Assessing the Potential Effects of Treponeme Associated Hoof Disease (TAHD) on Elk Population Dynamics in Southwest Washington

Various hoof diseases have been reported worldwide in numerous free-ranging ungulates, including elk (Cervus elaphus; Murie 1930, Gray et al. 2001, Thorne et al. 2002), mule deer (Odocoileus hemionus; Wobeser et al. 1975), white-tailed deer (O. virginianus; Sleeman et al. 2009), moose (Alces; Flynn et al. 1977, Clauss et al. 2009), fallow deer (Dama; Lavin et al. 2004), reindeer (Rangifer tarandus; Handeland et al. 2010), roe deer (Capreolus; Handeland and Vik.ren 2005), and mouflon (Ovis gmelini musimon; Volmer et al. 2008). Reports of elk in southwestern Washington with evidence of lameness or various hoof abnormalities were historically sporadic and infrequent. In early 2008, however, the number and geographic extent of elk displaying evidence of an apparently novel hoof disease significantly increased (Mansfield et al. 2011, WDFW unpublished data).

The emergence of this disease in southwest Washington elk herds is unique in that bacteria in the genus Treponema, (aka "treponemes"), never previously associated with hoof diseases in any free-ranging ungulate, have been identified as causal (Clegg et al., 2015). Treponemes are strongly associated with similar diseases of domestic livestock: bovine digital dermatitis of cattle (Evans et al. 2009), contagious ovine digital dermatitis (CODD) of domestic sheep (Sayers 2009), and a CODD-like disease of domestic goats (Sullivan et al., 2015).

Elk affected by treponeme-associated hoof disease (TAHD) often have severely overgrown and deformed hooves with sole ulcers and sloughed hoof walls (Han and Mansfield 2014). TAHD can occur in multiple limbs and can affect all age and sex classes (Clegg et al. 2015). The severity of clinical signs, coupled with the seemingly rapid expansion of impacted areas, have generated a great deal of concern for the Washington Department of Fish and Wildlife (WDFW), other resource management agencies, hunters, tribes, and local citizens. In response to these concerns, WDFW continues to work with several specialists to better understand the etiology of TAHD. In addition, WDFW established a Hoof Disease Technical Advisory Group (HDTAG) and a Hoof Disease Public Working Group (HDPWG). The HDTAG has guided the diagnostic effort, identified research needs, and provided review and input to management options. The HDPWG has provided input to management and research options and serves as a venue for WDFW to share information with the public. However, it is difficult to assess what implications TAHD will have for the management of affected elk herds because the effects of TAHD on elk vital rates (e.g., survival, reproduction, etc.) are unknown.

It is reasonable to assume that elk with advanced stages of TAHD may have a decreased probability of survival because their infirmities may predispose them to predation, harvest, severe weather events, or other types of disease (Bender et al. 2008). For example, mule deer with chronic wasting disease (CWD), prior to developing obvious clinical signs, have been shown to be more vulnerable to predation (Miller et al. 2008, Krumm et al. 2009), vehicle collisions (Krumm et al. 2005), and possibly harvest (Conner et al. 2000). This is an important consideration because the growth rate of large ungulate populations, such as elk, is highly sensitive to changes in adult female survival (Nelson and Peek 1982, Eberhardt 2002) and strongly correlated with the production and survival of juveniles (Gaillard et al. 2000; see also Smith and Anderson 1998, Raithel et al. 2007). When adult female and juvenile survival are concurrently reduced, populations would be expected to decline (Gaillard et al. 2000; see also Bender et al. 2007, McCorquodale et al. 2014). Consequently, if TAHD reduces the survival of affected cows and calves, it has the potential to have a negative effect on the population dynamics of impacted elk herds.

Although McCorquodale et al. (2014) monitored 16 adult female elk that had varying degrees of presumed TAHD (i.e., they had varying degrees of hoof deformities, but no lab samples were collected and tested) inferences from their work are limited. Twelve of 16 affected elk they monitored survived ≥ 1 year and of those that did not survive ≥ 1 year, all were harvest-related mortalities. In addition, 3 of 4 elk that were fitted with VHF collars that had a battery life of several years survived until radio contact was lost 3-4 years after they were captured. Anecdotally, this indicates that if TAHD negatively affects the natural survival of elk, it may take several years before it does so. We need to improve our understanding of how quickly TAHD progresses and if, and when, it may begin to predispose affected elk to mortality.

TAHD may also have the potential to affect the population dynamics of impacted elk herds because of its effect on energy dynamics of female elk. The nutritional condition of female ungulates can influence age at first breeding (Cook et al. 2004), timing of estrus and subsequent birth date (Andersen and Linnell 1998, Cook et al. 2004, Bishop et al. 2009), probability of conception (Cook et al. 2004, Cook et al. 2013), fetal development and survival (Verme 1969, Ozoga and Verme 1982), birth weight (Verme and Ullrey 1984, Keech et al. 2000, Lomas and Bender 2007), milk yield or composition (Landete-Castillejos et al. 2003, Tollefson 2007), and subsequent growth and survival of juveniles (Clutton-Brock et al. 1982, Bishop et al. 2009). For example, elk from the Mount St. Helens elk herd area (MSH) and other coastal regions of Washington are characterized by pregnancy rates for prime-aged females that are consistently depressed [Kuttel 1975 (74%), Smith 1980 (61%), Cook et al. 2013 (68-100%), McCorquodale et al. 2014 (71%)] because marginal summer-autumn nutrition limits the level of condition female elk are able to achieve during the breeding season (Cook et al. 2013). Due to the additional energetic requirements for mounting an immune response and for tissue repair (Deming 2009), TAHD may further limit the ability of affected elk to improve their condition during the breeding season and therefore has the potential to reduce overall pregnancy rates even further, which could reduce demographic vigor.

Some have attributed recent declines in the MSH elk herd to TAHD because the monitored portions of the MSH herd declined by 30-35% over a 4-year period (2009.2013; McCorquodale et al. 2014) that coincided with an increase in the prevalence and distribution of the disease (WDFW, unpublished data). However, this period of population decline also occurred concurrently with a directed effort by WDFW to reduce the elk population through substantial increases in antlerless harvest because of evidence that the MSH elk herd was above ecological carrying capacity (WDFW 2006, McCorquodale et al. 2014). Moreover, density independent severe winter weather that occurred in 2012 likely contributed to the documented decline (McCorquodale et al. 2014). Because these three events overlapped temporally and elk with presumed TAHD represented <15% of the adult females that were monitored, McCorquodale et al. (2014) were not able to conclude whether or not TAHD was a contributing factor in observed declines.

The number of elk that have TAHD and the effects of TAHD on elk vital rates, collectively, will determine what the long-term implications of TAHD are for the viability, and subsequent management, of impacted elk herds (Wobeser 2007). Consequently, our primary research goals are to quantify how TAHD may affect the survival, pregnancy rates, productivity, and nutritional condition of adult female elk. Our specific study objectives include:

1. Estimate the effects of TAHD on survival of adult (. 2 years old) female elk.
2. Determine cause-specific mortality rates for adult female elk that have TAHD.
3. Estimate the effects of TAHD on the pregnancy rates of adult female elk.
4. Estimate the effects of TAHD on elk productivity (i.e., survivorship of calves).
5. Estimate the effects of TAHD on the level of condition (i.e., IFBF) adult female elk are able to achieve in autumn.
6. Increase our understanding of how TAHD progresses in individual elk, and whether affected elk may recover from the disease.