Research Article

Distribution and Migration Chronology ofEastern Population Sandhill Cranes

DAVID L FRONCZAK1 United States Fish and Wildlife Service 5600 American Boulevard West Suite 960 Bloomington MN 55437 USA

DAVID E ANDERSEN United States Geological Survey Minnesota Cooperative Fish and Wildlife Research Unit 200 Hodson Hall 1980 FolwellAvenue St Paul MN 55108 USA

EVERETT E HANNA Department of Biology Western University 1151 Richmond Street London ON N6A 3K7 Canada

THOMAS R COOPER United States Fish and Wildlife Service 5600 American Boulevard West Suite 960 Bloomington MN 55437 USA

ABSTRACT The Eastern Population (EP) of greater sandhill cranes (Antigone canadensis tabida cranes) isexpanding in size and geographic range Little information exists regarding the geographic extent ofbreeding migration and wintering ranges migration chronology or use of staging areas for cranes in the EPTo obtain these data we attached solar global positioning system (GPS) platform transmitting terminals(PTTs) to 42 sandhill cranes and monitored daily locations from December 2009 through August 2014 Onaverage tagged cranes settled in summer areas during late-March in Minnesota (7) Wisconsin (29)Michigan USA (21) and Ontario Canada (38) and arrived at their winter terminus beginning mid-December in Indiana (15) Kentucky (3) Tennessee (45) Georgia (5) and Florida (32) Cranesinitiated springmigration beginningmid-February to their respective summer areas on routes similar to thoseused during fall migration Twenty-five marked cranes returned to the same summer area after a secondspring migration of which 19 (76) settledlt3 km from the estimated mean center of the summer area of theprevious year During the 2010ndash2012 United States Fish and Wildlife Service (USFWS) Cooperative FallAbundance Survey for cranes in the EP we estimated that approximately 29ndash31 of cranes that summered inboth Wisconsin and the Lower Peninsula of Michigan were not in areas included in the survey Theinformation we collected on crane movements provides insight into distribution and migration chronologythat will aid in assessment of the current USFWS fall survey In addition information on specific use sites canassist state and federal managers to identify and protect key staging and winter areas particularly duringcurrent and future recreational harvest seasons 2017 The Wildlife Society

KEY WORDS Antigone canadensis tabida distribution Eastern Population migration chronology sandhill cranesatellite telemetry

At the turn of the 20th century the Eastern Population (EP)of greater sandhill cranes (Antigone canadensis tabida cranes)was nearly extirpated from its historical breeding rangebecause of habitat alteration and unregulated hunting(Walkinshaw and Wing 1955 Lumsden 1971 Hunt et al1976) Following protection from unregulated hunting afterpassage of the Migratory Bird Treaty Act of 1916 (ieclosure of hunting in 1918) and in response to conservationefforts by state and federal agencies and non-governmentorganizations cranes in the EP increased in number andexpanded their breeding range (Tacha et al 1994Amundson and Johnson 2011) However an increase inpopulation size has also been accompanied by managementissues related to crop depredation public pressure to allowrecreational harvest and enhancing non-consumptive wild-life opportunities (Van Horn et al 2010) Similar to other

crane populations basic biological and annual life cycleinformation are needed to better manage cranes in the EPespecially information relating to spatial distribution of thepopulation current migration patterns potential overlapwith neighboring migratory and non-migratory populationsand identification of important use areas throughout theirannual life cycle (Case and Sanders 2009)In 2009 theMigratory Shore and Upland Game Bird Task

Force under the direction of the Association of Fish andWildlife Agencies identified priority information needs thatwere specific to developing a better monitoring program toreflect current distributions and migration patterns of cranesin the EP This effort identified documenting the geographicextent of breeding migration and wintering ranges asessential information needs so that appropriate changes tothe spatial-temporal design of the current United States FishandWildlife Service (USFWS) Cooperative Fall AbundanceSurvey (hereafter USFWS fall survey) could be imple-mented (Case and Sanders 2009) The USFWS fall survey isa long-term survey established in 1979 that consists of effortsby volunteers and state and federal agencies from the Atlantic

Received 15 April 2016 Accepted 10 March 2017

1E-mail dave_fronczakfwsgov

The Journal of Wildlife Management DOI 101002jwmg21272

Fronczak et al Eastern Population Sandhill Crane Migration 1

and Mississippi flyways (ie MN WI MI IN OH PATN GA and FL) The main goal of the survey is to providean estimate of the size and trend of the cranes in the EP andis focused on counting cranes that concentrate in IndianaMichigan and Wisconsin during fall migration The surveywas initially designed to begin the last week of October whencranes are concentrated in these states (Van Horn et al2010) The initial 1979 survey counted 14385 cranes and hasincreased to 83479 in 2014 with a recent 3-year average of78532 cranes for 2012ndash2014 (Kruse and Dubovsky 2015)To address crane priority information needs we combined

and analyzed data from 2 concurrent ecology studies forcranes in the EP One study focused on crane ecology onManitoulin Island Ontario (Hanna et al 2014) and theother focused on crane migration ecology (Fronczak 2014)both studies used global positioning system (GPS) satelliteplatform transmitting terminals (PTTs) Specifically wemonitored cranes from 2009 to 2014 and identified currentcrane staging areas travel routes and migration chronologyidentified current summer and wintering areas and durationsof stay and determined the proportion of cranes thatoccurred within the areas surveyed during the USFWS fallsurvey

STUDY AREA

Our study area included the breeding and wintering groundsand migration corridors of cranes in the EP as determined byGPS locations of PTT-marked cranes and consisted of statesand provinces within the Canadian Shield Great LakesMississippi Valley and Southeast geographic regions ofNorth America (Fig 1) These areas include borealsoftwoods prairie hardwoods eastern tall grass prairiecentral hardwoods and southeastern coastal plains Wecaptured and marked cranes throughout spring and fallmigration and winter at 7 sites within the known EP range(Tacha et al 1994) during 2009ndash2012We conducted the majority of our trapping at the Jasper-

Pulaski Fish and Wildlife Area (FWA) northwest Indiana(4128N 8698W) and the Hiwassee Wildlife Refuge(WR) southeast Tennessee (3548N 8508W) both areconsidered major staging areas for cranes in the EP(Fronczak 2014) In addition to these 3 primary sites wealso trapped and placed satellite transmitters on cranes atGoose Pond Fish and Wildlife Area Indiana (3908N8728W) Sherburne National Wildlife Refuge Minnesota(4558N 9388W) Crex Meadows Wildlife Area Wis-consin (4588N 9268W) and Hop-In Wildlife RefugeTennessee (3338N 8908W) We also captured andmarked cranes in the EP at a Canadian staging site atManitoulin Island Ontario Canada (4588N 8228W)Detailed area descriptions of trap locations have beendescribed previously (Fronczak 2014 Hanna et al 2014)

METHODS

TrappingCranes in the EP occur across broad geographic areasduring summer and winter and exhibit variation in timing of

migration across their range To mark a representativesample of cranes in the EP we used a strategy similar to thatused by Krapu et al (2011) in a comparable study ofMid-continent Population (MCP) cranes We identifiedJasper-Pulaski FWA and Hiwassee WR as the major stagingareas used during migration by cranes in the EP based onprevious studies that described migratory routes (Walkin-shaw 1960 Crete and Toepfer 1978) EP abundance surveys(eg Mississippi Flyway Cooperative Midwinter Survey[MWS] state surveys) and the subsequent behavior ofcranes we marked in this study For each stopover area weselected trapping periods before during and after estimatedpeak abundance based on periodic surveys of cranes duringfall migration (J Bergens Indiana Department of NaturalResources unpublished data and R Klippel TennesseeWildlife Resource Agency unpublished data) We employedthis capture protocol to mark a geographic representativesample of cranes and to capture the temporal variation incrane movements through these stopover areas In additionwe captured cranes on alternative breeding and staging areasincludingManitoulin Island Ontario (Hanna et al 2014) toinclude cranes from portions of their distribution that maynot have been represented by cranes we trapped at majorstaging areasWe used a rocket-propelled net assembly as the primary

method to capture cranes We identified potential trappingsites using the protocol developed for rocket netting cranes intheMCP by Krapu et al (2011) Their protocol gave priorityto daytime loafing sites withgt20 cranes present in pasture orother open land-cover types and incorporated decoys Wemodified their protocol by using bait instead of decoysWhen cranes responded to bait for 2 consecutive days weassembled a 131 197-m rocket-propelled net (Wheelerand Lewis 1972 D A Brandt US Geological Survey[USGS] Northern Prairie Wildlife Research Centerpersonal communication) at a potential capture locationWe conducted trapping primarily in the morning becausecranes consistently returned to baited sites after leavingnocturnal roosts In addition to using a rocket-propelled netwe used a Coda NetLauncher (Coda Enterprises Mesa AZUSA) to capture cranes in locations where using a rocket-propelled net was not feasible We followed the standardCoda NetLauncher protocol developed for cranes by theOhio Department of Natural Resources (D E ShermanOhio Department of Natural Resources personal commu-nication) We also used modified softcatch leghold traps(Victor no 3 Oneida Victor Euclid OH USA) as describedby King et al (1998)We targeted adult female cranes because they are more

likely than males to return to natal breeding grounds thefollowing spring (Walkinshaw 1949 Drewien 1973)Sandhill cranes are perennially monogamous (Tacha et al1994) will re-mate after death or permanent separation(Nesbitt 1989 Hayes and Barzen 2006) and pair bonds willremain after departure from summer areas (Tacha et al 1992Hayes 2015) We assume little difference in migrationbehavior among cohorts in our sample because we targetedadult cranes However we acknowledge that we did not

2 The Journal of Wildlife Management 9999()

know the breeding status of all of our marked cranes Weidentified adult female cranes based on the physical and socialcharacteristics described by Tacha (1998) as smaller body-sized cranes with red skin on the crown of the head morelikely to be viewed following larger-bodied male cranes andless likely to display agonistic behaviors However followingcapture if we could not identify family groups we selected asmaller-bodied adult crane that we presumed to be a femaleWe affixed a 30-g 3-solar-paneled GPS satellite PTT

(North Star Science and Technology Baltimore MD USA)to captured cranes via flanged leg bands We selected tarsus-mounted PTTs because similar transmitters and attachmentmethods have been used extensively on cranes withoutapparent negative effects on survival or movements incomparison to backpack mounted transmitters (Melvin et al

1983 Ellis et al 2001 Krapu et al 2011 Hayes 2015) Wemounted PTTs on a 76-cm 2-piece color-coated polyvinylchloride (PVC) alpha-numeric-coded engraved leg band(Haggie Engraving Crumpton MD USA) Satellite PTTsincorporated 2 additional solar panels to increase thelongevity of the transmitter compared with standardbattery-powered or single solar-paneled transmitters Wereinforced the connection between the antenna and thetransmitter housing as a precaution because it is a weak pointand a cause for loss of transmission (D A Brandt personalcommunication) For cranes not marked with PTTswe affixed a 76-cm 15-wrapped alpha-numeric-codedengraved PVC tarsus auxiliary leg band above the distaltibio-tarsus joint Flanged auxiliary markers and PTTstogether weighed approximately 80 g (2 of average body

Figure 1 Trapping locations by season where sandhill cranes were captured and marked with platform transmitting terminals Gray shading denotes thegeographic extent of movements of cranes during the annual cycle 2009ndash2014

Fronczak et al Eastern Population Sandhill Crane Migration 3

mass at capture) and were attached above the distal tibio-tarsus joint (Krapu et al 2011) All birds marked received astandard USGS aluminum bandFor each PTT-marked crane we drew blood from the vein

below the tibio-tarsus and placed samples in a lysis bufferanticoagulant solution (Jones et al 2005) Blood wassubsequently analyzed by Avian Biotech International(Tallahassee FL USA) to determine sex We did notcollect blood samples from cranes at Manitoulin IslandWhen we captured gt2 cranescrew member at one time wereleased cranes in excess of that number immediately withouttaking measurements or affixing markers to avoid detrimen-tal effects (eg myopathy) of extended handling times Wereleased all cranes held for processing as a group within30minutes of being captured We captured cranes underapproval of the University ofMinnesota Institutional AnimalCare and Use Committee (protocol no 1103A97333) andthe University of Western Ontario Animal Use Protocol(protocol no 2010ndash213)We programmed most PTTs on 5- and 6-hour intervals to

maximize the number of GPS locations per day whichafforded the ability to determine the status (ie sedentarylocations or continuous movement) of marked cranes Weprogrammed other PTTs to collect GPS locations every 6hours during migration and every 8 hours during non-migration periods in the summer and winter Spatial locationdata were transmitted to satellites each 72-hour duty cyclethrough standard Doppler technology Accuracy for GPSlocations was classified into 4 quality levelslt26m 26ndash50m51ndash75m and 76ndash100m by the manufacturerrsquos standards(K LeSage GeoTrak personal communication)

Data AnalysisWe used GPS locations for cranes in the EP that completedfull annual cycles beginning from the date of capture todescribe distribution and migration chronology For cranesthat did not complete an annual cycle because of either signalloss or death we used GPS data for any completed migrationor duration of stay at a winter or summer terminus Whennecessary we incorporated Argos Doppler system locationsas an alternative source of information to describe migrationroutes when GPS data locations were gt5 days betweensignals because of environmental conditions that resulted ininsufficient battery voltage and disrupted data transmissionto satellitesWe retrieved data transmitted by PTTs from Collecte

Localisation Satellites (CLS)-America (httpwwwargos-systemorg Largo MD USA) which we decoded into ageographic information system shapefile format throughsatellite data decoder software (DSDCODE version 4dNorth Star Science and Technologies) We filtered GPSlocations using an adapted location filtering system based onthe Douglas Argos-Filter Algorithm version 65 (Douglaset al 2012 Pearse et al 2015) We remained consistentwith Krapu et al (2011) and assigned attributes to GPSlocation records identifying 4 periods of the annual cycle(ie spring migration summer area fall migration winterarea) and classified summer and winter area locations as the

terminus of migration based on a major departure from theprevious location We assigned fall and spring migrationattributes to areas that we defined as a departuregt15 km forgt5 days from the summer or winter areas We definedstaging areas as locations where a crane stayed at a singlegeographic area for gt5 consecutive days during migrationWe considered areas where cranes stayed for a shorterduration (lt5 days) during migration as stopover sites(Warnock 2010)We estimated the proportion of marked cranes available for

detection during the USFWS fall surveys by spatiallymatching survey locations to GPS locations of marked cranesduring the same periods Fall survey location descriptionsranged from general names of townships nearest towns ornatural areas to precise GPS coordinates We determinedthat cranes were available during the fall survey if their GPScoordinates were located within the area described by theobserver or if the GPS locations of cranes were lt3 km froman observerrsquos GPS locationTo assess philopatry to summer area we compared centroids

of locations during the period PTT-marked cranes occupiedtheir summer range for consecutive summers We calculatedcentroids of locations for each summer area with ArcGIS(version92ESRIRedlandsCAUSA)Wesummarized thelinear distance between these centroids between or amongyears for individual cranes and report an overall mean

RESULTS

From 2009 through 2012 we captured 202 cranes 194 usingrocket-propelled nets 7 with a Coda NetLauncher and 1with a softcatch leghold trap We deployed 38 PTTs on 42cranes (ie we recovered 4 PTTs and deployed them on asecond crane) Of the 42 cranes 25 (60) were female 6(14) were male and 11 (26) were undetermined sexbecause we did not draw a blood sample or the sexing analysishad inconclusive results Seven cranes (17) died and 24(57) of the PTTs we deployed functionedgt24monthsWemonitored cranes between December 2009 and August 2014which generated 103882 GPS locations over 28677tracking days The average number of GPS locationsdaywas 34 (rangefrac14 17ndash40) where 55 and 32 of the locationquality levels were registered as lt26m and 26ndash50mrespectively

Summer AreasCrane summer areas were dispersed across 3 states and 1province of the Great Lakes region 3 (7) settled inMinnesota 12 (29) settled inWisconsin 9 (21) settled inMichigan and 16 (38) settled in Ontario (Fig 2A) Nine(21) cranes that were marked on Manitoulin IslandOntario remained on Manitoulin Island for the duration ofthe summer (Hanna et al 2014) Cranes spent summers ineast-central Minnesota (Benton and Sherburne counties)throughout Wisconsin (Brown Crawford Fond du LacGreen Lake Jefferson Lincoln Outagamie RacineWashburn Waupaca Waukesha andWinnebago counties)on the Upper Peninsula (Chippewa and Mackinac counties)and the Lower Peninsula (Ingham Jackson Kent and

4 The Journal of Wildlife Management 9999()

and Mississippi flyways (ie MN WI MI IN OH PATN GA and FL) The main goal of the survey is to providean estimate of the size and trend of the cranes in the EP andis focused on counting cranes that concentrate in IndianaMichigan and Wisconsin during fall migration The surveywas initially designed to begin the last week of October whencranes are concentrated in these states (Van Horn et al2010) The initial 1979 survey counted 14385 cranes and hasincreased to 83479 in 2014 with a recent 3-year average of78532 cranes for 2012ndash2014 (Kruse and Dubovsky 2015)To address crane priority information needs we combined

and analyzed data from 2 concurrent ecology studies forcranes in the EP One study focused on crane ecology onManitoulin Island Ontario (Hanna et al 2014) and theother focused on crane migration ecology (Fronczak 2014)both studies used global positioning system (GPS) satelliteplatform transmitting terminals (PTTs) Specifically wemonitored cranes from 2009 to 2014 and identified currentcrane staging areas travel routes and migration chronologyidentified current summer and wintering areas and durationsof stay and determined the proportion of cranes thatoccurred within the areas surveyed during the USFWS fallsurvey

STUDY AREA

Our study area included the breeding and wintering groundsand migration corridors of cranes in the EP as determined byGPS locations of PTT-marked cranes and consisted of statesand provinces within the Canadian Shield Great LakesMississippi Valley and Southeast geographic regions ofNorth America (Fig 1) These areas include borealsoftwoods prairie hardwoods eastern tall grass prairiecentral hardwoods and southeastern coastal plains Wecaptured and marked cranes throughout spring and fallmigration and winter at 7 sites within the known EP range(Tacha et al 1994) during 2009ndash2012We conducted the majority of our trapping at the Jasper-

Pulaski Fish and Wildlife Area (FWA) northwest Indiana(4128N 8698W) and the Hiwassee Wildlife Refuge(WR) southeast Tennessee (3548N 8508W) both areconsidered major staging areas for cranes in the EP(Fronczak 2014) In addition to these 3 primary sites wealso trapped and placed satellite transmitters on cranes atGoose Pond Fish and Wildlife Area Indiana (3908N8728W) Sherburne National Wildlife Refuge Minnesota(4558N 9388W) Crex Meadows Wildlife Area Wis-consin (4588N 9268W) and Hop-In Wildlife RefugeTennessee (3338N 8908W) We also captured andmarked cranes in the EP at a Canadian staging site atManitoulin Island Ontario Canada (4588N 8228W)Detailed area descriptions of trap locations have beendescribed previously (Fronczak 2014 Hanna et al 2014)

METHODS

TrappingCranes in the EP occur across broad geographic areasduring summer and winter and exhibit variation in timing of

migration across their range To mark a representativesample of cranes in the EP we used a strategy similar to thatused by Krapu et al (2011) in a comparable study ofMid-continent Population (MCP) cranes We identifiedJasper-Pulaski FWA and Hiwassee WR as the major stagingareas used during migration by cranes in the EP based onprevious studies that described migratory routes (Walkin-shaw 1960 Crete and Toepfer 1978) EP abundance surveys(eg Mississippi Flyway Cooperative Midwinter Survey[MWS] state surveys) and the subsequent behavior ofcranes we marked in this study For each stopover area weselected trapping periods before during and after estimatedpeak abundance based on periodic surveys of cranes duringfall migration (J Bergens Indiana Department of NaturalResources unpublished data and R Klippel TennesseeWildlife Resource Agency unpublished data) We employedthis capture protocol to mark a geographic representativesample of cranes and to capture the temporal variation incrane movements through these stopover areas In additionwe captured cranes on alternative breeding and staging areasincludingManitoulin Island Ontario (Hanna et al 2014) toinclude cranes from portions of their distribution that maynot have been represented by cranes we trapped at majorstaging areasWe used a rocket-propelled net assembly as the primary

method to capture cranes We identified potential trappingsites using the protocol developed for rocket netting cranes intheMCP by Krapu et al (2011) Their protocol gave priorityto daytime loafing sites withgt20 cranes present in pasture orother open land-cover types and incorporated decoys Wemodified their protocol by using bait instead of decoysWhen cranes responded to bait for 2 consecutive days weassembled a 131 197-m rocket-propelled net (Wheelerand Lewis 1972 D A Brandt US Geological Survey[USGS] Northern Prairie Wildlife Research Centerpersonal communication) at a potential capture locationWe conducted trapping primarily in the morning becausecranes consistently returned to baited sites after leavingnocturnal roosts In addition to using a rocket-propelled netwe used a Coda NetLauncher (Coda Enterprises Mesa AZUSA) to capture cranes in locations where using a rocket-propelled net was not feasible We followed the standardCoda NetLauncher protocol developed for cranes by theOhio Department of Natural Resources (D E ShermanOhio Department of Natural Resources personal commu-nication) We also used modified softcatch leghold traps(Victor no 3 Oneida Victor Euclid OH USA) as describedby King et al (1998)We targeted adult female cranes because they are more

likely than males to return to natal breeding grounds thefollowing spring (Walkinshaw 1949 Drewien 1973)Sandhill cranes are perennially monogamous (Tacha et al1994) will re-mate after death or permanent separation(Nesbitt 1989 Hayes and Barzen 2006) and pair bonds willremain after departure from summer areas (Tacha et al 1992Hayes 2015) We assume little difference in migrationbehavior among cohorts in our sample because we targetedadult cranes However we acknowledge that we did not

2 The Journal of Wildlife Management 9999()

know the breeding status of all of our marked cranes Weidentified adult female cranes based on the physical and socialcharacteristics described by Tacha (1998) as smaller body-sized cranes with red skin on the crown of the head morelikely to be viewed following larger-bodied male cranes andless likely to display agonistic behaviors However followingcapture if we could not identify family groups we selected asmaller-bodied adult crane that we presumed to be a femaleWe affixed a 30-g 3-solar-paneled GPS satellite PTT

(North Star Science and Technology Baltimore MD USA)to captured cranes via flanged leg bands We selected tarsus-mounted PTTs because similar transmitters and attachmentmethods have been used extensively on cranes withoutapparent negative effects on survival or movements incomparison to backpack mounted transmitters (Melvin et al

1983 Ellis et al 2001 Krapu et al 2011 Hayes 2015) Wemounted PTTs on a 76-cm 2-piece color-coated polyvinylchloride (PVC) alpha-numeric-coded engraved leg band(Haggie Engraving Crumpton MD USA) Satellite PTTsincorporated 2 additional solar panels to increase thelongevity of the transmitter compared with standardbattery-powered or single solar-paneled transmitters Wereinforced the connection between the antenna and thetransmitter housing as a precaution because it is a weak pointand a cause for loss of transmission (D A Brandt personalcommunication) For cranes not marked with PTTswe affixed a 76-cm 15-wrapped alpha-numeric-codedengraved PVC tarsus auxiliary leg band above the distaltibio-tarsus joint Flanged auxiliary markers and PTTstogether weighed approximately 80 g (2 of average body

Figure 1 Trapping locations by season where sandhill cranes were captured and marked with platform transmitting terminals Gray shading denotes thegeographic extent of movements of cranes during the annual cycle 2009ndash2014

Fronczak et al Eastern Population Sandhill Crane Migration 3

mass at capture) and were attached above the distal tibio-tarsus joint (Krapu et al 2011) All birds marked received astandard USGS aluminum bandFor each PTT-marked crane we drew blood from the vein

below the tibio-tarsus and placed samples in a lysis bufferanticoagulant solution (Jones et al 2005) Blood wassubsequently analyzed by Avian Biotech International(Tallahassee FL USA) to determine sex We did notcollect blood samples from cranes at Manitoulin IslandWhen we captured gt2 cranescrew member at one time wereleased cranes in excess of that number immediately withouttaking measurements or affixing markers to avoid detrimen-tal effects (eg myopathy) of extended handling times Wereleased all cranes held for processing as a group within30minutes of being captured We captured cranes underapproval of the University ofMinnesota Institutional AnimalCare and Use Committee (protocol no 1103A97333) andthe University of Western Ontario Animal Use Protocol(protocol no 2010ndash213)We programmed most PTTs on 5- and 6-hour intervals to

maximize the number of GPS locations per day whichafforded the ability to determine the status (ie sedentarylocations or continuous movement) of marked cranes Weprogrammed other PTTs to collect GPS locations every 6hours during migration and every 8 hours during non-migration periods in the summer and winter Spatial locationdata were transmitted to satellites each 72-hour duty cyclethrough standard Doppler technology Accuracy for GPSlocations was classified into 4 quality levelslt26m 26ndash50m51ndash75m and 76ndash100m by the manufacturerrsquos standards(K LeSage GeoTrak personal communication)

Data AnalysisWe used GPS locations for cranes in the EP that completedfull annual cycles beginning from the date of capture todescribe distribution and migration chronology For cranesthat did not complete an annual cycle because of either signalloss or death we used GPS data for any completed migrationor duration of stay at a winter or summer terminus Whennecessary we incorporated Argos Doppler system locationsas an alternative source of information to describe migrationroutes when GPS data locations were gt5 days betweensignals because of environmental conditions that resulted ininsufficient battery voltage and disrupted data transmissionto satellitesWe retrieved data transmitted by PTTs from Collecte

Localisation Satellites (CLS)-America (httpwwwargos-systemorg Largo MD USA) which we decoded into ageographic information system shapefile format throughsatellite data decoder software (DSDCODE version 4dNorth Star Science and Technologies) We filtered GPSlocations using an adapted location filtering system based onthe Douglas Argos-Filter Algorithm version 65 (Douglaset al 2012 Pearse et al 2015) We remained consistentwith Krapu et al (2011) and assigned attributes to GPSlocation records identifying 4 periods of the annual cycle(ie spring migration summer area fall migration winterarea) and classified summer and winter area locations as the

terminus of migration based on a major departure from theprevious location We assigned fall and spring migrationattributes to areas that we defined as a departuregt15 km forgt5 days from the summer or winter areas We definedstaging areas as locations where a crane stayed at a singlegeographic area for gt5 consecutive days during migrationWe considered areas where cranes stayed for a shorterduration (lt5 days) during migration as stopover sites(Warnock 2010)We estimated the proportion of marked cranes available for

detection during the USFWS fall surveys by spatiallymatching survey locations to GPS locations of marked cranesduring the same periods Fall survey location descriptionsranged from general names of townships nearest towns ornatural areas to precise GPS coordinates We determinedthat cranes were available during the fall survey if their GPScoordinates were located within the area described by theobserver or if the GPS locations of cranes were lt3 km froman observerrsquos GPS locationTo assess philopatry to summer area we compared centroids

of locations during the period PTT-marked cranes occupiedtheir summer range for consecutive summers We calculatedcentroids of locations for each summer area with ArcGIS(version92ESRIRedlandsCAUSA)Wesummarized thelinear distance between these centroids between or amongyears for individual cranes and report an overall mean

RESULTS

From 2009 through 2012 we captured 202 cranes 194 usingrocket-propelled nets 7 with a Coda NetLauncher and 1with a softcatch leghold trap We deployed 38 PTTs on 42cranes (ie we recovered 4 PTTs and deployed them on asecond crane) Of the 42 cranes 25 (60) were female 6(14) were male and 11 (26) were undetermined sexbecause we did not draw a blood sample or the sexing analysishad inconclusive results Seven cranes (17) died and 24(57) of the PTTs we deployed functionedgt24monthsWemonitored cranes between December 2009 and August 2014which generated 103882 GPS locations over 28677tracking days The average number of GPS locationsdaywas 34 (rangefrac14 17ndash40) where 55 and 32 of the locationquality levels were registered as lt26m and 26ndash50mrespectively

Summer AreasCrane summer areas were dispersed across 3 states and 1province of the Great Lakes region 3 (7) settled inMinnesota 12 (29) settled inWisconsin 9 (21) settled inMichigan and 16 (38) settled in Ontario (Fig 2A) Nine(21) cranes that were marked on Manitoulin IslandOntario remained on Manitoulin Island for the duration ofthe summer (Hanna et al 2014) Cranes spent summers ineast-central Minnesota (Benton and Sherburne counties)throughout Wisconsin (Brown Crawford Fond du LacGreen Lake Jefferson Lincoln Outagamie RacineWashburn Waupaca Waukesha andWinnebago counties)on the Upper Peninsula (Chippewa and Mackinac counties)and the Lower Peninsula (Ingham Jackson Kent and

4 The Journal of Wildlife Management 9999()

know the breeding status of all of our marked cranes Weidentified adult female cranes based on the physical and socialcharacteristics described by Tacha (1998) as smaller body-sized cranes with red skin on the crown of the head morelikely to be viewed following larger-bodied male cranes andless likely to display agonistic behaviors However followingcapture if we could not identify family groups we selected asmaller-bodied adult crane that we presumed to be a femaleWe affixed a 30-g 3-solar-paneled GPS satellite PTT

(North Star Science and Technology Baltimore MD USA)to captured cranes via flanged leg bands We selected tarsus-mounted PTTs because similar transmitters and attachmentmethods have been used extensively on cranes withoutapparent negative effects on survival or movements incomparison to backpack mounted transmitters (Melvin et al

1983 Ellis et al 2001 Krapu et al 2011 Hayes 2015) Wemounted PTTs on a 76-cm 2-piece color-coated polyvinylchloride (PVC) alpha-numeric-coded engraved leg band(Haggie Engraving Crumpton MD USA) Satellite PTTsincorporated 2 additional solar panels to increase thelongevity of the transmitter compared with standardbattery-powered or single solar-paneled transmitters Wereinforced the connection between the antenna and thetransmitter housing as a precaution because it is a weak pointand a cause for loss of transmission (D A Brandt personalcommunication) For cranes not marked with PTTswe affixed a 76-cm 15-wrapped alpha-numeric-codedengraved PVC tarsus auxiliary leg band above the distaltibio-tarsus joint Flanged auxiliary markers and PTTstogether weighed approximately 80 g (2 of average body

Figure 1 Trapping locations by season where sandhill cranes were captured and marked with platform transmitting terminals Gray shading denotes thegeographic extent of movements of cranes during the annual cycle 2009ndash2014

Fronczak et al Eastern Population Sandhill Crane Migration 3

mass at capture) and were attached above the distal tibio-tarsus joint (Krapu et al 2011) All birds marked received astandard USGS aluminum bandFor each PTT-marked crane we drew blood from the vein

below the tibio-tarsus and placed samples in a lysis bufferanticoagulant solution (Jones et al 2005) Blood wassubsequently analyzed by Avian Biotech International(Tallahassee FL USA) to determine sex We did notcollect blood samples from cranes at Manitoulin IslandWhen we captured gt2 cranescrew member at one time wereleased cranes in excess of that number immediately withouttaking measurements or affixing markers to avoid detrimen-tal effects (eg myopathy) of extended handling times Wereleased all cranes held for processing as a group within30minutes of being captured We captured cranes underapproval of the University ofMinnesota Institutional AnimalCare and Use Committee (protocol no 1103A97333) andthe University of Western Ontario Animal Use Protocol(protocol no 2010ndash213)We programmed most PTTs on 5- and 6-hour intervals to

maximize the number of GPS locations per day whichafforded the ability to determine the status (ie sedentarylocations or continuous movement) of marked cranes Weprogrammed other PTTs to collect GPS locations every 6hours during migration and every 8 hours during non-migration periods in the summer and winter Spatial locationdata were transmitted to satellites each 72-hour duty cyclethrough standard Doppler technology Accuracy for GPSlocations was classified into 4 quality levelslt26m 26ndash50m51ndash75m and 76ndash100m by the manufacturerrsquos standards(K LeSage GeoTrak personal communication)

Data AnalysisWe used GPS locations for cranes in the EP that completedfull annual cycles beginning from the date of capture todescribe distribution and migration chronology For cranesthat did not complete an annual cycle because of either signalloss or death we used GPS data for any completed migrationor duration of stay at a winter or summer terminus Whennecessary we incorporated Argos Doppler system locationsas an alternative source of information to describe migrationroutes when GPS data locations were gt5 days betweensignals because of environmental conditions that resulted ininsufficient battery voltage and disrupted data transmissionto satellitesWe retrieved data transmitted by PTTs from Collecte

Localisation Satellites (CLS)-America (httpwwwargos-systemorg Largo MD USA) which we decoded into ageographic information system shapefile format throughsatellite data decoder software (DSDCODE version 4dNorth Star Science and Technologies) We filtered GPSlocations using an adapted location filtering system based onthe Douglas Argos-Filter Algorithm version 65 (Douglaset al 2012 Pearse et al 2015) We remained consistentwith Krapu et al (2011) and assigned attributes to GPSlocation records identifying 4 periods of the annual cycle(ie spring migration summer area fall migration winterarea) and classified summer and winter area locations as the

terminus of migration based on a major departure from theprevious location We assigned fall and spring migrationattributes to areas that we defined as a departuregt15 km forgt5 days from the summer or winter areas We definedstaging areas as locations where a crane stayed at a singlegeographic area for gt5 consecutive days during migrationWe considered areas where cranes stayed for a shorterduration (lt5 days) during migration as stopover sites(Warnock 2010)We estimated the proportion of marked cranes available for

detection during the USFWS fall surveys by spatiallymatching survey locations to GPS locations of marked cranesduring the same periods Fall survey location descriptionsranged from general names of townships nearest towns ornatural areas to precise GPS coordinates We determinedthat cranes were available during the fall survey if their GPScoordinates were located within the area described by theobserver or if the GPS locations of cranes were lt3 km froman observerrsquos GPS locationTo assess philopatry to summer area we compared centroids

of locations during the period PTT-marked cranes occupiedtheir summer range for consecutive summers We calculatedcentroids of locations for each summer area with ArcGIS(version92ESRIRedlandsCAUSA)Wesummarized thelinear distance between these centroids between or amongyears for individual cranes and report an overall mean

RESULTS

From 2009 through 2012 we captured 202 cranes 194 usingrocket-propelled nets 7 with a Coda NetLauncher and 1with a softcatch leghold trap We deployed 38 PTTs on 42cranes (ie we recovered 4 PTTs and deployed them on asecond crane) Of the 42 cranes 25 (60) were female 6(14) were male and 11 (26) were undetermined sexbecause we did not draw a blood sample or the sexing analysishad inconclusive results Seven cranes (17) died and 24(57) of the PTTs we deployed functionedgt24monthsWemonitored cranes between December 2009 and August 2014which generated 103882 GPS locations over 28677tracking days The average number of GPS locationsdaywas 34 (rangefrac14 17ndash40) where 55 and 32 of the locationquality levels were registered as lt26m and 26ndash50mrespectively

Summer AreasCrane summer areas were dispersed across 3 states and 1province of the Great Lakes region 3 (7) settled inMinnesota 12 (29) settled inWisconsin 9 (21) settled inMichigan and 16 (38) settled in Ontario (Fig 2A) Nine(21) cranes that were marked on Manitoulin IslandOntario remained on Manitoulin Island for the duration ofthe summer (Hanna et al 2014) Cranes spent summers ineast-central Minnesota (Benton and Sherburne counties)throughout Wisconsin (Brown Crawford Fond du LacGreen Lake Jefferson Lincoln Outagamie RacineWashburn Waupaca Waukesha andWinnebago counties)on the Upper Peninsula (Chippewa and Mackinac counties)and the Lower Peninsula (Ingham Jackson Kent and

4 The Journal of Wildlife Management 9999()

mass at capture) and were attached above the distal tibio-tarsus joint (Krapu et al 2011) All birds marked received astandard USGS aluminum bandFor each PTT-marked crane we drew blood from the vein

below the tibio-tarsus and placed samples in a lysis bufferanticoagulant solution (Jones et al 2005) Blood wassubsequently analyzed by Avian Biotech International(Tallahassee FL USA) to determine sex We did notcollect blood samples from cranes at Manitoulin IslandWhen we captured gt2 cranescrew member at one time wereleased cranes in excess of that number immediately withouttaking measurements or affixing markers to avoid detrimen-tal effects (eg myopathy) of extended handling times Wereleased all cranes held for processing as a group within30minutes of being captured We captured cranes underapproval of the University ofMinnesota Institutional AnimalCare and Use Committee (protocol no 1103A97333) andthe University of Western Ontario Animal Use Protocol(protocol no 2010ndash213)We programmed most PTTs on 5- and 6-hour intervals to

maximize the number of GPS locations per day whichafforded the ability to determine the status (ie sedentarylocations or continuous movement) of marked cranes Weprogrammed other PTTs to collect GPS locations every 6hours during migration and every 8 hours during non-migration periods in the summer and winter Spatial locationdata were transmitted to satellites each 72-hour duty cyclethrough standard Doppler technology Accuracy for GPSlocations was classified into 4 quality levelslt26m 26ndash50m51ndash75m and 76ndash100m by the manufacturerrsquos standards(K LeSage GeoTrak personal communication)

Data AnalysisWe used GPS locations for cranes in the EP that completedfull annual cycles beginning from the date of capture todescribe distribution and migration chronology For cranesthat did not complete an annual cycle because of either signalloss or death we used GPS data for any completed migrationor duration of stay at a winter or summer terminus Whennecessary we incorporated Argos Doppler system locationsas an alternative source of information to describe migrationroutes when GPS data locations were gt5 days betweensignals because of environmental conditions that resulted ininsufficient battery voltage and disrupted data transmissionto satellitesWe retrieved data transmitted by PTTs from Collecte

Localisation Satellites (CLS)-America (httpwwwargos-systemorg Largo MD USA) which we decoded into ageographic information system shapefile format throughsatellite data decoder software (DSDCODE version 4dNorth Star Science and Technologies) We filtered GPSlocations using an adapted location filtering system based onthe Douglas Argos-Filter Algorithm version 65 (Douglaset al 2012 Pearse et al 2015) We remained consistentwith Krapu et al (2011) and assigned attributes to GPSlocation records identifying 4 periods of the annual cycle(ie spring migration summer area fall migration winterarea) and classified summer and winter area locations as the

terminus of migration based on a major departure from theprevious location We assigned fall and spring migrationattributes to areas that we defined as a departuregt15 km forgt5 days from the summer or winter areas We definedstaging areas as locations where a crane stayed at a singlegeographic area for gt5 consecutive days during migrationWe considered areas where cranes stayed for a shorterduration (lt5 days) during migration as stopover sites(Warnock 2010)We estimated the proportion of marked cranes available for

detection during the USFWS fall surveys by spatiallymatching survey locations to GPS locations of marked cranesduring the same periods Fall survey location descriptionsranged from general names of townships nearest towns ornatural areas to precise GPS coordinates We determinedthat cranes were available during the fall survey if their GPScoordinates were located within the area described by theobserver or if the GPS locations of cranes were lt3 km froman observerrsquos GPS locationTo assess philopatry to summer area we compared centroids

of locations during the period PTT-marked cranes occupiedtheir summer range for consecutive summers We calculatedcentroids of locations for each summer area with ArcGIS(version92ESRIRedlandsCAUSA)Wesummarized thelinear distance between these centroids between or amongyears for individual cranes and report an overall mean

RESULTS

From 2009 through 2012 we captured 202 cranes 194 usingrocket-propelled nets 7 with a Coda NetLauncher and 1with a softcatch leghold trap We deployed 38 PTTs on 42cranes (ie we recovered 4 PTTs and deployed them on asecond crane) Of the 42 cranes 25 (60) were female 6(14) were male and 11 (26) were undetermined sexbecause we did not draw a blood sample or the sexing analysishad inconclusive results Seven cranes (17) died and 24(57) of the PTTs we deployed functionedgt24monthsWemonitored cranes between December 2009 and August 2014which generated 103882 GPS locations over 28677tracking days The average number of GPS locationsdaywas 34 (rangefrac14 17ndash40) where 55 and 32 of the locationquality levels were registered as lt26m and 26ndash50mrespectively

Summer AreasCrane summer areas were dispersed across 3 states and 1province of the Great Lakes region 3 (7) settled inMinnesota 12 (29) settled inWisconsin 9 (21) settled inMichigan and 16 (38) settled in Ontario (Fig 2A) Nine(21) cranes that were marked on Manitoulin IslandOntario remained on Manitoulin Island for the duration ofthe summer (Hanna et al 2014) Cranes spent summers ineast-central Minnesota (Benton and Sherburne counties)throughout Wisconsin (Brown Crawford Fond du LacGreen Lake Jefferson Lincoln Outagamie RacineWashburn Waupaca Waukesha andWinnebago counties)on the Upper Peninsula (Chippewa and Mackinac counties)and the Lower Peninsula (Ingham Jackson Kent and

4 The Journal of Wildlife Management 9999()

Muskegon counties) of Michigan and the north shore ofLake Huron (Algoma District) Manitoulin Island (Man-itoulin District) and east-central (Algoma Cochrane andSudbury districts) OntarioTwenty-five cranes returned to their initial summer area

after a second spring migration of which 19 (76) settledlt3 km from the estimatedmean center of the summer area ofthe previous year For the 25 returning cranes the averagelinear distance between summer area centroids between yearswas 155 km (SDfrac14 218 km rangefrac14 003ndash783 km) Wewere unable to assess summer area philopatry for 17 cranesbecause of PTT failure or death (Fronczak et al 2015)

Fall MigrationCranes migrated to their respective winter termini usingmigration corridors on either side of Lake Michigan(Fig 3A) Cranes (nfrac14 14 33) that summered inMinnesota and Wisconsin migrated through Wisconsinand primarily used staging areas along the Wisconsin RiverValley (Iowa Juneau and Sauk counties) and areassurrounding Green Lake and Lake Poygon (Green LakeWaushara and Winnebago counties) All cranes summeringwest of Lake Michigan migrated to their winter terminithrough either the Kankakee River Valley (Jasper-PulaskiFWA nfrac14 12 86) or used routes through Illinois tosouthern Indiana staging areas (nfrac14 2 14)Cranes that summered on the North Channel of Lake

Huron and Manitoulin Island Ontario and the east sideof Lake Michigan in Michigan (nfrac14 17 40) migratedsouth through the central portion of southern Michiganand primarily staged in Barry Calhoun Eaton Jacksonand Kalamazoo counties (south-central MI) Of the 17

cranes that staged in south-central Michigan 8 departedsouth for routes through northeastern Indiana andwestern Ohio and 9 departed southwest to staging areasin IndianaCranes summering farther north in south-central Ontario

and the eastern Upper Peninsula of Michigan (nfrac14 6 14)usedmigration corridors onboth the east andwest side ofLakeMichigan These cranes traversed either west through theUpper Peninsula of Michigan to staging areas surroundingLake Poygon Wisconsin (Waushara and Winnebago coun-ties) and then followed similar routes as cranes summering ineastern Wisconsin or traversed southwest from the stagingarea northwest of Lake Huron (Algoma District) crossing atthe Straits of Mackinac Michigan to primarily use stagingareas in south-central Michigan or northwestern IndianaAs cranes continued south 27 (64) used the Kankakee

River Valley (specifically Jasper-Pulaski FWA) 1 year and11 (26) cranes bypassed the Kankakee River Valley1 yearduring fall migration These cranes left northern stagingareas in late November to early December and made briefstops in Indiana Illinois and Ohio as they moved south totheir winter termini (Fig 3A)Cranes that continued their migration south from Jasper-

Pulaski FWA and the Kankakee River Valley moved eitherthrough western or central Indiana and then converged onsimilar routes through central Kentucky Cranes thatcontinued south fromKentucky (nfrac14 18 42)moved throughcentral Tennessee used the Chickamauga Reservoir (specifi-cally Hiwassee WR) as a staging or stopover area prior tocompleting their migration One crane that summered inOntario migrated from south-central Indiana west throughnorth-centralKentuckyandeventuallywinteredat theHop-In

Figure 2 Summer (A) and winter (B) area locations (black stars) for platform transmitting terminal-marked greater sandhill cranes 2009ndash2014

Fronczak et al Eastern Population Sandhill Crane Migration 5

WA (Obion County) Tennessee Cranes that wintered inGeorgia and Florida made multiple stops in a migrationcorridor through northeastern Alabama and west-centralGeorgia to their respectivewinter areas (Fig3A)Therewere3(7) cranes that alternated their migration route around LakeMichigan to their respectivewinter termini at least onceduringthe study

Distribution of Winter AreasThe distribution of crane winter areas extended from Indianato Florida (Fig 2B) The majority of marked cranes (nfrac14 2662) wintered1 year in southeastern Tennessee within theHiwassee WR and surrounding Chickamauga Reservoir(Meigs Rhea and Hamilton counties) or in northwesternTennessee (Obion County) South of Tennessee 19 (45)cranes wintered throughout the Florida Peninsula (AlachuaBrevard Glades Highlands Lake Levy Marion OkaloosaOkeechobee Osceola Pasco Polk and Putnam counties)and 4 (10) wintered in Georgia (Brooks Crisp Floyd andPulaski counties) North of Tennessee 3 (7) craneswintered in Kentucky (Barren and Larue counties) and 12(29) wintered in Indiana (Greene Jackson JasperLaPorte and Starke counties) Eleven (26) cranes returnedto the same winter area in subsequent winters and 31 (74)wintered in 1 state at least once during the study period

Description of Spring MigrationMarked cranes used similar spring and fall migration routes(Fig 3)Cranes thatwintered inFlorida and southernGeorgiatraversed central and western Georgia and northeasternAlabama and staged either within theChickamaugaReservoir

(ie Hiwassee WR nfrac14 12 29) or migrated directly tostaging areas in north-central Tennessee central Kentuckyand southern Indiana Cranes that wintered in the Chick-amauga Reservoir used multiple staging areas in Kentuckyconcentrated in south-central Kentucky (nfrac14 17 40 Barrenand Metcalfe counties) and north-central Kentucky (nfrac14 2560 Hardin and Larue counties) Two cranes (5)that wintered in northwestern Tennessee (Obion County)migrated to Indiana staging areas either through southernIndiana or through eastern IllinoisCranes migrating north from central Kentucky moved to

southern Indiana staging areas in the White River Valley(nfrac14 27 64 Jackson County) and west-central Indianastaging area at Goose Pond FWA and surrounding area(nfrac14 13 31 Greene County) Most cranes (nfrac14 35 83)used stopover sites at the Jasper-Pulaski FWA andthroughout the Kankakee River Valley after departingsouthern Indiana and from there continued migration totheir respective summer areas Several cranes (nfrac14 10 24)moved from southern Indiana staging areas and bypassedJasper-Pulaski FWA and the Kankakee River Valley on totheir summer areasCranes that summeredoutsideofLowerPeninsulaMichigan

and north and east of LakeMichigan (nfrac14 16 38) followed aroute through centralMichigan continuing through either theStraits of Mackinaw or across Lake Huron (ie thosesummering on Manitoulin Island Ontario) Cranes thatsummered outside of Wisconsin and north and west of LakeMichigan migrated either through central Wisconsin (nfrac14 37) or through easternWisconsin and continued through the

Figure 3 Fall (A) and spring (B) migration routes for platform transmitting terminal-marked greater sandhill cranes 2010ndash2014

6 The Journal of Wildlife Management 9999()

Upper Peninsula of Michigan (nfrac14 7 17) Seven (17)cranes that summered on theUpper Peninsula ofMichigan innorth-central Ontario and east of Lake Michigan (NorthShore of Lake Huron and Manitoulin Island Ontario)migrated west of Lake Michigan and through the UpperPeninsula of Michigan at least once during the study periodThere were 6 (14) cranes that alternated their migrationroute aroundLakeMichigan to their respective summer area atleast once during the study

Chronology of Annual MovementsCranes on average arrived at their summer areas on 27March(nfrac14 97 rangefrac14 22 Febndash3 May Table 1) and stayed onsummer areas 189 days (nfrac14 75 SDfrac14 34 days) Averagedeparture from summer areas for staging areas was 5 October(nfrac14 85 rangefrac14 10 Augndash28 Dec) and cranes traveled onaverage 59 days (nfrac14 79 SDfrac14 37 days Table 2) fromsummer areas to winter termini Average cumulative distancetraveled between subsequent stops from summer to wintertermini during fall migration was 237 km (nfrac14 433 SDfrac14 177 km) Cranes used an average of 61 stops (nfrac14 75SDfrac14 28 stops) during fall migration on average 18(nfrac14 84 SDfrac14 10 stop) of these stops were at staging areasand 33 (nfrac14 89 SDfrac14 21 stops) were at stopover sites Weidentified 43 fall staging areas and 136 fall stopover sitesAverage number of days spent on individual staging areas was35 (nfrac14 79 SDfrac14 22 days) where 34 and 18 of fallmigration days were spent in the Kankakee River Valley andsouth-central Michigan respectively (Appendix A)Average arrival date of cranes at their winter areas was 11

December (nfrac14 86 rangefrac14 21 Sepndash27 Jan Table 3) and theyremained on winter areas an average of 67 days (nfrac14 85SDfrac14 33 days) Average departure date of cranes from winterareas was 16 February (nfrac14 102 rangefrac14 15 Janndash29Mar) and

cranes traveled on average 40 days (nfrac14 94 SDfrac14 18 daysTable 4) from winter areas to summer areas Averagedistance traveled between sequential stops prior to reachingsummer areas was approximately 195 km (nfrac14 652 SDfrac14 132 km) Cranes stopped on average 76 times (nfrac14 97SDfrac14 31 stops) during spring migration 46 (nfrac14 100SDfrac14 27 stops) of these stops were at stopover sites and 21(nfrac14 91 SDfrac14 10 stop) were at staging areas We identified43 spring staging areas and 172 spring stopover sitesAverage number of days spent on staging areas during springmigration was 15 (nfrac14 91 SDfrac14 6 days) where 34 and 9of the cumulative migration days were spent in the KankakeeRiver Valley northwestern Indiana and the White RiverValley southern Indiana respectively (Appendix B)

USFWS Fall SurveyApproximately 30 29 and 31 of crane locations wereoutside of areas surveyed during the 2010 2011 and 2012USFWS fall surveys respectively Crane locations that wereoutside surveyed areas included northern and east-centralWisconsin and central and north-central Michigan (Fig 4)However these cranes eventually moved to staging areas thatwould have been covered by observers after the fall surveyperiod All of the cranes that summered in Canada werepresent in surveyed locations during the survey period

DISCUSSION

Summer Area DistributionSummer areas used by marked cranes were widely distributedthroughout Minnesota Wisconsin Michigan and in theeastern portion of Ontario Canada and occurred within theprevious estimated breeding range of cranes in the EP (VanHorn et al 2010) Geographic distribution and migrationroutes that cranes used coincided with previously reported

Table 1 Mean arrival date length of stay and departure date on summer areas for platform transmitting terminal-marked greater sandhill cranes in theEastern Population North America 2010ndash2014

Arrival date Length of stay (days) Departure date

Yr n x SD (days) n x SD (days) n x SD (days)

2010 7 22 Mar 14 7 194 34 16 6 Oct 222011 32 30 Mar 16 30 185 32 30 6 Oct 282012 33 20 Mar 11 26 196 36 27 5 Oct 302013 17 3 Apr 15 9 176 34 9 2 Oct 252014 8 6 Apr 16 3 192 36 3 11Oct 31All years 97 27 Mar 15 75 189 34 85 5 Oct 27

Table 2 Average number of days number of stops and length of stay at staging areas (stop duration gt5 days) from summer areas to winter termini forplatform transmitting terminal-marked greater sandhill cranes in the Eastern Population North America 2010ndash2013

Total days No stops Length of stay (days)

Yra n x SD (days) n x SD (stops) n x SD (days)

2010 16 54 32 16 59 19 17 42 232011 30 58 36 29 59 26 29 36 222012 25 61 41 22 63 36 23 29 192013 8 72 40 8 68 32 7 33 13All years 79 59 37 75 61 28 79 35 22

a Summaries are unavailable for 2014

Fronczak et al Eastern Population Sandhill Crane Migration 7

geographic distributions (Walkinshaw 1949 Tacha et al1994) and migration routes (Walkinshaw 1960 Melvin andTemple 1982) However at smaller spatial scales some areasthought to support breeding cranes in the EP were notrepresented by our sample including northwestern Wis-consin-northeastern Minnesota (Cutright et al 2006Minnesota Breeding Bird Atlas Project 2014) and easternOntario including the eastern shore of Lake Huron and thesouthern peninsula of Ontario (Cadman et al 2007) Noneof the cranes in our sample overlapped the summer range ofcranes in theMCP identified by Krapu et al (2011) Summerarea locations of marked cranes were concentrated in east-central Wisconsin and the north shore of Lake HuronOntario which support relatively high densities of breedingcranes (Cadman et al 2007 Van Horn et al 2010)Cranes consistently returned to the same summer areas in

subsequent years similar to cranes in the Rocky MountainPopulation (Drewien 1973) and MCP (Krapu et al 2011)On average cranes in the EP and MCP (Krapu et al 2011)used subsequent summer areas within lt10 km of oneanother 65 of cranes in the EP used subsequent summerareas within 1 km of one another compared to 38 forcranes in the MCP (Krapu et al 2011) We observed 3instances when individual cranes moved an average lineardistance of 672 km (rangefrac14 251ndash989 km) among subse-quent summer areas and we speculate that these cranes mayhave been sub-adults (Bennett 1989 Nesbitt and Williams1990 Hayes 2015) However 2 of these cranes returned tothe same summer area in subsequent years and the thirdcrane died during the first summer

Migration Routes and ChronologyCranes followed migration routes similar to those identifiedin previous migration studies that used very high-frequency

transmitters in south-central Minnesota (Crete and Toepfer1978) and cranes marked with ARGOS Doppler PTTs onManitoulin Island Ontario Canada (S A Petrie BirdStudies Canada unpublished data) Unlike those studies wewere able to provide more precise information particularlyabout routes and locations where cranes stopped duringmigration We also observed that 8 (19) cranes withsummer areas in the Upper Peninsula of MichiganManitoulin Island Ontario and east-central Ontario usedmigration routes west of Lake Michigan instead of the moreintuitive route through the Lower Peninsula of MichiganSix of these cranes also alternated migration routes aroundLake Michigan during sequential migration (ie fall andspring) and in subsequent years Our study represents thefirst evidence of this behavior and use of both of thesemigration routes by individual cranes We also documentedcranes (nfrac14 8 19) that summered on Manitoulin Islandused migration routes over large bodies of water (LakeHuron and portions of Lake Michigan) similar to Krapuet al (2011) who described cranes crossing the Bering Straitof Alaska and the Arctic Ocean to and from breedinggroundsRecent observations of migratory cranes that wintered in

Louisiana led to speculation that EP cranes are potentiallyusing non-traditional migration corridors King et al(2010) monitored 2 PTT-marked cranes that wintered incentral Louisiana and migrated north through the Mis-sissippi Alluvial Valley toward summer areas within the EPsummer distribution One marked crane in their study useda migration route through west-central Illinois to south-western Wisconsin and then into the Upper Peninsula ofMichigan after which they were unable to receiveadditional location information from the transmitter Theother crane traveled a similar route used by some of the

Table 3 Mean arrival date length of stay and departure date on winter areas for platform transmitting terminal-marked greater sandhill cranes in theEastern Population North America 2009ndash2014

Arrival datea Length of stay (days)a Departure date

Yr n x SD (days) n x SD (days) n x SD (days)

2009ndash2010 6 16 Feb 62010ndash2011 24 10 Dec 13 26 66 17 35 18 Feb 92011ndash2012 31 14 Dec 28 31 65 32 34 15 Feb 152012ndash2013 22 6 Dec 34 20 74 49 19 19 Feb 222013ndash2014 8 14 Dec 20 7 63 29 8 14 Feb 17All years 86 11 Dec 26 85 67 33 102 16 Feb 14

a Summaries unavailable for 2009ndash2010 winter locations

Table 4 Average number of days number of stops and length of stay at staging areas (stop duration gt5 days) from winter termini to summer areas forplatform transmitting terminal-marked greater sandhill cranes in the Eastern Population North America 2010ndash2014

Total days No stops Length of stay (days)

Yr n x SD (days) n x SD (stops) n x SD (days)

2010 6 39 11 6 68 08 6 12 12011 33 42 16 34 84 30 33 14 42012 33 32 15 33 71 30 29 17 82013 16 41 22 17 71 40 15 17 72014 6 62 11 7 77 25 8 14 4All years 94 39 18 97 76 31 91 15 6

8 The Journal of Wildlife Management 9999()

cranes in our sample through west-central Indiana (GoosePond FWA) northwestern Indiana (Kankakee RiverValley) east-central Wisconsin and then through theUpper Peninsula of Michigan to a summer area in north-central OntarioA single PTT-marked crane in our sample from the

Manitoulin Island Ontario study site migrated west fromthe White River Valley Indiana (Jackson County) andwintered at the Hop-In WR Tennessee (Obion County)However that crane did not return to Hop-In WR in thesubsequent year In anticipation of identifying a potentialwestern migration corridor to Louisiana we affixed 2 PTTsto cranes that wintered at Hop-In WR However only 1returned to winter at the Hop-InWR and the other winteredin Florida the following year

Cranes consistently used staging areas along migrationroutes but made fewer stops for a longer duration during fallmigration than during spring migration Locations wherecranes stopped during migration along the north shore ofLake Huron Algoma District Ontario Lake Poygon east-central Wisconsin the Kankakee River Valley northwesternIndiana and counties in south-central Michigan werefrequented more by cranes during fall migration (Fig 3AAppendix A) Cranes used Hardin County north-centralKentucky White River Valley (Ewing Bottoms JacksonCounty) south-central Indiana and the Kankakee RiverValley northwestern Indiana more frequently during springthan during fall migration (Fig 3B Appendix B) Theseareas support shallow-marsh roosting habitat have anabundant food supply from agricultural production and

Figure 4 Locations of areas surveyed (gray squares) during the United States Fish and Wildlife Service Fall Eastern Population Sandhill Crane Survey alsoshown are locations (black triangles) where platform transmitting terminal-marked sandhill cranes were not observed during the annual surveys 2010ndash2012

Fronczak et al Eastern Population Sandhill Crane Migration 9

are more likely to be protected from human disturbances (iehunting and sporting activities) The Kankakee River Valleylocated in northwestern Indiana was the most frequented ofall staging areas during fall and spring migration wherecranes spent 34 of migration days

Winter Area DistributionCrane winter areas extended from Indiana to Florida wellnorth of the areas previously described by Tacha et al(1994) who reported that cranes in the EP winteredmainly in southeastern Georgia and in Florida Craneswintered in Florida Georgia Tennessee Kentucky andIndiana with the Chickamauga Reservoir (specificallyHiwassee WR) Tennessee having the highest proportionof use among all winter areas The proportion of cranesthat used the Chickamauga Reservoir Valley for thewinters of 2009 and 2010 may be biased high because wetrapped the majority of cranes (nfrac14 6 14 2009ndash2010nfrac14 12 28 2010ndash2011) at the Hiwassee WR duringDecember and January of those years However in 2010MWS estimates for the Chickamauga Reservoir indicateda record number of wintering cranes (nfrac14 48300 J RKelley USFWS-Migratory Bird Management personalcommunication) suggesting that a large portion of cranesin the EP likely used that area Furthermore many ofthe cranes we captured in other locations used theChickamauga Reservoir during migration in years follow-ing their capture The Chickamauga Reservoir is ideal forstaging and wintering cranes Agriculture crops (corn soybeans and winter wheat) are produced annually forwintering wildlife and adjacent sand bars and low waterlevels on Chickamauga Reservoir provided roostinghabitat and sanctuary for sandhill cranes during the falland winter months (R Klippel personal communication)Hunting for sandhill cranes is allowed throughout thereservoir however the Hiwassee WR is designated a non-hunting zoneRecent analysis of long-term Christmas Bird Count data

indicated that cranes in the EP wintered at staging areasnorth of Georgia and Florida in recent decades (Amundsonand Johnson 2011 Lacy et al 2015) Lacy et al (2015)suggested that climate and increased agriculture throughoutthe EP range may influence cranes to winter farther northHowever there has been no formal analysis to demonstratethat these factors are main contributors to a more northerlywinter strategy Thirty-one (74) cranes in our sample thatwintered north of Georgia and Florida used migrationstaging areas in Indiana Kentucky and Tennessee Winterdestinations for cranes varied among years where 11 (26)returned to the same winter area in subsequent winters Theremaining 31 (74) cranes wintered in1 state at least onceduring the study period indicating a variation in migrationstrategy possibly influencing cranes to settle in a particularwinter areaEight (19) cranes used the Kankakee River Valley in

northwestern Indiana as a winter area in1 year These cranesthat wintered in the Kankakee River Valley used the Jasper-Pulaski FWA until all available roosting sites froze

then roosted within the Schahfer Generating Stationproperty in Wheatfield Indiana about 10 km north-northwest of Jasper-Pulaski FWA The power stationprovided roosting areas that remained ice-free and thesurrounding agricultural landscape provided waste grainthroughout the winter (J Bergens personal communica-tion) These cranes then used areas throughout theKankakee River Valley in proximity to Jasper-PulaskiFWA as the winter progressedAdditional winter locations outside of traditional areas are

not well understood Location data from King et al (2010)suggested that cranes in the EP used winter areas in northernLouisiana and 3 cranes from our sample wintered at theHop-In WR Tennessee west of previously reportedwintering areas for cranes in the EP Observations duringthe MWS of cranes at the Wheeler NWR Alabama havesteadily increased since 2003 with an average of 4583 cranes(rangefrac14 293ndash12032) and a peak count of 12032 cranes in2013 Furthermore observations of cranes in Arkansas andMississippi along theMississippi Alluvial Valley are frequentduring the MWS (J R Kelley personal communication)These cranes likely are part of the EP and most likely usedthe Hop-In WR as a staging area

MANAGEMENT IMPLICATIONS

Location and chronology data from our study can be used toadjust timing of hunting seasons to potentially avoid highharvest pressure on local breeding adults and focus harvestpressure on migrants Location and chronology data canalso be used to identify specific locality information forpotential land conservation and opportunity for non-consumptive recreational activities (eg crane festivalsand bird watching)We estimated a proportion of cranes in the EP were located

in areas not currently covered by the USFWS fall surveyduring the survey period If the absence of these cranes in theUSFWS fall survey area is similar through time series ofcounts from surveys will likely provide a reasonable estimateof population trend Alternatively if the proportion of cranesnot present in survey areas changes it may be necessary toredesign the survey by moving the survey period later into thefall or early winter when a larger portion of cranes areassociated with known concentration areas Howeverthe effects of moving the survey period would need to beevaluated to avoid issues related to 1) comparability toprevious surveys and 2) cranes arriving on wintering areas inFlorida where it is difficult to distinguish cranes in theEP from Florida non-migratory greater sandhill cranes(A c pratensis)

ACKNOWLEDGMENTS

Any use of trade firm or product names is for descriptivepurposes only and does not imply endorsement by the USGovernment or the University of Minnesota We thankD A Brandt P Link R Klippel J Jackson B Swiney SShelby J Womac B Layton D E Sherman B Feaster APhelps R Ronk J Bergens J Gilbert E E Carroll RKnopick W Delks V Clarkston E Hockman C Trosen

10 The Journal of Wildlife Management 9999()

A Johnson C Kanke P Peterson S Zudrow TPaulson W Tamminga J Brunjes E Harper LArmstrong T White M Morse S Bossuyt M BossuytA Dhamorikar J E Hanna KWeaver and S A Petrie forfield assistance We thank A T Pearse for comments onprevious drafts of this manuscript The project was supportedthrough Research Work Order no 86 at the USGSMinnesota Cooperative Fish and Wildlife Research Unitat the University of Minnesota This research was supportedby the USFWS Webless Migratory Game Bird Programthe USGS Northern Prairie Wildlife Research Center theMinnesota Cooperative Fish and Wildlife Research Unitthe USFWS Region 3Migratory BirdManagement OfficeLong Point Waterfowl SC Johnsons and Son Ltdthe Ontario Federation of Anglers and Hunters theCanadian Wildlife Service Wildlife Habitat Canada andthe TD Friends of the Environment Foundation

LITERATURE CITEDAmundson C L and D H Johnson 2011 Assessment of the EasternPopulation of greater sandhill cranes (Grus canadensis tabida) FallMigrationSurvey 1979ndash2009 US Fish and Wildlife Service BloomingtonMinnesota USA

Bennett A J 1989 Movements and home ranges of Florida sandhill cranesJournal of Wildlife Management 53830ndash836

Cadman M D D A Sutherland G G Beck D Lepage and A RCouturier 2007 Ontario Breeding Bird Atlas website httpwwwbirdsontarioorgatlasindexjsp Accessed 14 Mar 2014

CreteRA andJEToepfer1978Migrationof radio-taggedeasterngreatersandhill cranes US Fish and Wildlife Service Twin Cities MinnesotaUSA

Cutright N J B R Harriman and R W Howe 2006 WisconsinBreeding Bird Atlas httpuwgbedubirdswbbaindexhtm Accessed 14Mar 2014

Case D J and S J Sanders editors 2009 Priority information needs forsandhill cranes a funding strategy Association of Fish and WildlifeAgencies Washington DC USA

Douglas D C R Weinzierl S C Davidson R Kays M Wikelski andG Bohrer 2012 Moderating ARGOS location errors in animal trackingdata Methods in Ecology and Evolution 3999ndash1007

Drewien R C 1973 Ecology of Rocky Mountain greater sandhill cranesDissertation University of Idaho Moscow USA

Ellis D H P W Howey and G L Krapu 2001 Recommendations forthe attachment of satellite transmitters to cranes North American CraneWorkshop Proceedings 8211ndash212

Fronczak D L 2014 Distribution migration chronology and survival ratesof Eastern Population sandhill crane Thesis University of Minnesota StPaul USA

Fronczak D L D E Andersen E E Hanna and T R Cooper 2015Annual survival rate estimate of satellite transmitter-marked EasternPopulation greater sandhill cranes Journal of Fish and WildlifeManagement 6464ndash471

Hanna E E M L Schummer and S A Petrie 2014 Migratorychronology autumn recruitment and population size of EasternPopulation sandhill crane (Grus canadensis) from the North Shore Regionof Lake Huron Ontario Canada CanadianWildlife Service and OntarioMinistry of Natural Resources Port Rowan Ontario Canada

Hayes M A and J A Barzen 2006 Dynamics of breeding and non-breeding sandhill cranes in south-central Wisconsin Passenger Pigeon68345ndash352

Hayes M A 2015 Dispersal and population genetic structure in twoflyways of sandhill cranes (Grus canadensis) Dissertation University ofWisconsin-Madison Madison USA

Hunt R A E A Gluesing and L E Nauman 1976 The sandhill crane inWisconsin Wisconsin Department of Natural Resources Madison USA

Jones K L G L Krapu D A Brandt andMV Ashley 2005 Populationgenetic structure in migratory sandhill cranes and the role of Pleistoceneglaciations Molecular Ecology 142645ndash2657

King D T J D Paulson D J Leblanc and K Bruce 1998 Two capturetechniques for American white pelicans and great blue herons ColonialWaterbirds 21258ndash260

King S L A R Pierce K R Hersey and N Winstead 2010 Migrationpatterns and movements of sandhill cranes wintering in central andsouthwest Louisiana Proceedings of the North American CraneWorkshop 1157ndash61

Krapu G L D A Brandt K L Jones and D H Johnson 2011Geographic distribution of the Mid-continent Population of sandhillcranes and related management applications Wildlife Monographs1751ndash38

Kruse K L and J A Dubovsky 2015 Status and harvests of sandhillcranes Mid-continent Rocky Mountain Lower Colorado River Valleyand Eastern Populations US Fish and Wildlife Service LakewoodColorado USA

Lacy A E J A Barzen D MMoore and K E Norris 2015 Changes inthe number and distribution of greater sandhill cranes in the EasternPopulation Journal Field Ornithology 86317ndash325

Lumsden H G 1971 The status of the sandhill crane in northern OntarioCanadian Field-Naturalist 85285ndash293

Melvin S M and S A Temple 1982 Migration ecology of sandhillcranes a review Pages 73ndash87 in J C Lewis editor Proceedings of the1981 Crane Workshop National Audubon Society Travernier FloridaUSA

Melvin S M R C Drewien S A Temple and E G Bizeau 1983 Leg-band attachment of radio transmitters for large birds Wildlife SocietyBulletin 11286ndash285

Minnesota Breeding Bird Atlas Project 2014 Minnesota Breeding BirdAtlas mnbbaorgcgi-binquerypl Accessed 14 Mar 2014

Nesbitt S A 1989 The significance of mate loss in Florida sandhill cranesWilson Bulletin 101648ndash651

Nesbitt S A and K S Williams 1990 Home range and habitat use ofFlorida sandhill cranes Journal of Wildlife Management 5492ndash96

Pearse A T D A Brandt W C Harrell K L Metzger D M Baaschand T J Hefley 2015 Whooping crane stopover site use intensitywithin the Great Plains US Geological Survey Open-File Report2015-1166 httpdxdoi org103133ofr20151166 Accessed 28 Dec2015

Tacha TC 1998 Social organization of sandhill cranes frommidcontinentalNorth America Wildlife Monographs 993ndash37

Tacha T C S A Nesbitt and P A Vohs 1992 Sandhill crane Account31 in A Poole P Stettenheim and F Gill editors The birds of NorthAmerica Academy of Natural Sciences Philadelphia Pennsylvania andAmerican Ornithological Union Washington DC USA

Tacha T C S A Nesbitt and P A Vohs 1994 Sandhill cranePages 77ndash94 in T C Tacha and C E Braun editors Migratoryshore and upland game bird management in North AmericaInternational Association of Fish and Wildlife Agencies Wash-ington DC USA

Van Horn K T White W Atkins T Cooper R Urbanek D HolmD Sherman D Aborn J Suckow K Cleveland and R Brook 2010Management plan for the Eastern Population of sandhill cranesMississippi and Atlantic Flyway Councils Madison Wisconsin USA

Walkinshaw L H 1949 Cranes of the worldWinchester Press NewYorkNew York USA

Walkinshaw L H 1960 Migration of the sandhill crane east of theMississippi River Wilson Bulletin 72358ndash384

Walkinshaw L H and H F Wing 1955 Censusing southern Michigansandhill crane Auk 72374ndash384

Warnock N 2010 Stopping vs staging the difference between a hop and ajump Journal of Avian Biology 41621ndash626

Wheeler R H and J C Lewis 1972 Trapping techniques for sandhillcrane studies in the Platte River Valley US Department of theInterior U S Fish and Wildlife Service Natural Resource Publication107 Washington DC USA

Associate Editor Courtney Amundson

Fronczak et al Eastern Population Sandhill Crane Migration 11

WA (Obion County) Tennessee Cranes that wintered inGeorgia and Florida made multiple stops in a migrationcorridor through northeastern Alabama and west-centralGeorgia to their respectivewinter areas (Fig3A)Therewere3(7) cranes that alternated their migration route around LakeMichigan to their respectivewinter termini at least onceduringthe study

Distribution of Winter AreasThe distribution of crane winter areas extended from Indianato Florida (Fig 2B) The majority of marked cranes (nfrac14 2662) wintered1 year in southeastern Tennessee within theHiwassee WR and surrounding Chickamauga Reservoir(Meigs Rhea and Hamilton counties) or in northwesternTennessee (Obion County) South of Tennessee 19 (45)cranes wintered throughout the Florida Peninsula (AlachuaBrevard Glades Highlands Lake Levy Marion OkaloosaOkeechobee Osceola Pasco Polk and Putnam counties)and 4 (10) wintered in Georgia (Brooks Crisp Floyd andPulaski counties) North of Tennessee 3 (7) craneswintered in Kentucky (Barren and Larue counties) and 12(29) wintered in Indiana (Greene Jackson JasperLaPorte and Starke counties) Eleven (26) cranes returnedto the same winter area in subsequent winters and 31 (74)wintered in 1 state at least once during the study period

Description of Spring MigrationMarked cranes used similar spring and fall migration routes(Fig 3)Cranes thatwintered inFlorida and southernGeorgiatraversed central and western Georgia and northeasternAlabama and staged either within theChickamaugaReservoir

(ie Hiwassee WR nfrac14 12 29) or migrated directly tostaging areas in north-central Tennessee central Kentuckyand southern Indiana Cranes that wintered in the Chick-amauga Reservoir used multiple staging areas in Kentuckyconcentrated in south-central Kentucky (nfrac14 17 40 Barrenand Metcalfe counties) and north-central Kentucky (nfrac14 2560 Hardin and Larue counties) Two cranes (5)that wintered in northwestern Tennessee (Obion County)migrated to Indiana staging areas either through southernIndiana or through eastern IllinoisCranes migrating north from central Kentucky moved to

southern Indiana staging areas in the White River Valley(nfrac14 27 64 Jackson County) and west-central Indianastaging area at Goose Pond FWA and surrounding area(nfrac14 13 31 Greene County) Most cranes (nfrac14 35 83)used stopover sites at the Jasper-Pulaski FWA andthroughout the Kankakee River Valley after departingsouthern Indiana and from there continued migration totheir respective summer areas Several cranes (nfrac14 10 24)moved from southern Indiana staging areas and bypassedJasper-Pulaski FWA and the Kankakee River Valley on totheir summer areasCranes that summeredoutsideofLowerPeninsulaMichigan

and north and east of LakeMichigan (nfrac14 16 38) followed aroute through centralMichigan continuing through either theStraits of Mackinaw or across Lake Huron (ie thosesummering on Manitoulin Island Ontario) Cranes thatsummered outside of Wisconsin and north and west of LakeMichigan migrated either through central Wisconsin (nfrac14 37) or through easternWisconsin and continued through the

Figure 3 Fall (A) and spring (B) migration routes for platform transmitting terminal-marked greater sandhill cranes 2010ndash2014

6 The Journal of Wildlife Management 9999()

Upper Peninsula of Michigan (nfrac14 7 17) Seven (17)cranes that summered on theUpper Peninsula ofMichigan innorth-central Ontario and east of Lake Michigan (NorthShore of Lake Huron and Manitoulin Island Ontario)migrated west of Lake Michigan and through the UpperPeninsula of Michigan at least once during the study periodThere were 6 (14) cranes that alternated their migrationroute aroundLakeMichigan to their respective summer area atleast once during the study

Chronology of Annual MovementsCranes on average arrived at their summer areas on 27March(nfrac14 97 rangefrac14 22 Febndash3 May Table 1) and stayed onsummer areas 189 days (nfrac14 75 SDfrac14 34 days) Averagedeparture from summer areas for staging areas was 5 October(nfrac14 85 rangefrac14 10 Augndash28 Dec) and cranes traveled onaverage 59 days (nfrac14 79 SDfrac14 37 days Table 2) fromsummer areas to winter termini Average cumulative distancetraveled between subsequent stops from summer to wintertermini during fall migration was 237 km (nfrac14 433 SDfrac14 177 km) Cranes used an average of 61 stops (nfrac14 75SDfrac14 28 stops) during fall migration on average 18(nfrac14 84 SDfrac14 10 stop) of these stops were at staging areasand 33 (nfrac14 89 SDfrac14 21 stops) were at stopover sites Weidentified 43 fall staging areas and 136 fall stopover sitesAverage number of days spent on individual staging areas was35 (nfrac14 79 SDfrac14 22 days) where 34 and 18 of fallmigration days were spent in the Kankakee River Valley andsouth-central Michigan respectively (Appendix A)Average arrival date of cranes at their winter areas was 11

December (nfrac14 86 rangefrac14 21 Sepndash27 Jan Table 3) and theyremained on winter areas an average of 67 days (nfrac14 85SDfrac14 33 days) Average departure date of cranes from winterareas was 16 February (nfrac14 102 rangefrac14 15 Janndash29Mar) and

cranes traveled on average 40 days (nfrac14 94 SDfrac14 18 daysTable 4) from winter areas to summer areas Averagedistance traveled between sequential stops prior to reachingsummer areas was approximately 195 km (nfrac14 652 SDfrac14 132 km) Cranes stopped on average 76 times (nfrac14 97SDfrac14 31 stops) during spring migration 46 (nfrac14 100SDfrac14 27 stops) of these stops were at stopover sites and 21(nfrac14 91 SDfrac14 10 stop) were at staging areas We identified43 spring staging areas and 172 spring stopover sitesAverage number of days spent on staging areas during springmigration was 15 (nfrac14 91 SDfrac14 6 days) where 34 and 9of the cumulative migration days were spent in the KankakeeRiver Valley northwestern Indiana and the White RiverValley southern Indiana respectively (Appendix B)

USFWS Fall SurveyApproximately 30 29 and 31 of crane locations wereoutside of areas surveyed during the 2010 2011 and 2012USFWS fall surveys respectively Crane locations that wereoutside surveyed areas included northern and east-centralWisconsin and central and north-central Michigan (Fig 4)However these cranes eventually moved to staging areas thatwould have been covered by observers after the fall surveyperiod All of the cranes that summered in Canada werepresent in surveyed locations during the survey period

DISCUSSION

Summer Area DistributionSummer areas used by marked cranes were widely distributedthroughout Minnesota Wisconsin Michigan and in theeastern portion of Ontario Canada and occurred within theprevious estimated breeding range of cranes in the EP (VanHorn et al 2010) Geographic distribution and migrationroutes that cranes used coincided with previously reported

Table 1 Mean arrival date length of stay and departure date on summer areas for platform transmitting terminal-marked greater sandhill cranes in theEastern Population North America 2010ndash2014

Arrival date Length of stay (days) Departure date

Yr n x SD (days) n x SD (days) n x SD (days)

2010 7 22 Mar 14 7 194 34 16 6 Oct 222011 32 30 Mar 16 30 185 32 30 6 Oct 282012 33 20 Mar 11 26 196 36 27 5 Oct 302013 17 3 Apr 15 9 176 34 9 2 Oct 252014 8 6 Apr 16 3 192 36 3 11Oct 31All years 97 27 Mar 15 75 189 34 85 5 Oct 27

Table 2 Average number of days number of stops and length of stay at staging areas (stop duration gt5 days) from summer areas to winter termini forplatform transmitting terminal-marked greater sandhill cranes in the Eastern Population North America 2010ndash2013

Total days No stops Length of stay (days)

Yra n x SD (days) n x SD (stops) n x SD (days)

2010 16 54 32 16 59 19 17 42 232011 30 58 36 29 59 26 29 36 222012 25 61 41 22 63 36 23 29 192013 8 72 40 8 68 32 7 33 13All years 79 59 37 75 61 28 79 35 22

a Summaries are unavailable for 2014

Fronczak et al Eastern Population Sandhill Crane Migration 7

geographic distributions (Walkinshaw 1949 Tacha et al1994) and migration routes (Walkinshaw 1960 Melvin andTemple 1982) However at smaller spatial scales some areasthought to support breeding cranes in the EP were notrepresented by our sample including northwestern Wis-consin-northeastern Minnesota (Cutright et al 2006Minnesota Breeding Bird Atlas Project 2014) and easternOntario including the eastern shore of Lake Huron and thesouthern peninsula of Ontario (Cadman et al 2007) Noneof the cranes in our sample overlapped the summer range ofcranes in theMCP identified by Krapu et al (2011) Summerarea locations of marked cranes were concentrated in east-central Wisconsin and the north shore of Lake HuronOntario which support relatively high densities of breedingcranes (Cadman et al 2007 Van Horn et al 2010)Cranes consistently returned to the same summer areas in

subsequent years similar to cranes in the Rocky MountainPopulation (Drewien 1973) and MCP (Krapu et al 2011)On average cranes in the EP and MCP (Krapu et al 2011)used subsequent summer areas within lt10 km of oneanother 65 of cranes in the EP used subsequent summerareas within 1 km of one another compared to 38 forcranes in the MCP (Krapu et al 2011) We observed 3instances when individual cranes moved an average lineardistance of 672 km (rangefrac14 251ndash989 km) among subse-quent summer areas and we speculate that these cranes mayhave been sub-adults (Bennett 1989 Nesbitt and Williams1990 Hayes 2015) However 2 of these cranes returned tothe same summer area in subsequent years and the thirdcrane died during the first summer

Migration Routes and ChronologyCranes followed migration routes similar to those identifiedin previous migration studies that used very high-frequency

transmitters in south-central Minnesota (Crete and Toepfer1978) and cranes marked with ARGOS Doppler PTTs onManitoulin Island Ontario Canada (S A Petrie BirdStudies Canada unpublished data) Unlike those studies wewere able to provide more precise information particularlyabout routes and locations where cranes stopped duringmigration We also observed that 8 (19) cranes withsummer areas in the Upper Peninsula of MichiganManitoulin Island Ontario and east-central Ontario usedmigration routes west of Lake Michigan instead of the moreintuitive route through the Lower Peninsula of MichiganSix of these cranes also alternated migration routes aroundLake Michigan during sequential migration (ie fall andspring) and in subsequent years Our study represents thefirst evidence of this behavior and use of both of thesemigration routes by individual cranes We also documentedcranes (nfrac14 8 19) that summered on Manitoulin Islandused migration routes over large bodies of water (LakeHuron and portions of Lake Michigan) similar to Krapuet al (2011) who described cranes crossing the Bering Straitof Alaska and the Arctic Ocean to and from breedinggroundsRecent observations of migratory cranes that wintered in

Louisiana led to speculation that EP cranes are potentiallyusing non-traditional migration corridors King et al(2010) monitored 2 PTT-marked cranes that wintered incentral Louisiana and migrated north through the Mis-sissippi Alluvial Valley toward summer areas within the EPsummer distribution One marked crane in their study useda migration route through west-central Illinois to south-western Wisconsin and then into the Upper Peninsula ofMichigan after which they were unable to receiveadditional location information from the transmitter Theother crane traveled a similar route used by some of the

Table 3 Mean arrival date length of stay and departure date on winter areas for platform transmitting terminal-marked greater sandhill cranes in theEastern Population North America 2009ndash2014

Arrival datea Length of stay (days)a Departure date

Yr n x SD (days) n x SD (days) n x SD (days)

2009ndash2010 6 16 Feb 62010ndash2011 24 10 Dec 13 26 66 17 35 18 Feb 92011ndash2012 31 14 Dec 28 31 65 32 34 15 Feb 152012ndash2013 22 6 Dec 34 20 74 49 19 19 Feb 222013ndash2014 8 14 Dec 20 7 63 29 8 14 Feb 17All years 86 11 Dec 26 85 67 33 102 16 Feb 14

a Summaries unavailable for 2009ndash2010 winter locations

Table 4 Average number of days number of stops and length of stay at staging areas (stop duration gt5 days) from winter termini to summer areas forplatform transmitting terminal-marked greater sandhill cranes in the Eastern Population North America 2010ndash2014

Total days No stops Length of stay (days)

Yr n x SD (days) n x SD (stops) n x SD (days)

2010 6 39 11 6 68 08 6 12 12011 33 42 16 34 84 30 33 14 42012 33 32 15 33 71 30 29 17 82013 16 41 22 17 71 40 15 17 72014 6 62 11 7 77 25 8 14 4All years 94 39 18 97 76 31 91 15 6

8 The Journal of Wildlife Management 9999()

cranes in our sample through west-central Indiana (GoosePond FWA) northwestern Indiana (Kankakee RiverValley) east-central Wisconsin and then through theUpper Peninsula of Michigan to a summer area in north-central OntarioA single PTT-marked crane in our sample from the

Manitoulin Island Ontario study site migrated west fromthe White River Valley Indiana (Jackson County) andwintered at the Hop-In WR Tennessee (Obion County)However that crane did not return to Hop-In WR in thesubsequent year In anticipation of identifying a potentialwestern migration corridor to Louisiana we affixed 2 PTTsto cranes that wintered at Hop-In WR However only 1returned to winter at the Hop-InWR and the other winteredin Florida the following year

Cranes consistently used staging areas along migrationroutes but made fewer stops for a longer duration during fallmigration than during spring migration Locations wherecranes stopped during migration along the north shore ofLake Huron Algoma District Ontario Lake Poygon east-central Wisconsin the Kankakee River Valley northwesternIndiana and counties in south-central Michigan werefrequented more by cranes during fall migration (Fig 3AAppendix A) Cranes used Hardin County north-centralKentucky White River Valley (Ewing Bottoms JacksonCounty) south-central Indiana and the Kankakee RiverValley northwestern Indiana more frequently during springthan during fall migration (Fig 3B Appendix B) Theseareas support shallow-marsh roosting habitat have anabundant food supply from agricultural production and

Figure 4 Locations of areas surveyed (gray squares) during the United States Fish and Wildlife Service Fall Eastern Population Sandhill Crane Survey alsoshown are locations (black triangles) where platform transmitting terminal-marked sandhill cranes were not observed during the annual surveys 2010ndash2012

Fronczak et al Eastern Population Sandhill Crane Migration 9

are more likely to be protected from human disturbances (iehunting and sporting activities) The Kankakee River Valleylocated in northwestern Indiana was the most frequented ofall staging areas during fall and spring migration wherecranes spent 34 of migration days

Winter Area DistributionCrane winter areas extended from Indiana to Florida wellnorth of the areas previously described by Tacha et al(1994) who reported that cranes in the EP winteredmainly in southeastern Georgia and in Florida Craneswintered in Florida Georgia Tennessee Kentucky andIndiana with the Chickamauga Reservoir (specificallyHiwassee WR) Tennessee having the highest proportionof use among all winter areas The proportion of cranesthat used the Chickamauga Reservoir Valley for thewinters of 2009 and 2010 may be biased high because wetrapped the majority of cranes (nfrac14 6 14 2009ndash2010nfrac14 12 28 2010ndash2011) at the Hiwassee WR duringDecember and January of those years However in 2010MWS estimates for the Chickamauga Reservoir indicateda record number of wintering cranes (nfrac14 48300 J RKelley USFWS-Migratory Bird Management personalcommunication) suggesting that a large portion of cranesin the EP likely used that area Furthermore many ofthe cranes we captured in other locations used theChickamauga Reservoir during migration in years follow-ing their capture The Chickamauga Reservoir is ideal forstaging and wintering cranes Agriculture crops (corn soybeans and winter wheat) are produced annually forwintering wildlife and adjacent sand bars and low waterlevels on Chickamauga Reservoir provided roostinghabitat and sanctuary for sandhill cranes during the falland winter months (R Klippel personal communication)Hunting for sandhill cranes is allowed throughout thereservoir however the Hiwassee WR is designated a non-hunting zoneRecent analysis of long-term Christmas Bird Count data

indicated that cranes in the EP wintered at staging areasnorth of Georgia and Florida in recent decades (Amundsonand Johnson 2011 Lacy et al 2015) Lacy et al (2015)suggested that climate and increased agriculture throughoutthe EP range may influence cranes to winter farther northHowever there has been no formal analysis to demonstratethat these factors are main contributors to a more northerlywinter strategy Thirty-one (74) cranes in our sample thatwintered north of Georgia and Florida used migrationstaging areas in Indiana Kentucky and Tennessee Winterdestinations for cranes varied among years where 11 (26)returned to the same winter area in subsequent winters Theremaining 31 (74) cranes wintered in1 state at least onceduring the study period indicating a variation in migrationstrategy possibly influencing cranes to settle in a particularwinter areaEight (19) cranes used the Kankakee River Valley in

northwestern Indiana as a winter area in1 year These cranesthat wintered in the Kankakee River Valley used the Jasper-Pulaski FWA until all available roosting sites froze

then roosted within the Schahfer Generating Stationproperty in Wheatfield Indiana about 10 km north-northwest of Jasper-Pulaski FWA The power stationprovided roosting areas that remained ice-free and thesurrounding agricultural landscape provided waste grainthroughout the winter (J Bergens personal communica-tion) These cranes then used areas throughout theKankakee River Valley in proximity to Jasper-PulaskiFWA as the winter progressedAdditional winter locations outside of traditional areas are

not well understood Location data from King et al (2010)suggested that cranes in the EP used winter areas in northernLouisiana and 3 cranes from our sample wintered at theHop-In WR Tennessee west of previously reportedwintering areas for cranes in the EP Observations duringthe MWS of cranes at the Wheeler NWR Alabama havesteadily increased since 2003 with an average of 4583 cranes(rangefrac14 293ndash12032) and a peak count of 12032 cranes in2013 Furthermore observations of cranes in Arkansas andMississippi along theMississippi Alluvial Valley are frequentduring the MWS (J R Kelley personal communication)These cranes likely are part of the EP and most likely usedthe Hop-In WR as a staging area

MANAGEMENT IMPLICATIONS

Location and chronology data from our study can be used toadjust timing of hunting seasons to potentially avoid highharvest pressure on local breeding adults and focus harvestpressure on migrants Location and chronology data canalso be used to identify specific locality information forpotential land conservation and opportunity for non-consumptive recreational activities (eg crane festivalsand bird watching)We estimated a proportion of cranes in the EP were located

in areas not currently covered by the USFWS fall surveyduring the survey period If the absence of these cranes in theUSFWS fall survey area is similar through time series ofcounts from surveys will likely provide a reasonable estimateof population trend Alternatively if the proportion of cranesnot present in survey areas changes it may be necessary toredesign the survey by moving the survey period later into thefall or early winter when a larger portion of cranes areassociated with known concentration areas Howeverthe effects of moving the survey period would need to beevaluated to avoid issues related to 1) comparability toprevious surveys and 2) cranes arriving on wintering areas inFlorida where it is difficult to distinguish cranes in theEP from Florida non-migratory greater sandhill cranes(A c pratensis)

ACKNOWLEDGMENTS

Any use of trade firm or product names is for descriptivepurposes only and does not imply endorsement by the USGovernment or the University of Minnesota We thankD A Brandt P Link R Klippel J Jackson B Swiney SShelby J Womac B Layton D E Sherman B Feaster APhelps R Ronk J Bergens J Gilbert E E Carroll RKnopick W Delks V Clarkston E Hockman C Trosen

10 The Journal of Wildlife Management 9999()

A Johnson C Kanke P Peterson S Zudrow TPaulson W Tamminga J Brunjes E Harper LArmstrong T White M Morse S Bossuyt M BossuytA Dhamorikar J E Hanna KWeaver and S A Petrie forfield assistance We thank A T Pearse for comments onprevious drafts of this manuscript The project was supportedthrough Research Work Order no 86 at the USGSMinnesota Cooperative Fish and Wildlife Research Unitat the University of Minnesota This research was supportedby the USFWS Webless Migratory Game Bird Programthe USGS Northern Prairie Wildlife Research Center theMinnesota Cooperative Fish and Wildlife Research Unitthe USFWS Region 3Migratory BirdManagement OfficeLong Point Waterfowl SC Johnsons and Son Ltdthe Ontario Federation of Anglers and Hunters theCanadian Wildlife Service Wildlife Habitat Canada andthe TD Friends of the Environment Foundation

LITERATURE CITEDAmundson C L and D H Johnson 2011 Assessment of the EasternPopulation of greater sandhill cranes (Grus canadensis tabida) FallMigrationSurvey 1979ndash2009 US Fish and Wildlife Service BloomingtonMinnesota USA

Bennett A J 1989 Movements and home ranges of Florida sandhill cranesJournal of Wildlife Management 53830ndash836

Cadman M D D A Sutherland G G Beck D Lepage and A RCouturier 2007 Ontario Breeding Bird Atlas website httpwwwbirdsontarioorgatlasindexjsp Accessed 14 Mar 2014

CreteRA andJEToepfer1978Migrationof radio-taggedeasterngreatersandhill cranes US Fish and Wildlife Service Twin Cities MinnesotaUSA

Cutright N J B R Harriman and R W Howe 2006 WisconsinBreeding Bird Atlas httpuwgbedubirdswbbaindexhtm Accessed 14Mar 2014

Case D J and S J Sanders editors 2009 Priority information needs forsandhill cranes a funding strategy Association of Fish and WildlifeAgencies Washington DC USA

Douglas D C R Weinzierl S C Davidson R Kays M Wikelski andG Bohrer 2012 Moderating ARGOS location errors in animal trackingdata Methods in Ecology and Evolution 3999ndash1007

Drewien R C 1973 Ecology of Rocky Mountain greater sandhill cranesDissertation University of Idaho Moscow USA

Ellis D H P W Howey and G L Krapu 2001 Recommendations forthe attachment of satellite transmitters to cranes North American CraneWorkshop Proceedings 8211ndash212

Fronczak D L 2014 Distribution migration chronology and survival ratesof Eastern Population sandhill crane Thesis University of Minnesota StPaul USA

Fronczak D L D E Andersen E E Hanna and T R Cooper 2015Annual survival rate estimate of satellite transmitter-marked EasternPopulation greater sandhill cranes Journal of Fish and WildlifeManagement 6464ndash471

Hanna E E M L Schummer and S A Petrie 2014 Migratorychronology autumn recruitment and population size of EasternPopulation sandhill crane (Grus canadensis) from the North Shore Regionof Lake Huron Ontario Canada CanadianWildlife Service and OntarioMinistry of Natural Resources Port Rowan Ontario Canada

Hayes M A and J A Barzen 2006 Dynamics of breeding and non-breeding sandhill cranes in south-central Wisconsin Passenger Pigeon68345ndash352

Hayes M A 2015 Dispersal and population genetic structure in twoflyways of sandhill cranes (Grus canadensis) Dissertation University ofWisconsin-Madison Madison USA

Hunt R A E A Gluesing and L E Nauman 1976 The sandhill crane inWisconsin Wisconsin Department of Natural Resources Madison USA

Jones K L G L Krapu D A Brandt andMV Ashley 2005 Populationgenetic structure in migratory sandhill cranes and the role of Pleistoceneglaciations Molecular Ecology 142645ndash2657

King D T J D Paulson D J Leblanc and K Bruce 1998 Two capturetechniques for American white pelicans and great blue herons ColonialWaterbirds 21258ndash260

King S L A R Pierce K R Hersey and N Winstead 2010 Migrationpatterns and movements of sandhill cranes wintering in central andsouthwest Louisiana Proceedings of the North American CraneWorkshop 1157ndash61

Krapu G L D A Brandt K L Jones and D H Johnson 2011Geographic distribution of the Mid-continent Population of sandhillcranes and related management applications Wildlife Monographs1751ndash38

Kruse K L and J A Dubovsky 2015 Status and harvests of sandhillcranes Mid-continent Rocky Mountain Lower Colorado River Valleyand Eastern Populations US Fish and Wildlife Service LakewoodColorado USA

Lacy A E J A Barzen D MMoore and K E Norris 2015 Changes inthe number and distribution of greater sandhill cranes in the EasternPopulation Journal Field Ornithology 86317ndash325

Lumsden H G 1971 The status of the sandhill crane in northern OntarioCanadian Field-Naturalist 85285ndash293

Melvin S M and S A Temple 1982 Migration ecology of sandhillcranes a review Pages 73ndash87 in J C Lewis editor Proceedings of the1981 Crane Workshop National Audubon Society Travernier FloridaUSA

Melvin S M R C Drewien S A Temple and E G Bizeau 1983 Leg-band attachment of radio transmitters for large birds Wildlife SocietyBulletin 11286ndash285

Minnesota Breeding Bird Atlas Project 2014 Minnesota Breeding BirdAtlas mnbbaorgcgi-binquerypl Accessed 14 Mar 2014

Nesbitt S A 1989 The significance of mate loss in Florida sandhill cranesWilson Bulletin 101648ndash651

Nesbitt S A and K S Williams 1990 Home range and habitat use ofFlorida sandhill cranes Journal of Wildlife Management 5492ndash96

Pearse A T D A Brandt W C Harrell K L Metzger D M Baaschand T J Hefley 2015 Whooping crane stopover site use intensitywithin the Great Plains US Geological Survey Open-File Report2015-1166 httpdxdoi org103133ofr20151166 Accessed 28 Dec2015

Tacha TC 1998 Social organization of sandhill cranes frommidcontinentalNorth America Wildlife Monographs 993ndash37

Tacha T C S A Nesbitt and P A Vohs 1992 Sandhill crane Account31 in A Poole P Stettenheim and F Gill editors The birds of NorthAmerica Academy of Natural Sciences Philadelphia Pennsylvania andAmerican Ornithological Union Washington DC USA

Tacha T C S A Nesbitt and P A Vohs 1994 Sandhill cranePages 77ndash94 in T C Tacha and C E Braun editors Migratoryshore and upland game bird management in North AmericaInternational Association of Fish and Wildlife Agencies Wash-ington DC USA

Van Horn K T White W Atkins T Cooper R Urbanek D HolmD Sherman D Aborn J Suckow K Cleveland and R Brook 2010Management plan for the Eastern Population of sandhill cranesMississippi and Atlantic Flyway Councils Madison Wisconsin USA

Walkinshaw L H 1949 Cranes of the worldWinchester Press NewYorkNew York USA

Walkinshaw L H 1960 Migration of the sandhill crane east of theMississippi River Wilson Bulletin 72358ndash384

Walkinshaw L H and H F Wing 1955 Censusing southern Michigansandhill crane Auk 72374ndash384

Warnock N 2010 Stopping vs staging the difference between a hop and ajump Journal of Avian Biology 41621ndash626

Wheeler R H and J C Lewis 1972 Trapping techniques for sandhillcrane studies in the Platte River Valley US Department of theInterior U S Fish and Wildlife Service Natural Resource Publication107 Washington DC USA

Associate Editor Courtney Amundson

Fronczak et al Eastern Population Sandhill Crane Migration 11

Upper Peninsula of Michigan (nfrac14 7 17) Seven (17)cranes that summered on theUpper Peninsula ofMichigan innorth-central Ontario and east of Lake Michigan (NorthShore of Lake Huron and Manitoulin Island Ontario)migrated west of Lake Michigan and through the UpperPeninsula of Michigan at least once during the study periodThere were 6 (14) cranes that alternated their migrationroute aroundLakeMichigan to their respective summer area atleast once during the study

Chronology of Annual MovementsCranes on average arrived at their summer areas on 27March(nfrac14 97 rangefrac14 22 Febndash3 May Table 1) and stayed onsummer areas 189 days (nfrac14 75 SDfrac14 34 days) Averagedeparture from summer areas for staging areas was 5 October(nfrac14 85 rangefrac14 10 Augndash28 Dec) and cranes traveled onaverage 59 days (nfrac14 79 SDfrac14 37 days Table 2) fromsummer areas to winter termini Average cumulative distancetraveled between subsequent stops from summer to wintertermini during fall migration was 237 km (nfrac14 433 SDfrac14 177 km) Cranes used an average of 61 stops (nfrac14 75SDfrac14 28 stops) during fall migration on average 18(nfrac14 84 SDfrac14 10 stop) of these stops were at staging areasand 33 (nfrac14 89 SDfrac14 21 stops) were at stopover sites Weidentified 43 fall staging areas and 136 fall stopover sitesAverage number of days spent on individual staging areas was35 (nfrac14 79 SDfrac14 22 days) where 34 and 18 of fallmigration days were spent in the Kankakee River Valley andsouth-central Michigan respectively (Appendix A)Average arrival date of cranes at their winter areas was 11

December (nfrac14 86 rangefrac14 21 Sepndash27 Jan Table 3) and theyremained on winter areas an average of 67 days (nfrac14 85SDfrac14 33 days) Average departure date of cranes from winterareas was 16 February (nfrac14 102 rangefrac14 15 Janndash29Mar) and

cranes traveled on average 40 days (nfrac14 94 SDfrac14 18 daysTable 4) from winter areas to summer areas Averagedistance traveled between sequential stops prior to reachingsummer areas was approximately 195 km (nfrac14 652 SDfrac14 132 km) Cranes stopped on average 76 times (nfrac14 97SDfrac14 31 stops) during spring migration 46 (nfrac14 100SDfrac14 27 stops) of these stops were at stopover sites and 21(nfrac14 91 SDfrac14 10 stop) were at staging areas We identified43 spring staging areas and 172 spring stopover sitesAverage number of days spent on staging areas during springmigration was 15 (nfrac14 91 SDfrac14 6 days) where 34 and 9of the cumulative migration days were spent in the KankakeeRiver Valley northwestern Indiana and the White RiverValley southern Indiana respectively (Appendix B)

USFWS Fall SurveyApproximately 30 29 and 31 of crane locations wereoutside of areas surveyed during the 2010 2011 and 2012USFWS fall surveys respectively Crane locations that wereoutside surveyed areas included northern and east-centralWisconsin and central and north-central Michigan (Fig 4)However these cranes eventually moved to staging areas thatwould have been covered by observers after the fall surveyperiod All of the cranes that summered in Canada werepresent in surveyed locations during the survey period

DISCUSSION

Summer Area DistributionSummer areas used by marked cranes were widely distributedthroughout Minnesota Wisconsin Michigan and in theeastern portion of Ontario Canada and occurred within theprevious estimated breeding range of cranes in the EP (VanHorn et al 2010) Geographic distribution and migrationroutes that cranes used coincided with previously reported

Table 1 Mean arrival date length of stay and departure date on summer areas for platform transmitting terminal-marked greater sandhill cranes in theEastern Population North America 2010ndash2014

Arrival date Length of stay (days) Departure date

Yr n x SD (days) n x SD (days) n x SD (days)

2010 7 22 Mar 14 7 194 34 16 6 Oct 222011 32 30 Mar 16 30 185 32 30 6 Oct 282012 33 20 Mar 11 26 196 36 27 5 Oct 302013 17 3 Apr 15 9 176 34 9 2 Oct 252014 8 6 Apr 16 3 192 36 3 11Oct 31All years 97 27 Mar 15 75 189 34 85 5 Oct 27

Table 2 Average number of days number of stops and length of stay at staging areas (stop duration gt5 days) from summer areas to winter termini forplatform transmitting terminal-marked greater sandhill cranes in the Eastern Population North America 2010ndash2013

Total days No stops Length of stay (days)

Yra n x SD (days) n x SD (stops) n x SD (days)

2010 16 54 32 16 59 19 17 42 232011 30 58 36 29 59 26 29 36 222012 25 61 41 22 63 36 23 29 192013 8 72 40 8 68 32 7 33 13All years 79 59 37 75 61 28 79 35 22

a Summaries are unavailable for 2014

Fronczak et al Eastern Population Sandhill Crane Migration 7

geographic distributions (Walkinshaw 1949 Tacha et al1994) and migration routes (Walkinshaw 1960 Melvin andTemple 1982) However at smaller spatial scales some areasthought to support breeding cranes in the EP were notrepresented by our sample including northwestern Wis-consin-northeastern Minnesota (Cutright et al 2006Minnesota Breeding Bird Atlas Project 2014) and easternOntario including the eastern shore of Lake Huron and thesouthern peninsula of Ontario (Cadman et al 2007) Noneof the cranes in our sample overlapped the summer range ofcranes in theMCP identified by Krapu et al (2011) Summerarea locations of marked cranes were concentrated in east-central Wisconsin and the north shore of Lake HuronOntario which support relatively high densities of breedingcranes (Cadman et al 2007 Van Horn et al 2010)Cranes consistently returned to the same summer areas in

subsequent years similar to cranes in the Rocky MountainPopulation (Drewien 1973) and MCP (Krapu et al 2011)On average cranes in the EP and MCP (Krapu et al 2011)used subsequent summer areas within lt10 km of oneanother 65 of cranes in the EP used subsequent summerareas within 1 km of one another compared to 38 forcranes in the MCP (Krapu et al 2011) We observed 3instances when individual cranes moved an average lineardistance of 672 km (rangefrac14 251ndash989 km) among subse-quent summer areas and we speculate that these cranes mayhave been sub-adults (Bennett 1989 Nesbitt and Williams1990 Hayes 2015) However 2 of these cranes returned tothe same summer area in subsequent years and the thirdcrane died during the first summer

Migration Routes and ChronologyCranes followed migration routes similar to those identifiedin previous migration studies that used very high-frequency

transmitters in south-central Minnesota (Crete and Toepfer1978) and cranes marked with ARGOS Doppler PTTs onManitoulin Island Ontario Canada (S A Petrie BirdStudies Canada unpublished data) Unlike those studies wewere able to provide more precise information particularlyabout routes and locations where cranes stopped duringmigration We also observed that 8 (19) cranes withsummer areas in the Upper Peninsula of MichiganManitoulin Island Ontario and east-central Ontario usedmigration routes west of Lake Michigan instead of the moreintuitive route through the Lower Peninsula of MichiganSix of these cranes also alternated migration routes aroundLake Michigan during sequential migration (ie fall andspring) and in subsequent years Our study represents thefirst evidence of this behavior and use of both of thesemigration routes by individual cranes We also documentedcranes (nfrac14 8 19) that summered on Manitoulin Islandused migration routes over large bodies of water (LakeHuron and portions of Lake Michigan) similar to Krapuet al (2011) who described cranes crossing the Bering Straitof Alaska and the Arctic Ocean to and from breedinggroundsRecent observations of migratory cranes that wintered in

Louisiana led to speculation that EP cranes are potentiallyusing non-traditional migration corridors King et al(2010) monitored 2 PTT-marked cranes that wintered incentral Louisiana and migrated north through the Mis-sissippi Alluvial Valley toward summer areas within the EPsummer distribution One marked crane in their study useda migration route through west-central Illinois to south-western Wisconsin and then into the Upper Peninsula ofMichigan after which they were unable to receiveadditional location information from the transmitter Theother crane traveled a similar route used by some of the

Table 3 Mean arrival date length of stay and departure date on winter areas for platform transmitting terminal-marked greater sandhill cranes in theEastern Population North America 2009ndash2014

Arrival datea Length of stay (days)a Departure date

Yr n x SD (days) n x SD (days) n x SD (days)

2009ndash2010 6 16 Feb 62010ndash2011 24 10 Dec 13 26 66 17 35 18 Feb 92011ndash2012 31 14 Dec 28 31 65 32 34 15 Feb 152012ndash2013 22 6 Dec 34 20 74 49 19 19 Feb 222013ndash2014 8 14 Dec 20 7 63 29 8 14 Feb 17All years 86 11 Dec 26 85 67 33 102 16 Feb 14

a Summaries unavailable for 2009ndash2010 winter locations

Table 4 Average number of days number of stops and length of stay at staging areas (stop duration gt5 days) from winter termini to summer areas forplatform transmitting terminal-marked greater sandhill cranes in the Eastern Population North America 2010ndash2014

Total days No stops Length of stay (days)

Yr n x SD (days) n x SD (stops) n x SD (days)

2010 6 39 11 6 68 08 6 12 12011 33 42 16 34 84 30 33 14 42012 33 32 15 33 71 30 29 17 82013 16 41 22 17 71 40 15 17 72014 6 62 11 7 77 25 8 14 4All years 94 39 18 97 76 31 91 15 6

8 The Journal of Wildlife Management 9999()

cranes in our sample through west-central Indiana (GoosePond FWA) northwestern Indiana (Kankakee RiverValley) east-central Wisconsin and then through theUpper Peninsula of Michigan to a summer area in north-central OntarioA single PTT-marked crane in our sample from the

Manitoulin Island Ontario study site migrated west fromthe White River Valley Indiana (Jackson County) andwintered at the Hop-In WR Tennessee (Obion County)However that crane did not return to Hop-In WR in thesubsequent year In anticipation of identifying a potentialwestern migration corridor to Louisiana we affixed 2 PTTsto cranes that wintered at Hop-In WR However only 1returned to winter at the Hop-InWR and the other winteredin Florida the following year

Cranes consistently used staging areas along migrationroutes but made fewer stops for a longer duration during fallmigration than during spring migration Locations wherecranes stopped during migration along the north shore ofLake Huron Algoma District Ontario Lake Poygon east-central Wisconsin the Kankakee River Valley northwesternIndiana and counties in south-central Michigan werefrequented more by cranes during fall migration (Fig 3AAppendix A) Cranes used Hardin County north-centralKentucky White River Valley (Ewing Bottoms JacksonCounty) south-central Indiana and the Kankakee RiverValley northwestern Indiana more frequently during springthan during fall migration (Fig 3B Appendix B) Theseareas support shallow-marsh roosting habitat have anabundant food supply from agricultural production and

Figure 4 Locations of areas surveyed (gray squares) during the United States Fish and Wildlife Service Fall Eastern Population Sandhill Crane Survey alsoshown are locations (black triangles) where platform transmitting terminal-marked sandhill cranes were not observed during the annual surveys 2010ndash2012

Fronczak et al Eastern Population Sandhill Crane Migration 9

are more likely to be protected from human disturbances (iehunting and sporting activities) The Kankakee River Valleylocated in northwestern Indiana was the most frequented ofall staging areas during fall and spring migration wherecranes spent 34 of migration days

Winter Area DistributionCrane winter areas extended from Indiana to Florida wellnorth of the areas previously described by Tacha et al(1994) who reported that cranes in the EP winteredmainly in southeastern Georgia and in Florida Craneswintered in Florida Georgia Tennessee Kentucky andIndiana with the Chickamauga Reservoir (specificallyHiwassee WR) Tennessee having the highest proportionof use among all winter areas The proportion of cranesthat used the Chickamauga Reservoir Valley for thewinters of 2009 and 2010 may be biased high because wetrapped the majority of cranes (nfrac14 6 14 2009ndash2010nfrac14 12 28 2010ndash2011) at the Hiwassee WR duringDecember and January of those years However in 2010MWS estimates for the Chickamauga Reservoir indicateda record number of wintering cranes (nfrac14 48300 J RKelley USFWS-Migratory Bird Management personalcommunication) suggesting that a large portion of cranesin the EP likely used that area Furthermore many ofthe cranes we captured in other locations used theChickamauga Reservoir during migration in years follow-ing their capture The Chickamauga Reservoir is ideal forstaging and wintering cranes Agriculture crops (corn soybeans and winter wheat) are produced annually forwintering wildlife and adjacent sand bars and low waterlevels on Chickamauga Reservoir provided roostinghabitat and sanctuary for sandhill cranes during the falland winter months (R Klippel personal communication)Hunting for sandhill cranes is allowed throughout thereservoir however the Hiwassee WR is designated a non-hunting zoneRecent analysis of long-term Christmas Bird Count data

indicated that cranes in the EP wintered at staging areasnorth of Georgia and Florida in recent decades (Amundsonand Johnson 2011 Lacy et al 2015) Lacy et al (2015)suggested that climate and increased agriculture throughoutthe EP range may influence cranes to winter farther northHowever there has been no formal analysis to demonstratethat these factors are main contributors to a more northerlywinter strategy Thirty-one (74) cranes in our sample thatwintered north of Georgia and Florida used migrationstaging areas in Indiana Kentucky and Tennessee Winterdestinations for cranes varied among years where 11 (26)returned to the same winter area in subsequent winters Theremaining 31 (74) cranes wintered in1 state at least onceduring the study period indicating a variation in migrationstrategy possibly influencing cranes to settle in a particularwinter areaEight (19) cranes used the Kankakee River Valley in

northwestern Indiana as a winter area in1 year These cranesthat wintered in the Kankakee River Valley used the Jasper-Pulaski FWA until all available roosting sites froze

then roosted within the Schahfer Generating Stationproperty in Wheatfield Indiana about 10 km north-northwest of Jasper-Pulaski FWA The power stationprovided roosting areas that remained ice-free and thesurrounding agricultural landscape provided waste grainthroughout the winter (J Bergens personal communica-tion) These cranes then used areas throughout theKankakee River Valley in proximity to Jasper-PulaskiFWA as the winter progressedAdditional winter locations outside of traditional areas are

not well understood Location data from King et al (2010)suggested that cranes in the EP used winter areas in northernLouisiana and 3 cranes from our sample wintered at theHop-In WR Tennessee west of previously reportedwintering areas for cranes in the EP Observations duringthe MWS of cranes at the Wheeler NWR Alabama havesteadily increased since 2003 with an average of 4583 cranes(rangefrac14 293ndash12032) and a peak count of 12032 cranes in2013 Furthermore observations of cranes in Arkansas andMississippi along theMississippi Alluvial Valley are frequentduring the MWS (J R Kelley personal communication)These cranes likely are part of the EP and most likely usedthe Hop-In WR as a staging area

MANAGEMENT IMPLICATIONS

Location and chronology data from our study can be used toadjust timing of hunting seasons to potentially avoid highharvest pressure on local breeding adults and focus harvestpressure on migrants Location and chronology data canalso be used to identify specific locality information forpotential land conservation and opportunity for non-consumptive recreational activities (eg crane festivalsand bird watching)We estimated a proportion of cranes in the EP were located

in areas not currently covered by the USFWS fall surveyduring the survey period If the absence of these cranes in theUSFWS fall survey area is similar through time series ofcounts from surveys will likely provide a reasonable estimateof population trend Alternatively if the proportion of cranesnot present in survey areas changes it may be necessary toredesign the survey by moving the survey period later into thefall or early winter when a larger portion of cranes areassociated with known concentration areas Howeverthe effects of moving the survey period would need to beevaluated to avoid issues related to 1) comparability toprevious surveys and 2) cranes arriving on wintering areas inFlorida where it is difficult to distinguish cranes in theEP from Florida non-migratory greater sandhill cranes(A c pratensis)

ACKNOWLEDGMENTS

Any use of trade firm or product names is for descriptivepurposes only and does not imply endorsement by the USGovernment or the University of Minnesota We thankD A Brandt P Link R Klippel J Jackson B Swiney SShelby J Womac B Layton D E Sherman B Feaster APhelps R Ronk J Bergens J Gilbert E E Carroll RKnopick W Delks V Clarkston E Hockman C Trosen

10 The Journal of Wildlife Management 9999()

A Johnson C Kanke P Peterson S Zudrow TPaulson W Tamminga J Brunjes E Harper LArmstrong T White M Morse S Bossuyt M BossuytA Dhamorikar J E Hanna KWeaver and S A Petrie forfield assistance We thank A T Pearse for comments onprevious drafts of this manuscript The project was supportedthrough Research Work Order no 86 at the USGSMinnesota Cooperative Fish and Wildlife Research Unitat the University of Minnesota This research was supportedby the USFWS Webless Migratory Game Bird Programthe USGS Northern Prairie Wildlife Research Center theMinnesota Cooperative Fish and Wildlife Research Unitthe USFWS Region 3Migratory BirdManagement OfficeLong Point Waterfowl SC Johnsons and Son Ltdthe Ontario Federation of Anglers and Hunters theCanadian Wildlife Service Wildlife Habitat Canada andthe TD Friends of the Environment Foundation

LITERATURE CITEDAmundson C L and D H Johnson 2011 Assessment of the EasternPopulation of greater sandhill cranes (Grus canadensis tabida) FallMigrationSurvey 1979ndash2009 US Fish and Wildlife Service BloomingtonMinnesota USA

Bennett A J 1989 Movements and home ranges of Florida sandhill cranesJournal of Wildlife Management 53830ndash836

Cadman M D D A Sutherland G G Beck D Lepage and A RCouturier 2007 Ontario Breeding Bird Atlas website httpwwwbirdsontarioorgatlasindexjsp Accessed 14 Mar 2014

CreteRA andJEToepfer1978Migrationof radio-taggedeasterngreatersandhill cranes US Fish and Wildlife Service Twin Cities MinnesotaUSA

Cutright N J B R Harriman and R W Howe 2006 WisconsinBreeding Bird Atlas httpuwgbedubirdswbbaindexhtm Accessed 14Mar 2014

Case D J and S J Sanders editors 2009 Priority information needs forsandhill cranes a funding strategy Association of Fish and WildlifeAgencies Washington DC USA

Douglas D C R Weinzierl S C Davidson R Kays M Wikelski andG Bohrer 2012 Moderating ARGOS location errors in animal trackingdata Methods in Ecology and Evolution 3999ndash1007

Drewien R C 1973 Ecology of Rocky Mountain greater sandhill cranesDissertation University of Idaho Moscow USA

Ellis D H P W Howey and G L Krapu 2001 Recommendations forthe attachment of satellite transmitters to cranes North American CraneWorkshop Proceedings 8211ndash212

Fronczak D L 2014 Distribution migration chronology and survival ratesof Eastern Population sandhill crane Thesis University of Minnesota StPaul USA

Fronczak D L D E Andersen E E Hanna and T R Cooper 2015Annual survival rate estimate of satellite transmitter-marked EasternPopulation greater sandhill cranes Journal of Fish and WildlifeManagement 6464ndash471

Hanna E E M L Schummer and S A Petrie 2014 Migratorychronology autumn recruitment and population size of EasternPopulation sandhill crane (Grus canadensis) from the North Shore Regionof Lake Huron Ontario Canada CanadianWildlife Service and OntarioMinistry of Natural Resources Port Rowan Ontario Canada

Hayes M A and J A Barzen 2006 Dynamics of breeding and non-breeding sandhill cranes in south-central Wisconsin Passenger Pigeon68345ndash352

Hayes M A 2015 Dispersal and population genetic structure in twoflyways of sandhill cranes (Grus canadensis) Dissertation University ofWisconsin-Madison Madison USA

Hunt R A E A Gluesing and L E Nauman 1976 The sandhill crane inWisconsin Wisconsin Department of Natural Resources Madison USA

Jones K L G L Krapu D A Brandt andMV Ashley 2005 Populationgenetic structure in migratory sandhill cranes and the role of Pleistoceneglaciations Molecular Ecology 142645ndash2657

King D T J D Paulson D J Leblanc and K Bruce 1998 Two capturetechniques for American white pelicans and great blue herons ColonialWaterbirds 21258ndash260

King S L A R Pierce K R Hersey and N Winstead 2010 Migrationpatterns and movements of sandhill cranes wintering in central andsouthwest Louisiana Proceedings of the North American CraneWorkshop 1157ndash61

Krapu G L D A Brandt K L Jones and D H Johnson 2011Geographic distribution of the Mid-continent Population of sandhillcranes and related management applications Wildlife Monographs1751ndash38

Kruse K L and J A Dubovsky 2015 Status and harvests of sandhillcranes Mid-continent Rocky Mountain Lower Colorado River Valleyand Eastern Populations US Fish and Wildlife Service LakewoodColorado USA

Lacy A E J A Barzen D MMoore and K E Norris 2015 Changes inthe number and distribution of greater sandhill cranes in the EasternPopulation Journal Field Ornithology 86317ndash325

Lumsden H G 1971 The status of the sandhill crane in northern OntarioCanadian Field-Naturalist 85285ndash293

Melvin S M and S A Temple 1982 Migration ecology of sandhillcranes a review Pages 73ndash87 in J C Lewis editor Proceedings of the1981 Crane Workshop National Audubon Society Travernier FloridaUSA

Melvin S M R C Drewien S A Temple and E G Bizeau 1983 Leg-band attachment of radio transmitters for large birds Wildlife SocietyBulletin 11286ndash285

Minnesota Breeding Bird Atlas Project 2014 Minnesota Breeding BirdAtlas mnbbaorgcgi-binquerypl Accessed 14 Mar 2014

Nesbitt S A 1989 The significance of mate loss in Florida sandhill cranesWilson Bulletin 101648ndash651

Nesbitt S A and K S Williams 1990 Home range and habitat use ofFlorida sandhill cranes Journal of Wildlife Management 5492ndash96

Pearse A T D A Brandt W C Harrell K L Metzger D M Baaschand T J Hefley 2015 Whooping crane stopover site use intensitywithin the Great Plains US Geological Survey Open-File Report2015-1166 httpdxdoi org103133ofr20151166 Accessed 28 Dec2015

Tacha TC 1998 Social organization of sandhill cranes frommidcontinentalNorth America Wildlife Monographs 993ndash37

Tacha T C S A Nesbitt and P A Vohs 1992 Sandhill crane Account31 in A Poole P Stettenheim and F Gill editors The birds of NorthAmerica Academy of Natural Sciences Philadelphia Pennsylvania andAmerican Ornithological Union Washington DC USA

Tacha T C S A Nesbitt and P A Vohs 1994 Sandhill cranePages 77ndash94 in T C Tacha and C E Braun editors Migratoryshore and upland game bird management in North AmericaInternational Association of Fish and Wildlife Agencies Wash-ington DC USA

Van Horn K T White W Atkins T Cooper R Urbanek D HolmD Sherman D Aborn J Suckow K Cleveland and R Brook 2010Management plan for the Eastern Population of sandhill cranesMississippi and Atlantic Flyway Councils Madison Wisconsin USA

Walkinshaw L H 1949 Cranes of the worldWinchester Press NewYorkNew York USA

Walkinshaw L H 1960 Migration of the sandhill crane east of theMississippi River Wilson Bulletin 72358ndash384

Walkinshaw L H and H F Wing 1955 Censusing southern Michigansandhill crane Auk 72374ndash384

Warnock N 2010 Stopping vs staging the difference between a hop and ajump Journal of Avian Biology 41621ndash626

Wheeler R H and J C Lewis 1972 Trapping techniques for sandhillcrane studies in the Platte River Valley US Department of theInterior U S Fish and Wildlife Service Natural Resource Publication107 Washington DC USA

Associate Editor Courtney Amundson

Fronczak et al Eastern Population Sandhill Crane Migration 11

geographic distributions (Walkinshaw 1949 Tacha et al1994) and migration routes (Walkinshaw 1960 Melvin andTemple 1982) However at smaller spatial scales some areasthought to support breeding cranes in the EP were notrepresented by our sample including northwestern Wis-consin-northeastern Minnesota (Cutright et al 2006Minnesota Breeding Bird Atlas Project 2014) and easternOntario including the eastern shore of Lake Huron and thesouthern peninsula of Ontario (Cadman et al 2007) Noneof the cranes in our sample overlapped the summer range ofcranes in theMCP identified by Krapu et al (2011) Summerarea locations of marked cranes were concentrated in east-central Wisconsin and the north shore of Lake HuronOntario which support relatively high densities of breedingcranes (Cadman et al 2007 Van Horn et al 2010)Cranes consistently returned to the same summer areas in

subsequent years similar to cranes in the Rocky MountainPopulation (Drewien 1973) and MCP (Krapu et al 2011)On average cranes in the EP and MCP (Krapu et al 2011)used subsequent summer areas within lt10 km of oneanother 65 of cranes in the EP used subsequent summerareas within 1 km of one another compared to 38 forcranes in the MCP (Krapu et al 2011) We observed 3instances when individual cranes moved an average lineardistance of 672 km (rangefrac14 251ndash989 km) among subse-quent summer areas and we speculate that these cranes mayhave been sub-adults (Bennett 1989 Nesbitt and Williams1990 Hayes 2015) However 2 of these cranes returned tothe same summer area in subsequent years and the thirdcrane died during the first summer

Migration Routes and ChronologyCranes followed migration routes similar to those identifiedin previous migration studies that used very high-frequency

transmitters in south-central Minnesota (Crete and Toepfer1978) and cranes marked with ARGOS Doppler PTTs onManitoulin Island Ontario Canada (S A Petrie BirdStudies Canada unpublished data) Unlike those studies wewere able to provide more precise information particularlyabout routes and locations where cranes stopped duringmigration We also observed that 8 (19) cranes withsummer areas in the Upper Peninsula of MichiganManitoulin Island Ontario and east-central Ontario usedmigration routes west of Lake Michigan instead of the moreintuitive route through the Lower Peninsula of MichiganSix of these cranes also alternated migration routes aroundLake Michigan during sequential migration (ie fall andspring) and in subsequent years Our study represents thefirst evidence of this behavior and use of both of thesemigration routes by individual cranes We also documentedcranes (nfrac14 8 19) that summered on Manitoulin Islandused migration routes over large bodies of water (LakeHuron and portions of Lake Michigan) similar to Krapuet al (2011) who described cranes crossing the Bering Straitof Alaska and the Arctic Ocean to and from breedinggroundsRecent observations of migratory cranes that wintered in

Louisiana led to speculation that EP cranes are potentiallyusing non-traditional migration corridors King et al(2010) monitored 2 PTT-marked cranes that wintered incentral Louisiana and migrated north through the Mis-sissippi Alluvial Valley toward summer areas within the EPsummer distribution One marked crane in their study useda migration route through west-central Illinois to south-western Wisconsin and then into the Upper Peninsula ofMichigan after which they were unable to receiveadditional location information from the transmitter Theother crane traveled a similar route used by some of the

Table 3 Mean arrival date length of stay and departure date on winter areas for platform transmitting terminal-marked greater sandhill cranes in theEastern Population North America 2009ndash2014

Arrival datea Length of stay (days)a Departure date

Yr n x SD (days) n x SD (days) n x SD (days)

2009ndash2010 6 16 Feb 62010ndash2011 24 10 Dec 13 26 66 17 35 18 Feb 92011ndash2012 31 14 Dec 28 31 65 32 34 15 Feb 152012ndash2013 22 6 Dec 34 20 74 49 19 19 Feb 222013ndash2014 8 14 Dec 20 7 63 29 8 14 Feb 17All years 86 11 Dec 26 85 67 33 102 16 Feb 14

a Summaries unavailable for 2009ndash2010 winter locations

Table 4 Average number of days number of stops and length of stay at staging areas (stop duration gt5 days) from winter termini to summer areas forplatform transmitting terminal-marked greater sandhill cranes in the Eastern Population North America 2010ndash2014

Total days No stops Length of stay (days)

Yr n x SD (days) n x SD (stops) n x SD (days)

2010 6 39 11 6 68 08 6 12 12011 33 42 16 34 84 30 33 14 42012 33 32 15 33 71 30 29 17 82013 16 41 22 17 71 40 15 17 72014 6 62 11 7 77 25 8 14 4All years 94 39 18 97 76 31 91 15 6

8 The Journal of Wildlife Management 9999()

cranes in our sample through west-central Indiana (GoosePond FWA) northwestern Indiana (Kankakee RiverValley) east-central Wisconsin and then through theUpper Peninsula of Michigan to a summer area in north-central OntarioA single PTT-marked crane in our sample from the

Manitoulin Island Ontario study site migrated west fromthe White River Valley Indiana (Jackson County) andwintered at the Hop-In WR Tennessee (Obion County)However that crane did not return to Hop-In WR in thesubsequent year In anticipation of identifying a potentialwestern migration corridor to Louisiana we affixed 2 PTTsto cranes that wintered at Hop-In WR However only 1returned to winter at the Hop-InWR and the other winteredin Florida the following year

Cranes consistently used staging areas along migrationroutes but made fewer stops for a longer duration during fallmigration than during spring migration Locations wherecranes stopped during migration along the north shore ofLake Huron Algoma District Ontario Lake Poygon east-central Wisconsin the Kankakee River Valley northwesternIndiana and counties in south-central Michigan werefrequented more by cranes during fall migration (Fig 3AAppendix A) Cranes used Hardin County north-centralKentucky White River Valley (Ewing Bottoms JacksonCounty) south-central Indiana and the Kankakee RiverValley northwestern Indiana more frequently during springthan during fall migration (Fig 3B Appendix B) Theseareas support shallow-marsh roosting habitat have anabundant food supply from agricultural production and

Figure 4 Locations of areas surveyed (gray squares) during the United States Fish and Wildlife Service Fall Eastern Population Sandhill Crane Survey alsoshown are locations (black triangles) where platform transmitting terminal-marked sandhill cranes were not observed during the annual surveys 2010ndash2012

Fronczak et al Eastern Population Sandhill Crane Migration 9

are more likely to be protected from human disturbances (iehunting and sporting activities) The Kankakee River Valleylocated in northwestern Indiana was the most frequented ofall staging areas during fall and spring migration wherecranes spent 34 of migration days

Winter Area DistributionCrane winter areas extended from Indiana to Florida wellnorth of the areas previously described by Tacha et al(1994) who reported that cranes in the EP winteredmainly in southeastern Georgia and in Florida Craneswintered in Florida Georgia Tennessee Kentucky andIndiana with the Chickamauga Reservoir (specificallyHiwassee WR) Tennessee having the highest proportionof use among all winter areas The proportion of cranesthat used the Chickamauga Reservoir Valley for thewinters of 2009 and 2010 may be biased high because wetrapped the majority of cranes (nfrac14 6 14 2009ndash2010nfrac14 12 28 2010ndash2011) at the Hiwassee WR duringDecember and January of those years However in 2010MWS estimates for the Chickamauga Reservoir indicateda record number of wintering cranes (nfrac14 48300 J RKelley USFWS-Migratory Bird Management personalcommunication) suggesting that a large portion of cranesin the EP likely used that area Furthermore many ofthe cranes we captured in other locations used theChickamauga Reservoir during migration in years follow-ing their capture The Chickamauga Reservoir is ideal forstaging and wintering cranes Agriculture crops (corn soybeans and winter wheat) are produced annually forwintering wildlife and adjacent sand bars and low waterlevels on Chickamauga Reservoir provided roostinghabitat and sanctuary for sandhill cranes during the falland winter months (R Klippel personal communication)Hunting for sandhill cranes is allowed throughout thereservoir however the Hiwassee WR is designated a non-hunting zoneRecent analysis of long-term Christmas Bird Count data

indicated that cranes in the EP wintered at staging areasnorth of Georgia and Florida in recent decades (Amundsonand Johnson 2011 Lacy et al 2015) Lacy et al (2015)suggested that climate and increased agriculture throughoutthe EP range may influence cranes to winter farther northHowever there has been no formal analysis to demonstratethat these factors are main contributors to a more northerlywinter strategy Thirty-one (74) cranes in our sample thatwintered north of Georgia and Florida used migrationstaging areas in Indiana Kentucky and Tennessee Winterdestinations for cranes varied among years where 11 (26)returned to the same winter area in subsequent winters Theremaining 31 (74) cranes wintered in1 state at least onceduring the study period indicating a variation in migrationstrategy possibly influencing cranes to settle in a particularwinter areaEight (19) cranes used the Kankakee River Valley in

northwestern Indiana as a winter area in1 year These cranesthat wintered in the Kankakee River Valley used the Jasper-Pulaski FWA until all available roosting sites froze

then roosted within the Schahfer Generating Stationproperty in Wheatfield Indiana about 10 km north-northwest of Jasper-Pulaski FWA The power stationprovided roosting areas that remained ice-free and thesurrounding agricultural landscape provided waste grainthroughout the winter (J Bergens personal communica-tion) These cranes then used areas throughout theKankakee River Valley in proximity to Jasper-PulaskiFWA as the winter progressedAdditional winter locations outside of traditional areas are

not well understood Location data from King et al (2010)suggested that cranes in the EP used winter areas in northernLouisiana and 3 cranes from our sample wintered at theHop-In WR Tennessee west of previously reportedwintering areas for cranes in the EP Observations duringthe MWS of cranes at the Wheeler NWR Alabama havesteadily increased since 2003 with an average of 4583 cranes(rangefrac14 293ndash12032) and a peak count of 12032 cranes in2013 Furthermore observations of cranes in Arkansas andMississippi along theMississippi Alluvial Valley are frequentduring the MWS (J R Kelley personal communication)These cranes likely are part of the EP and most likely usedthe Hop-In WR as a staging area

MANAGEMENT IMPLICATIONS

Location and chronology data from our study can be used toadjust timing of hunting seasons to potentially avoid highharvest pressure on local breeding adults and focus harvestpressure on migrants Location and chronology data canalso be used to identify specific locality information forpotential land conservation and opportunity for non-consumptive recreational activities (eg crane festivalsand bird watching)We estimated a proportion of cranes in the EP were located

in areas not currently covered by the USFWS fall surveyduring the survey period If the absence of these cranes in theUSFWS fall survey area is similar through time series ofcounts from surveys will likely provide a reasonable estimateof population trend Alternatively if the proportion of cranesnot present in survey areas changes it may be necessary toredesign the survey by moving the survey period later into thefall or early winter when a larger portion of cranes areassociated with known concentration areas Howeverthe effects of moving the survey period would need to beevaluated to avoid issues related to 1) comparability toprevious surveys and 2) cranes arriving on wintering areas inFlorida where it is difficult to distinguish cranes in theEP from Florida non-migratory greater sandhill cranes(A c pratensis)

ACKNOWLEDGMENTS

Any use of trade firm or product names is for descriptivepurposes only and does not imply endorsement by the USGovernment or the University of Minnesota We thankD A Brandt P Link R Klippel J Jackson B Swiney SShelby J Womac B Layton D E Sherman B Feaster APhelps R Ronk J Bergens J Gilbert E E Carroll RKnopick W Delks V Clarkston E Hockman C Trosen

10 The Journal of Wildlife Management 9999()

A Johnson C Kanke P Peterson S Zudrow TPaulson W Tamminga J Brunjes E Harper LArmstrong T White M Morse S Bossuyt M BossuytA Dhamorikar J E Hanna KWeaver and S A Petrie forfield assistance We thank A T Pearse for comments onprevious drafts of this manuscript The project was supportedthrough Research Work Order no 86 at the USGSMinnesota Cooperative Fish and Wildlife Research Unitat the University of Minnesota This research was supportedby the USFWS Webless Migratory Game Bird Programthe USGS Northern Prairie Wildlife Research Center theMinnesota Cooperative Fish and Wildlife Research Unitthe USFWS Region 3Migratory BirdManagement OfficeLong Point Waterfowl SC Johnsons and Son Ltdthe Ontario Federation of Anglers and Hunters theCanadian Wildlife Service Wildlife Habitat Canada andthe TD Friends of the Environment Foundation

LITERATURE CITEDAmundson C L and D H Johnson 2011 Assessment of the EasternPopulation of greater sandhill cranes (Grus canadensis tabida) FallMigrationSurvey 1979ndash2009 US Fish and Wildlife Service BloomingtonMinnesota USA

Bennett A J 1989 Movements and home ranges of Florida sandhill cranesJournal of Wildlife Management 53830ndash836

Cadman M D D A Sutherland G G Beck D Lepage and A RCouturier 2007 Ontario Breeding Bird Atlas website httpwwwbirdsontarioorgatlasindexjsp Accessed 14 Mar 2014

CreteRA andJEToepfer1978Migrationof radio-taggedeasterngreatersandhill cranes US Fish and Wildlife Service Twin Cities MinnesotaUSA

Cutright N J B R Harriman and R W Howe 2006 WisconsinBreeding Bird Atlas httpuwgbedubirdswbbaindexhtm Accessed 14Mar 2014

Case D J and S J Sanders editors 2009 Priority information needs forsandhill cranes a funding strategy Association of Fish and WildlifeAgencies Washington DC USA

Douglas D C R Weinzierl S C Davidson R Kays M Wikelski andG Bohrer 2012 Moderating ARGOS location errors in animal trackingdata Methods in Ecology and Evolution 3999ndash1007

Drewien R C 1973 Ecology of Rocky Mountain greater sandhill cranesDissertation University of Idaho Moscow USA

Ellis D H P W Howey and G L Krapu 2001 Recommendations forthe attachment of satellite transmitters to cranes North American CraneWorkshop Proceedings 8211ndash212

Fronczak D L 2014 Distribution migration chronology and survival ratesof Eastern Population sandhill crane Thesis University of Minnesota StPaul USA

Fronczak D L D E Andersen E E Hanna and T R Cooper 2015Annual survival rate estimate of satellite transmitter-marked EasternPopulation greater sandhill cranes Journal of Fish and WildlifeManagement 6464ndash471

Hanna E E M L Schummer and S A Petrie 2014 Migratorychronology autumn recruitment and population size of EasternPopulation sandhill crane (Grus canadensis) from the North Shore Regionof Lake Huron Ontario Canada CanadianWildlife Service and OntarioMinistry of Natural Resources Port Rowan Ontario Canada

Hayes M A and J A Barzen 2006 Dynamics of breeding and non-breeding sandhill cranes in south-central Wisconsin Passenger Pigeon68345ndash352

Hayes M A 2015 Dispersal and population genetic structure in twoflyways of sandhill cranes (Grus canadensis) Dissertation University ofWisconsin-Madison Madison USA

Hunt R A E A Gluesing and L E Nauman 1976 The sandhill crane inWisconsin Wisconsin Department of Natural Resources Madison USA

Jones K L G L Krapu D A Brandt andMV Ashley 2005 Populationgenetic structure in migratory sandhill cranes and the role of Pleistoceneglaciations Molecular Ecology 142645ndash2657

King D T J D Paulson D J Leblanc and K Bruce 1998 Two capturetechniques for American white pelicans and great blue herons ColonialWaterbirds 21258ndash260

King S L A R Pierce K R Hersey and N Winstead 2010 Migrationpatterns and movements of sandhill cranes wintering in central andsouthwest Louisiana Proceedings of the North American CraneWorkshop 1157ndash61

Krapu G L D A Brandt K L Jones and D H Johnson 2011Geographic distribution of the Mid-continent Population of sandhillcranes and related management applications Wildlife Monographs1751ndash38

Kruse K L and J A Dubovsky 2015 Status and harvests of sandhillcranes Mid-continent Rocky Mountain Lower Colorado River Valleyand Eastern Populations US Fish and Wildlife Service LakewoodColorado USA

Lacy A E J A Barzen D MMoore and K E Norris 2015 Changes inthe number and distribution of greater sandhill cranes in the EasternPopulation Journal Field Ornithology 86317ndash325

Lumsden H G 1971 The status of the sandhill crane in northern OntarioCanadian Field-Naturalist 85285ndash293

Melvin S M and S A Temple 1982 Migration ecology of sandhillcranes a review Pages 73ndash87 in J C Lewis editor Proceedings of the1981 Crane Workshop National Audubon Society Travernier FloridaUSA

Melvin S M R C Drewien S A Temple and E G Bizeau 1983 Leg-band attachment of radio transmitters for large birds Wildlife SocietyBulletin 11286ndash285

Minnesota Breeding Bird Atlas Project 2014 Minnesota Breeding BirdAtlas mnbbaorgcgi-binquerypl Accessed 14 Mar 2014

Nesbitt S A 1989 The significance of mate loss in Florida sandhill cranesWilson Bulletin 101648ndash651

Nesbitt S A and K S Williams 1990 Home range and habitat use ofFlorida sandhill cranes Journal of Wildlife Management 5492ndash96

Pearse A T D A Brandt W C Harrell K L Metzger D M Baaschand T J Hefley 2015 Whooping crane stopover site use intensitywithin the Great Plains US Geological Survey Open-File Report2015-1166 httpdxdoi org103133ofr20151166 Accessed 28 Dec2015

Tacha TC 1998 Social organization of sandhill cranes frommidcontinentalNorth America Wildlife Monographs 993ndash37

Tacha T C S A Nesbitt and P A Vohs 1992 Sandhill crane Account31 in A Poole P Stettenheim and F Gill editors The birds of NorthAmerica Academy of Natural Sciences Philadelphia Pennsylvania andAmerican Ornithological Union Washington DC USA

Tacha T C S A Nesbitt and P A Vohs 1994 Sandhill cranePages 77ndash94 in T C Tacha and C E Braun editors Migratoryshore and upland game bird management in North AmericaInternational Association of Fish and Wildlife Agencies Wash-ington DC USA

Van Horn K T White W Atkins T Cooper R Urbanek D HolmD Sherman D Aborn J Suckow K Cleveland and R Brook 2010Management plan for the Eastern Population of sandhill cranesMississippi and Atlantic Flyway Councils Madison Wisconsin USA

Walkinshaw L H 1949 Cranes of the worldWinchester Press NewYorkNew York USA

Walkinshaw L H 1960 Migration of the sandhill crane east of theMississippi River Wilson Bulletin 72358ndash384

Walkinshaw L H and H F Wing 1955 Censusing southern Michigansandhill crane Auk 72374ndash384

Warnock N 2010 Stopping vs staging the difference between a hop and ajump Journal of Avian Biology 41621ndash626

Wheeler R H and J C Lewis 1972 Trapping techniques for sandhillcrane studies in the Platte River Valley US Department of theInterior U S Fish and Wildlife Service Natural Resource Publication107 Washington DC USA

Associate Editor Courtney Amundson

Fronczak et al Eastern Population Sandhill Crane Migration 11

cranes in our sample through west-central Indiana (GoosePond FWA) northwestern Indiana (Kankakee RiverValley) east-central Wisconsin and then through theUpper Peninsula of Michigan to a summer area in north-central OntarioA single PTT-marked crane in our sample from the

Manitoulin Island Ontario study site migrated west fromthe White River Valley Indiana (Jackson County) andwintered at the Hop-In WR Tennessee (Obion County)However that crane did not return to Hop-In WR in thesubsequent year In anticipation of identifying a potentialwestern migration corridor to Louisiana we affixed 2 PTTsto cranes that wintered at Hop-In WR However only 1returned to winter at the Hop-InWR and the other winteredin Florida the following year

Cranes consistently used staging areas along migrationroutes but made fewer stops for a longer duration during fallmigration than during spring migration Locations wherecranes stopped during migration along the north shore ofLake Huron Algoma District Ontario Lake Poygon east-central Wisconsin the Kankakee River Valley northwesternIndiana and counties in south-central Michigan werefrequented more by cranes during fall migration (Fig 3AAppendix A) Cranes used Hardin County north-centralKentucky White River Valley (Ewing Bottoms JacksonCounty) south-central Indiana and the Kankakee RiverValley northwestern Indiana more frequently during springthan during fall migration (Fig 3B Appendix B) Theseareas support shallow-marsh roosting habitat have anabundant food supply from agricultural production and

Figure 4 Locations of areas surveyed (gray squares) during the United States Fish and Wildlife Service Fall Eastern Population Sandhill Crane Survey alsoshown are locations (black triangles) where platform transmitting terminal-marked sandhill cranes were not observed during the annual surveys 2010ndash2012

Fronczak et al Eastern Population Sandhill Crane Migration 9

are more likely to be protected from human disturbances (iehunting and sporting activities) The Kankakee River Valleylocated in northwestern Indiana was the most frequented ofall staging areas during fall and spring migration wherecranes spent 34 of migration days

Winter Area DistributionCrane winter areas extended from Indiana to Florida wellnorth of the areas previously described by Tacha et al(1994) who reported that cranes in the EP winteredmainly in southeastern Georgia and in Florida Craneswintered in Florida Georgia Tennessee Kentucky andIndiana with the Chickamauga Reservoir (specificallyHiwassee WR) Tennessee having the highest proportionof use among all winter areas The proportion of cranesthat used the Chickamauga Reservoir Valley for thewinters of 2009 and 2010 may be biased high because wetrapped the majority of cranes (nfrac14 6 14 2009ndash2010nfrac14 12 28 2010ndash2011) at the Hiwassee WR duringDecember and January of those years However in 2010MWS estimates for the Chickamauga Reservoir indicateda record number of wintering cranes (nfrac14 48300 J RKelley USFWS-Migratory Bird Management personalcommunication) suggesting that a large portion of cranesin the EP likely used that area Furthermore many ofthe cranes we captured in other locations used theChickamauga Reservoir during migration in years follow-ing their capture The Chickamauga Reservoir is ideal forstaging and wintering cranes Agriculture crops (corn soybeans and winter wheat) are produced annually forwintering wildlife and adjacent sand bars and low waterlevels on Chickamauga Reservoir provided roostinghabitat and sanctuary for sandhill cranes during the falland winter months (R Klippel personal communication)Hunting for sandhill cranes is allowed throughout thereservoir however the Hiwassee WR is designated a non-hunting zoneRecent analysis of long-term Christmas Bird Count data

indicated that cranes in the EP wintered at staging areasnorth of Georgia and Florida in recent decades (Amundsonand Johnson 2011 Lacy et al 2015) Lacy et al (2015)suggested that climate and increased agriculture throughoutthe EP range may influence cranes to winter farther northHowever there has been no formal analysis to demonstratethat these factors are main contributors to a more northerlywinter strategy Thirty-one (74) cranes in our sample thatwintered north of Georgia and Florida used migrationstaging areas in Indiana Kentucky and Tennessee Winterdestinations for cranes varied among years where 11 (26)returned to the same winter area in subsequent winters Theremaining 31 (74) cranes wintered in1 state at least onceduring the study period indicating a variation in migrationstrategy possibly influencing cranes to settle in a particularwinter areaEight (19) cranes used the Kankakee River Valley in

northwestern Indiana as a winter area in1 year These cranesthat wintered in the Kankakee River Valley used the Jasper-Pulaski FWA until all available roosting sites froze

then roosted within the Schahfer Generating Stationproperty in Wheatfield Indiana about 10 km north-northwest of Jasper-Pulaski FWA The power stationprovided roosting areas that remained ice-free and thesurrounding agricultural landscape provided waste grainthroughout the winter (J Bergens personal communica-tion) These cranes then used areas throughout theKankakee River Valley in proximity to Jasper-PulaskiFWA as the winter progressedAdditional winter locations outside of traditional areas are

not well understood Location data from King et al (2010)suggested that cranes in the EP used winter areas in northernLouisiana and 3 cranes from our sample wintered at theHop-In WR Tennessee west of previously reportedwintering areas for cranes in the EP Observations duringthe MWS of cranes at the Wheeler NWR Alabama havesteadily increased since 2003 with an average of 4583 cranes(rangefrac14 293ndash12032) and a peak count of 12032 cranes in2013 Furthermore observations of cranes in Arkansas andMississippi along theMississippi Alluvial Valley are frequentduring the MWS (J R Kelley personal communication)These cranes likely are part of the EP and most likely usedthe Hop-In WR as a staging area

MANAGEMENT IMPLICATIONS

Location and chronology data from our study can be used toadjust timing of hunting seasons to potentially avoid highharvest pressure on local breeding adults and focus harvestpressure on migrants Location and chronology data canalso be used to identify specific locality information forpotential land conservation and opportunity for non-consumptive recreational activities (eg crane festivalsand bird watching)We estimated a proportion of cranes in the EP were located

in areas not currently covered by the USFWS fall surveyduring the survey period If the absence of these cranes in theUSFWS fall survey area is similar through time series ofcounts from surveys will likely provide a reasonable estimateof population trend Alternatively if the proportion of cranesnot present in survey areas changes it may be necessary toredesign the survey by moving the survey period later into thefall or early winter when a larger portion of cranes areassociated with known concentration areas Howeverthe effects of moving the survey period would need to beevaluated to avoid issues related to 1) comparability toprevious surveys and 2) cranes arriving on wintering areas inFlorida where it is difficult to distinguish cranes in theEP from Florida non-migratory greater sandhill cranes(A c pratensis)

ACKNOWLEDGMENTS

Any use of trade firm or product names is for descriptivepurposes only and does not imply endorsement by the USGovernment or the University of Minnesota We thankD A Brandt P Link R Klippel J Jackson B Swiney SShelby J Womac B Layton D E Sherman B Feaster APhelps R Ronk J Bergens J Gilbert E E Carroll RKnopick W Delks V Clarkston E Hockman C Trosen

10 The Journal of Wildlife Management 9999()

A Johnson C Kanke P Peterson S Zudrow TPaulson W Tamminga J Brunjes E Harper LArmstrong T White M Morse S Bossuyt M BossuytA Dhamorikar J E Hanna KWeaver and S A Petrie forfield assistance We thank A T Pearse for comments onprevious drafts of this manuscript The project was supportedthrough Research Work Order no 86 at the USGSMinnesota Cooperative Fish and Wildlife Research Unitat the University of Minnesota This research was supportedby the USFWS Webless Migratory Game Bird Programthe USGS Northern Prairie Wildlife Research Center theMinnesota Cooperative Fish and Wildlife Research Unitthe USFWS Region 3Migratory BirdManagement OfficeLong Point Waterfowl SC Johnsons and Son Ltdthe Ontario Federation of Anglers and Hunters theCanadian Wildlife Service Wildlife Habitat Canada andthe TD Friends of the Environment Foundation

LITERATURE CITEDAmundson C L and D H Johnson 2011 Assessment of the EasternPopulation of greater sandhill cranes (Grus canadensis tabida) FallMigrationSurvey 1979ndash2009 US Fish and Wildlife Service BloomingtonMinnesota USA

Bennett A J 1989 Movements and home ranges of Florida sandhill cranesJournal of Wildlife Management 53830ndash836

Cadman M D D A Sutherland G G Beck D Lepage and A RCouturier 2007 Ontario Breeding Bird Atlas website httpwwwbirdsontarioorgatlasindexjsp Accessed 14 Mar 2014

CreteRA andJEToepfer1978Migrationof radio-taggedeasterngreatersandhill cranes US Fish and Wildlife Service Twin Cities MinnesotaUSA

Cutright N J B R Harriman and R W Howe 2006 WisconsinBreeding Bird Atlas httpuwgbedubirdswbbaindexhtm Accessed 14Mar 2014

Case D J and S J Sanders editors 2009 Priority information needs forsandhill cranes a funding strategy Association of Fish and WildlifeAgencies Washington DC USA

Douglas D C R Weinzierl S C Davidson R Kays M Wikelski andG Bohrer 2012 Moderating ARGOS location errors in animal trackingdata Methods in Ecology and Evolution 3999ndash1007

Drewien R C 1973 Ecology of Rocky Mountain greater sandhill cranesDissertation University of Idaho Moscow USA

Ellis D H P W Howey and G L Krapu 2001 Recommendations forthe attachment of satellite transmitters to cranes North American CraneWorkshop Proceedings 8211ndash212

Fronczak D L 2014 Distribution migration chronology and survival ratesof Eastern Population sandhill crane Thesis University of Minnesota StPaul USA

Fronczak D L D E Andersen E E Hanna and T R Cooper 2015Annual survival rate estimate of satellite transmitter-marked EasternPopulation greater sandhill cranes Journal of Fish and WildlifeManagement 6464ndash471

Hanna E E M L Schummer and S A Petrie 2014 Migratorychronology autumn recruitment and population size of EasternPopulation sandhill crane (Grus canadensis) from the North Shore Regionof Lake Huron Ontario Canada CanadianWildlife Service and OntarioMinistry of Natural Resources Port Rowan Ontario Canada

Hayes M A and J A Barzen 2006 Dynamics of breeding and non-breeding sandhill cranes in south-central Wisconsin Passenger Pigeon68345ndash352

Hayes M A 2015 Dispersal and population genetic structure in twoflyways of sandhill cranes (Grus canadensis) Dissertation University ofWisconsin-Madison Madison USA

Hunt R A E A Gluesing and L E Nauman 1976 The sandhill crane inWisconsin Wisconsin Department of Natural Resources Madison USA

Jones K L G L Krapu D A Brandt andMV Ashley 2005 Populationgenetic structure in migratory sandhill cranes and the role of Pleistoceneglaciations Molecular Ecology 142645ndash2657

King D T J D Paulson D J Leblanc and K Bruce 1998 Two capturetechniques for American white pelicans and great blue herons ColonialWaterbirds 21258ndash260

King S L A R Pierce K R Hersey and N Winstead 2010 Migrationpatterns and movements of sandhill cranes wintering in central andsouthwest Louisiana Proceedings of the North American CraneWorkshop 1157ndash61

Krapu G L D A Brandt K L Jones and D H Johnson 2011Geographic distribution of the Mid-continent Population of sandhillcranes and related management applications Wildlife Monographs1751ndash38

Kruse K L and J A Dubovsky 2015 Status and harvests of sandhillcranes Mid-continent Rocky Mountain Lower Colorado River Valleyand Eastern Populations US Fish and Wildlife Service LakewoodColorado USA

Lacy A E J A Barzen D MMoore and K E Norris 2015 Changes inthe number and distribution of greater sandhill cranes in the EasternPopulation Journal Field Ornithology 86317ndash325

Lumsden H G 1971 The status of the sandhill crane in northern OntarioCanadian Field-Naturalist 85285ndash293

Melvin S M and S A Temple 1982 Migration ecology of sandhillcranes a review Pages 73ndash87 in J C Lewis editor Proceedings of the1981 Crane Workshop National Audubon Society Travernier FloridaUSA

Melvin S M R C Drewien S A Temple and E G Bizeau 1983 Leg-band attachment of radio transmitters for large birds Wildlife SocietyBulletin 11286ndash285

Minnesota Breeding Bird Atlas Project 2014 Minnesota Breeding BirdAtlas mnbbaorgcgi-binquerypl Accessed 14 Mar 2014

Nesbitt S A 1989 The significance of mate loss in Florida sandhill cranesWilson Bulletin 101648ndash651

Nesbitt S A and K S Williams 1990 Home range and habitat use ofFlorida sandhill cranes Journal of Wildlife Management 5492ndash96

Pearse A T D A Brandt W C Harrell K L Metzger D M Baaschand T J Hefley 2015 Whooping crane stopover site use intensitywithin the Great Plains US Geological Survey Open-File Report2015-1166 httpdxdoi org103133ofr20151166 Accessed 28 Dec2015

Tacha TC 1998 Social organization of sandhill cranes frommidcontinentalNorth America Wildlife Monographs 993ndash37

Tacha T C S A Nesbitt and P A Vohs 1992 Sandhill crane Account31 in A Poole P Stettenheim and F Gill editors The birds of NorthAmerica Academy of Natural Sciences Philadelphia Pennsylvania andAmerican Ornithological Union Washington DC USA

Tacha T C S A Nesbitt and P A Vohs 1994 Sandhill cranePages 77ndash94 in T C Tacha and C E Braun editors Migratoryshore and upland game bird management in North AmericaInternational Association of Fish and Wildlife Agencies Wash-ington DC USA

Van Horn K T White W Atkins T Cooper R Urbanek D HolmD Sherman D Aborn J Suckow K Cleveland and R Brook 2010Management plan for the Eastern Population of sandhill cranesMississippi and Atlantic Flyway Councils Madison Wisconsin USA

Walkinshaw L H 1949 Cranes of the worldWinchester Press NewYorkNew York USA

Walkinshaw L H 1960 Migration of the sandhill crane east of theMississippi River Wilson Bulletin 72358ndash384

Walkinshaw L H and H F Wing 1955 Censusing southern Michigansandhill crane Auk 72374ndash384

Warnock N 2010 Stopping vs staging the difference between a hop and ajump Journal of Avian Biology 41621ndash626

Wheeler R H and J C Lewis 1972 Trapping techniques for sandhillcrane studies in the Platte River Valley US Department of theInterior U S Fish and Wildlife Service Natural Resource Publication107 Washington DC USA

Associate Editor Courtney Amundson

Fronczak et al Eastern Population Sandhill Crane Migration 11

are more likely to be protected from human disturbances (iehunting and sporting activities) The Kankakee River Valleylocated in northwestern Indiana was the most frequented ofall staging areas during fall and spring migration wherecranes spent 34 of migration days

Winter Area DistributionCrane winter areas extended from Indiana to Florida wellnorth of the areas previously described by Tacha et al(1994) who reported that cranes in the EP winteredmainly in southeastern Georgia and in Florida Craneswintered in Florida Georgia Tennessee Kentucky andIndiana with the Chickamauga Reservoir (specificallyHiwassee WR) Tennessee having the highest proportionof use among all winter areas The proportion of cranesthat used the Chickamauga Reservoir Valley for thewinters of 2009 and 2010 may be biased high because wetrapped the majority of cranes (nfrac14 6 14 2009ndash2010nfrac14 12 28 2010ndash2011) at the Hiwassee WR duringDecember and January of those years However in 2010MWS estimates for the Chickamauga Reservoir indicateda record number of wintering cranes (nfrac14 48300 J RKelley USFWS-Migratory Bird Management personalcommunication) suggesting that a large portion of cranesin the EP likely used that area Furthermore many ofthe cranes we captured in other locations used theChickamauga Reservoir during migration in years follow-ing their capture The Chickamauga Reservoir is ideal forstaging and wintering cranes Agriculture crops (corn soybeans and winter wheat) are produced annually forwintering wildlife and adjacent sand bars and low waterlevels on Chickamauga Reservoir provided roostinghabitat and sanctuary for sandhill cranes during the falland winter months (R Klippel personal communication)Hunting for sandhill cranes is allowed throughout thereservoir however the Hiwassee WR is designated a non-hunting zoneRecent analysis of long-term Christmas Bird Count data

indicated that cranes in the EP wintered at staging areasnorth of Georgia and Florida in recent decades (Amundsonand Johnson 2011 Lacy et al 2015) Lacy et al (2015)suggested that climate and increased agriculture throughoutthe EP range may influence cranes to winter farther northHowever there has been no formal analysis to demonstratethat these factors are main contributors to a more northerlywinter strategy Thirty-one (74) cranes in our sample thatwintered north of Georgia and Florida used migrationstaging areas in Indiana Kentucky and Tennessee Winterdestinations for cranes varied among years where 11 (26)returned to the same winter area in subsequent winters Theremaining 31 (74) cranes wintered in1 state at least onceduring the study period indicating a variation in migrationstrategy possibly influencing cranes to settle in a particularwinter areaEight (19) cranes used the Kankakee River Valley in

northwestern Indiana as a winter area in1 year These cranesthat wintered in the Kankakee River Valley used the Jasper-Pulaski FWA until all available roosting sites froze

then roosted within the Schahfer Generating Stationproperty in Wheatfield Indiana about 10 km north-northwest of Jasper-Pulaski FWA The power stationprovided roosting areas that remained ice-free and thesurrounding agricultural landscape provided waste grainthroughout the winter (J Bergens personal communica-tion) These cranes then used areas throughout theKankakee River Valley in proximity to Jasper-PulaskiFWA as the winter progressedAdditional winter locations outside of traditional areas are

not well understood Location data from King et al (2010)suggested that cranes in the EP used winter areas in northernLouisiana and 3 cranes from our sample wintered at theHop-In WR Tennessee west of previously reportedwintering areas for cranes in the EP Observations duringthe MWS of cranes at the Wheeler NWR Alabama havesteadily increased since 2003 with an average of 4583 cranes(rangefrac14 293ndash12032) and a peak count of 12032 cranes in2013 Furthermore observations of cranes in Arkansas andMississippi along theMississippi Alluvial Valley are frequentduring the MWS (J R Kelley personal communication)These cranes likely are part of the EP and most likely usedthe Hop-In WR as a staging area

MANAGEMENT IMPLICATIONS

Location and chronology data from our study can be used toadjust timing of hunting seasons to potentially avoid highharvest pressure on local breeding adults and focus harvestpressure on migrants Location and chronology data canalso be used to identify specific locality information forpotential land conservation and opportunity for non-consumptive recreational activities (eg crane festivalsand bird watching)We estimated a proportion of cranes in the EP were located

in areas not currently covered by the USFWS fall surveyduring the survey period If the absence of these cranes in theUSFWS fall survey area is similar through time series ofcounts from surveys will likely provide a reasonable estimateof population trend Alternatively if the proportion of cranesnot present in survey areas changes it may be necessary toredesign the survey by moving the survey period later into thefall or early winter when a larger portion of cranes areassociated with known concentration areas Howeverthe effects of moving the survey period would need to beevaluated to avoid issues related to 1) comparability toprevious surveys and 2) cranes arriving on wintering areas inFlorida where it is difficult to distinguish cranes in theEP from Florida non-migratory greater sandhill cranes(A c pratensis)

ACKNOWLEDGMENTS

Any use of trade firm or product names is for descriptivepurposes only and does not imply endorsement by the USGovernment or the University of Minnesota We thankD A Brandt P Link R Klippel J Jackson B Swiney SShelby J Womac B Layton D E Sherman B Feaster APhelps R Ronk J Bergens J Gilbert E E Carroll RKnopick W Delks V Clarkston E Hockman C Trosen

10 The Journal of Wildlife Management 9999()

A Johnson C Kanke P Peterson S Zudrow TPaulson W Tamminga J Brunjes E Harper LArmstrong T White M Morse S Bossuyt M BossuytA Dhamorikar J E Hanna KWeaver and S A Petrie forfield assistance We thank A T Pearse for comments onprevious drafts of this manuscript The project was supportedthrough Research Work Order no 86 at the USGSMinnesota Cooperative Fish and Wildlife Research Unitat the University of Minnesota This research was supportedby the USFWS Webless Migratory Game Bird Programthe USGS Northern Prairie Wildlife Research Center theMinnesota Cooperative Fish and Wildlife Research Unitthe USFWS Region 3Migratory BirdManagement OfficeLong Point Waterfowl SC Johnsons and Son Ltdthe Ontario Federation of Anglers and Hunters theCanadian Wildlife Service Wildlife Habitat Canada andthe TD Friends of the Environment Foundation

LITERATURE CITEDAmundson C L and D H Johnson 2011 Assessment of the EasternPopulation of greater sandhill cranes (Grus canadensis tabida) FallMigrationSurvey 1979ndash2009 US Fish and Wildlife Service BloomingtonMinnesota USA

Bennett A J 1989 Movements and home ranges of Florida sandhill cranesJournal of Wildlife Management 53830ndash836

Cadman M D D A Sutherland G G Beck D Lepage and A RCouturier 2007 Ontario Breeding Bird Atlas website httpwwwbirdsontarioorgatlasindexjsp Accessed 14 Mar 2014

CreteRA andJEToepfer1978Migrationof radio-taggedeasterngreatersandhill cranes US Fish and Wildlife Service Twin Cities MinnesotaUSA

Cutright N J B R Harriman and R W Howe 2006 WisconsinBreeding Bird Atlas httpuwgbedubirdswbbaindexhtm Accessed 14Mar 2014

Case D J and S J Sanders editors 2009 Priority information needs forsandhill cranes a funding strategy Association of Fish and WildlifeAgencies Washington DC USA

Douglas D C R Weinzierl S C Davidson R Kays M Wikelski andG Bohrer 2012 Moderating ARGOS location errors in animal trackingdata Methods in Ecology and Evolution 3999ndash1007

Drewien R C 1973 Ecology of Rocky Mountain greater sandhill cranesDissertation University of Idaho Moscow USA

Ellis D H P W Howey and G L Krapu 2001 Recommendations forthe attachment of satellite transmitters to cranes North American CraneWorkshop Proceedings 8211ndash212

Fronczak D L 2014 Distribution migration chronology and survival ratesof Eastern Population sandhill crane Thesis University of Minnesota StPaul USA

Fronczak D L D E Andersen E E Hanna and T R Cooper 2015Annual survival rate estimate of satellite transmitter-marked EasternPopulation greater sandhill cranes Journal of Fish and WildlifeManagement 6464ndash471

Hanna E E M L Schummer and S A Petrie 2014 Migratorychronology autumn recruitment and population size of EasternPopulation sandhill crane (Grus canadensis) from the North Shore Regionof Lake Huron Ontario Canada CanadianWildlife Service and OntarioMinistry of Natural Resources Port Rowan Ontario Canada

Hayes M A and J A Barzen 2006 Dynamics of breeding and non-breeding sandhill cranes in south-central Wisconsin Passenger Pigeon68345ndash352

Hayes M A 2015 Dispersal and population genetic structure in twoflyways of sandhill cranes (Grus canadensis) Dissertation University ofWisconsin-Madison Madison USA

Hunt R A E A Gluesing and L E Nauman 1976 The sandhill crane inWisconsin Wisconsin Department of Natural Resources Madison USA

Jones K L G L Krapu D A Brandt andMV Ashley 2005 Populationgenetic structure in migratory sandhill cranes and the role of Pleistoceneglaciations Molecular Ecology 142645ndash2657

King D T J D Paulson D J Leblanc and K Bruce 1998 Two capturetechniques for American white pelicans and great blue herons ColonialWaterbirds 21258ndash260

King S L A R Pierce K R Hersey and N Winstead 2010 Migrationpatterns and movements of sandhill cranes wintering in central andsouthwest Louisiana Proceedings of the North American CraneWorkshop 1157ndash61

Krapu G L D A Brandt K L Jones and D H Johnson 2011Geographic distribution of the Mid-continent Population of sandhillcranes and related management applications Wildlife Monographs1751ndash38

Kruse K L and J A Dubovsky 2015 Status and harvests of sandhillcranes Mid-continent Rocky Mountain Lower Colorado River Valleyand Eastern Populations US Fish and Wildlife Service LakewoodColorado USA

Lacy A E J A Barzen D MMoore and K E Norris 2015 Changes inthe number and distribution of greater sandhill cranes in the EasternPopulation Journal Field Ornithology 86317ndash325

Lumsden H G 1971 The status of the sandhill crane in northern OntarioCanadian Field-Naturalist 85285ndash293

Melvin S M and S A Temple 1982 Migration ecology of sandhillcranes a review Pages 73ndash87 in J C Lewis editor Proceedings of the1981 Crane Workshop National Audubon Society Travernier FloridaUSA

Melvin S M R C Drewien S A Temple and E G Bizeau 1983 Leg-band attachment of radio transmitters for large birds Wildlife SocietyBulletin 11286ndash285

Minnesota Breeding Bird Atlas Project 2014 Minnesota Breeding BirdAtlas mnbbaorgcgi-binquerypl Accessed 14 Mar 2014

Nesbitt S A 1989 The significance of mate loss in Florida sandhill cranesWilson Bulletin 101648ndash651

Nesbitt S A and K S Williams 1990 Home range and habitat use ofFlorida sandhill cranes Journal of Wildlife Management 5492ndash96

Pearse A T D A Brandt W C Harrell K L Metzger D M Baaschand T J Hefley 2015 Whooping crane stopover site use intensitywithin the Great Plains US Geological Survey Open-File Report2015-1166 httpdxdoi org103133ofr20151166 Accessed 28 Dec2015

Tacha TC 1998 Social organization of sandhill cranes frommidcontinentalNorth America Wildlife Monographs 993ndash37

Tacha T C S A Nesbitt and P A Vohs 1992 Sandhill crane Account31 in A Poole P Stettenheim and F Gill editors The birds of NorthAmerica Academy of Natural Sciences Philadelphia Pennsylvania andAmerican Ornithological Union Washington DC USA

Tacha T C S A Nesbitt and P A Vohs 1994 Sandhill cranePages 77ndash94 in T C Tacha and C E Braun editors Migratoryshore and upland game bird management in North AmericaInternational Association of Fish and Wildlife Agencies Wash-ington DC USA

Van Horn K T White W Atkins T Cooper R Urbanek D HolmD Sherman D Aborn J Suckow K Cleveland and R Brook 2010Management plan for the Eastern Population of sandhill cranesMississippi and Atlantic Flyway Councils Madison Wisconsin USA

Walkinshaw L H 1949 Cranes of the worldWinchester Press NewYorkNew York USA

Walkinshaw L H 1960 Migration of the sandhill crane east of theMississippi River Wilson Bulletin 72358ndash384

Walkinshaw L H and H F Wing 1955 Censusing southern Michigansandhill crane Auk 72374ndash384

Warnock N 2010 Stopping vs staging the difference between a hop and ajump Journal of Avian Biology 41621ndash626

Wheeler R H and J C Lewis 1972 Trapping techniques for sandhillcrane studies in the Platte River Valley US Department of theInterior U S Fish and Wildlife Service Natural Resource Publication107 Washington DC USA

Associate Editor Courtney Amundson

Fronczak et al Eastern Population Sandhill Crane Migration 11

A Johnson C Kanke P Peterson S Zudrow TPaulson W Tamminga J Brunjes E Harper LArmstrong T White M Morse S Bossuyt M BossuytA Dhamorikar J E Hanna KWeaver and S A Petrie forfield assistance We thank A T Pearse for comments onprevious drafts of this manuscript The project was supportedthrough Research Work Order no 86 at the USGSMinnesota Cooperative Fish and Wildlife Research Unitat the University of Minnesota This research was supportedby the USFWS Webless Migratory Game Bird Programthe USGS Northern Prairie Wildlife Research Center theMinnesota Cooperative Fish and Wildlife Research Unitthe USFWS Region 3Migratory BirdManagement OfficeLong Point Waterfowl SC Johnsons and Son Ltdthe Ontario Federation of Anglers and Hunters theCanadian Wildlife Service Wildlife Habitat Canada andthe TD Friends of the Environment Foundation

LITERATURE CITEDAmundson C L and D H Johnson 2011 Assessment of the EasternPopulation of greater sandhill cranes (Grus canadensis tabida) FallMigrationSurvey 1979ndash2009 US Fish and Wildlife Service BloomingtonMinnesota USA

Bennett A J 1989 Movements and home ranges of Florida sandhill cranesJournal of Wildlife Management 53830ndash836

Cadman M D D A Sutherland G G Beck D Lepage and A RCouturier 2007 Ontario Breeding Bird Atlas website httpwwwbirdsontarioorgatlasindexjsp Accessed 14 Mar 2014

CreteRA andJEToepfer1978Migrationof radio-taggedeasterngreatersandhill cranes US Fish and Wildlife Service Twin Cities MinnesotaUSA

Cutright N J B R Harriman and R W Howe 2006 WisconsinBreeding Bird Atlas httpuwgbedubirdswbbaindexhtm Accessed 14Mar 2014

Case D J and S J Sanders editors 2009 Priority information needs forsandhill cranes a funding strategy Association of Fish and WildlifeAgencies Washington DC USA

Douglas D C R Weinzierl S C Davidson R Kays M Wikelski andG Bohrer 2012 Moderating ARGOS location errors in animal trackingdata Methods in Ecology and Evolution 3999ndash1007

Drewien R C 1973 Ecology of Rocky Mountain greater sandhill cranesDissertation University of Idaho Moscow USA

Ellis D H P W Howey and G L Krapu 2001 Recommendations forthe attachment of satellite transmitters to cranes North American CraneWorkshop Proceedings 8211ndash212

Fronczak D L 2014 Distribution migration chronology and survival ratesof Eastern Population sandhill crane Thesis University of Minnesota StPaul USA

Fronczak D L D E Andersen E E Hanna and T R Cooper 2015Annual survival rate estimate of satellite transmitter-marked EasternPopulation greater sandhill cranes Journal of Fish and WildlifeManagement 6464ndash471

Hanna E E M L Schummer and S A Petrie 2014 Migratorychronology autumn recruitment and population size of EasternPopulation sandhill crane (Grus canadensis) from the North Shore Regionof Lake Huron Ontario Canada CanadianWildlife Service and OntarioMinistry of Natural Resources Port Rowan Ontario Canada

Hayes M A and J A Barzen 2006 Dynamics of breeding and non-breeding sandhill cranes in south-central Wisconsin Passenger Pigeon68345ndash352

Hayes M A 2015 Dispersal and population genetic structure in twoflyways of sandhill cranes (Grus canadensis) Dissertation University ofWisconsin-Madison Madison USA

Hunt R A E A Gluesing and L E Nauman 1976 The sandhill crane inWisconsin Wisconsin Department of Natural Resources Madison USA

Jones K L G L Krapu D A Brandt andMV Ashley 2005 Populationgenetic structure in migratory sandhill cranes and the role of Pleistoceneglaciations Molecular Ecology 142645ndash2657

King D T J D Paulson D J Leblanc and K Bruce 1998 Two capturetechniques for American white pelicans and great blue herons ColonialWaterbirds 21258ndash260

King S L A R Pierce K R Hersey and N Winstead 2010 Migrationpatterns and movements of sandhill cranes wintering in central andsouthwest Louisiana Proceedings of the North American CraneWorkshop 1157ndash61

Krapu G L D A Brandt K L Jones and D H Johnson 2011Geographic distribution of the Mid-continent Population of sandhillcranes and related management applications Wildlife Monographs1751ndash38

Kruse K L and J A Dubovsky 2015 Status and harvests of sandhillcranes Mid-continent Rocky Mountain Lower Colorado River Valleyand Eastern Populations US Fish and Wildlife Service LakewoodColorado USA

Lacy A E J A Barzen D MMoore and K E Norris 2015 Changes inthe number and distribution of greater sandhill cranes in the EasternPopulation Journal Field Ornithology 86317ndash325

Lumsden H G 1971 The status of the sandhill crane in northern OntarioCanadian Field-Naturalist 85285ndash293

Melvin S M and S A Temple 1982 Migration ecology of sandhillcranes a review Pages 73ndash87 in J C Lewis editor Proceedings of the1981 Crane Workshop National Audubon Society Travernier FloridaUSA

Melvin S M R C Drewien S A Temple and E G Bizeau 1983 Leg-band attachment of radio transmitters for large birds Wildlife SocietyBulletin 11286ndash285

Minnesota Breeding Bird Atlas Project 2014 Minnesota Breeding BirdAtlas mnbbaorgcgi-binquerypl Accessed 14 Mar 2014

Nesbitt S A 1989 The significance of mate loss in Florida sandhill cranesWilson Bulletin 101648ndash651

Nesbitt S A and K S Williams 1990 Home range and habitat use ofFlorida sandhill cranes Journal of Wildlife Management 5492ndash96

Pearse A T D A Brandt W C Harrell K L Metzger D M Baaschand T J Hefley 2015 Whooping crane stopover site use intensitywithin the Great Plains US Geological Survey Open-File Report2015-1166 httpdxdoi org103133ofr20151166 Accessed 28 Dec2015

Tacha TC 1998 Social organization of sandhill cranes frommidcontinentalNorth America Wildlife Monographs 993ndash37

Tacha T C S A Nesbitt and P A Vohs 1992 Sandhill crane Account31 in A Poole P Stettenheim and F Gill editors The birds of NorthAmerica Academy of Natural Sciences Philadelphia Pennsylvania andAmerican Ornithological Union Washington DC USA

Tacha T C S A Nesbitt and P A Vohs 1994 Sandhill cranePages 77ndash94 in T C Tacha and C E Braun editors Migratoryshore and upland game bird management in North AmericaInternational Association of Fish and Wildlife Agencies Wash-ington DC USA

Van Horn K T White W Atkins T Cooper R Urbanek D HolmD Sherman D Aborn J Suckow K Cleveland and R Brook 2010Management plan for the Eastern Population of sandhill cranesMississippi and Atlantic Flyway Councils Madison Wisconsin USA

Walkinshaw L H 1949 Cranes of the worldWinchester Press NewYorkNew York USA

Walkinshaw L H 1960 Migration of the sandhill crane east of theMississippi River Wilson Bulletin 72358ndash384

Walkinshaw L H and H F Wing 1955 Censusing southern Michigansandhill crane Auk 72374ndash384

Warnock N 2010 Stopping vs staging the difference between a hop and ajump Journal of Avian Biology 41621ndash626

Wheeler R H and J C Lewis 1972 Trapping techniques for sandhillcrane studies in the Platte River Valley US Department of theInterior U S Fish and Wildlife Service Natural Resource Publication107 Washington DC USA

Associate Editor Courtney Amundson

Fronczak et al Eastern Population Sandhill Crane Migration 11

APPENDIX A

Fall migration staging areas average number of cranes using the staging areas and percent of fall migration days at the stagingarea for platform transmitting terminal-marked sandhill cranes in the Eastern Population North America 2010ndash2013 Fallstaging areas are areas that had 1 of the total fall migration days in that area

APPENDIX B

Spring migration staging areas average number of cranes using the staging area and the percent of spring migration days at thestaging area for platform transmitting terminal-marked sandhill cranes in the Eastern Population North America 2010ndash2014Spring staging areas in the table are areas that had 1 of the total spring migration days spent in that area

State orProvince County or District Location names

No marked cranes(x)

Days in staging area()

IN LaPorte Jasper Pulaski Starke Kankakee River Valley 12 337MI Barry Calhoun Eaton Jackson

KalamazooSouth-central Michigan 9 179

WI Waushara Winnebago Lake Poygon 4 59WI Columbia Iowa Juneau Sauk Wisconsin River Valley 2 44ON Algoma Sudbury North Shore of Lake Huron 3 41WI Wood 2 33WI Lincoln 1 26TN Meigs Rhea Chickamauga Reservoir 7 25IN Jackson Ewing Bottoms 4 22MI Chippewa 2 20MN Sherburne Sherburne National Wildlife

Refuge1 19

WI Green Lake Green Lake 1 15MI Bay 1 12WI Oconto 1 10

State or Province County or District Location names No marked cranes (x) Days in staging area ()

IN LaPorte Jasper Pulaski Starke Kankakee River Valley 19 339IN Jackson White River Valley 9 87MI BarryCalhoun Eaton Jackson Kalamazoo South-central Michigan 5 66KY Hardin Larue Cecilia 8 64IN Greene Goose Pond FWA 3 38WI Waushara Winnebago Lake Poygon 2 29MI Missaukee 2 24KY Barren Metecalf Barren River Valley Reservoir 4 23MI Cass 1 23WI Green Lake Green Lake 2 21IN Morgan 1 20MI Oceana 2 19ON Algoma Sudbury North Shore of Lake Huron 4 17TN Meigs Rhea Chickamauga Reservoir 3 14MI Mason 1 11

12 The Journal of Wildlife Management 9999()