Revisiting the Historic Distribution and Habitats of the Whooping Crane Jane E

University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln

USGS Northern Prairie Wildlife Research Center US Geological Survey

2019 Revisiting the Historic Distribution and Habitats of the Whooping Crane Jane E. Austin

Matthew A. Hayes

Jeb A. Barzen

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This Article is brought to you for free and open access by the US Geological Survey at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in USGS Northern Prairie Wildlife Research Center by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. CHAPTER 3 Revisiting the Historic Distribution and Habitats of the Whooping Crane Jane E. Austin*, Matthew A. Hayes**, Jeb A. Barzen** *U.S. Geological Survey, Northern Prairie Wildlife Research Center, Jamestown, ND, United States **International Crane Foundation, Baraboo, WI, United States

INTRODUCTION habitat (Prince, 1997). Many endangered species persist only on the periphery of their historic Understanding the historic range and habitats range because of habitat loss or modification in of an endangered species can assist in conserva- their core range (Channell and Lomolino, 2000), tion and reintroduction efforts for that species. which can bias our understanding of the spe- Individuals reintroduced into a species’ historic cies’ habitat preferences. Further, habitat models core range have a higher survival rate compared based on locations where species persist neces- to individuals introduced near the periphery or sarily emphasize local conditions rather than outside the historic range (Falk and Olwell, 1992; historical conditions (Kuemmerle et al., 2011). Griffith et al., 1989). Individuals on the periph- For example, habitat models for the European ery of a species’ range tend to occupy less favor- bison (Bison bonasus) suggested it was a wood- able habitats and have lower and more variable land species, but assessment of the bison’s densities than those near the core of their range historic range indicated it preferred mosaic- (Brown, 1984; Brown et al., 1995, 1996). Such type landscapes and had a more eastern and conclusions, however, presume that historic northern distribution than previously reported habitats have not changed since a species was (Kuemmerle et al., 2011, 2012). Hence, accurate extirpated from core areas – a difficult assump- determination of the historic range and habitat tion for many areas, and particularly for wetland conditions for endangered species can improve

Whooping Cranes: Biology and Conservation http://dx.doi.org/10.1016/B978-0-12-803555-9.00003-7 25 2019 Published by Elsevier Inc. 26 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane our understanding of their ecology and assist habitat use descriptions, taking advantage of in conservation and reintroduction efforts. newly discovered information, new tools such Examining the historic range from an ecological as geographic information systems (GIS), and perspective can also help identify where appro- contemporary knowledge of wetland and spe- priate habitat still exists that could sustain a cies ecology. We review the historic range, population. adjust the ecological niche for Whooping Cranes The main source of information on the his- described by Allen by linking historic habitat use toric distribution of Whooping Cranes is a patterns to the life history of Whooping Cranes monograph, The Whooping Crane, a detailed (most of which was developed after Allen’s pub- report by Allen (1952). Allen collected and ana- lication), and discuss how our analysis could lyzed all available information about Whooping help to direct current and future reintroduction Cranes, including historic observations. From efforts. Finally, we hypothesize that the ecologi- these observations, Allen constructed a historic cal niche for Whooping Cranes is defined pri- Whooping Crane distribution map (Allen, 1952, marily by wetland productivity, which meets p. 2) to illustrate historic breeding and winter- three primary needs in the annual cycle: rapid ing distribution and habitat use. Allen’s maps chick growth in spring and summer, habitat for and information have been utilized by many molting birds in summer, and acquisition of sources since their initial publication (e.g., large energy reserves in winter. We focus our Doughty, 1989; Meine and Archibald, 1996; results and discussion on breeding, summering, Walkinshaw, 1973), including the International and wintering records; we do not consider habi- Whooping Crane Recovery Plan (Canadian tat use during migration. Wildlife Service and U.S. Fish and Wildlife Ser- vice, 2005), and Allen’s report remains a cru- cial source for Whooping Crane biology today. METHODS However, it has never been updated and ques- tions remain about some areas identified by All location records contained in The Whooping Allen. For example, when comparing recorded Crane (Allen 1952, pp. 51–64), as well as records observations and the historical principal range published after that document was published, map in Allen (1952, p. 2), many observations were entered into a tabular database (available listed are not included in the map (e.g., southern on USGS ScienceBase; doi:10.5066/F7QZ282R). Michigan), and some observations indicated on We used online search tools to find observations the map are not found in the table (Allen 1952, of Whooping Cranes in the literature published pp. 51–64). Also, some areas encompassed by after 1952; some records were also found in early the former breeding (e.g., northern Illinois) or publications that are now available electroni- wintering ranges that Allen described do not cally that may not have been available to Allen. appear to be supported by his tabular data. We used only observations up to 1941, the low Given the influence ofAllen’s (1952) report point of 15 or 16 Whooping Cranes in the AWBP in planning reintroduction efforts and the need (Allen, 1952), to determine their historic distri- to manage for a growing Aransas-Wood Buffalo bution. Thus, our evaluation of Allen’s work was Population (AWBP), Allen’s original analyses constrained to the period before the intensifica- deserved a review. In this chapter, we update tion of agricultural practices that occurred after Allen’s work with records found subsequent the 1940s, but encompassed the years of rapid to the publication of The Whooping Crane, and settlement and cropland expansion across the evaluate the accuracy of Allen’s range maps and Great Plains during the 1870s–1930 (Waisanen

Results 27 and Bliss, 2002). Records were organized by or unknown (biome = 0), we assigned the loca- date, location (place, city, county, state, country, tion to the nearest adjacent biome and ecoregion. and latitude/longitude), type of record (nesting, We also replicated the hand-drawn map ranges observation, specimen, or captured bird), and from Allen (1952, pp. 2, 19) in the GIS to the best life-history stage (breeding, summering, migrat- of our ability, given uncertainty about the map ing, or wintering). We categorized observations projection used by Allen. or specimens of a nest, eggs, or flightless chick as breeding; other observations between mid-May and 30 August were categorized as summering. RESULTS For observations or specimens that included a year but no date, we categorized life-history Updating Allen’s Historic Distribution stage as unknown. We included observer iden- tity, if known, source for each record, and, for We compiled a total of 884 records from birds that were killed or collected, current loca- Allen (1952), dating from 1722 to 1941: 25 (2.8%) tion of existing specimens. Some records lacked records preceded 1850, 390 (44.1%) occurred information on multiple aspects (e.g., observer from 1850 to 1899, 221 (25.0%) from 1900 to 1919, name, specific location, season) and, without and 201 (22.7%) from 1920 to 1941; 47 (5.3%) had other supporting information, were included no known date specified. We added 74 records in the database but not included in analyses. to those listed in Allen (1952): 20 breeding loca- Records were exported to a GIS, ArcMap10 tions (total 87), 21 summering locations (total (ESRI, Redlands, CA, USA), and sorted by life- 169), 18 migrating locations (total 367), 5 winter- history stage and season. ing locations (total 121), and 10 unknown life- Landscape features and patterns of habitat history stages (total 140) (Table 3.1; Appendix). use were derived by overlaying locations and The additional records were from 28 published range polygons on the World Wildlife Fund sources, including eight that had been published (WWF) terrestrial biomes and ecoregions (Abell before 1952 but either may not have been avail- et al., 2008). These data layers were selected able to Allen or we felt deserved to be reinter- because of their use of a comprehensive set preted. Among the most striking additions were of characteristics at the ecoregion level (wet- the 13 breeding records, 14 summering records, lands, climate, topography, and vegetation). and 7 migration records reported for Saskatch- Additional information was obtained from ewan by Hjertaas (1994) and 4 breeding records two other ecoregion classification systems: the in southern Texas (Oberholser, 1938, 1974). Also Canadian Ecological Framework (CEF), which notable were six records for Wisconsin and six closely matches the WWF ecoregion boundaries each for Florida and Indiana. (ESWG, 1995), and the Prairie Pothole Region Verifying the location of some records was (PPR) in the northern Great Plains (Euliss problematic. For example, in a report of his et al., 1999). Finally, we used descriptions of hab- explorations of the Canadian north, Samuel itat types in Allen (1952), landscape and wetland Hearne indicated that “[t]his bird visits Hud- information from WWF global habitat types son’s Bay in the Spring, though not in great and ecoregions, and contemporary information numbers. They are generally seen only in pairs, about landscape and wetland characteristics in and then not very often… It seldom has more those locations to assess landscape and habitat than two young and retires Southward early in features of Whooping Crane locations. Where the fall” (Hearne, 1795, pp. 271–272). Unfortu- locations were identified as a “lake” (biome = 98) nately, Hearne did not give specific locations of

28 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane % 65.0 7.1 17.1 15.0 1.4 4.3 2.9 6.4 3.6 2.1 4.3 0.7 0 10.7 10.7 9.3 2.1 2.1 otal No. 91 10 24 21 2 6 4 9 5 3 6 1 0 15 15 13 3 3 Unknown 0 0 0 0 0 0 0 0 0 0 0 0 0 72.5 72.5 3.3 0 0 % 0 0 0 0 0 0 0 0 0 0 0 0 0 87 87 4 0 0 Wintering No. 92.9 29.7 18.0 6.5 7.6 4.9 6.0 7.4 5.7 3.0 1.4 2.2 0.5 0.5 0.5 5.4 1.6 1.1 % 341 109 66 24 28 18 22 27 21 11 5 8 2 2 2 20 6 4 No. Migrating 72.8 0.6 4.1 25.4 13.6 18.3 7.1 2.4 0 0 0.6 0 0.6 4.1 4.1 5.9 3.0 1.2 % 123 1 7 43 23 31 12 4 0 0 1 0 1 7 7 10 5 2 No. Summering 70.9 0 21.2 20.0 9.4 7.1 14.1 0 0 0 0 0 0 10.6 10.6 5.8 4.7 0 % No. 61 0 18 17 8 6 12 0 0 0 0 0 0 9 9 5 4 0 Breeding % 69.8 13.6 13.1 11.9 6.9 6.9 5.7 4.5 2.9 1.6 1.4 1.0 0.3 13.6 13.6 5.9 2.1 1.0 616 120 115 105 61 61 50 40 26 14 12 9 3 120 120 52 18 9 Total No.

Distribution of Historic Whooping Crane L ocations by (WW F ) Ecoregions (Indented) within Biomes (Bold), in O rder T T hose hat Had O bservations or S pecimens of a N est, Eggs, F lightless Chick; ther N umber of Records. Breeding Records Were Categorized as S ummering. O bservation Records in eptember–Mid-May or O bservations between Mid-May and 30 A ugust Were Records L acking a Date Were Classified as Migrating and T hose of December–Mid-March Wintering. L ate March–Mid-May Were Classified as U nknown. Grasslands Parklands Transition Grasslands Grasslands Transition

Central and Southern Mixed Grasslands Central Tall and Aspen Forests Canadian Northern Mixed Grasslands Northern Short Grasslands Grasslands Northern Tall Central Forest-Grasslands Blackland Prairies Texas Nebraska Sand Hills Mixed Short Grasslands Western Grasslands Flint Hills Tall Plateau Savanna Edwards Gulf Coastal Western Upper Midwest Forest-Savanna Lakes Forests Southern Great Ecoregion Shrublands Savannas, and Shrublands Temperate Grasslands, Savannas, and Temperate and Subtropical Grasslands, Tropical Broadleaf and Mixed Forest Temperate Biome TABLE 3.1 TABLE

Results 29 ) % 1.4 1.4 0 0 0 0.7 0 1.4 7.9 0.7 5.7 0.7 0.7 0 0 0.7 0 0 0.7 0 5.7 2.9 2.9 Continued ( No. 2 2 0 0 0 1 0 2 11 1 8 1 1 0 0 1 0 0 1 0 8 4 4 Unknown 0 1.7 0 1.7 0 0 0 0 14.2 12.5 0 0.8 0 0.8 0 0 0 0 0 0 5.8 4.2 1.7 % 0 2 0 2 0 0 0 0 17 15 0 1 0 1 0 0 0 0 0 0 7 4 2 Wintering No. 1.1 0.8 0.8 0 0 0 0 0 1.1 0.3 0.5 0 0 0 0.3 0 0 0 0 0 0 0 0 % 4 3 3 0 0 0 0 0 4 1 2 0 0 0 1 0 0 0 0 0 0 0 0 No. Migrating 0.6 0 0.6 0 0 0 0.6 0 0.6 0 0.6 0 0 0 0 8.3 4.7 2.4 0.6 0.6 3.0 0 0 % 1 0 1 0 0 0 1 0 1 0 1 0 0 0 0 14 8 4 1 1 5 0 0 No. Summering 0 0 0 0 1.2 0 0 0 0 0 0 0 0 0 0 10.6 9.3 1.2 0 0 1.2 0 0 % No. 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 9 8 1 0 0 1 0 0 Breeding % 0.8 0.8 0.4 0.2 0.1 0.1 0.1 0.2 3.7 1.9 1.2 0.2 0.1 0.1 0.1 2.7 1.8 0.6 0.2 0.1 2.4 1.0 0.7 7 7 4 2 1 1 1 2 33 17 11 2 1 1 1 24 16 5 2 1 21 7 6 Total No. Forests Transition Forests Mesophytic Forests Forests Forests Central U.S. Hardwood Central U.S. Hardwood Mississippi Lowland Forests Southeastern Mixed Forests Northeastern Coastal Forests Forests East Central Texas Eastern Forest-Boreal Lakes Lowland Eastern Great Appalachian Mixed Mezquital Tamaulipan Steppe Basin Shrub Wyoming Chihuahuan Desert Steppe Basin Shrub Great Meseta Central Matorral Steppe Snake-Columbia Shrub Mid-Continental Canadian Muskwa-Slave Lake Forests Southern Hudson Bay Taiga North Canadian Shield Taiga Southeastern Conifer Forests Middle Atlantic Coastal Ecoregion Deserts and Xeric Shrublands Boreal Forest/Taiga Coniferous Forests Temperate Biome

30 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane % 0 0 0 0 0 0 0 0 0.7 0.7 15.9 otal ) No. 0 0 0 0 0 0 0 0 1 1 140 Unknown cont. % 0 0 0 1.7 1.7 0 0 0 2.5 2.5 13.5 No. 0 0 0 2 2 0 0 0 3 3 119 Wintering % 0 0 0 0 0 0 0 0 0 0 41.7 0 0 0 0 0 0 0 0 0 0 367 No. Migrating % 1.8 0.6 0.6 0 0 5.3 4.7 0.6 0 0 19.2 3 1 1 0 0 9 8 1 0 0 169 No. Summering 1.2 0 0 0 0 0 0 0 0 0 9.7 % 1 0 0 0 0 0 0 0 0 0 85 No. Breeding 0.4 0.1 0.1 0.2 0.2 1.0 0.9 0.1 0.4 0.4 100.0 % 4 1 1 2 2 9 8 1 4 4 880 No. Total Distribution of Historic Whooping Crane L ocations by (WW F ) Ecoregions (Indented) within Biomes (Bold), in O rder T Pine-Oak Forests South Central Rockies Forests North Central Rockies Forests Forests Piney Woods Belt Volcanic Trans-Mexican Coastal Tundra Arctic Low Arctic Tundra Baja Dry Forests Ecoregion Forests Forests Tropical and Subtropical Coniferous Tropical Tundra and Subtropical Dry Broadleaf Tropical Total Biome TABLE 3.1 TABLE T hose hat Had O bservations or S pecimens of a N est, Eggs, F lightless Chick; ther N umber of Records. Breeding Records Were Categorized as S ummering. O bservation Records in eptember–Mid-May or L ate March–Mid-May Were between Mid-May and 30 A ugust Were Classified as U nknown. ( Records L acking a Date Were Classified as Migrating and T hose of December–Mid-March Wintering.

Results 31

Whooping Crane nests or chicks, so his com- Our revised set of records alters the historic ments could apply to the broader region he distribution of Whooping Cranes from that of explored, which extended from the Churchill Allen (Fig. 3.1). Additional breeding records in River to the Great Slave Lake, then north to Iowa and North Dakota fall within the former the Coppermine River (Hearne 1795). There- breeding area Allen delineated, and the breed- fore, Allen (1952) centralized Hearne’s findings ing area in Saskatchewan now appears larger. at Churchill, Manitoba, as we did. Whooping There is no direct evidence to support Allen’s Cranes did occur near Hudson Bay; records extension of the breeding range into northern show a Whooping Crane adult collected in 1771 Illinois but Allen clearly believed northern at York Factory, located south of Churchill, and Illinois was breeding habitat for Whooping others observed along the Hudson Bay coast Cranes, a contradiction that was unresolvable (Houston et al., 2003). Migrating Whooping with data that we could evaluate. Four addi- Cranes were also observed around Churchill in tional breeding records were found for Texas, 1953 and 1964 (Jehl and Smith, 1970). Nelson’s possibly associated with the original nonmi- (1876) reference of breeding pairs in central Illi- gratory population. New records extend the nois was similarly problematic, as it provided occurrence of summering birds into central no date, specific location, observer, or note Saskatchewan, southern Wisconsin, and north- of actual nest, eggs, or chicks. An authorita- eastern Illinois. Some of the summer observa- tive reference of the era (Ridgway, 1889) listed tions may well suggest breeding birds (e.g., Whooping Crane only as a summer resident of failed breeders that lacked evidence of eggs or Illinois. We considered but excluded a nesting chicks) or they may have included birds that report in Brown County, Wisconsin, reported were not yet breeding but ranging widely from in Carr (1890) but not cited in Allen (1952), their natal areas in summer (see Barzen et al., and two observations in Florida noted in Bent Chapter 14, this volume). Our additional sum- (1926) but never supported in Allen (1952) or mer records in Wisconsin and Illinois provide Nesbitt’s (1981) review of Whooping Cranes somewhat more support for Whooping Cranes in Florida. We chose to exclude Nelson (1876), breeding in that area, but no actual records of Carr (1890), and Bent’s (1926) records from our nests, eggs, or chicks exist. updated map and biome and ecoregion assess- We accepted one record as breeding around ments because of their vague nature, but those the Greater Yellowstone-Teton area, coinciding records are included in the Appendix. We were with summering records for that area (Fig. 3.1). unable to determine specific locations for two Kemsies (1930) noted that “regular reports private ranches listed in Allen (1952, p. 56), but are received of the breeding of the Whooping placed them near La Barca, Jalisco (west-central Crane,” including observation of two young Mexico), as described in Allen’s text. Allen’s and later two adults in the southwest part extension of the wintering range into northern of Yellowstone National Park by a reputable Mexico is not supported by any listed loca- observer. Allen (1952) included Kemsies’ record tions. However, discussions with biologists with but suspected his observations to be summer- extensive knowledge of northern Mexico sug- ing subadult birds in association with Sand- gest that including that area within the winter- hill Cranes. In this particular case, we believe ing range may have been accurate (R. Drewein, the breeding records from Kemsies (1930) are U.S. Fish and Wildlife Service [retired], and J. viable, given the dates of observations and Taylor, U.S. Fish and Wildlife Service [deceased], size differential between Whooping Cranes personal communications). chicks near fledging (100–120 cm; Urbanek

32 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane

FIGURE 3.1 Distribution of historic Whooping Crane locations 1722–1941, by life-history stage. Records from Allen (1952) and additional records (this chapter), with breeding and wintering ranges delineated by Allen. Multiple records may be represented by a single symbol. Scale = 1:25,000,000.

Results 33 and Lewis, 2015) and adult greater Sandhill summering ecoregions also were the Aspen Cranes in this region (Grus canadensis tabida, Parkland and Northern Mixed Grasslands, as 130–150 cm). well as adjacent Northern Short Grasslands. Our revised distribution of wintering birds These ecoregions coincide with the PPR, a gla- along the Gulf of Mexico is similar to that cially formed region in the northern midcon- described by Allen (1952) but extended further tinent grasslands that is largely defined by its east into Mississippi. Along the Atlantic coast high wetland densities (Kantrud et al., 1989; we assess the range to have been much broader Smith et al., 1964). Clearly once an important and to have extended from North Carolina into area for Whooping Cranes, the PPR encom- central Florida (Fig. 3.1). Most of the Atlantic passed 57.6% of all breeding records, 52.1% of coast records occurred before 1880, but several all summering records, and 26.7% of migrat- additional records reporting Whooping Cranes ing records. Collectively, the PPR accounted for in Florida as late as 1936 suggest that a few 30.1% of all 880 records. The highest density of cranes continued to winter in the state into the breeding records occurred in the Central Tall early 20th century. Grasslands of Iowa, especially that portion of northern Iowa that corresponds to the southern- Biome Affiliations most extent of the PPR. This area was identified by Allen as likely the historic center of breeding. Historic observations of Whooping Cranes The Upper Midwest Forest-Savanna Transition occurred across nine biomes, extending from ecoregion was used to a lesser extent; it is more the Tundra and Boreal Forest/Taiga of interior mesic and found at lower latitudes than the PPR Canada to the Tropical/Subtropical Grasslands, or other grasslands. Savannas, and Shrublands along the Gulf coast, Within the Boreal Forest/Taiga biome, the and to the Desert and Xeric Shrublands of inte- majority of breeding and summering cranes rior Mexico (Table 3.1, Fig. 3.2). Most (69.8%) were observed in the Midcontinental Canadian of the locations, however, occurred within the forests. Four summer observations and one Temperate Grasslands, Savannas, and Shrub- breeding observation fall within the Muskwa- lands biome, which extends from the northern Slave Lake Forests ecoregion, which encom- Great Plains to the Gulf coast. This biome also passes the Hay River, Slave River, Peace, and encompassed more than two-thirds of both Wabasca Lowlands, Boreal Transition, and the breeding and summering records. All of the Mid-Boreal Upland CEF ecoregions. Locations southern (nonmigratory) breeding records and fall in areas of high wetland density, or along two-thirds of the wintering locations occurred rivers or large lakes. in the Tropical/Subtropical Grasslands, Savan- Records of nonmigratory breeding and nas, and Shrublands biome along the Gulf southern summering cranes were located pri- coast. marily in the Western Gulf Coast Grasslands. The last remnants of the nonmigratory popula- Ecoregion Affiliations tion at White Lake, Louisiana, were observed in this ecoregion as late as the 1930s. The breeding Breeding and summering locations – Most location at Eagle Lake in Wharton County, Texas, breeding records were affiliated with the Central appears to fall within the East Central Texas For- Tall Grasslands, Canadian Aspen Forest and est ecoregion but more likely is at the edge of the Parkland (hereafter Aspen Parkland), and Western Gulf Coast Grasslands. Northern Tall Grasslands (primarily in northern Other records of breeding or summering Iowa) ecoregions (Table 3.1, Fig. 3.3). Primary cranes are scattered across various ecoregions

34 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane

FIGURE 3.2 Distribution of historic Whooping Crane locations, 1722–1941, by life-history stage, overlain on WWF biomes. Records from Allen (1952) and additional records (this chapter). Multiple records may be represented by a single symbol. Scale = 1:28,000,000.

Results 35

FIGURE 3.3 Locations of historic Whooping Crane records, 1722–1941, by life-history stage, in northern breeding and summering areas, overlaid on WWF ecoregions. The core breeding range was delineated by Allen (1952). Records from Allen (1952) and additional records (this chapter). Only those ecoregions having at least two breeding or summering crane records are identified in the legend. Multiple records may be represented by a single symbol. Scale = 1:15,000,000. from the Arctic tundra and northern forest territory (Kuyt, 1993; Novakowski, 1966; Van ecoregions to southern forests and grasslands. Schmidt et al., 2014) and can appear in late sum- Like Allen (1952), we believe many summer mer as adults in small groups or as pairs. records were wandering subadult or nonbreed- Wintering locations – The Western Gulf Coast ing birds; such wandering has been documented Grasslands also was the most important win- by satellite tracking of subadult and nonbreed- tering ecoregion, encompassing 72.5% of the ing cranes in the Eastern Migratory Population wintering records, with records dispersed (Mueller et al., Chapter 11, this volume; Urbanek along much of its length (Fig. 3.4). As noted by et al., 2014). Unlike Allen, however, we suspect Allen (1952, p. 28), 20 specimens were taken some of the summering records and records of in the area between 1889 and 1904. Today, the unknown life-history stage were breeding birds AWBP Whooping Cranes still winter in this because we now know that, in any one year, ecoregion. Allen identified another impor- cranes that fail at nesting still retain their nest tant wintering area (12.5% of winter records)

36 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane

FIGURE 3.4 Distribution of historic Whooping Crane locations, 1722–1941, in southern breeding and wintering areas, with ranges delineated by Allen, overlain on WWF ecoregions. Records from Allen (1952) and additional records (this chapter). Only those ecoregions having at least two breeding, summering, or wintering crane records are identified in the legend. Multiple records may be represented by a single symbol. Scale = 1:18,000,000. along the mouth of the Rio Grande River near locations in Florida were likely associated with Brownsville, Texas, located within the Tam- wetlands embedded in prairie (e.g., Kissim- paulipan/Mezquital ecoregion, just inland mee prairie). from the Western Gulf Coast Grasslands ecore- Allen (1952) listed 10 records for wintering gion. Whooping cranes were found wintering cranes in the interior of Mexico; we found no in the interior of Florida and along the Atlan- additional records. None of the records sup- tic coast in association with the Southeastern ported Allen’s extension of the wintering range Coniferous Forest and the Northern, Middle into the northern interior highlands of Mexico. Atlantic, and Southeastern coastal forests. All Whooping cranes were observed at Bolson coastal locations were associated with coastal del Mapimi, Chihuahua (Chihuahan Desert); wetlands or river deltas (e.g., Savannah River on the Plains of Silao, Guanajuato (Trans- and Waccamaw River, South Carolina). Inland Mexican Volcanic Belt Pine-oak Forests); near

Discussion 37

Lerdo, Durago (Mesata Central Matorral); and well recognized (Canadian Wildlife Service and in the state of Jalisco (Bajio Dry Forests; the U.S. Fish and Wildlife Service, 2005; Urbanek southernmost locations), all of which appear and Lewis, 2015; Walkinshaw, 1973). The spe- to be associated with large playa lake or ripar- cies’ large body size, energetic demands, and ian systems. The latter included three records flightlessness during wing molt Urbanek( and somewhere near La Barca, possibly along the Lewis, 2015) necessitate a reliance on produc- Rio Lerma or Rio Santiago, and one on marshes tive wetland habitat that maintains permanent of Lake Chapala, the largest freshwater lake in water for safety from predators and ensures Mexico (1,112 km2). sufficient food resources for themselves and, during breeding, for their chicks (Welling- ton et al., 1996, p. 82). Foods consumed by DISCUSSION Whooping Cranes during breeding and wintering periods tend toward higher trophic Allen’s (1952) description of the historic levels, such as crabs, snakes, minnows, and breeding and wintering distribution of Whoop- small mammals; energy-rich tubers, berries, ing Cranes appears to be largely accurate. The corn, wheat, and acorns are also seasonally additional 70 locations, however, combined important (Allen, 1952; Pugesek et al., 2013; with GIS-based information about landscape Urbanek and Lewis, 2015; International Crane and wetland features within ecoregions, pro- Foundation, unpublished data). Abundance of vide new insights about the landscapes and high-energy foods on the wintering grounds habitats used by the species before some of the is critical to the survival of over-wintering major environmental alterations of habitats Whooping Cranes (Chavez-Ramirez, 1996; occurred in the mid to late 1900s. We discuss Hunt and Slack, 1989; Pugesek et al., 2013), the life-history strategies of Whooping Cranes as well as for the acquisition of sufficient fat as understood under contemporary conditions, reserves to provision energy demands to sup- then reexamine and redefine Allen’s descrip- port their 4,000-km spring migration (Chavez- tion of niche based on habitat features of the Ramirez, 1996; Urbanek and Lewis, 2015) and ecoregions and ecological processes. Here we egg formation upon arrival. Territorial fidelity define the species’ ecological niche in terms of in breeding and winter periods confers ben- habitat – that is, as the noninteractive (sceno- efits through greater familiarity with habitat poetic) variables and environmental conditions quality (e.g., nest sites, food resources) and on broad scales, relevant to understanding risks of predation or disturbance (Piper, 2011; coarse-scale ecological and geographic proper- Switzer, 1993). ties of the species (Soberón, 2007, p. 1117), such Periodic drawdown is important to sustain- as climate, terrain, and land cover features. ing wetland productivity (Mitsch and Gos- selink, 2000) and hence important to sustain Habitat Attributes Important to Life long-term habitat quality for cranes. However, Strategies drawdown conditions may limit both food resources and safe feeding areas and con- Historic and current patterns of Whooping tribute to poor chick survival (Kuyt, 1981a; Crane distribution demonstrate that breed- Spalding et al., 2009). Drawdown conditions ing and wintering populations use a diversity may be similarly detrimental to adult Whoop- of wetland and wet grassland habitats. The ing Cranes in summer, as adults become dependence of Whooping Cranes on fresh and flightless during a synchronous remigial brackish wetlands across life-history stages is molt every 2–3 years (Folk et al., 2008; Lacy

38 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane and McElwee, 2014). Wetlands or wetland sedge marshes and swales, covered with stand- complexes would therefore need to be large ing water and having the biotic characteristics enough to maintain some water in a substan- found in the willow communities of the Aspen tial portion of the cranes’ territory so as to Parkland. There must be a great abundance of provide safe areas for foraging and molting small animal life, including basic invertebrate habitat (i.e., shallow open water and flooded forms. The entire area must be several hundred emergent vegetation) in summer, while also (or even several thousand) acres in size and having the ecological processes in place to completely isolated from human disturbance support high productivity (Mitsch and Gosse- of any sort.” These features have been used to link, 2000). Areas where most wetlands dry out inform selection of new sites for reintroduc- entirely annually or in most years would not tion efforts (Canadian Wildlife Service and U.S. provide suitable habitat for breeding or flight- Fish and Wildlife Service, 2005, p. 150; Can- less Whooping Cranes. For deltaic or coastal non, 1999). Allen’s general features are consis- wetlands, productivity is sustained by inflow tent with findings of contemporary habitat-use of nutrients via flooding or tidal influxes but studies at Wood Buffalo National Park (WBNP) may similarly be reduced during low-water (Kuyt, 1981b, 1993; Olson and Olson, 2003; conditions (Mitsch and Gosselink, 2000). Timoney et al., 1997; Timoney, 1999), ANWR Given these life history features, areas with (Labuda and Butts, 1979), and southwestern extensive productive wetlands would be most Lousiana (Kang and King, 2014). likely to support crane territories on the breed- Based on our evaluation of the habitat fea- ing grounds in most years, provide safe feed- tures of the historic locations and contemporary ing and molting sites in summer (molt cycles knowledge of wetland ecology and Whooping are variable, and thus unpredictable, among Crane life history (see Barzen, Chapter 15, this individual birds; Folk et al., 2008), and provide volume), we identify four features that are impor- adequate food to support cranes during winter. tant for nesting and wintering habitats. First, Further, shallow flooded habitat provides for habitats used by Whooping Cranes have gentle isolation of nests, chicks, flightless cranes, and to rolling topography with an interspersion of roost sites (all seasons) from mammalian preda- wetland and low meadow or prairie habitats and tors. Habitats that are visually open (i.e., little relatively sparse cover of trees and shrubs. Sec- vertical vegetation obstructing view) enhance ond, Whooping Cranes were commonly found cranes’ ability to detect threats and have long in areas having high densities of wetlands or been recognized as an important feature of areas wetland complexes that provide open, shal- used by cranes throughout their life cycle (Arm- lowly flooded habitat. Primary examples of such bruster, 1990; Howe, 1989; Timoney, 1999). conditions are found in the PPR, portions of the southern boreal-taiga region, inland river deltas, Redefining Allen’s Niche Description large playa lakes in the high plains of Mexico, the Chenier Plain of coastal Louisiana and Texas, Allen (1952) provided the first detailed and estuarine or deltaic systems along the Gulf assessment of habitats used by whooping Coast. Third, wetland complexes collectively cranes. He frequently referred to their “biotic provide reliable habitat conditions important for niche” – habitat features that were similar across nesting, brood rearing, molt, or feeding in most breeding and wintering locations. Allen (1952, years, even though hydrological variation within p. 48) described the “preferred niche, especially an individual wetland might be large. Fourth, for nesting, as a flat or slightly rolling, open historical locations were usually in areas that are area interspersed with bulrush, cattail, and considered highly productive due to fertile soils,

Discussion 39 hydrological pulsing, periodic inflow of nutri- which Allen considered the historic center of ents, or other periodic perturbations such as fire breeding, corresponds to the southernmost or storms (Euliss et al., 1999; Mitsch and Gosse- extent of the PPR. Before European settle- link, 2000; Prince, 1997). Allen (1952) touched on ment, wetlands in that portion of Iowa were many of the features we examine in more detail predominantly temporarily flooded to satu- below but lacked the depth of knowledge about rated wetlands (84% of wetland area; Miller ecological processes important to sustaining et al., 2012), embedded in tallgrass prairie. By these conditions. In the following sections, we the 1870s, much of Iowa was settled, and most discuss the significance of these four features in wetlands were drained by the 1930s (Prince, the three primary regions used by breeding and 1997; Schrader, 1955). Severe drought during wintering Whooping Cranes: PPR and northern the 1930s, combined with hunting and other grasslands, boreal forest and taiga, and southern persecution, appeared to be key factors in the region. extirpation of Whooping Cranes from the Sas- PPR and northern grasslands – As recognized katchewan prairies (Hjertaas, 1994) and likely by Allen (1952, pp. 24–25), the greatest num- elsewhere in the PPR. ber of historic nesting records occurred in the Crane observations in the Northern Tall Canadian Aspen Parkland, Central Tall Grass- Grasslands largely fall within the area of rich, lands, and Northern Tall Grasslands, largely deep lake deposits of glacial Lake Agassiz, corresponding to the PPR (Fig. 3.3). Com- which once extended along the Red River Val- pared to boreal regions, frequent nest records ley from northern South Dakota, across the from the PPR and northern grasslands may be northern third of Minnesota, and extending explained in part by the greater opportunity into Manitoba and western Ontario. This flat, for detection by explorers and early settlers open prairie region once had a fractured sys- in the late 1800s and the growth of interest in tem of low intersecting ridges and shallow wet- ornithology in that era. However, the density lands (Augustadt, 1955). The deep rich soils, of records also indicates these regions were relatively high annual precipitation, hydro- historically important for breeding and sum- logical pulses of wet/dry years, and wildfires mering Whooping Cranes. would have supported highly productive wet- The PPR is broadly delineated by its gla- lands and grasslands, and periodic extensive cial history, with high wetland densities, high flooding and wildfires would have severely soil fertility, warm summers, and seasonal limited woody growth. Most wetlands in this and interannual hydrological pulsing (Acton region were lost to extensive drainage efforts in et al., 1998; Euliss et al., 1999; Mitsch and Gosse- the 1920s and 1930s (Prince, 1997). link, 2000; van der Valk, 1998). Precipitation in Extensive wet prairies and wetlands also the PPR increases to the east and north (Millett occurred in northern and central Illinois and et al., 2009); therefore, the more northern and northeastern Indiana (Prince, 1997; Robertson eastern portions of the ecoregions tend to have et al., 1997), which fall within the Central Forest- more reliable water conditions across years and Grassland Transition ecoregion. Whooping are less prone to extended periods of drought cranes were known to frequent that area, largely than the more westerly PPR ecoregions (Acton as migrants (Butler, 1898; Woodruff, 1907). The et al., 1998; Smith et al., 1964). This corresponds Upper Midwest Forest-Savanna Transition sup- to Whooping Crane locations in the PPR, which ported some breeding and summering cranes, were largely in the Aspen Parkland and the largely in locations having extensive lake, Northern Tall Grasslands (Table 3.2, Fig. 3.3). wetland, and bog complexes. The relatively few A cluster of nesting records in northern Iowa, records suggest these two ecoregions were of

40 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane

TABLE 3.2 Distribution of Historic Whooping Crane Breeding and Summering Locations within the PPR by WWF Ecoregion. Breeding Records Were Those That Had Observations or Specimens of a Nest, Eggs, or Flightless Chick; Other Observations between Mid-May and 30 August Were Categorized as Summering. Observation Records in September–Mid-May or Late March–Mid-May Were Classified as Migrating and Those of December–Mid-March as Wintering. Records Lacking a Date Were Classified as Unknown.

Total Breeding Summering Migrating Wintering Unknown

Ecoregion No. % No. % No. % No. % No. % No. %

Canadian Aspen 88 32.6 12 24.5 39 44.3 17 17.3 0 0 20 57.1 forests and parklands Northern mixed 58 21.5 8 16.3 22 25.0 26 26.3 0 0 2 5.7 grasslands Central tall grass- 53 19.6 13 26.5 4 4.5 31 31.6 0 0 5 14.3 lands Northern tall 40 14.8 10 20.4 7 7.9 19 19.4 0 0 4 11.4 grasslands Northern short 29 10.7 6 12.4 15 17.0 5 5.1 0 0 3 8.6 grasslands Upper Midwest 2 0.7 0 0 1 1.1 0 0 0 0 1 2.9 Forest-Savanna Transition Total 270 100.0 49 18.1 88 32.6 98 36.3 0 0 35 13.0

less importance when compared to Northern waterbird communities (Austin and Pyle, 2004; Tall Grasslands, and cranes in some of these far- Cody, 1996; Debinski et al., 1999), including flung locations may have been summer wan- breeding Sandhill Cranes in the Rocky Mountain derers or birds that were unable to complete Population (Gerber et al., 2014). Allen consid- migration. Only one observation (summering) ered the Whooping Cranes reported in south- occurred at Horicon Marsh, a 12,950-ha extinct eastern Idaho and northwestern Wyoming to be glacial lake within the Upper Midwest Forest- summer wanderers rather than breeding birds. Savanna Transition of southeastern Wiscon- However, the frequency of reports of Whooping sin. Horicon Marsh had been manipulated by Cranes in the Greater Yellowstone Area during damming, drainage, and restoration during the 1906–35 and information in Kemsies (1930) sug- 1800s, coinciding with much of the period when gest there might have been a small population Whooping Crane records were acquired. in the central Rocky Mountains. Moreover, the The westernmost cluster of Whooping historic locations of Whooping Cranes are simi- Crane observations falls within the Greater Yel- lar to areas used by breeding and summering lowstone Ecosystem, located in the juncture Greater Sandhill Cranes (Grus canadensis tabida) of Montana, Idaho, and Wyoming (Ricketts of the Rocky Mountain Population (Drewien et al., 1999). This region has extensive riparian et al., 1999). Breeding in this region may have wet meadow systems and some large shallow been constrained, however, by cool climate and wetlands, fed largely by spring run-off from short growing season. The disappearance of mountain snowpack, which support diverse Whooping Cranes from the Greater Yellowstone

Discussion 41

Area coincides with the decline of Sandhill with sedge (Carex) meadows and sparse shrubs Crane numbers as the region was settled during and conifers. The area is typified by long sum- the early 1900s (Drewien and Bizeau, 1974), both mer day length and short growing season, but likely the victims of shooting. climate varies among these CEF ecoregions from Boreal forest and taiga – Observations of cool to warm summers (ESWG, 1995). The cur- Whooping Cranes in the huge expanse of the rent breeding area in WBNP is in an area with Boreal Forest/Taiga biome and its ecoregions high densities of shallow wetland complexes were much scarcer. Observations are likely and sparse woody cover (Novakowski, 1966; biased toward locations near large rivers and Olson and Olson, 2003; Timoney, 1999). Many lakes, as these were the main travel routes of wetlands currently used by breeding Whoop- explorers and settlers who recorded biological ing Cranes are associated with smaller creeks or information, and these geographic features were rivers. Riparian flow, periodic drought, and fire among the relatively few named features of the likely support wetland productivity, although it landscape. Despite imprecise locations, the fea- is relatively low. During dry years, cranes tended tures for existing records correspond to our pro- to shift use areas closer to riparian habitat, which posed ecological niche. Broadly, the landscapes would provide more stable water conditions and where cranes were observed have flat or rolling provided foods, such as minnows (Kuyt, 1993). topography and high wetland densities – low- In this system, cranes likely need large breed- lands, river deltas, and shoreline wetlands on ing territories (averaging 4.1 km2; Kuyt, 1993) large lakes. Ten locations occurred in associa- that encompass both the more stable riparian- tion with river deltas or large riparian-wetland supported wetlands and those that dry more fre- complexes. Such areas have a high density and quently and may be more productive. interspersion of wetlands, extensive shore- Interestingly, the current breeding area lies lines, diverse wildlife, rich growth of emergent just north of the Peace-Athabasca Delta, the and submergent vegetation supported by river largest freshwater inland delta in the world, rec- inflows, sediments, and annual spring floods, ognized for high productivity, ecological diver- relatively warm summers, and long summer sity, and high densities of breeding waterfowl day length, which maximize the growing season (Mitsch and Gosselink, 2000). Despite European for that latitude (Mitsch and Gosselink, 2000; settlements starting in the late 1700s and active Peters et al., 2006). Allen (1952) also often rec- research in the area over the last 50 years or ognized the importance of marshes associated more, only one Whooping Crane observation with river deltas. was recorded in those deltaic wetlands (Fort Lakes used by Whooping Cranes were Chipewyan in 1898). We speculate that breed- located in areas with extensive wetland and lake ing cranes may be unable to establish territories complexes. Four records were reported on two or successfully nest in the large delta due to the huge lakes in central Manitoba (three on Lake magnitude or timing of flood pulses and lim- Winnipeg and one on Lake Winnipegosis) in the ited availability of unflooded habitat during the late 1800s. While vague in location, these two early breeding season, but it doesn’t explain the lakes are remnants of glacial Lake Agassiz and absence of summering birds. Traditional ecolog- lie within large lowland areas of extensive shal- ical knowledge of the aboriginal communities low palustrine wetlands and peatlands. of the area may yield further insights into this Crane locations in and around WBNP fall at the uncertainty. edge of the Hay River Lowlands and Northern A few summering cranes were observed Alberta Uplands CEF ecoregions, in an area hav- in the far north in the Taiga Plains of the ing a high density of small ponds interspersed Northwest Territories. This is an expansive area

42 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane

encompassing several major river deltas and habitats for foraging, drinking water, and safe other areas that have high densities of small roost sites appears to be an important aspect lakes and fens, dominated by sedges and Sphag- of this ecoregion for wintering birds. Allen num moss, interspersed with shrubby tundra (1952, pp. 28–39) identified 10 different habi- (ESWG, 1995; Ricketts et al., 1999). The area has tat types within this ecoregion, ranging from a very short growing season, long summer day tallgrass prairie inland to coastal lagoons and length, a subarctic climate, and continuous per- beaches, and noted differences in use by win- mafrost that would support limited productiv- tering versus breeding birds (discussed later). ity of larger animal food resources important to Much of the inland portion was converted to Whooping Cranes. rice production by 1900 (Gomez et al., 2003), Southern areas for wintering and nonmigratory but remaining freshwater marshes in the breeding cranes – Records added to Allen’s (1952) region sustain a high biodiversity (Esslinger wintering range indicate a more extensive if and Wilson, 2001). spotty distribution along the south Atlantic and Eight of the 10 southern breeding records also Gulf coast (e.g., North Carolina and Florida), as fall within the Western Gulf Coast Grasslands. well as three inland sites in Florida. Most of the Four of the added breeding records were away eastern records are coastal locations associated from the coast in Texas, possibly in Allen’s prai- with river mouths (deltas) or coastal freshwater rie swale and prairie marsh type. Five of the to brackish wetland systems. The majority of breeding records were in the vicinity of White winter records fall within the Western Gulf Coast Lake, an area of extensive and highly productive Grasslands ecoregion, which also sustained the shallow marshes located in the Chenier Plain of only known nonmigratory breeding population western Louisiana (see King et al., Chapter 22, at White Lake, Louisiana. This ecoregion is one this volume). These marshes were described in of the most wetland-rich regions of the world, the 1930s as 16,000 ha in area, 12–20 cm deep, with high productivity sustained through tidal and largely inaccessible until the Intracoastal pulsing and nutrient-rich inflows of fresh water Waterway was extended in 1929–30 (J. Lynch, (Esslinger and Wilson, 2001; Mitsch and Gosse- in Drewien et al., 2001). The freshwater, maid- link, 2000; Smith et al., 1989). It is characterized encane (Panicum hemitomon) marshes such as by flat topography, high soil fertility, high aver- those found at White Lake were considered the age rainfall, and a long growing season. Peri- most important to breeding birds (Allen, 1952; odic disturbance from wildfire, grazing (bison Lynch in Drewien et al., 2001) but little used by historically, domestic cattle since the early wintering birds. White Lake today is relatively 1900s), and hurricanes, and nutrient inflow intact ecologically and is one of the largest unde- from river inflows, storm surges, and daily tidal veloped freshwater marshes in the southeast- fluctuations sustain the high productivity of the ern United States (Gomez et al., 2003). Recent wetland and grassland systems of the region assessment of the coastal Louisiana habitats and (Frost, 1995). Importantly, the inland portion of food resources demonstrated that the freshwa- this ecoregion encompasses what was once tall- ter marshes could support Whooping Cranes, grass prairie, interspersed with numerous small particularly in spring and summer (Kang and wetlands and scattered oak savanna (Esslinger King, 2014). The final one of the 10 southern and Wilson, 2001), where cranes could forage breeding records was located on Boca del Rio and find freshwater water when food availability (mouth of the Rio Grande), a once-extensive in coastal wetlands was low (Blankenship, 1976; coastal delta of the Rio Grande near Bagdad Labuda and Butts, 1979). Juxtaposition of these Matamoros, Tamaulipas, Mexico.

Summary and outlook 43

The wetlands in the interior of Mexico once speculate on the source of the Whooping Cranes supported some wintering Whooping Cranes. in interior Mexico, as he did for birds winter- Deltiac areas associated with large interior lakes, ing in Louisiana and Texas. We speculate that a such as the marshes of Lake Chapala, were likely small, western group of Whooping Cranes once important wintering sites. River inflows would migrated from the Idaho-Wyoming area through have provided nutrient inputs, periodic flood- Great Salt Lake, Utah (Allen, 1952, p. 17) down ing disturbance, and reliable water conditions the Front Range of the Rocky Mountains to win- in most years. Other observations in the Mexi- ter along the Rio Grande, New Mexico, and on can interior were in areas of semiarid to arid interior Mexican wetlands, similar to the path shrub/grasslands with small to large ephemeral now taken by the Rocky Mountain Population lakes that were often brackish or saline (Hen- of Sandhill Cranes. Riparian and shallow lake rickson and Johnston, 1986; Saunders and Saun- habitats along that path (e.g., San Luis Valley, ders, 1981). Under good water conditions, such Colorado, Rio Grande in New Mexico) would basins provided safe roost sites and high produc- have provided appropriate habitats for migrant tivity of aquatic plant foods (Goldman, 1951). cranes. Additional evidence suggesting a western However, interannual variability in water condi- flyway and wintering area includes the depiction tions of these playa-like basins may have been of a Whooping Crane on Kiva murals in New too large to support consistent winter use. The Mexico (Hibben, 1975), observations of Whoop- Whooping Crane’s wintering distribution in ing Cranes at Fort Thorn on the Rio Grande in interior Mexico may therefore have shifted annu- southern New Mexico in 1855 (Bailey, 1928), ally in response to changing water and wetland and observations of “G. americantus ” in New conditions and food availability, as observed for Mexico during migration in March and October Sandhill Cranes (Drewien et al., 1996; Saunders (Henry, 1859). Further research into historic and Saunders, 1981). Intensified land and water Spanish and Native American resources may use have degraded many of Mexico’s interior yield additional support for this possible migra- wetlands, including those at Lake Chapala (Cer- tion path. vantes and Abarca, 1996; Drewien et al., 1996; Most sources used by Allen (1952) for his Limón and Lind, 1990; Perez-Arteaga et al., 2002; historic distribution map came from English, Saunders and Saunders, 1981). French, and American explorers and ornitholo- Allen’s winter distribution extended into the gists. There are various sources that have yet to interior highlands of northcentral Mexico but be examined that may yield new information, he provided no specific records associated with particularly older Spanish sources for Mexico, that area. That delineation appears to be based southwestern United States, Florida, and possi- on communications about Sandhill Crane migra- bly Cuba. Tapping into the traditional ecological tion and wintering areas described in communi- knowledge of aboriginal societies also may yield cations to Allen from George B. Saunders, a U.S. greater insights into distribution and the ecology Fish and Wildlife Service biologist who spent of areas historically used by Whooping Cranes. many years surveying waterfowl and other birds across Mexico (Allen, 1952, pp. 37–38). Saun- ders’s descriptions of wetlands used by winter- SUMMARY AND OUTLOOK ing Sandhill Cranes match those noted above for Whooping Cranes (broad, shallow lake basins, Ideally, priority for reintroduction sites should surrounded by grasslands, or extensive marshes be placed within areas where Whooping Cranes and wet meadows in river deltas). Allen did not historically bred and wintered. Unfortunately,

44 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane this option is often not feasible due to habi- core or peripheral area are (1) subtle to rolling tat loss or degradation. Wetland and grassland topography with an interspersion of wetland habitats through much of the central portion of and low meadow or prairie habitats, with rela- the Whooping Crane’s historic breeding range tively sparse cover of trees and shrubs; (2) high have been dramatically altered, primarily by densities of shallow, open wetlands or wetland agriculture (Dahl and Gifford, 1996; Prince, 1997; complexes; (3) hydrological regimes providing Samson et al., 1998). Agriculture and industrial reliable conditions for nesting, brood rearing, development (chiefly timber harvest and oil and flightless adults; and (4) high productiv- development; Foote and Krogman, 2006) also ity due to fertile soils, hydrological pulsing, are expanding into the southern boreal forest periodic inflow of nutrients, or other periodic (midcontinent Canadian forest ecoregion). In perturbations. Some important wetland areas the boreal plain of Saskatchewan, Hobson et al. within the historic range but outside of ANWR (2002) reported 73% of the forest in the historic and WBNP remain relatively intact and have boreal transition zone was converted to agricul- been protected under state, provincial, or fed- ture between 1966 and 1994. Human impacts on eral ownership or wetland regulations (e.g., habitats in other boreal ecoregions are substan- Quill Lakes, Saskatchewan; Hails, 1997). Both tially less (Kerr and Deguise, 2004). Although traditional and more innovative approaches northern areas unaffected by habitat change (e.g., conservation agreements with private may be promising for alternative reintroduction landowners and industry) may be needed to sites, they conflict with the current recovery goal protect additional reintroduction sites that have of establishing biologically separate Whooping potential. Reintroduction evaluations addition- Crane populations (Canadian Wildlife Service ally should consider potential impacts of chang- and U.S. Fish and Wildlife Service, 2005). Breed- ing climate, in particular the potential impacts ing Whooping Cranes from new reintroduction on wetland hydrology and food resources. Fur- sites in northern areas would likely overlap with ther research into the hydrology regimes and the current AWBP outside the breeding season. wetland productivity of current reintroduction Further, many currently intact habitats are at risk sites would be valuable to deepen our under- on the main wintering grounds. Coastal wetlands standing of the ecological niche and seasonal continued to be threatened by development, pol- habitat needs of cranes. lution, invasive species, and erosion (Dahl and This study provides new insights into the his- Gifford, 1996; Güneralp et al., 2013; Montagna toric distribution and habitats used by Whoop- et al., 2011). ing Cranes during the 1700s to mid-1900s. We These challenges suggest that the most prom- believe site-specific studies of current habitat ising opportunities for reintroduction or resto- use by Whooping Cranes would benefit from ration may lie in the periphery of the historic greater consideration of hydrological regime and range, where suitable habitats are more intact site productivity, and the ecological processes and less likely to be impacted in the near future that sustain productivity at higher trophic lev- (Channell and Lomolino, 2000). Iowa has lost els (e.g., seasonal hydrological cycles, nutrient 89% of its original wetlands, but immediately dynamics, vulnerability to long-term drought). to the northeast, Wisconsin – most of which is These historic data and insights should stimu- in the Upper Midwest Forest-Savanna Transi- late new discussions about habitats assessments tion ecoregion – has retained just over 50% of used for selection of reintroduction locations its wetlands (Dahl, 1990). Those key habitat fea- and ultimately enhance the long-term success of tures that should be present in any remaining recovery efforts.

REFERENCES 45

Acknowledgments Barzen, J.A., Lacy, A., Thompson, H.L., Gossens, A.P., 2018. Habitat use by the reintroduced Eastern Migratory Popu- We would like to recognize the late Robert P. Allen for the lation of Whooping Cranes (Chapter 14). In: French, Jr., extensive time and effort put into collecting and organizing J.B., Converse, S.J., Austin, J.E. (Eds.), Whooping Cranes: information in his 1952 monograph about the Whooping Biology and Conservation. Biodiversity of the World: Crane. The Canadian and U.S. Whooping Crane Recovery Conservation from Genes to Landscapes. Academic Teams, and all those involved, have worked diligently to Press, San Diego, CA. protect the Whooping Crane and the habitats it utilizes. Ad- Bent, A. C., 1926. Life histories of North American marsh ditional historical Whooping Crane records were collected birds. Bull. U.S. Natl. Mus. 135, Smithsonian Institution. through the assistance of George Archibald, Ann Burke, Mike Blankenship, D.R., 1976. Studies of whooping cranes on the Putnam, and Betsy Didrickson. Dorn Moore provided guid- wintering grounds. In: Lewis, J.C. (Ed.), Proceedings of the ance in building the database. Hugh Britten and Bill Mitsch 1975 International Crane Workshop. Oklahoma State Uni- provided useful comments in the earlier development of versity Publishing and Printing, Stillwater, pp. 197–207. the manuscript. Rod Drewien, Felipe Chavez-Ramirez, and Brown, J.H., 1984. On the relationship between abundance the late John Taylor provided useful comments concerning and distribution of species. Am. Nat. 124 (2), 255–279. the distribution map. We thank Leigh Fredrickson, Sammy Brown, J.H., Mehlman, D.W., Stevens, G.C., 1995. Spatial King, and an anonymous reviewer for their comments on variation in abundance. Ecology 76 (7), 2028–2043. later versions of this manuscript. Any use of trade, firm, or Brown, J.H., Stevens, G.C., Kaufman, D.M., 1996. The geo- product names is for descriptive purposes only and does not graphic range: size, shape, boundaries, and internal imply endorsement by the U.S. Government. structure. Annu. Rev. Ecol. Syst. 27, 597–623. Butler, A. W., 1898. The Birds of Indiana. Twenty-second an- References nual report, Indiana Department of Geology and Natural Abell, R., Thieme, M.L., Revenga, C., Bryer, M., Kottelat, Resources, 575–1187. M., Bogutskaya, N., Coad, B., Mandrak, N., Contreras, Canadian Wildlife Service and U.S. Fish and Wildlife Ser- S.B., Bussing, W., Stiassny, M.L.J., Skelton, P., Allen, G.R., vice, 2005. International Recovery Plan for the Whoop- Unmack, P., Naseka, A., Ng, R., Sindorf, N., Robertson, ing Crane. Recovery of Nationally Endangered Wildlife J., Armijo, E., Higgins, J.V., Heibel, T.J., Wikramanayake, (RENEW), Ottawa, and U.S. Fish and Wildlife Service, E., Olson, D., López, H.L., Reis, R.E., Lundberg, R.J., Sa- Albuquerque, NM. baj, M.H.P., Petry, P., 2008. 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Appendix 49 (Continued) Observer Farley F.L. Bobbie Herron George Dr. Farley F.L. Sidney Stansell Sidney Stansell Sidney Stansell Farley F.L. Sidney Stansell Park keeper Farley F.L. Farley F.L. C. H. Davis C. H. Davis J. Dewey Soper C. E. Mills Gillese J. P. King Richard H. Jones Seton and Preble Sidney Stansell Farley F.L. Farley F.L. Life stage Breed Breed Breed Breed Breed Breed Breed Migr Migr Migr Migr Migr Migr Migr Summ Summ Summ Summ Summ Summ Day 11 15 22 13 16 21 Month Oct Oct Oct Apr Jun May Oct May May Year 1905 1917 1904 1906 1907 1908 1909 1909 1914 1886 1886 1906 1910 1919 1922 1933 1835 1905 1907 1909 1913 1919 111.850 110.620 114.150 111.850 113.470 113.470 113.470 112.250 113.470 118.460 112.830 112.830 111.000 111.000 110.550 114.530 114.250 111.130 113.920 111.380 114.000 115.300 112.670 Longitude − − − − − − − − − − − − − − − − − − − − − − − Latitude 52.780 54.420 52.070 52.780 53.550 53.550 53.550 53.870 53.550 59.320 53.020 53.020 52.770 52.770 53.400 53.750 54.080 58.700 55.970 56.730 53.530 55.430 53.370 Country Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can APPENDIX State/ province AB AB AB AB AB AB AB AB AB AB AB AB AB AB AB AB AB AB AB AB AB AB AB County Chipewyan McMurray City Killam Killam Edmonton Edmonton Edmonton Edmonton Wainright Camrose Camrose Islay Fort Fort Stony Plain Tofield River National Park Location Jackfish Creek Reed Deer Lake Whitford Buffalo Wood Battle River Battle River Lake Buffalo Lake Buffalo Lake Birch Paddle River River Willow Swan River

(Historical Records of Whooping Crane O bservations, rdered by S tate/Province and L ife-History tage Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 50 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane and E. R. Kalmbach Rockwell Gordon Observer Farley F.L. Farley F.L. Farley F.L. Thomas Hook Unknown Captain Sprinkle D. B. Wiler Spencer Smith Smith Poley F. F. Hershey and Mrs. Clara Fischer Dr. Dyke S. Van T. Life stage Summ Summ Summ Summ Migr Migr Wint Migr Migr Migr Migr Migr Migr Summ Wint Wint Day 13 4 22 30 5 22 8 16 13 20 Month Sep Jun Sep Nov Nov Apr Apr Apr Oct Jun ) Year 1920 1923 1927 1899 1882 1914 1880 1889 1890 1941 1931 1889 1894 1722 1723 cont. 112.000 112.830 113.080 113.920 86.506 87.751 88.184 91.220 90.580 104.821 105.074 105.074 102.789 104.383 105.084 103.320 103.850 81.323 81.323 Longitude − − − − − − − − − − − − − − − − − − − Latitude 52.000 53.020 53.050 55.970 32.379 32.661 30.249 34.444 36.408 38.834 40.398 40.398 38.761 39.867 40.585 25.330 25.817 29.895 29.895 States Country Can Can Can Can US US US US US US US US US US US Mex Mex United US State/ province AB AB AB AB AL AL AL AR AR CO CO CO CO CO CO DR DR FL FL County Autauga Hale Mobile Arkansas Clay El Paso Larimer Larimer Cheyenne Adams Larimer Saint Johns Saint Johns Bluff Springs Augustine Augustine City Camrose Camrose Prattville Millwood Crockett’s Corning Colorado Loveland Loveland Lerdo Saint Saint Creek Refuge Mapimi Location Sullivan Lake Bittern Lake River Willow Mulberry Slough Cypress Dauphin Island White River Kit Carson Fort Collins Bolson del Rivers near city Rivers near city Niedrach (1965)

(Source Hallman (1965) (1954) Sprunt (1940) Shaffer Harmon (1954) Bent (1926) Bent (1926) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Griffin (1957) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 52 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane Observer Trostler Trostler Hall Johnson S. Tiberghin Blackburn J. H. Bowles John Krider E. Dickinson Rice W. John Krider C. Reice W. S. Howland C. M. Jones C. M. Jones Farmer Preston J. W. Life stage Breed Breed Breed Breed Breed Breed Breed Breed Breed Breed Breed Breed Breed Breed Breed Day 25 12 15 6 8 12 2 4 14 8 2 4 Month Apr May May May May May May May May May May May ) Year 1868 1868 1871 1871 1874 1877 1877 1879 1880 1880 1881 1881 1882 1883 1883 cont. 90.583 95.533 95.617 94.377 93.583 94.215 93.734 95.117 90.867 92.317 92.317 95.397 95.617 94.684 93.538 93.267 93.386 94.217 93.717 93.386 91.697 93.734 Longitude − − − − − − − − − − − − − − − − − − − − − − Latitude 42.167 41.333 41.033 42.015 41.628 42.721 43.129 42.383 42.483 42.467 42.467 41.310 42.733 42.551 43.420 42.733 43.143 43.200 42.733 43.143 43.005 43.129 Country US US US US US US US US US US US US US US US US US US US US US US State/ province IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA - - tamie Hawk Hawk mie Gordo Gordo County Jackson Pottawat Mills Greene Polk Humboldt Hancock Sac Dubuque Black Black Pottawata Cherokee Calhoun Winnebago Franklin Cerro Kossuth Wright Cerro Floyd Hancock City Jefferson Humboldt Pomeroy Midway Location Eagle Lake Oakland Valley Lake Mills Clear Lake Clear Lake Eagle Lake (1994)

(Historical Records of Whooping Crane O bservations, rdered by S tate/Province and L ife-History tage ( Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Dinsmore Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 Appendix 53 (Continued) Observer R. M. Anderson Local farmers R. M. Anderson Parker D. O. of B. Sandpiper Bond Preston Cline Scougal Cline Scougal Peck Mosker Bond Talbot Scougal Talbot Scougal Bond Bond Bond Life stage Breed Breed Breed Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Day 26 5 5 10 30 13 26 29 1 5 23 30 30 7 22 3 8 8 25 3 25 Month May Jun May Nov Apr Oct Oct Apr Mar Mar Apr Apr Mar Mar Mar Apr Mar Apr Apr Apr Apr Mar Apr Mar Year 1894 1897 1897 1871 1873 1876 1877 1884 1884 1884 1885 1885 1885 1885 1885 1886 1886 1887 1887 1887 1887 1887 1888 1888 93.734 93.734 93.734 93.783 92.192 95.110 94.500 96.033 95.202 91.530 94.674 96.400 94.674 96.400 92.192 95.110 95.202 96.078 96.400 96.078 96.400 95.202 95.202 95.202 Longitude − − − − − − − − − − − − − − − − − − − − − − − − Latitude 43.129 43.129 43.129 40.751 42.493 43.424 42.495 42.383 42.643 41.661 43.108 42.500 43.108 42.500 42.315 43.424 42.643 42.271 42.500 42.271 42.500 42.643 42.643 42.643 Country US US US US US US US US US US US US US US US US US US US US US US US US State/ province IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA Hawk Vista Hawk Vista Vista Vista Vista County Hancock Hancock Hancock Decatur Black Dickinson Calhoun Woodbury Buena Johnson Palo Alto Woodbury Palo Alto Woodbury Black Dickinson Buena Woodbury Woodbury Woodbury Woodbury Buena Buena Buena City Waterloo Iowa City Emmetsburg Sioux City Emmetsburg Sioux City La Porte City Holly Springs Sioux City Holly Springs Sioux City Slough Location Eagle Lake Eagle Lake Eagle Lake Spirit Lake Cedar Creek Creek Wolf Storm Lake Spirit Lake Storm Lake Storm Lake Storm Lake Storm Lake

(Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 54 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane Gabrielson and Thomas Nuttall Observer Bond Byington Johnson Collett Jeffrey Bigelow Jeffrey Jeffrey Jeffrey Jeffrey J. A. Spurrell Rich Ira N. Scougal Berry Anderson Anderson Rich J. K. Townsend Danby Nelson 1877 Life stage Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Summ Summ Summ Summ Summ Summ Summ Breed Day 30 2 3 1 2 2 29 30 23 21 10 9 7 5 15 5 8 8 Month Mar Apr Apr Apr Apr Apr Mar Mar Sep Mar Apr Apr Apr May Jun May May Jul Apr ) Year 1888 1889 1890 1891 1900 1900 1900 1900 1900 1900 1904 1906 1911 1922 1885 1890 1897 1897 1909 1834 1899 1870s cont. 95.202 93.602 96.485 94.729 93.557 93.376 93.557 93.557 93.557 93.557 95.093 96.400 95.012 94.715 96.400 96.485 93.633 93.633 96.400 111.309 116.896 89.630 Longitude − − − − − − − − − − − − − − − − − − − − − − Latitude 42.643 43.104 42.996 42.057 41.358 42.520 41.358 41.358 41.358 41.358 42.271 42.500 42.949 43.304 42.500 42.996 43.262 43.262 42.500 42.297 47.813 40.620 Country US US US US US US US US US US US US US US US US US US US US US US State/ province IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA ID ID IL Vista County Buena Hancock Sioux Carroll Warren Harding Warren Warren Warren Warren Sac Woodbury Clay Emmet Woodbury Sioux Winnebago Winnebago Woodbury Bear Lake Kootenai Peoria City Garner Hawarden Glidden Indianola Alden Indianola Indianola Indianola Indianola Sioux City Webb Sioux City Hawarden City Forest City Forest Sioux City Montpelier Rathdrum Peoria marshes Location Storm Lake Lake Wall High Lake Bear River Cemtral Illinois * (1994) Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Dinsmore Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) ) Historical Records of Whooping Crane O bservations, rdered by S tate/Province and L ife-History tage Appendix 55 (Continued) Bogardus R. and Dr. Mathews Observer Fuchs Adam H. Blackney T. Gault B. T. Baczkowski F. Charles Dury Stein Dr. Trouslot Jouy Wilson E. W. D. E. Lantz H. D. Burchell C. B. Heinricks Life stage Migr Migr Migr Migr Summ Summ Summ Summ Migr Summ Wint Wint Wint Day 27 23 6 15 6 28 25 Month Mar Oct Mar Jun Apr Apr Aug Mar Apr Jan Jan Year 1880 1871 1877 1879 1858 1881 1891 1905 1881 1887 1894 1890 1903 1903 1878 1907 1912 89.186 88.200 88.251 91.434 87.761 87.650 88.622 90.183 87.650 87.061 86.526 88.027 87.365 86.723 87.061 102.570 102.950 102.570 102.570 96.571 97.450 97.669 Longitude − − − − − − − − − − − − − − − − − − − − − − Latitude 36.990 40.133 40.583 40.359 38.411 41.850 40.747 42.350 41.850 41.473 39.165 37.798 41.417 41.611 41.473 20.280 20.190 20.280 20.280 39.184 37.717 38.024 Country US US US US US US US US US US US US US US US Mex Mex Mex Mex US US US State/ province IL IL IL IL IL IL IL IL IL IN IN IN IN IN IN JL JL JL JL KS KS KS County Alexander Champaign Ford Hancock Wabash Cook McLean Jo Daviess Cook Porter Monroe Posey Lake La Porte Porter Riley Sedgwick Harvey City Cairo Warsaw Mount Carmel Chicago Weston Chicago Bloomington La Barca La Barca La Barca Manhattan Burton River Valley Chapala Location Old Apple Lower Wabash Lago de (1878) (1884) (1907) (1955)

(Source Allen (1952) Allen (1952) Bogardus Allen (1952) Allen (1952) et al. Baird Allen (1952) Allen (1952) Woodruff Baczkowski Butler (1898) Butler (1898) Coale (1912) Allen (1952) Butler (1898) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 56 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane Observer A. Crocker Blachly D. E. Lantz Kellogg Kellogg Smith D. E. Lantz D. E. Lantz J. A. Bryant Jaques F.L. Monahan B. Peabody P. M. McClom W. Hal G. Everts Crevecoeur Crevecoeur Hal G. Everts Hal G. Everts Life stage Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Day 15 8 18 27 18 21 15 8 28 7 18 18 6 18 31 13 18 20 Month Apr Apr Mar Mar Mar Mar Apr Apr Mar Mar Mar Apr Apr Apr Mar Mar Oct Oct Oct ) Year 1867 1882 1884 1884 1885 1885 1887 1887 1887 1887 1890 1891 1893 1896 1903 1904 1905 1906 1906 1907 1907 1907 1919 1922 1929 cont. 97.506 95.743 96.571 96.571 96.181 96.181 95.254 95.254 95.254 95.254 95.254 96.571 96.571 95.833 96.571 97.228 95.186 99.326 96.659 99.900 98.600 96.170 96.170 98.600 98.600 99.883 Longitude − − − − − − − − − − − − − − − − − − − − − − − − − − Latitude 38.858 38.194 39.184 39.184 38.404 38.404 38.403 38.403 38.403 38.403 38.403 39.184 39.184 39.400 39.184 38.486 38.775 38.879 39.682 37.700 37.962 39.489 39.489 37.962 37.962 39.350 Country US US US US US US US US US US US US US US US US US US US US US US US US US US State/ province KS KS KS KS KS KS KS KS KS KS KS KS KS KS KS KS KS KS KS KS KS KS KS KS KS KS - - mie mie County Saline Coffey Riley Riley Lyon Lyon Franklin Franklin Franklin Franklin Franklin Riley Riley Jackson Riley Marion Douglas Ellis Marshall Ford Stafford Pottawato Pottawato Stafford Stafford Graham City Burlington Emporia Emporia Richmond Richmond Richmond Richmond Richmond Durham Baldwin Hays Blue Rapids Onaga Onaga Location Saline River

(Historical Records of Whooping Crane O bservations, rdered by S tate/Province and L ife-History tage ( Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 Appendix 57 (Continued) and A. Wilson Wilson Nunez Nunez Nunez Observer J. J. Audubon Audubon and Audubon Duncan Crain Chote Grevillen O’Neil and L. J. Merovka E. A. McIlhenny Ulyese Veazey Duncan Crain Mrs. Gaspard O’Neil and O’Neil and J. Lynch Duncan Crain E. A. McIlhenny E. A. McIlhenny J. J. Audubon Life stage Migr Migr Migr Migr Breed Breed Breed Breed Breed Summ Summ Wint Day 20 28 14 20 15 5 5 Month Mar Oct Nov Aug May Jun Jun Apr Year 1810 1810 1820 1886 1895 1895 1901 1934 1883 1890 1900 1939 1937 1937 1821 85.759 87.590 88.563 89.186 92.975 92.300 93.111 93.111 91.906 92.433 92.403 92.403 93.111 93.111 93.111 93.111 93.111 91.906 89.949 Longitude − − − − − − − − − − − − − − − − − − − Latitude 38.254 37.836 37.065 36.571 29.767 29.817 29.717 29.717 29.898 29.642 29.858 29.858 29.717 29.717 29.717 29.717 29.717 29.898 29.986 Country US US US US US US US US US US US US US US US US US US US State/ province KY KY KY KY LA LA LA LA LA LA LA LA LA LA LA LA LA LA LA County Jefferson Henderson Livingston Fulton Cameron Vermilion Cameron Cameron Iberia Vermilion Vermilion Vermilion Cameron Cameron Cameron Cameron Cameron Iberia Orleans City Louisville Henderson Hickman Pecan Island New Orleans Tennessee Tennessee River to Johnson’s Bayou Marsh Marsh Marsh Marsh Marsh Marsh Marsh Marsh Marsh Location Mouth of Grand Chenier Hellhole White Lake White Lake Island Avery White Lake White Lake White Lake White Lake White Lake White Lake White Lake Island Avery

(Source Allen (1952) Mengel (1965) Mengel (1965) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 58 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane S. C. Arthur and Ulyese Veazey Observer Delcambre Delcambre Delcambre Bailey S. C. Arthur A. M. Bailey and A. M. Bailey Alcie Daigle S. C. Arthur Nelson E. W. Ulyese Veazey Ralph Sagrera Life stage Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Day 11 7 Month Feb Nov Mar Jan ) Year 1895 1895 1895 1899 1916 1916 1916 1917 1918 1928 1928 1929 1929 1930 1931 1931 cont. 92.744 91.989 91.846 93.350 92.300 92.744 92.300 92.300 93.350 92.744 92.433 92.300 92.433 92.300 92.300 92.363 Longitude − − − − − − − − − − − − − − − − Latitude 29.894 29.948 29.566 30.233 29.817 29.894 29.817 29.817 30.233 29.894 29.642 29.817 29.642 29.817 29.817 29.547 Country US US US US US US US US US US US US US US US US State/ province LA LA LA LA LA LA LA LA LA LA LA LA LA LA LA LA County Cameron Iberia Iberia Calcasieu Vermilion Cameron Vermillion Vermillion Calcasieu Cameron Vermilion Vermillion Vermilion Vermillion Vermillion Vermilion City Delcambre Marsh Island Pecan Island Pecan Island Constance Croix Prairies Refuge Constance Tigre Tigre Prairies Constance Tigre Tigre Tigre Island Location Big Bayou Chenier la Grass Tall Louisiana State Big Bayou Chenier au Chenier au Grass Tall Big Bayou Chenier au Chenier au Chenier au Mulberry

(Historical Records of Whooping Crane O bservations, rdered by S tate/Province and L ife-History tage ( Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 Appendix 59 (Continued) and Ovid Abshire and Ovid Abshire and Richard Gordon and Gordon Schexnayder Observer Ralph Sagrera Ralph Sagrera R. B. Worthen Trappers Ralph Sagrera Lionel LeBlanc Ralph Sagrera Jno. Gaspard Daigre Ambrose Welch George Jno. Gaspard C. E. Gillham J. H. Baker A. Simmons, R. J. Lynch J. Lynch Life stage Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Day 16 23 16 27 30 Month Jan Dec Dec Nov Apr Jan Jan Jan Year 1932 1933 1933 1934 1934 1934 1935 1935 1935 1936 1936 1937 1937 1937 1939 1940 92.363 92.363 93.111 92.300 92.363 92.300 92.363 93.111 93.111 93.111 93.111 93.111 93.111 93.111 93.111 93.111 Longitude − − − − − − − − − − − − − − − − Latitude 29.547 29.547 29.717 29.817 29.547 29.817 29.547 29.717 29.717 29.717 29.717 29.717 29.717 29.717 29.717 29.717 Country US US US US US US US US US US US US US US US US State/ province LA LA LA LA LA LA LA LA LA LA LA LA LA LA LA LA County Vermilion Vermilion Cameron Vermillion Vermilion Vermillion Vermilion Cameron Cameron Cameron Cameron Cameron Cameron Cameron Cameron Cameron City Island Island Marsh Tigre Island Tigre Island Marsh Marsh Marsh Marsh Marsh Marsh Marsh Marsh Marsh Location Mulberry Mulberry White Lake Chenier au Mulberry Chenier au Mulberry White Lake White Lake White Lake White Lake White Lake White Lake White Lake White Lake White Lake

(Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 60 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane Committee Observer A. O. U. E. A. McIlhenny Jacobs F. Cresswell F. Hine William E. Thompson E. Thompson brothers Ward Brothers Ward brothers Ward C. Ward F. L. D. Schultz Small A. T. Life stage Wint Wint Wint Breed Breed Breed Breed Breed Breed Day 18 9 Month Aug May May ) Year 1941 1771 1884 1890 1890 1890 1895 1901 1904 1916 1917 1924 1871 1877 1883 1885 1885 1886 cont. 92.975 92.300 93.350 92.310 98.300 97.150 99.370 100.620 100.360 100.620 100.360 100.620 100.620 100.620 97.150 97.270 97.270 98.030 99.920 100.620 Longitude − − − − − − − − − − − − − − − − − − − − Latitude 29.767 29.817 30.233 57.000 49.970 49.880 49.870 50.400 50.000 50.400 50.000 50.400 50.400 50.400 49.880 52.130 52.130 50.500 52.450 50.400 Country US US US Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can State/ province LA LA LA MB MB MB MB MB MB MB MB MB MB MB MB MB MB MB MB MB County Cameron Vermillion Calcasieu Prairie City Portage la Winnipeg Carberry Winnipeg

to Johnson’s Bayou Tigre Prairies Lake Lake Winnipegosis Location Grand Chenier Chenier au Grass Tall Factory York Shoal Lake Whitewater Shoal Lake Whitewater Shoal Lake Shoal Lake Shoal Lake Lake Winnipeg Lake Winnipeg Oak Point Lake Shoal Lake al. (2003)

(Historical Records of Whooping Crane O bservations, rdered by S tate/Province and L ife-History tage ( Source Allen (1952) Allen (1952) Allen (1952) Houston et Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 Appendix 61 (Continued) Observer Raine W. Raine Walter Raine Walter Raine Walter Forge C. P. Small A. T. E. Calcutt Criddle Talbot Wemyss Wemyss Wemyss Wemyss Wemyss Criddle Talbot Wemyss C. K. Worthen Criddle Talbot Isham E. Thompson Small A. T. E. Calcutt E. Calcutt Nash C. W. Nash C. W. Wemyss Wemyss Life stage Breed Breed Breed Breed Breed Breed Breed Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Summ Summ Summ Summ Summ Summ Summ Summ Summ Day 17 21 13 16 18 30 15 16 12 10 17 15 19 19 6 9 19 12 19 1 30 3 23 27 Month Jun May May May May May Apr Apr Apr Apr Apr Apr Apr Apr Apr Apr Oct Oct Apr May Apr May May May Year 1891 1891 1893 1894 1900 1900 1906 1885 1890 1895 1896 1896 1897 1897 1900 1900 1900 1900 1904 1748 1882 1884 1885 1885 1890 1890 1898 1898 97.270 98.400 98.400 98.400 98.400 98.400 97.350 98.030 101.400 99.600 99.470 99.470 97.870 97.870 97.870 99.600 97.870 97.350 99.600 94.080 99.370 98.030 101.400 101.400 98.580 98.580 97.870 97.870 Longitude − − − − − − − − − − − − − − − − − − − − − − − − − − − − Latitude 52.130 49.700 49.700 49.700 49.700 49.700 49.350 50.500 50.970 49.720 50.230 50.230 50.080 50.080 50.080 49.720 50.080 49.350 49.720 58.750 49.870 50.500 50.970 50.970 50.130 50.130 50.080 50.080 Country Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can State/ province MB MB MB MB MB MB MB MB MB MB MB MB MB MB MB MB MB MB MB MB MB MB MB MB MB MB MB MB County City Aweme Neepawa Neepawa Reaburn Reaburn Reaburn Aweme Reaburn Aweme Churchill Carberry Westbourne Westbourne Reaburn Reaburn Location Lake Winnipeg Oak Lake Oak Lake Oak Lake Oak Lake Oak Lake Morris River Oak Point Shell River Morris River Oak Point Shell River Shell River

(Historical Records of Whooping Crane O bservations, rdered by S tate/Province and L ife-History tage ( Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) (1912) Barrows Bailey (1881) (1912) Barrows Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 Appendix 63 (Continued) Observer Miller J. C. Huoslef B. Peabody P. B. Peabody P. B. Peabody P. B. Peabody P. Schrooten Schrooten Schrooten B. Peabody P. B. Peabody P. Schrooten B. Peabody P. Schrooten Schrooten Schrooten Schrooten Schrooten B. Peabody P. B. Peabody P. B. Peabody P. B. M. Slee C. B. Miller Albert Lano Oium G. T. B. Peabody P. Life stage Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Day 30 31 3 31 13 12 11 14 15 3 7 7 20 28 30 7 28 30 17 6 25 12 15 10 10 19 Month Mar Mar Apr Mar Nov Apr Apr Apr Apr Apr Apr Apr Mar Mar Mar Apr Mar Mar Apr Apr Mar Apr Apr Apr Apr Apr Year 1884 1884 1885 1885 1885 1887 1888 1888 1888 1888 1888 1889 1889 1889 1889 1890 1890 1890 1891 1891 1891 1892 1892 1893 1895 1895 95.286 91.977 95.286 95.286 95.286 95.286 94.079 94.079 94.079 95.286 95.286 93.966 95.286 93.966 93.966 94.079 94.079 94.079 95.286 95.286 95.286 96.773 96.773 96.054 96.194 95.286 Longitude − − − − − − − − − − − − − − − − − − − − − − − − − − Latitude 43.798 43.721 43.798 43.798 43.798 43.798 44.291 44.291 44.291 43.798 43.798 45.076 43.798 45.076 45.076 44.291 44.291 44.291 43.798 43.798 43.798 48.197 48.197 44.993 44.809 43.798 Country US US US US US US US US US US US US US US US US US US US US US US US US US US State/ province MN MN MN MN MN MN MN MN MN MN MN MN MN MN MN MN MN MN MN MN MN MN MN MN MN MN Parle Parle County Jackson Fillmore Jackson Jackson Jackson Jackson Nicollet Nicollet Nicollet Jackson Jackson Wright Jackson Wright Wright Nicollet Nicollet Nicollet Jackson Jackson Jackson Marshall Marshall Lac Qui Lac Qui Jackson City Lanesboro North Star North Star North Star Waverly Waverly Waverly North Star North Star North Star Warren Warren Dawson of Madison Location Lake Heron Lake Heron Lake Heron Lake Heron Lake Heron Lake Heron Lake Heron Lake Heron Lake Heron Lake Heron Lake Heron 14 miles south Lake Heron

(Historical Records of Whooping Crane O bservations, rdered by S tate/Province and L ife-History tage ( Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 Appendix 65 (Continued) Mershon Observer Charles Dankers Cameron Thomas Coubeaux H. Williams W. H. Williams W. Williams H. V. Mershon B. William Delos Hatch Eastgate Eastgate Hibbard Judd E. T. Lewis and Clark C. E. McChesney C. E. McChesney Life stage Migr Wint Migr Migr Summ Wint Breed Breed Breed Breed Breed Breed Migr Migr Migr Day 27 15 5 8 1 22 3 18 18 11 Month Mar Apr Oct Apr May Apr Jun May May Apr Year 1913 1902 1904 1918 1903 1875 1899 1908 1919 1935 1871 1894 1908 1909 1915 1917 1805 1876 1876 95.421 89.330 105.313 108.500 110.113 77.945 97.388 100.889 98.336 97.298 98.935 99.751 99.751 99.814 97.626 98.336 97.862 100.405 100.066 102.323 102.533 102.533 Longitude − − − − − − − − − − − − − − − − − − − − − − Latitude 40.439 30.309 46.794 45.783 48.179 34.226 48.854 46.827 48.043 48.485 48.853 46.869 46.869 48.783 47.907 48.043 48.497 48.346 48.942 47.597 48.767 48.767 Country US US US US US US US US US US US US US US US US US US US US US US State/ province MO MS MT MT MT NC ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND Hanover Forks County Atchison Hancock Prairie Yellowstone Chouteau New Pembina Morton Nelson Walsh Cavalier Kidder Kidder Rolette Grand Nelson Walsh McHenry Rolette Dunn Burke Burke Louis Prairies Prairies City Corning Bay Saint Terry Billings Big Sandy Wilmington Big Slough Mandan Lakota Cashel Calvin Dawson Dawson Ina Larimore Lakota Adams Towner Couteau des Couteau des the Little Missouri River Location Mouth of

(Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Bailey (1881) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) (1956) Hibbard Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 66 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane McCumber Observer C. E. McChesney C. E. McChesney C. E. McChesney C. E. McChesney Tyler Tyler Louis A. Yorke Bowen Judd E. T. Eastgate Judd E. T. Bowen Bowen H. Williams W. Bowman Colling D. Warren H. Judd E. T. G. Bruening Life stage Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Day 12 5 11 18 5 8 10 11 11 18 12 11 13 18 Month Apr Apr Oct Apr Oct Apr Apr Apr Apr Apr Oct Apr Oct Apr Oct ) Year 1877 1877 1878 1878 1885 1885 1889 1890 1890 1893 1894 1894 1895 1899 1903 1904 1910 1912 1913 1917 1919 1920 cont. 102.533 102.533 102.533 102.533 100.531 100.531 99.751 97.476 99.200 97.626 100.066 97.476 97.476 97.447 98.931 97.644 99.489 97.453 99.443 100.066 97.752 99.916 Longitude − − − − − − − − − − − − − − − − − − − − − − Latitude 48.767 48.767 48.767 48.767 46.821 46.821 46.869 48.877 48.683 47.907 48.942 48.877 48.877 48.706 48.028 48.151 46.733 48.809 47.009 48.942 48.923 46.855 Country US US US US US US US US US US US US US US US US US US US US US US State/ province ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND Forks Forks County Burke Burke Burke Burke Burleigh Burleigh Kidder Pembina Towner Grand Rolette Pembina Pembina Pembina Ramsey Grand Kidder Pembina Stutsman Rolette Pembina Kidder Prairies Prairies Prairies Prairies City Couteau des Couteau des Couteau des Couteau des Menoken Menoken Dawson Bathgate Larimore Bathgate Bathgate Glasston Inkster Hamilton LeRoy Steele Location Devils Lake Lake George Chase Lake

(Historical Records of Whooping Crane O bservations, rdered by S tate/Province and L ife-History tage ( Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 Appendix 67 (Continued) McCumber McCumber Observer C. E. Boardman Russell Reid H. B. Williams H. B. Williams H. B. Williams C. E. Boardman Williams H. V. Maximilian Elliot Coues L. Abbott W. J. Hoffman W. E. S. Gaylord Eastgate Bowen Bowen Edwards Bowman R. Ross W. H. Currie A. J. Clark J. D. Allan Hibbard H. Life stage Migr Migr Migr Migr Migr Migr Migr Migr Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Day 2 8 1 15 22 6 2 21 6 17 28 16 20 1 3 19 14 2 7 Month Oct Apr Apr Oct Oct Oct Sep Jun Jul Sep Aug Jun Apr May Apr Apr Apr Apr May May Sep May May May Year 1921 1922 1923 1923 1923 1923 1924 1933 1833 1874 1879 1881 1883 1891 1893 1893 1894 1894 1903 1907 1908 1908 1908 1912 1912 1913 100.174 100.783 97.862 97.900 97.410 100.174 97.410 100.279 103.433 101.300 97.243 101.300 98.762 100.889 97.626 97.476 97.476 96.934 98.931 98.935 99.443 101.282 101.632 100.889 98.733 99.443 Longitude − − − − − − − − − − − − − − − − − − − − − − − − − − Latitude 46.709 46.808 48.497 48.500 48.412 46.709 48.412 46.485 48.303 47.642 48.966 47.642 46.209 46.827 47.907 48.877 48.877 47.052 48.028 48.853 47.009 48.971 48.960 46.827 48.233 47.009 Country US US US US US US US US US US US US US US US US US US US US US US US US US US State/ province ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND ND Forks County Kidder Burleigh Walsh Walsh Walsh Kidder Walsh Emmons Williams McLean Pembina McLean Dickey Morton Grand Pembina Pembina Cass Ramsey Cavalier Stutsman Bottineau Renville Morton Ramsey Stutsman Stevenson City Bismark Edinburg Grafton Grafton Hazelton Fort Pembina Fort Berthold Mandan Larimore Bathgate Bathgate Argusville Calvin Antler Sherwood Mandan Edmone Edinburg Location Long Lake Slough west of Long Lake Fort Union Devils Lake Chase Lake Chase Lake

(Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Bent (1926) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) (1956) Hibbard Allen (1952 68 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane and Russell Reid Skow Observer Russell Reid N. A. Wood C. E. Boardman Bernie Maurek Bernie Maurek G. Pearson T. Bernie Maurek J. S. Hunter hunter A G. Caldwell F. J. Brezee F. J. M. Bates L. Bruner A. Colby F. L. Bruner L. Bruner and L. L. Bruner Thomas Say Powell Life stage Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Migr Migr Day 16 24 1 15 8 13 13 29 30 19 24 Month Sep Jun Sep Sep Sep Sep Aug Aug Sep Sep May Mar Mar ) Year 1920 1921 1922 1922 1922 1923 1930 1930 1931 1932 1900 1918 1918 1936 1820 1884 cont. 100.783 99.299 100.174 100.174 100.174 100.174 100.174 100.711 99.916 99.916 96.686 95.931 100.498 100.468 98.342 100.551 98.767 96.683 96.708 96.364 95.931 95.931 96.026 98.468 Longitude − − − − − − − − − − − − − − − − − − − − − − − − Latitude 46.808 46.894 46.709 46.709 46.709 46.709 46.709 47.491 46.855 46.855 40.833 41.251 42.565 41.027 40.926 42.873 42.450 40.267 41.842 41.786 41.251 41.251 41.456 40.869 Country US US US US US US US US US US US US US US US US US US US US US US US US State/ province ND ND ND ND ND ND ND ND ND ND NE NE NE NE NE NE NE NE NE NE NE NE NE NE County Burleigh Stutsman Kidder Kidder Kidder Kidder Kidder McLean Kidder Kidder Lancaster Douglas Cherry Lincoln Hall Cherry Holt Gage Cuming Burt Douglas Douglas Washington Hall City Bismark Medina Mercer Steele Steele Lincoln Omaha Brady Island Grand Island Valentine Point West Craig Omaha Omaha Alda Location Long Lake Long Lake Long Lake Long Lake Long Lake Red Deer Lake

(Historical Records of Whooping Crane O bservations, rdered by S tate/Province and L ife-History tage ( Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 Appendix 69 (Continued) Observer Miller Miller Thatcher Wilson Powell Earl J. M. Bates J. M. Bates Dickinson J. Kennedy Bates Bunney Loren Bunney Loren Gunther Fred Merritt Cary Colt Merritt Cary Wolcott Colt Colt Colt Colt Colt Colt Colt Life stage Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Day 7 24 2 9 10 16 19 11 19 1 1 11 31 20 12 15 27 18 23 8 12 6 14 1 22 Month Nov Oct Apr Apr Apr Mar Mar Apr Sep Oct Apr Oct Mar Mar Oct Oct Oct Apr Mar Oct Apr Apr May Oct Oct Year 1884 1884 1887 1887 1889 1890 1890 1891 1891 1893 1894 1896 1897 1897 1898 1898 1899 1899 1899 1899 1899 1900 1900 1900 1901 1901 1901 1901 1901 99.372 98.600 98.783 98.783 98.844 95.602 98.844 98.749 95.602 100.551 100.551 97.402 100.163 99.699 99.369 99.369 98.342 98.029 96.595 98.029 96.686 96.595 96.595 96.595 99.081 96.595 96.595 96.595 96.595 Longitude − − − − − − − − − − − − − − − − − − − − − − − − − − − − − Latitude 40.719 40.819 42.900 42.900 40.749 40.061 40.749 42.205 40.061 42.873 42.873 41.028 40.926 42.534 40.436 40.436 40.926 42.129 40.088 42.129 40.833 40.088 40.088 40.088 40.695 40.088 40.088 40.088 40.088 Country US US US US US US US US US US US US US US US US US US US US US US US US US US US US US State/ province NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE County Buffalo Hall Boyd Boyd Buffalo Richardson Buffalo Boyd Richardson Cherry Cherry York Dawson Brown Harlan Harlan Hall Antelope Gage Antelope Lancaster Gage Gage Gage Buffalo Gage Gage Gage Gage City O’Neill O’Neill Gibbon Falls City Gibbon Chambers Falls City Valentine Valentine Gresham Gothenburg Long Pine Holdrege Holdrege Grand Island Neligh Badger Neligh Lincoln Badger Badger Badger Kearney Badger Badger Badger Badger Location Elm Creek River Wood

(Historical Records of Whooping Crane O bservations, rdered by S tate/Province and L ife-History tage ( Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 Appendix 71 (Continued) A. Black, Guy Smith Hicks William and Oscar Blevens Black, Garvin Observer Hunter Harry Connor C. A. Black C. A. Black C. A. Black C. A. Black Farm boy C. A. Black C. A. Black A. R. Golay C. A. Black A. Nooka C. A. Black C. A. Black C. A. Black C. L. Pitcarthley, R. Swanson and Tracy George Hicks William J. Q. Holmes A. Webert Roy Knapp Lilga, Golay, J. C. Chapman Life stage Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Day 15 29 2 15 7 2 14 20 14 1 20 22 13 2 16 5 12 19 25 25 18 26 4 5 8 Month Mar Mar Apr Oct Oct Apr Oct Oct Apr May Oct Oct Sep Apr Apr Oct Apr Oct Oct Oct Oct Apr Apr Apr Apr Apr Year 1918 1919 1920 1920 1920 1921 1921 1921 1922 1922 1922 1922 1923 1924 1924 1924 1925 1925 1925 1925 1925 1926 1926 1926 1926 1926 99.081 99.081 99.081 99.081 99.081 99.081 100.498 99.081 98.844 99.081 99.081 100.498 98.224 99.081 99.269 99.081 99.081 99.257 99.081 99.257 99.537 99.257 99.257 99.081 98.847 99.170 Longitude − − − − − − − − − − − − − − − − − − − − − − − − − − Latitude 40.695 40.695 40.695 40.695 40.695 40.695 42.565 40.695 40.749 40.695 40.695 42.565 40.594 40.695 40.769 40.695 40.695 40.702 40.695 40.702 40.739 40.702 40.702 40.695 40.648 40.365 Country US US US US US US US US US US US US US US US US US US US US US US US US US US State/ province NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE County Buffalo Buffalo Buffalo Buffalo Buffalo Buffalo Cherry Buffalo Buffalo Buffalo Buffalo Cherry Clay Buffalo Buffalo Buffalo Buffalo Buffalo Buffalo Buffalo Dawson Buffalo Buffalo Buffalo Kearney Kearney City Kearney Kearney Kearney Kearney Kearney Kearney Kearney Gibbon Kearney Kearney Inland Kearney Amherst Kearney Kearney Odessa Kearney Odessa Overton Odessa Odessa Kearney Lowell Wilcox Location Red Deer Lake Red Deer Lake

Mensinger H. Ligga, C. A. Black. Mrs. Frances Garvin Burmwood Observer C. A. Black A. R. Golay Farmer Keller A. R. Marsteller C. A. Black B. Armitage R. Kingsley F. L. and H. Brown Charles Radborn A. R. Golay Phillip B. Armitage C. A. Black A. R. Marsteller A. R. Golay, hunter A Mrs. Will Kennedy J. P. Anthony Roeser A. R. Marsteller Kennedy J. P. B. Armitage Life stage Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Day 8 20 1 15 9 6 9 9 24 22 10 18 27 12 19 2 6 25 5 4 Month Apr Oct Apr Oct Oct Apr Apr Apr Oct Sep Apr Apr Mar Mar Oct Apr Apr Oct Oct Apr Apr ) Year 1926 1926 1927 1927 1927 1928 1928 1928 1928 1928 1929 1929 1929 1929 1929 1930 1930 1930 1931 1931 1931 1932 1933 1933 cont. 99.126 99.257 99.081 98.963 102.582 99.170 98.844 98.844 98.963 98.963 98.847 99.081 101.700 99.257 99.081 99.170 99.081 101.491 98.600 100.163 99.372 99.170 100.163 99.537 Longitude − − − − − − − − − − − − − − − − − − − − − − − − Latitude 40.478 40.702 40.695 40.641 42.068 40.365 40.749 40.749 40.641 40.641 40.648 40.695 42.920 40.702 40.695 40.365 40.695 42.944 40.819 40.926 40.719 40.365 40.926 40.739 Country US US US US US US US US US US US US US US US US US US US US US US US US State/ province NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE NE County Kearney Buffalo Buffalo Kearney Sheridan Kearney Buffalo Buffalo Kearney Kearney Kearney Buffalo Cherry Buffalo Buffalo Kearney Buffalo Cherry Hall Dawson Buffalo Kearney Dawson Dawson City Axtel Odessa Kearney Newark Antioch Wilcox Gibbon Gibbon Newark Newark Lowell Kearney Merriman Odessa Kearney Wilcox Kearney Eli Gothenburg Wilcox Gothenburg Overton Location River Wood Elm Creek

(Historical Records of Whooping Crane O bservations, rdered by S tate/Province and L ife-History tage ( Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 Appendix 73 (Continued) Richardson Procter Foley Eaglestrom Wilson Turnbull Observer Herbert J. Flannery A. Black and C. Chris Zwink G. H. Phillips Conober and Adams C. Lanphear George J. Shields C. Swanson Baker Wells S. W. Alexander P. William C. Henry T. Dr. G. Vinzant W. J. Lockhart J. Lockhart B. R. Ross R. MacFarlane R. MacFarlane Life stage Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Summ Summ Wint Wint Breed Breed Summ Summ Summ Day 6 1 1 15 17 1 1 31 10 2 22 19 24 2 10 Month Apr Oct Apr Apr Apr Apr Apr Mar Apr Apr Oct Apr Jun May Jun Year 1933 1933 1934 1934 1934 1936 1936 1936 1937 1937 1939 1941 1889 1934 1810 1857 1853 1938 1864 1861 1862 1862 99.372 99.257 99.081 98.847 98.600 99.740 98.468 99.537 99.081 98.844 99.740 100.163 98.047 98.600 74.928 74.637 106.778 103.334 113.670 112.230 121.350 129.000 129.000 Longitude − − − − − − − − − − − − − − − − − − − − − − − Latitude 40.719 40.702 40.695 40.648 40.819 40.775 40.869 40.739 40.695 40.749 40.775 40.926 42.776 40.819 38.959 39.277 32.312 34.186 61.170 60.120 61.870 69.700 69.700 Country US US US US US US US US US US US US US US US US US US Can Can Can Can Can State/ province NE NE NE NE NE NE NE NE NE NE NE NE NE NE NJ NJ NM NM NT NT NT NT NT County Buffalo Buffalo Buffalo Kearney Hall Dawson Hall Dawson Buffalo Buffalo Dawson Dawson Knox Hall Cape May Cape May Dona Ana Roosevelt Resolution City Odessa Kearney Lowell Cozad Alda Overton Kearney Gibbon Cozad Gothenburg Portales Fort Fort Simpson Fort Anderson Fort Anderson River River Location Elm Creek River Wood Niobrara River River Wood Cape May Beesley’s Point Fort Thorn Salt River Anderson Anderson

(Historical Records of Whooping Crane O bservations, rdered by S tate/Province and L ife-History tage ( Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 Appendix 75 (Continued) Observer E. Lewis W. Latrobe A. Mayer W. hunter A Loane William Handy Potter Wesley Blair G. S. Agersborg G. S. Agersborg G. S. Agersborg Oliver C. K Worthen Cheney Blanchard Colt Colt Life stage Migr Migr Migr Migr Summ Summ Wint Migr Migr Migr Migr Migr Day 29 7 27 9 19 13 6 18 26 Month Oct Oct Apr Oct Sep May Oct Oct Apr Oct Oct Year 1909 1910 1910 1911 1911 1912 1912 1832 1899 1907 1928 1880 1895 1871 1885 1850 1885 1885 1885 1888 1905 1883 1887 1888 1889 1889 99.683 100.483 99.683 100.483 99.683 100.483 99.683 97.300 98.390 98.800 97.058 79.380 79.320 82.080 79.420 79.257 96.958 96.694 97.383 99.736 96.723 100.428 99.200 98.214 99.297 98.527 98.527 Longitude − − − − − − − − − − − − − − − − − − − − − − − − − − − Latitude 36.767 36.767 36.767 36.767 36.767 36.767 36.767 35.200 34.609 36.767 36.116 43.650 45.530 42.580 44.680 33.358 42.917 42.817 43.050 45.355 43.556 45.537 45.400 44.363 43.644 43.430 43.430 Country US US US US US US US US US US US Can Can Can Can US US US US US US US US US US US US State/ province OK OK OK OK OK OK OK OK OK OK OK ON ON ON ON SC SD SD SD SD SD SD SD SD SD SD SD County Harper Beaver Harper Beaver Harper Beaver Harper Cleveland Comanche Woods Payne Georgetown Clay Union Yankton Walworth Minnehaha Walworth Edmunds Bradle Brule Douglas Douglas City Stillwater Toronto Emsdale Camden Ardtrea Sioux Falls Mobridge Huron Grandview Harrison Harrison River Bowdle Location Waccamaw 12 miles SW of

(Historical Records of Whooping Crane O bservations, rdered by S tate/Province and L ife-History tage ( Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 Appendix 77 (Continued) and John Macoun Richardson Demaine and Swann Fred Observer Hind H. Y. Hind H. Y. Hind Christy Thomas McKay C. G. Harold farmer A Indian Indian Framk Fleming Macoun Macdonald Macoun Hugh Cowboy Brown Fernie William Arnold Edward Ernest J. R. Lloyd Life stage Breed Breed Breed Breed Breed Breed Breed Breed Breed Breed Breed Day 12 6 10 16 21 Month Aug Jul Jun Jun May May Year 1858 1858 1858 1884 1892 1908 1913 1919 1920 1925 1872 1880 1884 1895 1896 1900 1900 1901 1902 1907 1911 104.380 103.670 104.270 101.930 105.770 103.670 103.430 103.500 103.550 103.500 104.370 102.580 108.320 106.230 105.770 102.470 102.550 103.430 101.900 107.250 105.980 Longitude − − − − − − − − − − − − − − − − − − − − − Latitude 50.970 50.330 51.570 51.570 53.200 50.330 51.300 51.170 50.800 51.170 51.920 49.780 52.730 49.480 53.200 51.220 51.400 51.300 49.600 50.900 51.270 Country Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can State/ province SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK County City Prince Albert Ituna Ituna Balcarres Ituna Battleford Prince Albert Yorkton Ituna Wauchope Demaine Davidson - Hills River Mountain katchewan River Lake Location Loon Creek Indian Head Touchwood White Sand Indian Head Beaver Hills Big Quill Lake Moose North Sas Mile Twelve Cussed Creek Beaver Hills

(Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Hjertaas (1994) Allen (1952) Allen (1952) Allen (1952) Hjertaas (1994) Allen (1952) Hjertaas (1994) Hjertaas (1994) Hjertaas (1994) Hjertaas (1994) Allen (1952 78 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane Wytoski Wytoski Wytoski Wytoski Observer L. G. Moore Smith Archie H. M. Dahl L. G. Moore Smith W. W. Neil Gilmour Bradshaw Fred Smith Archibald Frank Miller Steve West Steve West Lang Thomas L. James Kingsley N. T. farmer A Andrew Andrew Andrew Andrew Life stage Breed Breed Breed Breed Breed Breed Breed Breed Breed Breed Breed Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Day 19 28 2 8 3 29 Month Jun May May Aug Oct Oct Oct Oct Nov Oct ) Year 1911 1911 1911 1912 1921 1922 1922 1928 1929 1929 1929 1904 1913 1914 1919 1927 1927 1927 1927 1928 1928 cont. 106.230 108.300 104.500 106.230 108.300 109.280 109.100 108.300 107.080 107.080 107.080 103.670 104.730 102.870 104.980 105.430 102.980 104.020 104.020 104.020 104.020 Longitude − − − − − − − − − − − − − − − − − − − − − Latitude 51.950 54.620 50.930 51.950 54.620 52.020 52.320 54.620 51.080 51.080 51.080 50.330 51.420 49.650 52.400 51.130 49.130 53.530 53.530 53.530 53.530 Country Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can State/ province SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK County City Bradwell Southey Bradwell Baliol Semans Forget Lake Lenore Liberty Estevan River River River River Location Shallow Lake Shallow Lake Kiyiu Lake Shallow Lake Luck Lake Luck Lake Luck Lake Indian Head White Fox White Fox White Fox White Fox

(Historical Records of Whooping Crane O bservations, rdered by S tate/Province and L ife-History tage ( Source Allen (1952) Hjertaas (1994) Hjertaas (1994) Allen (1952) Hjertaas (1994) Allen (1952) Allen (1952) Hjertaas (1994) Hjertaas (1994) Hjertaas (1994) Hjertaas (1994) Allen (1952) Allen (1952) Allen (1952) Hjertaas (1994) Allen (1952) Allen (1952) Hjertaas (1994) Hjertaas (1994) Hjertaas (1994) Hjertaas (1994 Appendix 79 (Continued) Wytoski Wytoski Estlin Observer Andrew Andrew R. H. Carruthers Guernsey MacDonald Colt Colt Cates Lang Lang Lang Lang Lang H. Hanson Ferry Lang H. C. Grose E. Margaret Life stage Migr Migr Migr Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Day 7 28 13 1 25 20 19 24 25 26 1 27 Month May Apr Jun May May Apr Apr Apr Sep Apr May Sep May Apr Apr Year 1929 1929 1827 1884 1891 1893 1893 1901 1904 1904 1904 1905 1905 1905 1908 1909 1909 1909 1910 1910 1913 104.020 104.020 108.570 105.680 103.670 107.400 106.550 106.550 103.670 103.670 103.670 103.670 103.670 103.670 104.170 104.150 104.530 104.370 105.230 103.670 103.620 109.130 Longitude − − − − − − − − − − − − − − − − − − − − − − Latitude 53.530 53.530 51.080 53.250 50.330 50.400 52.370 52.370 50.330 50.330 50.330 50.330 50.330 50.330 49.800 50.230 52.730 51.920 51.080 50.330 51.720 51.920 Country Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can State/ province SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK County City Isham Osler Osler Grass Yellow Lajord Resource Kerrobert River River River Lake and vicinity Location White Fox White Fox Saskatchewan Indian Head Rush Lake Indian Head Indian Head Indian Head Indian Head Indian Head Indian Head Big Quill Lake Last Mountain Indian Head Foam Lake

(Source Hjertaas (1994) Hjertaas (1994) Allen (1952) Allen (1952) Allen (1952) Hjertaas (1994) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Hjertaas (1994) Allen (1952 80 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane Estlin Joe Perry Dickson Observer H. Sharpe Lang J. R. Garden E. Margaret Finley and J. V. Hoyes Lloyd H. L. Felt Neil Gilmour Miss B. M. Neil Gilmour G. H. Cartwright Bradshaw Fred Bradshaw Fred John N. Hachett Emil Lestin Bauman George H. E. Dresser H. E. Dresser H. E. Dresser Life stage Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Summ Breed Summ Day 15 30 3 18 10 10 13 15 27 12 Month Jun Apr Sep May May Apr Jun Aug Jun May May Aug Aug Jun Jun ) Year 1916 1920 1921 1921 1921 1922 1923 1924 1925 1926 1926 1927 1930 1930 1930 1932 1935 1936 1863 1863 1860 1860 1863 cont. 107.020 103.670 103.670 109.130 109.400 108.300 103.670 105.400 107.080 107.650 107.080 107.000 106.100 107.080 107.080 105.900 107.080 107.750 97.260 97.300 95.378 94.909 97.517 Longitude − − − − − − − − − − − − − − − − − − − − − − − Latitude 53.830 50.330 50.330 51.920 52.080 54.620 50.330 50.780 51.080 51.320 51.080 53.830 52.320 51.080 51.080 53.600 51.080 55.430 26.000 25.520 28.876 29.222 26.117 Country Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Can Mex Mex US US US State/ province SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK SK TM TM TX TX TX County Brazoria Galveston Cameron City Big River Kerrobert Luseland Findlater Wiseton Vonda Bagdad Matamoros Port Isabel Grande Brazos River Island Location Indian Head Indian Head Shallow Lake Indian Head Luck Lake Luck Lake Ladder Lake Luck Lake Luck Lake Emma Lake Luck Lake Beaver River Boca del Rio Mouth of Galveston

(Historical Records of Whooping Crane O bservations, rdered by S tate/Province and L ife-History tage ( Source Hjertaas (1994) Allen (1952) Allen (1952) Allen (1952) Hjertaas (1994) Hjertaas (1994) Allen (1952) Allen (1952) Hjertaas (1994) Allen (1952) Hjertaas (1994) Hjertaas (1994) Hjertaas (1994) Hjertaas (1994) Hjertaas (1994) Hjertaas (1994) Hjertaas (1994) Hjertaas (1994) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 Appendix 81 (Continued) Strandtmann Observer H. E. Dresser J. A. Brundett Elanoides Elanoides Elanoides R. L. More Thomas Webb Jno. B. Litsey Reid Mrs. Bruce R. W. J. O. Stevenson J. O. Stevenson J. O. Stevenson J. O. Stevenson J. D. Mitchell J. D. Mitchell J. D. Mitchell Oliver Davie Life stage Breed Breed Breed Breed Day Month Year 1863 1885 1886 1888 1889 1908 1910 1920 1927 1928 1938 1939 1940 1941 1867 1869 1878 98.493 96.946 97.146 97.146 97.146 98.195 96.946 97.414 97.414 97.793 96.640 96.640 96.640 96.640 99.190 96.275 96.275 96.275 96.342 Longitude − − − − − − − − − − − − − − − − − − − Latitude 29.424 28.124 31.549 31.549 31.549 33.817 28.124 32.791 32.791 29.881 28.227 28.227 28.227 28.227 34.215 29.027 29.027 29.027 29.564 Country US US US US US US US US US US US US US US US US US US US State/ province TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX Wharton Wharton Wharton County Bexar Aransas McLennan McLennan McLennan Clay Aransas Tarrant Tarrant Caldwell Calhoun Calhoun Calhoun Calhoun Wilbarger Southern Southern Southern Colorado Worth Worth City San Antonio Waco Waco Waco Henrietta Dallas/Lake Dallas/Lake Maxwell Vernon Peninsula Ranch, 8 miles east of Henrietta Peninsula Island Island Island Island Location Blackjack Worsham Blackjack Matagorda Matagorda Matagorda Matagorda Red River Eagle Lake (1974) (1938) (1938)

(Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Oberholser Oberholser Oberholser Oberholser (1938 82 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane McCall Observer Colonel G. A. Lloyd W. G. B. Benners G. B. Benners G. B. Benners G. B. Benners G. B. Benners Lloyd W. Peters Peters Peters Peters Ragsdale Lloyd W. Peters Peters Peters Peters Peters Lloyd W. Peters Peters Peters Peters Peters B. Armstrong F. Life stage Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Migr Day 4 23 27 30 31 5 16 9 2 9 25 5 22 15 8 18 8 2 Month Nov Apr Apr Mar Mar May Apr Mar Mar Mar Mar Mar Nov Nov Apr Apr Mar Mar Nov Oct Oct Nov Nov Apr ) Year 1845 1884 1884 1884 1884 1884 1884 1885 1885 1885 1885 1885 1885 1885 1885 1885 1886 1886 1886 1886 1888 1888 1888 1889 1889 1890 cont. 98.493 100.437 97.600 98.300 97.600 98.300 97.600 100.437 96.178 96.178 96.178 96.178 97.133 100.437 96.178 96.178 96.178 96.178 96.178 100.437 96.178 96.178 96.178 96.178 96.178 97.497 Longitude − − − − − − − − − − − − − − − − − − − − − − − − − − Latitude 29.424 31.464 30.650 29.817 30.650 29.817 30.650 31.464 33.577 33.577 33.577 33.577 33.626 31.464 33.577 33.577 33.577 33.577 33.577 31.464 33.577 33.577 33.577 33.577 33.577 25.901 Country US US US US US US US US US US US US US US US US US US US US US US US US US US State/ province TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX County Bexar Green Tom Williamson Comal Williamson Comal Williamson Green Tom Fannin Fannin Fannin Fannin Cooke Green Tom Fannin Fannin Fannin Fannin Fannin Green Tom Fannin Fannin Fannin Fannin Fannin Cameron City San Antonio San Angelo San Angelo Bonham Bonham Bonham Bonham Gainesville San Angelo Bonham Bonham Bonham Bonham Bonham San Angelo Bonham Bonham Bonham Bonham Bonham Brownsville Location

(Historical Records of Whooping Crane O bservations, rdered by S tate/Province and L ife-History tage ( Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 Appendix 83 (Continued) McCall Sennett Sennett Observer Peters Peters Reid Mrs. Bruce H. Bauer W. G. B. Benners Lloyd W. Lloyd W. Lloyd W. Bailey Vernon Gaut Colonel G. A. B. George B. George H. Nehrling H. Nehrling Lloyd W. H. Nehrling Lloyd W. Worthen Life stage Migr Migr Migr Migr Migr Summ Summ Summ Summ Summ Summ Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Day 2 23 2 20 30 7 6 23 1 20 23 23 Month Apr Mar Apr Nov Nov May Aug Aug Oct May Apr Apr Mar Mar Dec Nov Feb Jan Mar Jan Dec Year 1890 1890 1899 1913 1932 1884 1891 1891 1891 1900 1905 1845 1877 1877 1878 1881 1881 1882 1882 1882 1885 1886 1889 96.178 96.178 97.146 93.929 96.809 98.300 97.368 97.368 97.396 97.368 94.965 97.396 97.497 97.497 97.368 95.363 95.363 95.363 95.363 95.363 96.018 94.102 97.497 Longitude − − − − − − − − − − − − − − − − − − − − − − − Latitude 33.577 33.577 31.549 29.899 28.468 29.817 26.844 26.844 27.800 26.418 30.324 27.800 25.901 25.901 26.844 29.763 29.763 29.763 29.763 29.763 28.585 30.086 25.901 Country US US US US US US US US US US US US US US US US US US US US US US US State/ province TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX County Fannin Fannin McLennan Jefferson Calhoun Comal Kennedy Kennedy Nueces Cameron Liberty Nueces Cameron Cameron Kennedy Harris Harris Harris Harris Harris Matagorda Jefferson Cameron City Bonham Bonham Waco Port Arthur Corpus Christi Cleveland Corpus Christi Brownsville Brownsville Houston Houston Houston Houston Houston Beaumont Brownsville Island Island Prairie Island Peninsula, Kanes Landing Location Mission Lake Head of Padre Head of Padre Tarkington Head of Padre Matagorda

(Historical Records of Whooping Crane O bservations, rdered by S tate/Province and L ife-History tage ( Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 Appendix 85 (Continued) and Richard and Richard Kleberg and Maunsell Crosby and Maunsell Crosby Observer E. Hough Worthen H. C. Oberholser H. C. Oberholser B. Armstrong F. B. Armstrong F. L. R. Cowen L. R. Cowen L. R. Cowen Kleberg Richard G. Pearson T. Ludlow Griscom Ludlow Griscom R. D. Camp Baker J. G. Fuller H. Clegg T. H. C. Oberholser Life stage Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Wint Day 29 20 2 29 10 3 16 15 22 27 23 12 4 12 8 10 Month Nov Feb Apr Mar Feb Feb Jan Feb Feb Feb Dec Jan Jan Mar Feb Feb Jan Year 1895 1896 1897 1900 1900 1904 1904 1904 1911 1911 1911 1915 1921 1923 1923 1924 1926 1931 1933 1933 96.946 97.368 97.368 96.640 96.640 97.368 97.396 97.368 97.497 97.497 97.497 97.398 97.398 97.498 97.398 97.497 97.398 96.946 97.581 96.526 Longitude − − − − − − − − − − − − − − − − − − − − Latitude 28.124 26.844 26.844 28.227 28.227 26.844 27.800 26.844 25.901 25.901 25.901 27.521 27.521 27.233 27.521 25.901 27.521 28.124 26.938 28.347 Country US US US US US US US US US US US US US US US US US US US US State/ province TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX TX County Aransas Kennedy Kennedy Calhoun Calhoun Kennedy Nueces Kennedy Cameron Cameron Cameron Kleberg Kleberg Kennedy Kleberg Cameron Kleberg Aransas Kennedy Calhoun City Island Padre Island Padre Island Padre Corpus Christi Island Padre Brownsville Brownsville Brownsville Laguna Larga Laguna Larga Laguna Larga Brownsville Laguna Larga Peninsula Island Island Bay Peninsula north of Brownsville Bay Location Blackjack Matagorda Matagorda South of Baffin Blackjack 63 miles Espiritu Santo

(Historical Records of Whooping Crane O bservations, rdered by S tate/Province and L ife-History tage ( Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952 Appendix 87 (Continued) Wade Observer Lewis and Clark R. Hoy P. R. Hoy P. Douglas E. Snyder W. Richard William Richard William Richard William Richard William Richard William Richard William Richard William Skinner M. P. Richard William Life stage Summ Migr Breed Migr Migr Summ Breed Migr Day 21 19 4 Month Jun Oct Oct Apr Aug Year 1876 1805 1840 1840 1854 1878 1935 1900 1906 1907 1908 1909 1910 1911 1912 1915 1914 1906 79.143 118.910 91.139 87.783 89.660 87.967 89.600 88.912 88.656 109.056 109.056 109.056 109.056 109.056 109.056 109.056 108.204 110.233 107.339 Longitude − − − − − − − − − − − − − − − − − − − Latitude 37.414 46.058 43.046 42.726 43.033 44.467 42.833 43.081 43.550 44.526 44.526 44.526 44.526 44.526 44.526 44.526 44.862 44.767 42.437 Country US US US US US US US US US US US US US US US US US US US State/ province VA WA WI WI WI WI WI WI WI WY WY WY WY WY WY WY WY WY WY Walla County Lynchburg Walla Crawford Racine Dane Brown Green Jefferson Dodge Park Park Park Park Park Park Park Bighorn Park Carbon Chien City Lynchburg Prairie du Racine Cody Cody Cody Cody Cody Cody Cody Rapids— Mouth of the Walla Walla River Wisconsin Park Location Umatilla Southwest Lake Michigan Sugar River Lake Mills Horicon Marsh Shoshone River Yellowstone Jackson Lake * (1953) Hollister (1951) Hollister (1951) Hollister (1951) Hollister (1951) Gluesing (1976)

(Source Bailey (1881) Jewett et al. Kumlien and Kumlien and Allen (1952) Carr (1890) Kumlien and Kumlien and Hunt and Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Bent (1926) Allen (1952 88 3. Revisiting the Historic Distribution and Habitats of the Whooping Crane and Ranger Bicknell Observer Richard William G. Pearson T. Bock J. F. Ray Wolfley Theone Wolfley Life stage Migr Summ Summ Summ Summ Day 23 Month Feb Apr May May ) Year 1912 1930 1934 1934 1935 cont. 107.339 110.996 104.421 111.027 111.027 Longitude − − − − − Latitude 42.437 44.148 43.987 43.123 43.123 Country US US US US US State/ province WY WY WY WY WY ) killed in St. Johns County, Florida. Florida Natur. 38, 23. Florida. Florida Natur. ) killed in St. Johns County, wintering. County Carbon Teton Weston Lincoln Lincoln Wint, . Redstart 7, 65. Grus americana , migrating; City Osage Migr Grus americana were questionable and not included in maps or in biome and ecoregion summaries. questionable and not included in maps or biome ecoregion were asterisk , summering; Location Jackson Lake Bechler River Star Valley Star Valley Summ , breeding; , breeding;

,Historical Records of Whooping Crane O bservations, rdered by S tate/Province and L ife-History tage ( Source Allen (1952) Allen (1952) Allen (1952) Allen (1952) Allen (1952) Breed marked with an Records Allen 1952, pp. 51–64: in addition to those listed Added sources Audubon Q, 33, 43–44. 1955. Whooping Cranes in Porter County 50 years ago. Indiana Baczkowski, F., I–XII, 70–71. and Stream Forest notes: an index and summary. H.B., 1881. Bird Bailey, CO. Denver, of Colorado. Denver Museum Natural History, A.M., Niedrach, R.J., 1965. Birds Bailey, Boston, MA. America, 2 vols. Little, Brown, of North R., 1884. The water birds Ridgway, T.M., Brewer, S.F., Baird, Agricultural College, Lansing, 822 pp. Life. Michigan 1912. Michigan Bird W.B., Barrows, DC. Smithsonian Institution, United States National Museum Bulletin 135, Washington, American Marsh Birds. A.C., 1926. Life Histories of North Bent, AZ. Publishing Co., Prescott, Shooting. Wolfe and Trap geese, cranes, and swans (Chapter 12). In: Field, Cover, A.H., 1878. Wild Bogardus, Annual Report 22, pp. 575–1187. of Indiana. Indiana Department Geology and Natural Resources 1898. The Birds A.W., Butler, 1, 78. Natur. 1890. Wisconsin C.F., Carr, pp. 353–370. Illinois. In: History of Lake County, of Lake County, Coale, H.K., 1912. The Birds Iowa City. in Iowa. University of Iowa Press, Country So Full of Game: The Story Wildlife A J.J., 1994. Dinsmore, Macon. Oriole 22. A Whooping Crane from 1957. W.W., Griffin, of Whooping Crane ( Hallman, R.C., 1965. Record 27, 22. 1954. Notes on cranes. Florida Natur. Harmon, W.Z., Bull. 68, 73–74. for the Whooping Crane in North Dakota. Wilson An old nesting record E.A., 1956. Hibbard, of the Whooping Crane in Saskatchewan. Blue Jay 52 (2), 99–115. records Hjertaas, D.G., 1994. Summer and breeding Montreal. 34. McGill-Queen’s University Press, McGill-Queen’s Native and Northern Series, Vol. Houston, M., 2003. Eighteenth-century naturalists of Hudson Bay. Houston, C. S., Ball, T., International Crane Foundation, Baraboo, WI, pp. 19–34. of the International Crane Workshop, In: Proceedings Hunt, R.A., Gluesing, E.A., 1976. The sandhill crane in Wisconsin. Seattle. Press, State. University of Washington of Washington 1953. Birds Aldrich, J.W., W.T., Shaw, W.P., Jewett, S.G., Taylor, Bull. 42 (3), 198–210. additions. Wilson National Park, with some recent Yellowstone of the Kemsies, E., 1930. Birds Madison. Society for Ornithology, Wisconsin of Wisconsin. N., 1951. The Birds Kumlien, L., Hollister, AOU Monograph 3. of Kentucky. Mengel, R.M., 1965. The Birds Life of Louisiana. Bulletin 28, Department Conservation, State Louisiana, Baton Rouge. H.C., 1938. The Bird Oberholser, Austin. Press, University of Texas Life of Texas. H.C., 1974. The Bird Oberholser, C., 1940. Whooping Crane, Shaffer, 527 pp. York, New Audubon Society, and National Life. Coward-McCann A., 1954. Florida Bird Jr., Sprunt 264 pp. Cornell University Library, Area. of the Chicago 1907. The Birds F.M., Woodruff