4215(115-131).pdf ALCES VOL. 42, 2006 HOFFMAN ET AL. - LONG-DISTANCE MOOSE DISPERSAL 115 LONG-DISTANCE DISPERSAL AND POPULATION TRENDS OF MOOSE IN THE CENTRAL UNITED STATES Justin D. Hoffman 1, Hugh H. Genoways1, and Jerry R. Choate2 1School of Natural Resources and University of Nebraska State Museum, W436 Nebraska Hall, University of Nebraska-Lincoln, Lincoln, NE 68588, USA; 2Sternberg Museum of Natural History, Fort Hays State University, Hays, KS 67601, USA ABSTRACT: Dispersal is a basic feature of the natural history of moose. Most information about moose dispersal pertains to short-distance movements because long-distance movements are uncom- and North Dakota. This may have contributed to several long-distance dispersal events for moose that recently were reported in the central United States. These dispersal events provide an opportunity to investigate both the causes and the biological implications of this rare phenomenon. Herein, we review long-distance dispersal events based on information obtained from a variety of sources. Dispersal routes that could be measured included two with minimal distances of 1,511 and 367 km, plus several others that were shorter. These dispersal events and recent evidence of moose reproducing outside the current range of the species could be the result of increasing population trends of moose in the central United States. We suggest that the dispersing moose are founder individuals that are dispersing naturally from established populations in search of suitable habitats and mates in areas to the south. We hypothesize that this type of geographic range expansion is similar to that of moose when they dispersed across North America during the early Holocene. As moose continue to move south, wildlife managers should be aware of habitats within their respective states that might sustain populations of moose. ALCES VOL. 42: 115-131 (2006) Key words: Alces alces, central United States, dispersal, moose, range expansion Dispersal is a basic feature of the life his- tory of most species. Local dispersal occurs within established populations of a species. It is important because it regulates population is movement into suitable habitats adjacent to the currently occupied range of the species. This type of dispersal can lead to a gradual expansion of the range of a species. Long- distance dispersal differs from diffusion dis- persal in that it may occur across large areas of unsuitable habitat. This type of dispersal can result in the discovery and colonization of isolated unoccupied habitats, and potentially the rapid expansion of a species’ geographic range (Ricklefs and Miller 2000). Long-dis- The moose (Alces alces) is the largest member of the family Cervidae and occurs pri- marily in the boreal forests of North America overall geographic distribution of moose ex- panded in the late Pleistocene, with individuals dispersing from eastern Asia into North Ameri- can via the Bering land bridge approximately 14,000-11,000 years ago (Hundertmark et al. 2002). At that time, glaciers in North America had begun to retreat, producing an abundance of the early successional habitats favored by moose and facilitating rapid expansion of the geographic range of the moose across North LONG-DISTANCE MOOSE DISPERSAL – HOFFMAN ET AL. ALCES VOL. 42, 2006 116 are highly mobile with a strong propensity for - ity, they can disperse hundreds of kilometers in a short period of time. Dispersers are often, but not exclusively, young individuals (Hun- and juveniles tend to disperse only a short distance from their natal range (Gasaway et al. tend to disperse farther from their natal range than juvenile females, and the percentage of overlap of home ranges between juvenile and dam is less for juvenile males than for juvenile Although moose usually disperse short distances, long-distance dispersal events have been documented. For example, Mytton and for 4 young moose and a 250-km dispersal distance for a young bull moose in Alberta, adult cow that dispersed a distance of 177 km in southern Alaska. These accounts rep- resent dispersal events within the geographic range of moose. To our knowledge, there have been few published accounts of long- distance dispersal of moose outside their that a dead moose was found approximately 500 km north of traditional moose range in the Northwest Territories, Canada. In the central tracked the dispersal route of a bull moose from Minnesota through Iowa and into Mis- souri. They estimated that the total distance recently, there have been several instances of moose dispersing outside their known range suggested that moose are still expanding their geographic range into areas that they have not occupied since the last glaciation. These long-distance dispersal records are impor- tant because they document a phenomenon that rarely is observed and potentially can provide insight into moose movements and biogeography. As discussed above, technical reports on long-distance dispersal events by moose outside their normal geographic range are few. Most such information is found in popular media and newspaper articles, which, by them- selves, provide little biological information. Herein, we present a summary of recent trends in moose populations in the central United States and of long-distance dispersal events by moose in this region. We discuss possible explanations for long distance dispersal by moose and potential biological implications METHODS Information from primary literature, government documents, and Minnesota and North Dakota moose harvest reports were used to summarize recent population trends of moose in the central United States. To describe long-distance moose dispersal, we collected information regarding moose sight- ings and potential dispersal routes for North Dakota, South Dakota, Minnesota, Iowa, Nebraska, Missouri, Kansas, Oklahoma, and Texas. The sources consulted included newspapers, popular journals, books, primary literature, and communication with employees of the Kansas Department of Wildlife and Parks (KDWP), Nebraska Game and Parks Commission (NGPC), North Dakota Game and Fish Department (NDGFD), and South Dakota Game, Fish, and Parks (SDGFP). We such as newspapers and popular journals, to document the movements of moose because moose are not likely to be confused with any other species. In this paper, we consider the “histori- cal” geographic range of moose in the central United States to be northeastern Minnesota ALCES VOL. 42, 2006 HOFFMAN ET AL. - LONG-DISTANCE MOOSE DISPERSAL 117 range consists of the “historical” range plus areas outside this region occupied in subse- quent years as part of the recent expansion of moose in the central United States. We calculated dispersal distances from localities at the southern edge of the current geographic because it is likely that most long-distance dispersers came from populations in those regions. We obtained potential dispersal routes by connecting chronologically ordered localities and calculated dispersal distances considered to be minimal distances travelled because they were measured as straight lines between localities. RESULTS Population Status When the north-central United States was settled by European immigrants and their descendents, moose occurred in northern Minnesota and northeastern North Dakota. been extirpated from North Dakota and Moose continued to inhabit the boreal forests of northeastern Minnesota although they un- doubtedly were less abundant than they had been in pre-settlement periods. Because of this decline in moose numbers, Minnesota closed numbers began to rise and moose began to reclaim their former range in northwestern - gan moving back into North Dakota and, by From there, they spread westward along the Canadian border to the Turtle Mountains and southward along the Red River Valley, where they inhabit the rugged lands of prehistoric resident moose populations again occurred Because of the recent increase of moose, both North Dakota and Minnesota have estab- lished hunting seasons. Minnesota reopened Two units were available to hunters, in the Northwest and the Northeast, with hunting harvested in Minnesota continued to increase with considerably more moose being taken in the northwestern unit (Minnesota Department northwestern unit and 737 moose were har- vested, as compared with 523 permits sold in the northeastern unit and 442 moose harvested (Minnesota Department of Natural Resources was noticed in the northwestern unit. As a result, restrictions on moose hunting were implemented throughout the northwestern Northwest was closed. Since then, population numbers have remained low in the Northwest stable in the Northeast (Minnesota Department of Natural Resources 2005). North Dakota implemented a moose hunt- harvest rates and hunting units have increased. Today, hunting is permitted in the Pembina Hills, the Turtle Mountains, and the Red River Valley, which encompass the north-central, northeastern, and extreme eastern parts of the state extending as far south as the South areas where moose occur in North Dakota, population estimates continue to be highest in the Pembina Hills area, followed by the Turtle Mountains, and then the Red River Valley LONG-DISTANCE MOOSE DISPERSAL – HOFFMAN ET AL. ALCES VOL. 42, 2006 of moose in North Dakota are more-or-less Dispersal Moose recently have been reported outside the current geographic range of the species in the central United States (Fig. 1). Some of those individuals dispersed over long distanc- es. The most notable dispersal was undertaken in South Dakota and eventually dispersed as was of 2 young bulls seen near Dell Rapids, separated, because there were no reports of a second moose beyond Dell Rapids. In Sep- across southeastern South Dakota, where it was reported near the towns of Alexandria, crossed the Missouri River near Avon, South a few days later in Page, Nebraska (Omaha southward across Nebraska, passing near the towns of Elba, Palmer, and Chapman (Horn- of Grand Island and the Platte River for the remainder of the year and was spotted near Saronville, approximately 45 km southeast of reported about 101 km to the southwest near Agra, Kansas, in late February (Kleinschmidt account of the moose’s movement in Kansas. The moose remained in north-central Kansas, reportedly being seen near Kirwin Reservoir on the North Fork of the Solomon River (which is just south of Agra), for the remainder of it was seen west of Stockton, approximately 45 km southwest of Agra. The moose re- from the summer heat at Webster Reservoir on the South Fork of the Solomon River (which is just west of Stockton), until September, when it was seen approximately 104 km farther south near Rush Center. From Rush Center, the moose quickly moved through south-central State Reference Locality Day Month South Dakota Dell Rapids (1) 15 Sept Alexandria (2) 27 Sept Dimock (3) Sept Parkston (4) Sept Avon (5) 1 Oct Nebraska Page (6) 5 Oct Elba (7) 13 Oct 16 Oct Oct Grand Island (10) Oct Phillips (11) 30 Oct Trumball (12) 12 Nov Harvard (13) 26 Dec Saronville (14) Nelson (15) 15 Guide Rock (16) 21 Kansas Agra (17) 22 Feb Sept Kinsley (20) 20 Sept Ashland (21) 22 Sept Englewood (22) Sept Texas Perrytown (23) Nov Dalhart (24) Nov Kansas Ulysses (25) Dec Sublette (26) 5 Feb Table 1. Time and locality information for the dispersal of a moose through South Dakota, Nebraska, Kansas, Oklahoma, and Texas. Numbers in paraenthesis correspond to those shown in Figure 1. ALCES VOL. 42, 2006 HOFFMAN ET AL. - LONG-DISTANCE MOOSE DISPERSAL Fig. 1. Records of occurrences and potential dispersal routes of moose in the central United States. Symbols connected by solid lines represent potential dispersal routes described in this study. The symbols connected by a dashed line represent a dispersal route described by Bowles and Gladfelter and diamond represent instances in which a moose remained in the area for an extended period of time. Numbers correspond to locality information listed in Tables 1 and 2. LONG-DISTANCE MOOSE DISPERSAL – HOFFMAN ET AL. ALCES VOL. 42, 2006 120 Kansas, passing near Kinsley, Ashland, and any reports of the moose having been seen in Oklahoma, it presumably passed through the Oklahoma Panhandle because it was next seen near Perrytown, Texas, in November and later location, the moose reversed directions and returned to southwestern Kansas, where it area, a local veterinarian obtained permission from KDWP to tranquilize the moose. She claimed that the moose was in poor health and in need of medical attention. On 5 Feb- near Sublette, Kansas, and transported to the veterinarian’s facility (Associated Press whether or not the moose was, in fact, sick. After treating the animal, it apparently was released somewhere in Colorado (the exact - served near Dell Rapids, South Dakota, the moose travelled an estimated straight-line distance of 1,511 km. The time it took to travel area from which this moose dispersed, and thus the total distance it moved, is impossible to know. However, the distance this moose travelled as measured from the southern edge of the current geographic range of the species In Iowa and Missouri, similar dispersal events by moose have been documented. distance dispersal by a bull moose that began in southwestern Minnesota, continued through Iowa, and ended in the vicinity of Bowling Green, Missouri, near the Mississippi River (Fig. 1). A similar dispersal event took place in the same area a few years later (Fig. 1). In observed in south-central Minnesota near St. reported in Iowa near the town of Fertile in of sightings, refer to Table 2). From Fertile, the moose travelled directly south across Iowa, passing near Clear Lake, Thornton, Latimer, observed in Iowa, the moose was reported From there, the moose veered southeastward and passed near Pleasantville, northeast of in early December was reported for the last time in Iowa in the vicinity of Georgetown and Table 2. Time and locality information for the dis- persal of a moose through Minnesota, Iowa, and Missouri. Numbers in paraentheses correspond to those shown in Figure 1. State Reference Locality Day Month Minnesota Oct Oct Iowa 2 Nov Clear Lake (30) 2 Nov Thornton (31) Nov Latimer (32) Nov Alden (33) 7 Nov Ames (34) Nov Nevada (35) 11 Nov NE of Des Moines (36) Nov Altoona (37) Nov 30 Nov 30 Nov 4 Dec Missouri Omaha (41) 31 Dec E of Pollock (42) 20 25 River (44) 4 Feb Dalton (45) Feb ALCES VOL. 42, 2006 HOFFMAN ET AL. - LONG-DISTANCE MOOSE DISPERSAL 121 near Fertile, it took a little over a month for the bull moose to travel 335 km across Iowa. The moose then moved into Missouri, where it was spotted near Omaha in late December again until approximately a month later, when it was seen east of Pollock, which is southwest moose was observed near the junction of High- there, it moved southwestward into Chariton County, where it reportedly was seen near the The last report of this moose in Missouri was - ing observed in south-central Minnesota, the moose travelled an approximate distance of 650 km. Because the exact dates when this average dispersal speed for the entire trip was not calculated. However, we were able to calculate an average dispersal speed from in Missouri, near Omaha. The approximate distance travelled by the moose was 365 km in 60 days, giving it an average dispersing from which this moose dispersed is unknown; however, the distance this moose travelled from the southern edge of the geographic range of the species was about 600 km. Several other, shorter dispersal events South Dakota, Nebraska, Iowa, and Minne- sota (Fig. 1). In South Dakota a young bull moose was reported near Mobridge (in the north-central part of the state), in November Nebraska, a bull moose was reported in the fall of 2000 near Verdigre (Tom Welstad, NGPC, personal communication). By December the moose had moved farther south to the vicinity had settled on the Elkhorn River near Battle Creek. For the next few months, the moose stayed south of Battle Creek in Madison County where it was observed feeding in a However, in the fall of 2001, it appeared 40 km back to the north near Osmond. Reportedly, the moose was in poor condition and having trouble standing. Shortly after the moose died, the NGPC transported the carcass to the University of Nebraska-Lincoln Veterinary diagnostic lab for necropsy (Associated Press 2001a). The report concluded that cause of death was pneumonia and that, otherwise, the moose was in good physical condition (Dave Oates, NGPC, personal communication). Several additional moose have been reported in Iowa in the past few decades, including 2 individuals that appeared in the northwestern part of the state and dispersed moose seen in Sheldon, Iowa, on 23 September travelled southward near Paulina (Bullard A day later, the moose was reported northeast near Woodbine (World-Herald News Service reported near Underwood, where the moose Another moose was seen in Rock County, for about a month until it was reported again where it was hit by a car near Worthington, LONG-DISTANCE MOOSE DISPERSAL – HOFFMAN ET AL. ALCES VOL. 42, 2006 122 As moose increasingly occur in the central United States, it is of interest to ascertain if any individuals have found suitable habits in which to reside for extended periods of time. In this regard, a cow moose was seen near Crawford, Nebraska (indicated by the hollow star in Fig. moose was seen again by hunters in the same the moose had resided in this area for more than 4 years. In another instance, a cow moose was reported inhabiting Oahe Wildlife Manage- ment Area (WMA), which is located along the Missouri River south of Bismark, North Dakota (indicated by the hollow diamond in cow moose were sighted wandering along the Missouri River just south of Bismark, North Dakota. Soon after, only 1 of the cows was - ers found the moose lying down and unable to stand up, and a NDGFD biologist was called to Oahe WMA to euthanize the moose. From WMA until it died, the moose had lived in the moose in North Dakota encompasses the entire extreme eastern part of the state. However, reproductive populations currently exist only as far south as Cass County, where, in 2001, - munication). Because no reproductive records have been reported south of this location, we conclude that populations located to the south of Fargo in Ransom, Richland, and Sargent counties consist only of vagrant individuals and no permanent populations. These vagrants appear to have spread into Roberts County in northeasternmost South Dakota. Reports of wandering moose in Roberts County have be- come common, with at least one moose being observed in that area each year (Higgins et al. 2000; Will Morlock, SDGFP, personal com- munication). Other reports indicate that moose are establishing reproductive pairs in central - tion). This report represents the westernmost reproductive record of moose in the central North Dakota have been reported from Steele, NDGFD, personal communication). moose have been seen in the central United States (Fig. 1). We report noteworthy records in Nebraska, South Dakota, and North Dakota in Table 3. These accounts are not a compre- hensive list of extralimital records of moose in the central United States. In fact, moose have become so common outside their current geographic range in North Dakota that biolo- gists are only keeping reproductive records and have stopped tracking non-reproductive communication). DISCUSSION Dispersal Patterns Our results suggest that certain dispersal patterns exist with regard to demographics and dispersal distance. For instance, the ma- jority of long-distance dispersal events that occurred throughout the central United States appear to have been undertaken by young bulls in accordance with the process known as “jump dispersal”. However, as proximity to regions inhabited by moose increases, the demographic composition of extralimital records changes. In areas of northeastern South Dakota, records of cow moose become more common and in western North Dakota ALCES VOL. 42, 2006 HOFFMAN ET AL. - LONG-DISTANCE MOOSE DISPERSAL 123 were reported. Female moose, especially diffusion dispersal in that they are dispersing relatively short distances from the established populations within the current geographic range of moose in North Dakota. Causes of Dispersal The increase in moose sightings outside their geographic range could be a direct or indirect consequence of an increase in popu- 341,700 moose inhabited North America, but the number of moose in North America had and to 1,000,000 by 2000 (Timmermann 2003). Harvest population index records indicate that moose numbers in Minnesota and North Da- kota have been at least sporadically increasing, and we suspect, based on published accounts of dispersal routes, that the populations in North Dakota and Minnesota are the source of most dispersing individuals in the central United States. However, it is possible that moose are dispersing into the central United States from other regions as well. The next closest populations of moose are those in the mountainous areas of Montana, Wyoming, in western Nebraska (Fig. 1) might have dis- persed from those populations. For example, a yearling moose was sighted in Laramie, Wyoming, which is situated between the Lara- mie and Medicine Bow mountain ranges, in 2001 (Associated Press 2001b). The straight line distance from Laramie to Scottsbluff, Nebraska, where a bull moose had been seen moose are capable of travelling long distances State Reference Locality Date seen Citation North Dakota 7-Oct-71 7 mi E of Bismark 10-Aug-72 Crown Butte Dam 7-Nov-72 northern Billings County notheastern McLean County southern McLean County southeastern Morton County South Dakota near Flandreau Sioux Falls Mobridge Will Morlock, SDGFP, pers. comm. Will Morlock, SDGFP, pers. comm. 0.5 mi N, and 10.5 mi E of Eureka Will Morlock, SDGFP, pers. comm. 3 mi N of Watertown Will Morlock, SDGFP, pers. comm. 6 mi W of Conde Will Morlock, SDGFP, pers. comm. Nebraska west of Scottsbluff 3 mi NE of Rose Apr-06 Aherns 2002 Table 3. Additional records of moose in North Dakota, South Dakota, and Nebraska. Dates with as- observation dates were given. LONG-DISTANCE MOOSE DISPERSAL – HOFFMAN ET AL. ALCES VOL. 42, 2006 124 possible that individuals can disperse from these areas into adjacent states. When moose began appearing far to the south of their historical geographic range, several hypotheses were proposed by wildlife agencies and media to explain this phenomenon. The idea that a parasite, Pare- laphostrongylus tenuis, causes long-distance dispersal in moose was suggested, primarily by the popular media, as the reason for this unusual behavior. P. tenuis is a nematode that causes a neurological disease known as “moose sickness” (Lankester and Samuel P. tenuis by incidentally ingesting infected gastropods. In many accounts, witnesses claimed that dispersing moose appeared disoriented and lost, thus ultimately leading to the perception that the moose were sick. However, to our sickness in any of the dispersing individuals. on moose of the disease that P. tenuis causes. Symptoms included walking in circles, hold- ing head and ears in abnormal positions, fearlessness, stumbling, deafness, blindness, paraplegia, and, in most cases, death. None of these symptoms suggest that long distance movements are a characteristic of this disease. Given the debilitating effects of this disease, especially on moose locomotory functions, it seems unlikely that infected moose would be physically able to disperse long distances. Another possible explanation is that moose are leaving areas of high population density in search of other suitable habitat and mates. In presumed marginal habitats, such as along the periphery of their geographic range, densities of moose are lower than in areas located toward the center of their geographic range (Telfer areas in marginal habitat cannot support large populations of moose, and that it is necessary for moose to disperse to new habitats. Given their preference for early successional habitats, it seems plausible that moose would have evolved a dispersal behavior that would allow them to travel long distances in search of such with his discussion of transient and permanent habitats are those created by disturbance and are unstable and short-lived. Moose invade these areas shortly after disturbance and, as climax forest reestablishes itself, moose popu- lations decline. Typically, disturbed habitats have a patchy distribution. Once an area is disturbed, species turnover rate is quite rapid and decreases as the community approaches successional habitats have relatively short life spans as compared with climax communities. In order to access these areas, moose would need to be very mobile. Moose were probably one of the last spe- cies to immigrate to North America from Asia via the Bering land bridge (Reeves and Mc- across northern North America (Hundertmark et al. 2002). Different mechanisms by which species expand their range include jump dis- persal, diffusion, and secular migration (Peilou the range expansion of moose by examining the genetic diversity of moose throughout their geographic range. They noted an overall lack of variation in mtDNA; however, haplotype composition was different between peripheral populations and populations inhabiting the central geographic range in North America. From this, they hypothesized that range expansion of moose occurred through a few founding individuals that dispersed from a pre-expansion population. Diffusion dispersal likely would not diminish genetic variation in - lations in previously occupied areas. Rather, diminished genetic variation suggests that range expansion of the moose was the result described as long-distance (i.e., leptokurtic) dispersal, where a few successful, long-dis- ALCES VOL. 42, 2006 HOFFMAN ET AL. - LONG-DISTANCE MOOSE DISPERSAL 125 tance dispersers founded new populations. well, suggesting that founder effects were the cause of genetic homogeneity among the dif- ferent subspecies of moose. Finally, this type of dispersal process agrees with Simpson’s in which an individual disperses from an es- tablished population across a major barrier to another suitable habitat patch. Although, over time, numerous individuals attempt this sort of dispersal event, few are ever successful. Our results favor the hypothesis that re- cent long-distance dispersal events by moose simply are the result of natural dispersal, rather than being induced by disease or other causes. If this is the case, the question then becomes what is the purpose for these dispersal events. We suggest that moose are dispersing from occupied habitats and searching for other suitable habitats. Consequently, we believe that moose are in the process of attempting to expand their geographic range southward and that this dynamic process is similar for most mammals. For example, Genoways et al. (2000) reported on extralimital records of the Mexican free-tailed bat (Tadarida brasil- iensis) throughout the central United States. They concluded that pioneering individuals of T. brasiliensis occurring in areas outside their normal reproductive range are primarily foraging and exploring for new roost sites. They suggested that this is a natural process by which species may extend their geographic recognizable species. An example is the nine- banded armadillo (Dasypus novemcinctus), which recently has dispersed northward from Texas to the central Great Plains (Choate and The likelihood that long-distance dispersal will result in an expansion of range is small. Long-distance movements are rare and gener- ally involve only one animal--often a male. The chance that both a bull and a cow (or a bull calf and heifer) will make a long move across unsuitable habitat to colonize the same new area is poor. The likelihood that diffusion dispersal will result in an expansion of range is much greater because of better reproduc- tive opportunities. Nevertheless, it may be an evolutionary strategy of moose to send out Then, as the more reproductively valuable females disperse more gradually, they will - tat. This would prevent loss of reproductive potential by cows wandering around looking for a mate. In conclusion, during the past 30 years there have been repeated records of moose occurring beyond their southern range bound- ary in the central United States. Most of these individuals consisted of solitary juveniles or young adults; however, there were some exceptions. We were able to track dispersal routes for some of the moose, whereas others were indicated by single locality records. This report documents the longest known distance a moose has dispersed from an established population. We conclude that these events were the result of natural dispersal that could lead to further expansion of their geographic range to the south. Further, we suggest that these dispersal events accurately represent the means by which moose expanded their range through North America during the early Holocene, as described by Hundertmark et al. (2003). Based on the assumption that most species share similar characteristics of range expansion, we believe that this phenomenon can serve as a model to illustrate how other species expand their geographic range. Management Implications The southward movement of moose in the central United States has management implications. Herein, we report two instances of moose inhabiting areas well outside the LONG-DISTANCE MOOSE DISPERSAL – HOFFMAN ET AL. ALCES VOL. 42, 2006 126 current range of moose for an extended pe- riod of time—one in western Nebraska and one in southern North Dakota (Fig 1). These records are noteworthy because they indicate that there are areas to the south of the current geographic range that may sustain permanent moose populations. The primary limiting factor for moose in the southern parts of their geographic range reportedly is climate, par- ticularly high temperatures (Kelsall and Telfer moose experience heat stress at temperatures above 14-20° C. When heat-stressed, moose actively seek areas that provide them with shade and water for cooling (Schwab and Pitt factor, although not as important as climate. Moose can adapt to a variety of forage, but in general they prefer shoots and other woody plants, such as willows (Salix spp.) and other early successional vegetation resulting from - factors may include the density of deer in an area and human impacts, such as urbanization, factors may work together to prevent dispersal events from resulting in an expansion of geo- graphic distribution. Moreover, dispersal can cause expansion only if dispersers eventually likelihood that a long-distance dispersal event will succeed is limited. In central North Dakota, a cow moose in- WMA consists of approximately 6,475 ha of Missouri River bottomland with good moose habitat that provides nearly continuous access to water and shade. Further, Oahe WMA is subject to frequent disturbances that enhance WMA (NDGFD 2005b). Because river bot- tomland usually has a large fuel load, it is disturbed habitat for moose. In northwestern Nebraska, a cow moose was reported living near Crawford for more than 4 years. Crawford is located within the Pine Ridge area where the habitat consists primarily of rolling prairies interspersed with pine forest. Several streams and ponds are located in the area, along with the White and Niobrara river valleys, which provide access to in this area has been converted to agriculture because much of it belongs to the state and that moose do not fare well in areas where human intolerance for moose is high, thus moose tend to occur more often in areas that have not been highly developed. The Pine Ridge area is subject to frequent disturbances resulting from timber harvesting (Blyth et al. along the White and Niobrara rivers. Because harvesting is currently the most important factor in that it stimulates production of early and increases the amount of edge next to these food sources. These areas, that combine edge and food, are favored by moose (Courtois and Beaumont 2002). Given that Pine Ridge is dominated by coniferous forest with a good supply of water and low human development, and given that a lone cow moose was able to survive in this area for more than 4 summers, it is possible that areas in the Pine Ridge area and the Niobrara River Valley may be capable of sustaining small populations of moose. amounts of shade and access to water, supple- mented by frequent disturbance that promotes new plant growth, potentially could serve as suitable habitats for moose populations south of their current geographic range. Areas simi- lar to those described above that are located in adjacent states, such as the Black Hills of South ALCES VOL. 42, 2006 HOFFMAN ET AL. - LONG-DISTANCE MOOSE DISPERSAL 127 Dakota and areas located along the Missouri River and its tributaries in South Dakota, Iowa, Missouri, Nebraska, and Kansas, also might serve as suitable habitat for moose. ACKNOWLEDGEMENTS We would like to thank Mark Sexson of Kansas Department of Wildlife and Parks, Dakota Game, Fish, and Parks, Bismark, North Dakota, Tom Welstad, Nebraska Game and Parks Commission, Norfolk, Nebraska, Dave Oates, Nebraska Game and Parks Commission, Lincoln, Nebraska, and Will Morlock, South Dakota Game, Fish, and Parks, Watertown, South Dakota, for providing us with moose information from their respective states, as well as their personal insight. We especially appreciate the insightful review of an earlier draft of this manuscript by Gordon Eason of the Ontario Ministry of Natural Resources. REFERENCES AHERNS, D. 2002. 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A., and UDSON Seasonal energy expenditures and thermo- regulatory responses of moose. Canadian RICKLEFS, R. E., and G. L. MILLER. 2000. Ecol- ogy. Fourth edition. W. H. Freeman and RINS - well seen near Melrose. The Des Moines Register, SCHWAB, F. E., and M. D. PITT selection of canopy cover types related to operative temperature, forage, and snow 3071-3077. SEABLOOM, R. W., R. D. CRAWFORD, and M. G. MCKENNA southwestern North Dakota: amphibians, reptiles, birds, mammals. Institute for Ecological Studies, University of North ND: Northern Prairie Wildlife Research (accessed 1 Oct 2005). SHEILDS and the evolution of philopatry. Pages in Greenwood, editors. The Ecology of Animal Movement. Clarendon Press, Oxford, UK. SHUGART ETT - cession: similarities of turnover rates. SIMPSON - emy of Sciences 30: 137-163. SIOUX FALLS ARGUS-LEADER on the run may be gone with a bang. Sioux STONE near Larrabee. The Des Moines Registar, SVIHOVEC day in area north of Mobridge. Mobridge TAULMAN OBBINS Recent range expansion and distributional limits of the nine-banded armadillo (Dasy- pus novemcinctus) in the United States. TELFER and habitat requirements of moose (Alces alces in R. Olson, R. Hastings, and F. Geddes, editors. Northern Ecology and Resource Management. The University of Alberta Press, Edmonton, Alberta, Canada. TIMMERMANN, H. R. 2003. The status and management of moose in North America- TREGO - kota Outdoors Vol. LVII(6): 2-4 pp. (UPI) UNITED PRESS INTERNATIONAL moose land, but… The Lincoln Star, 20 south. Omaha World-Herald, 3 March UNRUH ousted…again. Garden City Telegram. VANCE Dateline… ALCES VOL. 42, 2006 HOFFMAN ET AL. - LONG-DISTANCE MOOSE DISPERSAL 131 VOSBURGH, M. R., and R. PETERS of the wandering moose has sad end- ing. Worthington Daily Globe, 1 March WAGNER Dell Rapids. Sioux Falls Argus Leader, WHITE, T. 2001. Wandering moose makes stop in Madison County. Nebraskaland WORLD-HERALD NEWS SERVICE - bond moose draws crowd, stars in Wood- bine home video. Omaha World-Herald, WOSTER OST heads south. The Des Moines Register,