted on the migration ecology of this declining species (Keppie and
Whitting 1994). Although the timing of departure from the breeding grounds and
arrival on the wintering grounds has been documented, little is known about what
69
happens during the migration period between these areas. Knowledge of woodcock
migration has been limited to anecdotal evidence, interpretations of a limited sample
of band returns (Glasgow 1958, Sheldon 1967, Krohn et al. 1977), monitoring the
onset of migration in small samples of radio-marked birds (Godfrey 1974, Coon et al.
1976, Gregg 1984, Sepik and Derleth 1993), and harvest data (Roberts 1978, Murphy
1983). Minimal data have been collected in the Central Region (Fig. 1) on migration
habitat use, stopover duration, or fall migration routes of woodcock.
Woodcock breed primarily in dense early successional habitats in the Great
Lakes states and provinces (Keppie and Whitting 1994). They initiate fall migration
in the Central Region in late-October and early-November (Keppie and Whitting
1994). Their nocturnal migratory flights often coincide with the passage of cold
fronts. Owen (1977) documented the winter distribution of woodcock, with the
northern extent ending in northern Louisiana. Glasgow (1958) reported that most
woodcock have arrived on the wintering grounds by 15 December, but their
distribution in Louisiana depended on winter severity. During mild winters,
woodcock were spread throughout Louisiana, while during cool winters they were
absent from the northern half of the state and concentrated in the Atchafalaya and
Mississippi River basins (Glasgow 1958). Straw et al. (1994) analyzed Christmas
Bird Count data and reported that woodcock winter in low densities as far north as
Missouri.
Glasgow (1958) used a sample (n=175) of woodcock band returns to predict
fall migration routes across the woodcock’s range (Fig. 2). Glasgow hypothesized
that woodcock migrating from Minnesota and Wisconsin might follow the
70
Mississippi River south to Missouri where they then travel through central Missouri
and Arkansas on their way to south central Louisiana. Sheldon (1967) used ten
additional years of band return data (n=400) and hypothesized that woodcock
migrating from Minnesota and Wisconsin might follow the Mississippi River south
through the lower Mississippi Alluvial Valley (MAV) to the winter grounds (Figure
2). During the 1950s-1970s, the MAV lost 120,000 hectares per year of bottomland
hardwood forest (MacDonald et al. 1979). Woodcock may no longer use the MAV
during migration due to a lack of suitable migration habitat.
Changes in habitat availability might have caused a change in migration flight
distance and stopover duration, but little is known about these two components of
woodcock migration. A woodcock, radio-marked by Sepik and Derleth (1993) was
shot in New York on 9 November, two days after it was last located 680 km away in
Maine. Coon et al. (1976) documented the distance and stopover duration of 2 fall
migration flights. Two radio-marked woodcock were tracked up to 201 km SSW of
the study area on their first two nights of migration. Both birds traveled 53-56 km on
their first night’s flight. On the second evening of migration, one resumed migration
at 1845 hr and the other departed at 1910 hr. Other than Coon et al. (1976), there is
no other data on the length of each leg of woodcock fall migration, the duration of
stopovers, and the time at which woodcock resume migration during migration
stopovers.
Migration stopover locations are crucial links between breeding and wintering
grounds where avian migrants rest and/or refuel for the next leg of migration (Farmer
and Parent 1997). Many species of shorebirds use traditional stopover locations
71
where a significant portion of the population might stop in a given year (Farmer and
Parent 1997). Stopover duration is the length of time that a migrating bird remains
at a stopover location during fall migration. Stopover duration may be directly
related to the availability of high quality foraging sites (Piersma 1987). Making a
series of short flights is always energetically cheaper than covering the same distance
in one long flight (Piersma 1987). Short stopover durations would provide evidence
that woodcock are migrating in a series of short "hops." Longer stopover durations
would suggest that birds are refueling after migrating in several "jumps," where
jumps are long flights followed by long stopovers. The availability and distribution
of woodcock habitat in part of the Central Region could affect stopover duration.
The extent and distribution of woodcock habitat in the U.S. remains largely
unknown. Advancements in Geographic Information System (GIS) technologies
have potential applications in mapping national woodcock habitat distribution
(Duncan 2000). Currently, nation-wide, spatial data are not sufficient to map
woodcock habitat availability. Suitable woodcock habitat is determined by
vegetation physiognomy rather than species composition (Bourgeois 1977, Cade
1985, Straw et al. 1994), which is the primary classification method of current spatial
data.
In fall 2001, I began a 3-year study of woodcock fall migration ecology in the
Central Region. My project took advantage of a concurrent study on the effects of
hunting mortality on woodcock populations in the Great Lakes region. That study
radio-marked up to 360 woodcock on the breeding grounds each fall and monitored
their location daily until they began migration. Once those birds migrated, I relocated
72
them throughout the Central Region. The objectives of my research were to: 1)
document woodcock fall migration routes in the Central Region, 2) determine the
stopover duration of woodcock during fall migration, and 3) document woodcock
habitat use during fall migration.
STUDY AREA
In North America, the range of woodcock has been divided into 2
management units: Eastern and Central regions (Coon et al. 1977) (Fig. 1). My study
focused on the Central Region, which includes the woodcock’s range west of the
Appalachian Mountains. Woodcock were radio-marked at 3 northern study sites (Fig.
3): 1) Mille Lacs and Four Brooks Wildlife Management Area in east-central
Minnesota (45.93º N, 93.55º W), 2) Lincoln County Forest and Tomahawk
Timberlands in north-central Wisconsin (45.34º N, 89.94º W), and 3) Copper Country
State Forest on the west-central border of Upper Peninsula Michigan (46.15º N,
87.83º W). My aerial search efforts for radio-marked birds were focused in Arkansas,
Illinois, Iowa, Kentucky, Louisiana, Mississippi, Missouri, Oklahoma, Tennessee,
and Texas.
Based on the Palmer Drought Index, fall of 2001 was abnormally dry in
central Minnesota and northeast Arkansas during late-October and early-November
(N.O.A.A. 2004). Otherwise, there were no other drought conditions that year in the
Central Region. In fall 2002, abnormally dry and moderate drought conditions were
present during late-October and early-November in western Missouri and northwest
Arkansas. Later that fall, drought conditions spread throughout Missouri, southern
73
Iowa, northern Illinois, and northern Indiana. There were no abnormally dry
conditions observed in fall 2002 on the wintering or breeding grounds. In fall 2003,
Minnesota, Wisconsin, Michigan, Iowa, and northern Illinois experienced abnormally
dry to moderate drought conditions. During late-October and early-November
northern Louisiana and southern Arkansas also experienced abnormally dry
conditions. During late-November and December, there were abnormally dry
conditions in Oklahoma, northeast Texas, and southeastern Louisiana.
METHODS
Capture
In 2001, woodcock were captured only at the Minnesota study site, while in
August 2002 and 2003, woodcock were captured at each of the 3 northern study sites.
Capture efforts ended 1 October each year to reduce the probability of marking non-
resident woodcock. Woodcock were mist-netted during crepuscular flights between
diurnal habitat and nocturnal feeding fields. Capture by spotlighting from all terrain
vehicles, trucks, and on foot was also used when conditions permitted (Reiffenberger
and Kletzly 1967, McAuley et al. 1993).
After capture, birds were aged and sexed using wing plumage characteristic
(Martin 1964). Birds were outfitted with 4.4-g radio transmitters using all-weather
livestock tag cement and a belly-loop wire harness (McAuley et al. 1993). In 2001
and 2002, active radios had a pulse rate of 55 pulses per minute (ppm) and were
powered by a 1.5-volt silver-oxide battery. Based on the pilot’s observations, the
74
ground to air range of the radios was variable with a mean distance of about 8 km and
a maximum distance of 20 km.
Each radio transmitter had a frequency unique to the capture state, but there
were some frequency overlaps among the 3 states. I considered a bird to be uniquely
identifiable if the frequency of the radio transmitter was >0.007 MHz apart from all
other birds. In 2003, the problem of overlapping frequencies was corrected by setting
half of the radios at 55 ppm and the other half at 70 ppm. Using pulse rate and
frequency, all radio-marked woodcock should have been uniquely identifiable in
2003.
Telemetry
Each day northern field crews confirmed the presence of all radio-marked
birds until they were censored. Once several birds were missing from the study area,
aerial searches from fixed-wing aircraft were performed at 3- to 7-day intervals to
relocate missing birds. After two telemetry flights, if birds were not found within
approximately 24 km of the study area, they were classified as having possibly
migrated. The assumption that a bird had migrated when it could not be located was
problematic due to the possibility of radio failure, temporary emigration, or
unreported harvest, so migration dates and numbers of marked birds may be over
estimated. I communicated weekly with northern field crews to determine which
birds were missing from the study areas and the probable dates when migration
began.
After 50% of the radio-marked woodcock from the northern study areas were
censored, I began diurnal searches for the birds from fixed-wing aircraft. The
75
primary plane that I used was outfitted with two 2-element, directional antennas
(Gilmer et al. 1981). To a lesser extent, I used a different plane outfitted with 2 omni-
directional aircraft antennas (McAuley et al. 1993). I searched at an average airspeed
of 200 km/hr and an altitude of 1000-3000 m. Transmitter frequencies were
programmed into a receiver and scanned on 2-sec intervals for the duration of the
flight.
Before each field season, I contacted wildlife researchers in my study area to
determine the existence of other radio telemetry projects within my frequency range.
When flying over known telemetry projects, the pilot had a list of the frequencies and
locations of radio-marked animals unrelated to my project.
Each season, I flew the first flights in the northern portion of my study area
and then moved flights south as migration progressed. Areas searched and times of
search were based on current literature, historic band return data, wing receipt data,
and relief maps of the Central Region. In the first half of 2002 and in 2001, I flew
along major rivers, bluff edges of the Mississippi River, and randomly over areas of
possible habitat. In 2003 and the last half of 2002, I located more birds by flying 160
km x 144 km blocks with transects spaced 24 km apart. The size of these blocks was
occasionally altered due to time constraints, but the distance between transects was
consistent throughout the remainder of the study. After locating a signal, my pilot
recorded the location and immediately relayed the information to me on the ground.
The pilot was experienced in wildlife telemetry, so he screened each signal heard as a
good possibility or a questionable signal (e.g., due to signal strength, range, or pulse
rate.)
76
Once a radio signal was located from the air, I attempted to locate and flush
the bird to confirm that the signal was a radio-marked woodcock. Woodcock often
move up to 60 m in front of a researcher or bird dog before flushing (Dyer and
Hamilton 1977). To eliminate any difference between the initial location of the bird
and the flushing location, I triangulated the bird’s location from 10 m away while my
bird dog was kept at my side, then I sent the dog in to flush the radio-marked bird.
Habitat data, site description (see below), and the number of unmarked woodcock
flushed were recorded at the point of flush. Search effort for unmarked woodcock
was not equal between locations due to time constraints, property access, and cover
type. When time permitted, I monitored marked birds daily to determine stopover
duration. I considered radio-marked woodcock located south of 33° N latitude
(northern border of Louisiana) to be on the winter grounds, while birds located north
of this line were considered to be in migration.
If I was unable to relocate the signal when I arrived at the pilot's recorded
coordinates, then I searched a 5-km radius area with a truck-mounted, 5-element,
yagi-style antenna. If the signal was not found in the area, then the bird was
presumed to have resumed migration and the location was recorded as unconfirmed.
I used all radio-marked woodcock locations to create a fall migration route
map. I then used the migration routes, coupled with stopover duration, to identify
priority areas for future woodcock management in the Central Region.
Habitat
I recorded habitat data and site characteristics in a 30-m radius area centered at the
point of flush for each radio-marked bird. Cover type was classified using the
77
National Vegetation Classification System (Anderson et al. 1998). Size classes of
over-story trees were grouped into sapling (<7.5 cm diameter at breast height
(diameter at breast height (DBH)), pole (7.5 cm - 15 cm DBH), and mature (>15 cm
DBH). I recorded the 4 most prevalent species of ground vegetation (0 - 0.5 m) and
midstory vegetation (0.5 - 2 m). I recorded the horizontal density of the ground and
midstory vegetation in 4 cardinal directions by ranking the density on a scale of 1-5.
This equal interval scale was classified as: 1 -little or no standing vegetation, 2 - 20 –
40% cover, 3 - 40 – 60% cover, 4 - 60 – 80% cover, and 5 - nearly impenetrable
thicket with dense standing vegetation. I estimated canopy coverage, in 4 cardinal
directions, with a convex spherical densiometer (Lemmon 1957). Distance to the
nearest edge, edge type, and visual estimates of habitat patch size were recorded. I
defined edge as an abrupt change in habitat structure, such as a road, river, or clearcut
edge.
I determined soil texture by feel (Tinner 1999). In 2002, I ranked soil
moisture as dry, moist, or wet. In 2003, I collected soil samples with a cylinder 12.7
cm diameter by 8 cm deep; the maximum woodcock bill length (Keppie and Whitting
1994). After oven drying, the percentage moisture of the soil sample was calculated
by dividing the dry weight (g) by the wet weight (g).
Earthworm densities were estimated using a hot mustard extraction technique
(Gunn 1992, Lawrence and Bowers 2002). I removed all vegetation and leaf litter
from a 35 x 35 cm plot and evenly applied a solution of 78 ml oriental hot mustard
powder and 1 L of water. All earthworms that surfaced within 5 min of application
were collected and preserved in 4% formalin for 48 hr and then transferred to a 70%
78
alcohol solution. Earthworms are not evenly distributed in the soil (Poier and Richter
1992), so I collected three samples spaced 5 m apart at each woodcock location.
These samples were averaged to determine the earthworm density per m².
I was unable to find any documentation on the use of a soil penetrometer in
woodcock research. When studying woodcock, many biologists collect soil data such
as texture, moisture, and porosity. These are indirect measures of how easily a
woodcock can penetrate the soil. I used a Lang soil penetrometer to determine the
amount of pressure that is needed to penetrate the soil with a probe roughly the size
and shape of a woodcock beak. I collected 3 penetrometer readings adjacent to each
earthworm sampling plots. The values were converted from pounds per in² to kg/cm²
and then averaged to determine the average kg/cm² of earthworms for each sampling
plot.
I used the 1992 National Land Cover Data (NLCD) (U.S. Geological Survey
2003) in a Geographic Information System (GIS) to map potential woodcock habitat
in the Central Region. The NLCD was created from early to mid-1990s Landsat
Thematic Mapper satellite data. The NLCD is classified into 21 classes (Table 1) and
has a spatial resolution of 30 m. Using available spatial data, I was not able to map
habitat availability, but I was able to identify areas of no diurnal habitat (Table 1)
resulting in a map of potential woodcock habitat.
After mapping potential habitat, I overlaid 4 hypothesized migration routes
and 2 migration routes that I observed radio-marked birds using. The first 3
hypothesized migration routes were the shortest lines from each of the 3 northern
study areas to the winter grounds of central Louisiana. The other hypothesized route
79
followed the Mississippi River from the breeding grounds to central Louisiana. An
ArcView script was used to create a point every 5 km along these routes. I then
calculated the percent potential habitat within a 50 km buffer of each point. Values
were then graphically displayed to show the change in potential habitat availability
along each migration route.
RESULTS
Sample Size
In Fall 2001, 64 radio-marked woodcock were censored from the Minnesota
field site. In Fall 2002, 274 radio-marked woodcock were censored from Michigan
(n=92), Minnesota (n=94), and Wisconsin (n=91). In Fall 2003, 244 radio-marked
woodcock were censored from Michigan (n=63), Minnesota (n=102), and Wisconsin
(n=79).
Telemetry
During Fall 2001 pilot season, I located 4 radio-marked woodcock while
scanning 64 frequencies on 3 flights (Fig. 4) totaling 24 hours. Three of the locations
were confirmed by flushing the radio-marked woodcock. During Fall 2002, I located
29 radio-marked woodcock while scanning for 235 frequencies on 16 flights totaling
125 hours (Fig. 5). Twenty-two of the locations were confirmed. During Fall 2003, I
located 9 radio-marked woodcock while scanning for 162 frequencies on 10 flights
totaling 75 hours (Fig. 6). Seven of the locations were confirmed. I flew a greater
number of flights in 2002 than in 2003 due to budgetary constraints.
80
During 2001, woodcock were radio-marked only in Minnesota, so I know the
origin of birds located that year (Fig. 7). The origin of 18 of the relocated radio-
marked birds from 2002 was unknown because multiple birds were marked with
radios ≤ 0.007 MHz apart. Of the remaining relocated birds, 4 were from Wisconsin,
1 was from Michigan, and 5 were from Minnesota (Fig. 7). In 2003, the origin of 2 of
the relocated radio-marked birds was unknown due to drift in the pulse rate of our
radio transmitters. Of the remaining relocated birds, 3 were from Wisconsin, 1 from
Michigan, and 3 from Minnesota (Fig. 7). I often, flushed unmarked woodcock, up to
8, from the same forest stand as the radio-marked bird. Radio-marked woodcock
were never located again during migration or later on the winter grounds.
Of the possible radio-marked woodcock found in 2001, two were males and
two were females. Three of these were hatch year birds and one was an after
hatching year bird (Fig. 2). Of the locations in 2002, 10 were males, 14 were
females, and the sex of five was unknown. Eleven of the locations were hatch year
birds, five were after hatching year, and the age of 13 was unknown. In 2003, seven
of the marked-woodcock locations were females and two were unknown sex. Of the
nine locations, seven were after hatch year birds and 2 were unknown.
Migration Distance
I was able to record distance and estimated flight duration for 13 radio-marked
woodcock (Table 4). Two birds were found in Missouri and Illinois, 16 and 21 days
after they were last located on the breeding ground. Birds were found in southern
Arkansas 20 - 48 days after they were last located on the breeding ground. One bird
traveled from Wisconsin to northeastern Texas in <16 days, a total distance of 1406
81
km. Five birds on the winter ground were found 22-41 days after they were last
located on the breeding ground.
Stopover Duration
I documented stopover durations of 24 radio-marked woodcock (Table 5). I
know the exact stopover duration of seven birds, possible stopover range of six, and
minimum stopover duration for 11. Four birds resumed migration on the day of
initial location, but 13 birds stopped over for > four days. The longest stopover
observed was a bird in northern Arkansas that stayed for 10-16 days before resuming
migration. The exact stopover duration of this bird was unknown because I was
unable to visit the site for 6 days. I observed the timing of migration initiation after
stopover in four cases. Two birds migrated before 1945, one before 2100, and one
after 2000.
Migration Habitat
I collected habitat data at 22 confirmed locations. Marked woodcock located
in or north of the Ozark Mountains (35º N) were found more often in upland
hardwood cover types (n=6) than in bottomland cover types (n=2) (Table 6). Marked
woodcock locations south of the Ozark Mountains were found more often in upland
pine or pine/hardwood cover types (n=9) than in bottomland cover types (n=4) (Table
6). Locations were found more often in mature (n=8) and sapling (n=10) size classes
than in pole (n=3) size class. Median habitat block size was 0.108 (SD 3.754, n=18)
km² with a range of 0.0007-15.37 km². Median distance of marked woodcock from
habitat edge was 14 m (SD 52.236, n=18) with a range of 0-230 m.
82
Woodcock used habitat with ground densities ranging from 1-3.25 with a
mean of 1.98 (SD 0.778, n=18) and mode of 1 (Fig. 8). Ground vegetation was most
often herbaceous plant species, blackberry (Rubus spp.), greenbrier (Smilax spp.), and
Japanese honeysuckle (Lonicera japonica) (Table 7). Midstory densities ranged from
1-5 with a mean of 2.86 (SD 1.272, n=18) and median of 2.82 (Fig. 8). Blackberry,
loblolly pine (Pinus taeda), greenbrier, and herbaceous plant species were most often
found in the midstory vegetation (Table 7 ). Percentage canopy cover ranged from 0-
99.22% with a mean of 58.45% (SD 37.798) and median of 77.64% (Fig. 9).
I located marked woodcock in 7 different soil types including sandy loam
(n=5), clay loam (n=4), loam (n=3), silty clay loam (n=2), silt loam (n=2), silty clay
(n=1), and sandy clay loam (n=1). In 2003, percent soil moisture ranged from 13.2-
30.69% with a mean of 22.58% (SD 7.107, n=5). Earthworm densities ranged from
0-3.07 g/m² with a median density of 0.84 g/m² and mean of 1.37 g/m² (SD 1.509,
n=5). Soil hardness ranged from 2.41-4.37 kg/cm² with a mean hardness of 3.20
kg/cm² (SD 1.333, n=5) and median of 3.09 kg/cm².
Winter Habitat
Habitat data were collected at 10 confirmed winter locations. Wintering birds
were found more often in upland cover types (n=8) than in bottomland cover types
(n=2) (Table 6). Locations were more common in mature (n=5) and sapling (n=4)
size classes than in pole (n=1). I found blackberry, greenbrier, and oak species most
often in the midstory vegetation (Table 8) and blackberry and Japanese honeysuckle
most often in the ground vegetation (Table 8). Horizontal midstory densities ranged
83
from 2.5-5 with a mean and median of 3.5 (SD 0.792). Ground densities ranged from
1-3, with a mean of 1.37 (SD 0.891) and a median of 1.25.
Wintering marked woodcock were present in 4 soil types: loamy sand (n=3),
sand (n=2), sandy loam (n=2), and silty clay loam (n=1). In 2003, percent soil
moisture at the two winter locations were 23.06% and 28.35%. Earthworm densities
were 1.58 g/m² and 0 g/m². Soil hardness was 1.25 kg/cm² and 2.40 kg/cm².
Fall Migration Routes
Using my sample of radio-marked woodcock locations, I mapped 2 possible
fall migration routes: 1) Ozark Route and 2) Mississippi Route (Fig. 10). The 2
possible routes from Minnesota, Wisconsin, and Upper Peninsula Michigan
converged on the Mississippi River as they headed south into Iowa and Illinois. In
east-central Missouri, the Ozark Route birds headed south through the Ozark
Mountains until they reached the pinelands of the Gulf Coastal Plain. Once they
reached the pinelands, they eventually spread out and worked their way south
throughout the pinelands of western and central Louisiana and eastern Texas. A
smaller percentage of the birds used the Mississippi Route through the Bootheel of
Missouri, over the northern portion of the lower Mississippi Alluvial Valley (MAV),
and then followed the bluff edge of the MAV south into Mississippi.
Potential Habitat Map
Several areas in the Central Region had a limited amount of potential diurnal
woodcock habitat (Fig. 11). The first area was the agriculture/grassland-dominated
areas of southern Minnesota, Iowa, northern Illinois, northern Indiana, and western
Ohio. On the western edge of the woodcock’s range there was limited potential
84
habitat in the Dakotas, Nebraska, and Kansas, but potential habitat increased in
Oklahoma and Texas. There was also limited potential habitat throughout the MAV.
Within the MAV, the amount of potential habitat increased from north to south.
Extensive potential habitat existed in the northern half of the Great Lakes states, in
the Ozark Highlands of Missouri and Arkansas, the West Gulf Coastal Plain of
southern Arkansas, western Louisiana, and eastern Texas, and throughout Kentucky,
Tennessee, Mississippi, and Alabama. On the winter grounds in Louisiana, potential
habitat was highest in the upland areas in the west central, north central, and areas
north of Baton Rouge and Lake Pontchatrain.
DISCUSSION
Woodcock use early successional habitat almost exclusively on the breeding
grounds and a wider range of early successional and mature forests on the wintering
grounds (Cade 1985, Keppie and Whitting 1994, Straw et al. 1994), but the point in
migration at which they start using mature forest is unknown. The first woodcock
that I found using mature forests were in Illinois and Missouri where mature oak
forest was used. I also found woodcock in mature forest further south in Arkansas,
Mississippi, Louisiana, and Texas.
Historically, mature forests used on the wintering ground are thought to have
a distinct understory (Keppie and Whitting 1994). Understories least prefer by
woodcock are extremely dense or open habitats (Cade 1985). The southern locations
I found were usually in pinelands or hardwoods with a developed understory, whereas
the mature forest locations in Missouri and Illinois were void of an understory.
85
Although the majority of radio-marked woodcock locations were in moderately dense
habitats, some locations were found in extremely dense or open habitats.
On the winter grounds, woodcock are thought to primarily use bottomland
areas and secondarily use pineland areas (Roberts 1993, Straw et al. 1994). During
migration and on the winter grounds, I found most locations in upland oak, pine, or
pine/hardwood forests. Managers could have underestimated the importance of these
habitats.
When moisture is limited, pineland areas, mixed pine-hardwood stands,
hardwood drainages, and bottomlands provide more suitable habitat than
predominantly pine areas (Boggus and Whiting 1982). Researchers studying
woodcock habitat selection on the winter grounds have found that woodcock using
pinelands are often associated with riparian hardwood drainages located within a pine
stand (Roberts 1993). Eight of my locations were associated with these riparian strips,
while 9 locations were not associated with any type of drainage. During my study,
the Palmer Drought Index (N.O.A.A. 2004) never fell into significant drought
conditions in the southern Central Region. Thus, the habitat types used by marked
birds were probably reflective of normal weather conditions. Use of bottomland
hardwood stands may be restricted to periods of drought (Krementz and Pendleton
1994, Krementz 2000) or during later winter (M. Olinde, Louisiana Department of
Wildlife and Fisheries, personal communication).
During spring and summer, woodcock mostly feed during diurnal hours
(Keppie and Whitting 1994), however during winter they feed extensively at night
(Krementz et al. 1995). Some birds were located during the day in dry soil habitats
86
with no available earthworms, so migratory woodcock might be feeding primarily in
their nocturnal habitats.
On the winter grounds, woodcock prefer nocturnal block sizes from 0.05 – 0.4
km² (Krementz 2000). I found median stand size at diurnal migrating woodcock
locations of 0.108 km². Within these stands, woodcock were most often found
associated with a habitat edge. Of 28 confirmed locations, 36% were ≤ 10 m from an
edge and 86% were ≤ 50 m from an edge. Possible explanations of the association
with edge might have been caused by highly fragmented landscapes, differences in
vegetation structure near edges, or due to my consideration of a stream as an edge.
Habitat availability might have been limited in certain parts of the migration
route. Making a series of short flights is energetically cheaper than covering the same
distance in one long flight (Piersma 1987). Migrating shorebirds are thought to make
longer flights when the availability of quality habitat is limited (Piersma 1987).
Expansive areas of limited habitat could have been “ecological barriers” requiring
non-stop flights. Further research is needed to determine if woodcock are migrating
over areas of low habitat availability.
During fall migration, woodcock initially encountered an expansive area of
minimal potential habitat in Iowa, northern Illinois, and northern Indiana. Birds that
migrated from the 3 northern study sites, potentially had to travel 600-800 km before
reaching areas of expansive potential habitat in central Missouri and southern Illinois.
The MAV is another expansive area of limited potential habitat, especially compared
with the amount of habitat that was historically available.
87
Coon et al. (1976) documented 2 woodcock stopping over after an initial
nocturnal migration flight and then resuming fall migration the following evening.
The stopover durations I observed were much longer than Coon et al.’s observations.
There are two possible explanations for the difference in stopover durations. Coon et
al’s (1976) observations were both on the initial leg of fall migration that might not
be representative of true migration. An alternative explanation would be that these
birds were experiencing migratory restlessness. Krementz et al. (1994) documented
migratory restlessness when woodcock moved out of their study area before
commencing spring migration.
I was unable to monitor all birds until they resumed migration and my
stopover durations did not include the time that the radio-marked bird spent at the
location before my locating it. So, stopover durations are most likely longer than I
observed (Kaiser 1999, Lehnen 2004). Average stopover is probably greater than a
few days and often over a week.
There are two competing hypothesis proposed to explain the time that a
migrant bird spends at a stopover location: 1) time-selection (Alerstam and
Lindstrom 1990) and 2) energy-selection (Gudmundsson et al. 1991). Under the
time-selection hypothesis, the migrant minimizes its total migration time by passing
lower quality stopover sites. Under the energy-selection hypothesis, the migrant will
travel to the next stopover site as soon as it has the energy to do so, regardless of the
quality of the site. Migrants operating under the time-selection hypothesis would
spend less time at high quality sites, where as migrants would spend more time at
88
high quality sites under the energy-selection hypotheses. Further research is needed
to determine which mechanisms determine the stopover duration of woodcock.
Woodcock have the shortest migration distance of 37 shorebird species found
in the central U.S. (Skagen 1997). Furthermore, they are the only North American
shorebird with rounded rather than pointed wings (Sheldon 1967), making
continuous, cross-continent flights unlikely. I found radio-marked woodcock in the
mid-latitude states of the Central Region, so they were not migrating from the
breeding grounds to the wintering grounds in one large jump. I observed long
stopover durations, so woodcock were probably not migrating in many short flights
followed by short stopovers. Long stopover durations suggests that woodcock are
migrating in several long flights followed by extended periods of refueling.
Unlike migrating songbirds, shorebirds are thought to follow fixed routes
during migration, although these routes are more defined in coastal areas (Skagen
1997). Woodcock are found throughout the Central Region during migration, but
certain migration routes have much higher densities of migrants (Glasgow 1958). I
proposed 2 possible migration routes (Ozark and Mississippi Route) used by radio-
marked woodcock due to the spatial pattern of marked woodcock locations. After
flying 224 hours of telemetry flights in the Central Region, certain areas were
continuously void of radio-marked woodcock while other areas contained radio-
marked birds. My possible woodcock fall migration routes differed from those of
Glasgow (1958) and Sheldon (1967), mainly due to the lack of radio-marked bird
locations in the MAV. Loss of bottomland habitats within this area apparently has
89
caused a shift in woodcock migration routes. Migration routes that historically
passed through the MAV might have shifted to the valley edges to avoid this area.
The majority of radio-marked woodcock locations were along the Ozark
Route through central Missouri and central Arkansas. After crossing Iowa and
Illinois, potential habitat availability increased continuously to near 90% on the
winter grounds (Fig. 12). The Mississippi Route peaked near 90% in southeastern
Missouri but then dropped temporarily while crossing the MAV (Fig. 13). The two
observed migration routes (Ozark and Mississippi Routes) had greater densities of
potential habitat than straight lines between the 3 study sites and south central
Louisiana (Fig. 14 -16) and the route following the Mississippi River (Fig. 17).
I used the straight-line distance from each northern study area as hypothesized
routes because many migrant shorebirds take the shortest route between breeding and
winter grounds (Farmer and Parent 1997). I used the Mississippi River as the fourth
hypothesized migration route because Coon et al. (1976) observed radio-marked
woodcock migrating in a direction that coincided with a river valley. The 4
hypothesized migration routes ended in south central Louisiana because that is where
the highest densities of overwintering woodcock were historically thought to occur
(Glasgow 1958). My 2 observed routes (Ozark and Mississippi Routes) ended in the
pinelands of Louisiana and Mississippi because I did not locate any radio-marked
woodcock south of those locations. Due to the battery life of my radios, I could not
monitor the location of my birds through the winter, so I cannot be certain that the
birds in Texas, Louisiana, and Mississippi had arrived at their final winter locations.
90
Regardless of whether or not they were wintering, I have shown that these pineland
locations are important to woodcock.
Sheldon (1967) reported that woodcock mainly winter in southeastern
Arkansas, Louisiana, and south-western Mississippi. Owen et al. (1977) reported
that woodcock breeding west of the Appalachian Mountains winter in Arkansas,
Louisiana, Mississippi, and Alabama. Root (1988) analyzed Christmas Bird Count
data and found the greatest densities of wintering woodcock in east Texas near Sam
Rayburn and Toledo Bend Reservoirs. Straw et al. (1994) analyzed Christmas Bird
Count Data and reported that wintering woodcock are common to abundant in south
Louisiana and east Texas, scattered to common in southeast Mississippi, north
Louisiana, south and east Arkansas, east Texas, southeast Missouri, and west
Kentucky and Tennessee. Straw et al. (1994) concluded that woodcock wintering
range in the central region extends to central Missouri. I used 33º N latitude as the
northern extent of the primary wintering range due to banding records and
observations of radio-marked woodcock in Arkansas (see below). There is no
definitive boundary to the winter range. Woodcock probably winter in low densities
in southern Arkansas and northern Mississippi in some years and are absent in others.
Despite these few birds that overwinter north of traditional areas, most woodcock
winter farther south and arrive at their wintering location by 15 December. Of the
woodcock banded in Minnesota, Michigan, and Wisconsin and directly recovered
from 16-31 December 1929-2001 (n=26), 92.3% were south of 33º N latitude. This
same pattern is seen when looking at all Central Region direct recoveries during this
period (93.1%, n=98)) and all woodcock direct recoveries west of the Atlantic Coastal
91
States (89.7%). Of the 8 birds in southern Arkansas that I was able to monitor for
about a week after location, 6 resumed migration. The other 2 were present after 1
week.
In Louisiana, woodcock densities were historically thought to be highest in the
bottomland areas of south-central Louisiana (Glasgow 1958, Britt 1971, Straw et al.
1994, M. Olinde, Louisiana Department of Wildlife and Fisheries, personal
communications). In 2002 and 2003, I searched for radio-marked birds throughout
Louisiana and east Texas (6 - 20 December), but my only locations were in the
pinelands and associated bottomlands of the Gulf Coastal Plain (Fig. 7). Managers
could have underestimated the importance of this region to migrating and over-
wintering woodcock.
MANAGEMENT IMPLICATIONS
Many species of shorebirds in the central U.S. use traditional stopover
locations where a large percent of the population may stop in a given year (Farmer
and Parent 1997). From my telemetry data it appears that woodcock are not
concentrating at major stopover locations. They are most likely opportunistically
selecting stopover locations at the end of each night’s flight. Therefore, in the Central
Region, I identified large geographic areas of importance to migrating woodcock.
Based on densities of radio-marked woodcock locations, I identified four
locations as priority areas for consideration of future woodcock management in the
Central Region (Fig. 18). To determine high priority areas, I located areas that had the
highest densities of radio-marked woodcock locations and areas with moderate
92
densities to determine medium level priority areas. The first area was the southern
pinelands of Arkansas. This high priority area was centered on the Saline River and
dominated by industrial pinelands with lowlands of bottomland hardwoods.
Surrounding this area was a medium level priority area in which I located a few
radio-marked woodcock and the habitat was similar to that of the high priority area.
A medium level priority area was located in northeastern Missouri and west
central Illinois where the Mississippi, Illinois, and Missouri Rivers converge. This
funnel area concentrated birds from Minnesota, Wisconsin, and Upper Peninsula
Michigan. This area was a potential stopover site after a migration flight over
predominantly grassland/agriculture areas of Iowa and Illinois.
Another medium level priority area was the pinelands of northern Mississippi
along the bluff edge of the MAV. I selected this area because it was a possible funnel
area between the agricultural dominated lands of the MAV to the west and the
Tombigbee River Valley to the east. This area was a potential stopover after birds
crossed the primarily agricultural area of the MAV. Woodcock migrating from the
eastern half of the Central Region may have passed through this area also.
Two high priority areas were on the wintering grounds. One was the
pinelands of north-central Louisiana between the MAV and the Red River and the
second was the pinelands of western Louisiana and eastern Texas between the Red
River and Sam Rayburn Reservoir. A medium level priority area that covers the
pinelands of western Louisiana and eastern Texas surrounded these two areas.
Although I did not find many marked woodcock within this area, habitat conditions
are similar to those of the wintering high priority areas.
93
Expansive areas of no potential habitat should also be considered as
management priority areas, especially those that lie along possible migration routes.
Two of these areas were lands adjacent to the Mississippi River in Iowa and Illinois
and parts of the MAV in southeastern Missouri and northeastern Arkansas.
My observations of long stopover durations suggest that woodcock
populations would greatly benefit from habitat acquisition and proper management
within priority areas and along migration routes. Further research is needed to
develop best management practices for woodcock during fall migration in the Central
Region. Managers should also consider further investigations of woodcock wintering
distribution and the importance of southern pineland habitats to migrating and
wintering woodcock.
LITERATURE CITED
Anderson, M., P. Bourgeron, M. T. Bryer, R. Crawford, L. Engelking, D. Faber- Langendoen, M. Gallyoun, K. Goodin, D. H. Grossman, S. Landaal, K. Metzler, K. D. Patterson, M. Pyne, M. Reid, L. Sneddon, and A. S. Weakley. 1998. International Classification of Ecological Communities: Terrestrial Vegetation of the Unites States. Volume II. The National Vegetation Classification System: List of Types. The Nature Conservancy, Arlington, Virginia, USA.
Alerstam, T., and A. Lindstrom. 1990. Optimal bird migration: the relative
importance of time, energy, and safety. Pages 331-351 in E. Gwinner, editor. Bird migration: physiology and ecophysiology. Springer-Verlag, Berlin, Germany.
Boggus, T. G., and R. M. Whiting, Jr. 1982. Effects of habitat variables on foraging
of American woodcock wintering in East Texas. U.S. Fish and Wildlife Service Wildlife Research Report 14:148-153
Bourgeois, A. 1977. Quantitative analysis of American woodcock nest and brood
habitat. Proceedings of the Woodcock Symposium 6:108-118. Britt, T. L. 1971. Studies of woodcock on the Louisiana wintering ground. Thesis,
Louisiana State University, Baton Rouge, Louisiana, USA.
94
Cade, B. S. 1985. Habitat suitability index models: American woodcock (wintering).
Biological Report 82. U. S. Fish and Wildlife Service, Washington, D.C., USA.
Coon, R. A., P. D. Caldwell, and G. L. Storm. 1976. Some characteristics of fall
migration of female woodcock. Journal of Wildlife Management 40:91-95. ______, T. J. Dwyer, and J. W. Artmann. 1977. Identification of potential
harvest units in the United States for the American woodcock. Proceedings of the Woodcock Symposium 6:147-153
Duncan, P. S. 2000. American woodcock management, past, present, and future.
Pages 1-2 in D. G. McAuley, J. G. Bruggink, and G. F. Sepik, editors. Proceedings of the Ninth American Woodcock Symposium. U.S. Geological Survey, Biological Resources Division Information and Technology Report USGS/BRD/ITR-2000-0009, Patuxent Wildlife Research Center, Laurel, Maryland, USA.
Dyer, J. M., and R. B. Hamilton. 1977. Analyses of several site components of
diurnal woodcock habitat in southern Louisiana. Proceedings of the Woodcock Symposium 6:51-62.
Farmer, A. H. and A. H. Parent. 1997. Effects of the landscape on shorebird
movements at spring migration stopovers. The Condor 99:698-707. Gilmer, D. S., L. M. Cowardin, R. L. Duval, L. M. Mechlin, and C. W. Shaiffer.
1981. Procedures for the use of aircraft in wildlife biotelemetry studies. Resource Publication 140, U. S. Fish and Wildlife Service, Washington, D.C., USA.
Glasgow, L. L. 1958. Contributions to the knowledge of the ecology of the American
Woodcock, Philohela minor (Gmelin), on the wintering range in Louisiana. Dissertation, Texas A & M College, College Station, Texas, USA.
Godfrey, G. A. 1974. Behavior and ecology of American woodcock on the breeding
range in Minnesota. Dissertation, University of Minnesota, Minneapolis, Minnesota, USA.
Gregg, L. 1984. Population ecology of woodcock in Wisconsin. Wisconsin Department of Natural Resources. Technical Bulletin 144. Guild, W. F. M. L. 1948. The effects of soil type on the structure of earthworm
populations. Annals of Applied Biology 35:181-192. Gunmundsson, G. A., A. Lindstrom, and T. Alerstam. 1991. Optimal fat loads and
95
long-distance flights by migrating knots Calidris cantus, sanderlings, Calidris alba, and turnstones Arenaria interores. Ibis 144:140-152.
Gunn, A. 1992. The use of mustard to estimate earthworm populations.
Pedobiologia 36:65-67. Gutzwiller, K. J., K. R. Kinsley, G. L. Storm, W. M. Tzilkowski, and J. S. Wakeley.
1983. Relative value of vegetation structure and species composition for identifying American woodcock breeding habitat. Journal of Wildlife Management 47:535-540.
Kaiser, A. 1999. Stopover strategies in birds: a review of methods for estimating stopover length. Bird Study 46:S299-S308. Keppie, D. M. and R. M. Whiting, Jr. 1994. American woodcock (Scolopax minor).
The birds of North America, no. 100. The American Ornithologists’ Union, Washington, D.C., USA, and The Academy of Natural Sciences, Philadelphia, Pennsylvania, USA.
Krementz, D. G. and G. W. Pendleton. 1994. Diurnal habitat use of American
woodcock wintering along the Atlantic coast. Canadian Journal of Zoology 72:1945-1950.
_____, J. T. Seginak, and G. W. Pendleton. 1995. Habitat use at night by
wintering American woodcock in coastal Georgia and Virginia. Wilson Bulletin 107:686-697
_____. 2000. Habitat management for wintering American woodcock in the southeastern United States. Proceedings of the Woodcock Symposium 9:50-
54. Krohn, W. B., J. C. Rieffenberger, and F. Ferrigno. 1977. Fall migration of
woodcock at Cape May, New Jersey. Journal of Wildlife Management 41:104-111.
Lawrence, A. P., and M. A. Bowers. 2002. A test of the hot mustard extraction
method of sampling earthworms. Soil Biology and Biochemistry 34:549-552. Lehnen, S. E. 2004. Turnover rates of fall migration pectoral and least sandpipers
throughout the Lower Mississippi Alluvial Valley. Thesis, University of Arkansas, Fayetteville, Arkansas, USA.
Lemmon, P. E. 1957. A new instrument for measuring forest overstory density.
Journal of Forestry 55:2. MacDonald, P. O., W. E. Frayer, and J. K. Clauser. 1979. Documentation,
96
chronology, and future projections of bottomland hardwood habitat loss in the lower Mississippi Alluvial Plain. Volume 1, Basic Report. HRB-Singer, Incorporated, State College, Pennsylvania, USA.
Martin, F. W. 1964. Woodcock age and sex determination from wings. Journal of
Wildlife Management 28:287-293. McAuley, D. G., J. R. Longcore, and G. F. Sepik. 1993. Techniques for research into
woodcock: experiences and recommendations. Pages 5 –11 in J. R. Longcore and G. F. Sepik, editors. Eighth Woodcock Symposium. U.S. Fish and Wildlife Service, Wildlife Resource Report.
______, J. R. Goldsberry, J. R. Longcore. 1993. Omnidirectional aircraft antennas
for aerial telemetry. Wildlife Society Bulletin 21:487-491. Murphy, D. W. 1983. Ecology of American woodcock in central Missouri. Thesis, University of Missouri, Columbia, Missouri, USA. National Oceanic and Atmospheric Administration. 2004. Drought Information
Center. Washington, D. C., USA. Newling, C. J. 1990. Restoration of bottomland hardwood forests in the Lower
Mississippi Valley. Restoration and Management 8:23-28. Owen, R. B., Jr. 1977. American woodcock. Pages 149-186 in G. C. Sanderson,
editor. Management of migratory shore and upland game birds in North America. International Association of Fish and Wildlife Agencies, Washington, D.C., USA.
Piersma, T. 1987. Hop, skip, or jump? Constraints on migration of arctic waders by feeding, fattening, and flight speed. Limosa 60:185-194. Poier, K. R., and J. Richter. 1992. Spatial distribution of earthworms and soil
properties in an arable loess soil. Soil Biology and Biochemistry 24:1601-1608.
Rieffenberger, J. C., and R. C. Kletzly. 1967. Woodcock night-lighting techniques
and equipment. U.S. Sport Fisheries Bureau and Wildlife Special, Scientific Report- Wildlife 101:33-35.
Roberts, T. H. 1978. Migration, distribution, and breeding of American woodcock. Tennessee Wildlife Resources Agency, Technical report 79. _____. 1993. The ecology and management of wintering woodcocks. Biological Report 16:87-97.
97
Root, T. 1988. Atlas of wintering North American birds. An analysis of Christmas Bird Count data. University of Chicago Press, Chicago, Illinois, USA.
Sepik, G. F., and E. L. Derleth. 1993. Premigratory dispersal and fall migration of American woodcock in Maine. Biological Report 16:36-40. Sheldon, W. G. 1967. The book of the American woodcock. University of Massachusetts Press, Amherst, Massachusetts, USA. Skagen, S. K. 1997. Stopover ecology of transitory populations: The case of
migrant shorebirds. Ecological Studies 125:244-269. Straw, J. A., Jr., D. G. Krementz, M. W. Olinde, and G. F. Sepik. 1994. American
woodcock. Pages 96-116 in T. C. Tacha and C. E. Braun, editors. Migratory shore and upland game bird management in North America. International Association of Fish and Wildlife Agencies, Washington, D.C., USA.
Tinner, R. W. 1999. Wetland indicators: A guide to wetland identification,
delineation, classification and mapping. Lewis Publishers, New York, USA. U.S. Geological Survey. 2003. National Land Cover Data 1992.
http://landcover.usgs.gov/natllandcover.asp. U.S. Department of Interior, U.S. Geological Survey, Washington, D.C., USA.
98
Figure 1. Woodcock management regions determined by Coon et al. (1977) using band recovery data.
99
Figure 2. Possible American woodcock fall migration route proposed by Glasgow (1958) and Sheldon (1967) using band return data.
100
Figure 3. Location of study areas where woodcock were radio-marked by the University of Minnesota, University of Wisconsin, and Northern Michigan State University from 2001-2003.
101
Figure 4. Flight paths of 3 radio-telemetry searches flown in Fall of 2001 for woodcock radio-marked in Minnesota. See Table 3 for corresponding flight dates and durations. All flights originated in Bald Knob, AR.
102
Figure 5. Flight paths of 16 radio-telemetry searches flown in Fall of 2002 for woodcock radio-marked in Michigan, Minnesota, and Wisconsin. See Table 2 for corresponding flight dates and durations. All flights originated in Bald Knob, AR.
103
Figure 6. Flight paths of 10 radio-telemetry searches flown in Fall of 2003 for woodcock radio-marked in Michigan, Minnesota, and Wisconsin. See Table 2 for corresponding flight dates and durations. Flights 1 and 2 originated in Ames, IA, all other flights originated in Bald Knob, AR.
104
Figure 7. Location and origin of all radio-marked woodcock locations from 2001-2003. The origin of some radio-marked birds was unknown due to multiple birds being marked with radio transmitters ≤0.007 MHz apart. Confirmed locations are those where I was able to locate and flush the bird on the ground. Locations north of 33º N latitude were classified as migratory while those south of this line were considered to be on the winter grounds
105
0
1
2
3
4
5
6
7
1 2 3 4 5
Mid-story Density
Freq
uenc
y
0
1
2
3
4
5
6
7
8
9
10
1 2 3 4 5
Ground Density
Freq
uenc
y
Figure 8. Frequency of ground and mid-story vegetation horizontal density at confirmed migrating marked woodcock locations (n=18) in 2002 and 2003. Density was visually estimated on a scale of 1 (sparse) - 5 (dense).
106
0
1
2
3
4
5
6
7
8
9
10
0 - 20 20 - 40 40 - 60 60 - 80 80 - 100
Canopy (%)
Freq
uenc
y
Figure 9. Frequency of percent canopy cover at all confirmed migrating marked woodcock locations (n=18) in 2002 and 2003.
107
Figure 10. Possible woodcock fall migration routes used by woodcock radio-marked from 2001-2003 in Minnesota, Wisconsin, and Upper Peninsula Michigan. The route through northeast Arkansas and the route through southern Illinois are two routes that I suspected but only found minimal evidence.
108
Figure 11. Central Region potential woodcock habitat determined from 1992 National Land Cover Data (NLCD). Potential woodcock habitat includes NLCD classifications: woody wetlands, shrubland, deciduous forest, evergreen forest, mixed forest, orchard/vineyard/other, and transitional.
109
0
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
0.9
1
42.5
41.6
40.8
39.9
39.0
38.1
37.3
36.4
35.5
34.6
33.7
32.9
32.0
Latitude
Perc
enta
ge
Figure 12. Percent potential woodcock habitat availability along a 50-km buffered fall migration route (inset) that I observed radio-marked woodcock using from 2001 - 2003. Potential woodcock habitat included 1992 National Land Cover Data classifications: woody wetlands, shrubland, deciduous forest, evergreen forest, mixed forest, orchard/vineyard/other, and transitional.
110
0
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
0.9
1
42.5
41.6
40.8
39.9
39.0
38.1
37.3
36.4
35.5
34.6
33.8
32.9
32.0
Latitude
Perc
enta
ge
Figure 13. Percent potential woodcock habitat availability along a 50-km buffered fall migration route (inset) that I observed radio-marked birds using from 2001 to 2003. Potential woodcock habitat included 1992 National Land Cover Data classifications: woody wetlands, shrubland, deciduous forest, evergreen forest, mixed forest, orchard/vineyard/other, and transitional.
111
0
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
0.9
45.7
44.8
44.0
43.1
42.2
41.3
40.5
39.6
38.7
37.9
37.0
36.1
35.2
34.4
33.5
32.6
31.7
30.9
Latitude
Perc
enta
ge
Figure 14. Percent potential woodcock habitat availability along a 50-km buffered fall migration route (inset) directly from the Michigan field site to central Louisiana. Potential woodcock habitat included 1992 National Land Cover Data classifications: woody wetlands, shrubland, deciduous forest, evergreen forest, mixed forest, orchard/vineyard/other, and transitional.
112
0
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
0.9
1
45.7
44.8
43.9
43.0
42.2
41.3
40.4
39.5
38.6
37.7
36.8
35.9
35.1
34.2
33.3
32.4
31.5
Latitude
Perc
enta
ge
Figure 15. Percent potential woodcock habitat availability along a 50-km buffered fall migration route (inset) directly from the Minnesota field site to central Louisiana. Potential woodcock habitat included 1992 National Land Cover Data classifications: woody wetlands, shrubland, deciduous forest, evergreen forest, mixed forest, orchard/vineyard/other, and transitional.
113
0
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
0.9
1
45.4
44.5
43.6
42.7
41.9
41.0
40.1
39.2
38.3
37.4
36.5
35.6
34.7
33.9
33.0
32.1
31.2
Latitude
Perc
enta
ge
Figure 16. Percent potential woodcock habitat availability along a 50-km buffered fall migration route (inset) directly from the Wisconsin field site to central Louisiana. Potential woodcock habitat included 1992 National Land Cover Data classifications: woody wetlands, shrubland, deciduous forest, evergreen forest, mixed forest, orchard/vineyard/other, and transitional.
114
0
0.1
0.2
0.3
0.4
0.5
0.6
42.5
41.7
41.0
40.2
39.4
38.7
37.9
37.1
36.3
35.6
34.8
34.0
33.2
32.5
31.7
30.9
Latitude
Perc
enta
ge
Figure 17. Percent potential woodcock habitat availability along a 50-km buffered fall migration route (inset) following the Mississippi River to central Louisiana. Potential woodcock habitat included 1992 National Land Cover Data classifications: woody wetlands, shrubland, deciduous forest, evergreen forest, mixed forest, orchard/vineyard/other, and transitional.
115
Figure 18. Woodcock management priority areas in the Central Region that I designated based on locations of radio-marked woodcock. High priority areas were identified because of the high density of radio-marked woodcock locations from 2001-2003. Medium priority areas had fewer radio-marked woodcock locations but they contained similar habitat conditions as high priority areas, were funnel areas along migration routes, or were areas of suitable habitat after large expanses of limited potential habitat.
116
Table 1. 1992 National Land Cover Data land cover classifications used in mapping potential woodcock habitat in the Central Region. Potential habitats are those that possibly provide diurnal woodcock habitat.
Potential Habitat Cover-types
Shrubland
Woody Wetland
Deciduous Forest
Evergreen Forest
Mixed Forest
Orchards/Vineyards/Other
Transitional
No Potential Habitat Cover-types
Open Water
Perennial Ice/Snow
Bare Rock/Sand/Clay
Quarries/Strip Mines/Gravel Pits
Grassland/Herbaceous
Emergent Herbaceous Wetland
Low Intensity Residential
High Intensity Residential
117
Commercial/Industrial/Transportation
Pasture/Hay
Row Crops
Small Grain
Fallow
Urban/Recreational Grasses
118
Table 2. Sex and age of confirmed and unconfirmed radio-marked woodcock locations from 2001-2003. The sex and age of some birds was unknown due to multiple radio transmitters with frequencies ≤ .007 MHz apart. Age was recorded as hatch year (HY) or after hatching year (AHY).
Year Recovery Location Sex Age Confirmed
2001 Crossett, AR F HY Yes
Felsenthal, AR F HY No
Hope, AR M AHY Yes
Prescott, AR M HY Yes
2002 Adona, AR M AHY Yes
Atlanta, TX F HY Yes
Batchtown, IL M AHY Yes
Belfast, AR F Unknown Yes
Brookeland, TX M Unknown Yes
Burkeville, TX M HY No
Clayton, TX M HY Yes
Curtis, TX F HY No
De Ridder, LA F HY Yes
Fordyce, AR Unknown Unknown Yes
Fort Polk, LA F Unknown No
Fredricktown, MO M Unknown Yes
Grapevine, AR Unknown Unknown Yes
Grenada, MS F AHY Yes
Gunn, MS F Unknown Yes
Hannibal, MO Unknown Unknown No
Iberia, MO F AHY Yes
Jasper, TX F Unknown No
Lebanon, MO M HY Yes
119
Montrose, AR F HY No
Natchitoches, LA M HY Yes
Paragould, AR Unknown HY Yes
Ruston, LA F HY Yes
Sikes, LA Unknown Unknown Yes
Tarry, AR F HY Yes
Warren, AR F Unknown Yes
West, MS M Unknown Yes
Winchester, IL F Unknown No
Winnfield, LA M AHY Yes
2003 Alexandria, LA F AHY Yes
Columbia, MO F AHY No
Dennard, AR F AHY Yes
Jefferson City, MO Unknown AHY Yes
Pickneyville, IL Unknown Unknown Yes
Pollock, LA F AHY Yes
Rison, AR F AHY Yes
Star City, AR F Unknown Yes
Farber, MO F AHY No
120
Table 3. Date and duration of radio-telemetry flights flown from 2001-2003 searching for radio-marked woodcock. Flight numbers correspond to flight routes maps (Fig. 2-4).
Year Flight
Number Date Duration (hours)
2001 1 9 Nov 4.2
2 20 Nov 5.8
3 4 Dec 10
2002 1 1 Nov 7
2 2 Nov 6
3 7 Nov 6.7
4 8Nov 6.3
5 17 Nov 6.8
6 18 Nov 5
7 20 Nov 5.9
8 22 Nov 5.3
9 25 Nov 9.8
10 27 Nov 8.8
11 29 Nov 9.8
12 6 Dec 8.8
13 7 Dec 9
14 16 Dec 9.5
15 17 Dec 9.8
16 20 Dec 10.5
121
2003 1 1 Nov 6.5
2 7 Nov 7.2
3 10 Nov 8.6
4 19 Nov 6.4
5 20 Nov 7.1
6 21 Nov 8
7 26 Nov 6.9
8 6 Dec 8.7
9 7 Dec 6.7
10 9 Dec 8.9
122
Table 4. Straight-line migration distances (km) and flight duration of all confirmed, known origin radio-marked woodcock from 2001-2003. Estimated flight duration was the time between the date that the bird was last located on the breeding ground and the date that the bird was located during migration.
Nearest Town Origin Trip
Length (km)
Max. Flight Duration (days)
km/days
Batchtown, IL MN 784 21 37.33
Iberia, MO MI 978 16 61.13
Dennard, AR WI 1096 15 73.07
Prescott, AR MN 1346 48 28.04
Hope, AR MN 1368 34 40.24
Rison, AR MI 1410 20 70.50
Crossett, AR MN 1429 30 47.63
Atlanta, TX WI 1406 16 87.88
Clayton, TX WI 1528 30 50.93
Natchitoches, LA WI 1536 41 37.46
Alexandria, LA WI 1554 32 48.56
Pollock, LA MN 1607 22 73.05
De Ridder, LA MN 1693 39 43.41
123
Table 5. Stopover durations of radio-marked woodcock located during migration from 2001-2003. Stopover data were recorded only for birds that we were able to monitor for ≥1 day after locating them.
Recovery Site Date LocatedStopover Duration
Montrose, AR* 6 Dec, 2002 1
Prescott, AR 20 Nov, 2001 1
Star City, AR 20 Nov, 2003 1
Winchester, IL* 1 Nov, 2002 1
Grenada, MS 29 Nov, 2002 >1
Paragould, AR 17 Nov, 2002 >1
Adona, AR 20 Nov, 2002 1 - 2
Grapevine, AR 22 Nov, 2002 1 - 3
Hannibal, MO 17 Nov, 2002 2
Crossett, AR 1 Dec, 2001 >2
Hope, AR 4 Dec, 2001 >2
Fredericktown, MO 8 Nov, 2002 >3
West, MS 29 Nov, 2002 >3
Iberia, MO 7 Nov, 2002 4
Batchtown, IL 1 Nov, 2002 >4
Jefferson City, MO 7 Nov, 2003 >4
Pinckneyville, IL 10 Nov, 2003 >4
124
Pollock, AR 7 Dec, 2003 >4
Tarry, AR 28 Nov, 2002 4 - 6
Lebanon, MO 7 Nov, 2002 5
Fordyce, AR 27 Nov, 2002 5 - 7
Belfast, AR 26 Nov, 2002 >8
Rison, AR 20 Nov, 2003 6 - 16
Dennard, AR 19 Nov, 2003 10 - 14
* Radio-marked woodcock locations where the radio signal was not confirmed by flushing the woodcock due to the bird migrating between the time the pilot located the signal and when I arrived at the location.
125
Table 6. National Vegetation Classification System (NVCS) forest alliances at all confirmed marked woodcock locations from 2001-2003. Migration locations were divided between the oak-hickory dominated areas in and north of the Ozark Mountains (35º N) and predominately pine areas south of the Ozark Mountains, while wintering locations were south of 33º N latitude.
Migration locations in or north of the Ozark Mountains Frequency Eastern Red Cedar (Juniperus virginiana)- Quercus Forest Alliance 2
Post Oak (Quercus stellata)- Blackjack Oak (Quercus marilandica) Forest Alliance 2
Regenerating Old Field- Sumac¹ (Rhus spp.) 1
White Oak (Quercus alba) Forest Alliance 1
Overcup Oak (Quercus lyrata) Seasonally Flooded Forest Alliance 1
Pin Oak (Quercus palustris) Seasonally Flooded Forest Alliance 1
Migration Locations south of the Ozark Mountains Loblolly Pine (Pinus taeda)- Shortleaf Pine (Pinus echinata) Forest Alliance 8
Quercus Temporary Flooded Forest Alliance 2
Loblolly Pine (Pinus taeda)- Quercus Forest Alliance 2
Sweetgum (Liquidambar spp.) Temporary Flooded Forest Alliance 1
Loblolly Pine (Pinus taeda)- Temporary Flooded Forest Alliance 1
Winter Locations Loblolly Pine (Pinus taeda)- Shortleaf Pine (Pinus echinata) Forest Alliance 5
Loblolly Pine (Pinus taeda)- Quercus Forest Alliance 1
Old Field- Chinese Privet² (Lingustrum sinense) 1
Southern Red Oak (Quercus falcata) Forest Alliance 1
Quercus Temporary Flooded Forest Alliance 1
Sweetgum (Liquidambar spp.) Temporary Flooded Forest Alliance 1
Two locations did not fit into a NVCS vegetation class: ¹One radio-marked bird located in west-central Illinois was located in an old field that had regenerated into sumac (Rhus spp.). ²One radio-marked bird located in north-east Texas was located in an old field that had regenerated in a dense stand of Chinese Privet (Lingustrum sinense). Table 7. Dominant ground and mid-story vegetation species at migrating confirmed marked woodcock locations north of 33º N latitude from 2001-2003.
126
Table 7. Dominant ground and mid-story vegetation species at confirmed marked woodcock locations north of 33º N latitude from 2001-2003.
Ground Species Frequency Herbaceous spp. (non-woody vegetation) 10
Blackberry (Rubus spp.) 5
Greenbrier (Smilax spp.) 4
Japanese Honeysuckle (Lonicera japonica) 3
Coral berry (Symphoricarpos orbiculatus) 2
Oak (Quercus spp.) 2
Rose (Rosa spp.) 2
American Holly (Ilex opaca) 1
Chinese Privet (Ligustrum sinense) 1
Loblolly Pine (Pinus taeda) 1
Mid-story Species Frequency Blackberry (Rubus spp.) 7
Loblolly Pine (Pinus taeda) 6
Greenbrier (Smilax spp.) 4
Herbaceous spp. (non-woody vegetation) 4
Oak (Quercus spp.) 4
Elm (Ulmus spp.) 3
Sumac (Rhus spp.) 3
Chinese Privet (Ligustrum sinense) 2
Deciduous Holly (Ilex decidua) 2
Japanese Honeysuckle (Lonicera japonica) 2
127
Rose (Rosa spp.) 2
Sweetgum (Liquidambar styraciflua) 2
American Holly (Ilex opaca) 1
Autumn Olive (Elaegnus umbellata) 1
Eastern Redcedar (Juniperus virginiana) 1
Persimmon (Diospyros virginianas) 1
Red Maple (Acer rubrum) 1
Sweetbay (Ilex spp.) 1
Switchcane (Arundinaria gigantea) 1
Blueberry (Viburnum spp.) 1
128
Table 8. Dominant ground and mid-story vegetation species at confirmed marked woodcock locations south of 33º N latitude from 2001-2003.
Mid-story Species Frequency Oak (Quercus spp.) 6
Blackberry (Rubus spp.) 4
Greenbrier (Smilax spp.) 4
Chinese Privet (Ligustrum sinense) 3
Elderberry (Sambucus spp.) 3
Japanese Honeysuckle (Lonicera japonica) 2
Loblolly Pine (Pinus taeda) 2
Trumpet Creeper (Campsis radicans) 2
Herbaceous spp. 1
Sweetgum (Liquidambar styraciflua) 1
Longleaf Pine (Pinus palustris) 1
Grapevine (Vitis spp.) 1
Eastern Baccharis (Baccharis halimifolia) 1
Red Maple (Acer rubrum) 1
Ground Species Frequency Blackberry (Rubus spp.) 6
Japanese Honeysuckle (Lonicera japonica) 4
Chinese Privet (Ligustrum sinense) 2
Greenbrier (Smilax spp.) 2
Herbaceous spp. 2
129
130
Loblolly Pine (Pinus taeda) 1
Oak (Quercus spp.) 1
Elderberry (Sambucus spp.) 1