2 May 2014 Chelsie Beck, Jena Lucas, Garrett Shaw, Daniel Picone Pennsylvania State University University Park,

PA 16803, USA RH: Beck et al. Coyote Population Dynamics Coyote Population Dynamics and Management Concerns CHELSIE BECK, Pennsylvania State University, Wildlife Fisheries and Sciences major, Senior, University Park, PA 16802 JENA LUCAS, Pennsylvania State University, Wildlife Fisheries and Sciences major, Senior, University Park, PA 16802 GARRETT SHAW, Pennsylvania State University, Wildlife Fisheries and Sciences major, Senior, University Park, PA 16802 DANIEL PICONE, Pennsylvania State University, Wildlife Fisheries and Sciences major, Junior, University Park, PA 16802 ABSTRACT Coyote (Canis latrans) are responsible for numerous nuisance encounters with humans, livestock, and domestic animals. It is necessary that management practices regarding coyotes should focus on reducing their population. The objective is to lower the total population by 30 percent. All of the management practices focused on the population within Pennsylvania. With the use of multiple Leslie matrices, the population was projected over a number of years for four different harvest rates, and once for the current harvest rate. From this a recommendation of increasing the harvest rate to 30 percent was made, by means of hunter recruitment and programs to increase hunter awareness. This harvest rate was the best option because it lowered the population, while maintaining a healthy population of coyote. Any management action that is

used will require ongoing monitoring to ensure that the population neither increases nor decreases to an unstable size. KEY WORDS Canis Latrans, coyote, suburbs, population dynamics, population management. INTRODUCTION The coyote (Canis latrans) is found in nearly all parts of North America. Although there is some debate on the subject, it is thought that eastern coyotes are actually coyote/wolf hybrids. Coyotes and are also long-term monogamous breeders (Mastro 2011). Eastern coyotes tend to be much larger than their western counterparts, which is thought to be a result of their hybridization with wolves. This also leads to an increase in coyote and human interactions. They are highly adaptable to many habitats, and have even been found in heavily populated suburbs. Coyote are generalists, but tend to prey upon small mammals and deer. Despite many attempts to regulate coyote dispersal and abundance, their populations continue to expand, and collide with human populations (Hayden 2014). Coyotes are somewhat social and tend to live in groups of two to four, although there are records of larger groups that can include more individuals. The size of their territories and home ranges does fluctuate with age, sex, reproductive status, and environmental conditions (Mastro 2011). Coyotes in the eastern part of the United States tend to feed on deer and small mammals, whereas those found further north and west sustain themselves on jackrabbits (Clark 1972). The coyote population in North America is continuing to increase at a very fast rate. As the population continues to increase, there is an increase in the number of coyote interactions with humans. These interactions lead to an increase in human injury by coyotes, loss of livestock, and a loss of game species; therefore, management decisions should focus on lowering coyote populations. Our objective is to decrease the abundance of coyotes by 30 percent based

on the number of coyotes within one square mile. Any effective management practices for ten years, and then it will be determined if the practices need to continue this management based on accounts of human interactions with coyotes. To determine if the management practices are effective, we will compare harvest rates in the previous season to harvest rates the season after we put our management actions into practice. Three different management actions were considered. The first action is to extend the statewide trapping season to six months. Currently, the season runs for four months (midOctober to mid-February), the new season would run from mid-September to mid-March. This action would allow for a greater harvest of coyotes, as well as for coyotes to be harvested during their breeding season. This action would have the desired effect of lowering the overall population abundance; however, there is an issue of partial controllability due to issues of collecting harvest rates from hunters and trappers. The second management action is to have the statewide trapping associations work with local sportsmen clubs to host coyote hunts/trapping contests. These contests could occur in areas where coyotes come into contact with humans more often. This action may allow us to receive better counts on the number of coyotes that are actually harvested. The final option to consider is for state game commission offices to conduct coyote removals or culls in ten square mile patches where coyotes have had increasingly negative encounters with humans. METHODS The objective for this management action of eastern coyote populations are the following: to decrease the total abundance of coyotes by 30%, which is reflective of the problem description (areas where coyote/human interactions are greater). Comparison of harvest rates in the previous season to the season following the implementation of our management actions; but, there is

uncertainty related to partial controllability. The alternative management actions are firstly to extend the statewide trapping season by 2 months (one before and one after). The benefits should lower the overall populations and begin to see a stabilization below lambda equaling 1, see figures 7-10. Also, work with PGC and work in congruency with the Pennsylvania Trappers Association and local sportsmen clubs to host coyote hunts/trapping contests. Coyote removals or culls in one acre patches, all actions focus on areas where there are greater coyote/human interactions. The matrix was created by using known values for reproduction and survival (see Appendix A). In order to project the population size through time with an increase in the harvest rate, the survival rates of juveniles and adults were changed by 10 percent for each harvest rate increase. A formula was ten added to account for density dependence: =(slope*N+2)/3*Number of Juveniles+(slope*N+2)*Number of Adults, where slope was found to be -0.000035, the number of juveniles and adults at year one was arbitrary, 2 was the y-intercept of the most optimistic values for fecundity and survival, and 3 was the number of females born per female. RESULTS The figures are based upon our objective obligations and possible scenarios in concern with density dependence and type of harvest rate projections. When making calculations for the coyote population in excel, a different pattern was found from 30% harvest rates (Figure 3: A)to all other increasing harvest rates (Figure 3: B,C,D). This pattern was made due to the coyotes type of defense against lowering numbers within the population. They tend to fix the lowering numbers of coyotes by increasing the litter (4 to 6 pups 6 to 8 pups), which increases the

number of females in the sex ratio of 1:1. This leads to a density dependence factor within the population dynamics and is depicted in Figure 1. Additionally, within Figure 1, the graph shows the best case scenario for coyote populations (point 0, 2). Meaning that there is no competition in survival and every male finds a female to mate. As the population grows to 40,000 individuals the fecundity drops from 2.0 and is now placed at 0.6. This helped find the relationship among those two living cases, giving the equation for the density dependent scenario, -4E-0.5x +2. This equation was used within the excel formulas to predict the following population pattern at 30%, 40%, 50%, and 60% harvest rates. DISCUSSION In order to meet the objective of reducing the total population of coyote by 30 percent, annual harvest rate should be increased to 30 percent. This is ten percent higher than the current harvest rate, which is 20 percent. This is reinforced by figure 10, which shows that a 30 percent harvest rate does lower the population size, but also normalizes by year 50. It not only allows the goal to be met, but also maintains a healthy population of coyote. This is in contrast to the greater increases in harvest rate, of which a 50 and 60 percent harvest rates result in extinction by year 50. It is anticipated that a 40 percent rate will reach extinction by year 100. Figure 9 also supports the decision because, it shows that at a 30 percent harvest rate, lambda does not reach or exceed one. Also, it does not drop too far below one, resulting in extinction. This value for lambda will continue to decrease the population size; therefore, it is important to continue monitoring the coyote population to ensure that management and conservation practices change as the population changes.

As shown in figure 1, the bar graph continues the calculations in excel and compared them with the types of Harvest Rate Scenarios (HRS). Adding the density dependent factor, the desired information was obtained and concluded what type of HRS should be set in place. The 30% HRS is the targeted desire to lower the numbers of coyotes but allow them to stay below when lambda equals one. This is important because, when lambda is less than one it decreases the population which supports our objective having these results, the coyotes will not feel pressure to expand the litter size, become over populated, or endangered (40% and 50% HRS), or extinct (60% HRS). LITERATURE CITED Clark, Frank W. 1972. Influence of jackrabbit density on coyote population change. Journal of Wildlife Management. 36:343-356 Gommper, Matthew. 2002. Top carnivores in the suburbs? Ecological and conservation issues raised by colonization of north-eastern North America by coyotes. BioScience. 52:2. Hayden, Arnold. Pennsylvania Game Commision: Eastern Coyote. <http://www.portal.state.pa. us/portal/server.pt/community/coyote/14636. Accessed February 15, 2014. Mastro, Lauren L. 2011. Life history and ecology of coyotes in the mid-Atlantic states: A summary of the scientific literature. Southeastern Naturalist. 10(4):721-730. Prugh, L.R. 2005. Monitoring coyote population dynamics by genotyping faeces. Molecular Ecology. 14:1585-1596. Schrecengost, Joshua D. 2009. Home range, habitat use and survival of coyotes in western South Carolina. Am. Midl. Nat. 162:346-355. Windberg, Lamar. 1995. Demography of a high-density coyote population. Canada Journal of Zoology. 73:942-954.

Appendix A. Pre-Juvenile Pre-Juvenile Juvenile Adult 0 0.42 0 Juvenile 0.2 0 0.84 Adult 0.6 0 0.72

Table 1: This shows our initial matrix. The values for reproduction and survival were gathered from past research (Mastro 2011). 2.5 Fecundity (F) 2 1.5 1 0.5 0 0 10000 20000 30000 40000 Popultion Abundance (N) 50000 y = -4E-05x + 2

Figure 1: We assume there was density dependence. The relationship between Fecundity (F) and Population Abundance (N). We predicted fecundity when population size was small to be ~2 and that there would be a linear relationship between fecundity and N crossing at 0.68 for the current population size. 1.2 1.15 1.1 1.05 1 0.95 0.9 0.85 0.8 2000 2010 2020 2030 Time 2040 2050 2060

Figure 2: This figure displays a basic relationship between the lambda and time from 2000 to 2060. As the relationship depicts, around 2010 the lambda begins to have a deep drop (from 1.15 to .9). From 2010 to 2060 the lambda seems to stay constant, depicting the carrying capacity of the population.

Figure 3: (A) This figure depicts the basic relationship between lambda and time, within the 30% harvest rate of coyotes. In early 2005-2010 the lambda seems to have a gradual rise with a low dip of 0.8 within year 2010. After 2010, the lambda has a minor increase from 0.9 to 1.05. Overall, it seems to level off and remain constant. (B) This figure depicts the basic relationship between lambda and time, within the 40% harvest rate of coyotes. In early 20052010 the lambda seems to have a gradual rise with a low dip of 0.8 within year 2010. After 2010, the lambda has a minor increase from 0.9 to 1.0. Overall, it seems to level off and remain constant. (C) This figure depicts the basic relationship between lambda and time, within the 50% harvest rate of coyotes. In early 2005-2010 the lambda seems to have a sharp peak with a low dip following (0.75) within year 2010. After 2010, the lambda has a minor increase from 0.8 to 0.9. Overall, it seems to level off and remain constant. (D) This figure depicts the basic relationship between lambda and time, within the 60% harvest rate of coyotes. In early 2005-2010 the lambda seems to have a sharp peak with a low dip of 0.6 within year 2010. After 2010, the lambda has a minor increase from 0.8 to 0.9. Overall, it seems to level off and remain constant.

1.05 1 0.95 0.9 0.85 0.8 0.75 0.7

Figure 4: This figure depicts a relationship between lambda and change in abundance within a 30% harvest rate. The lambda seems to have a gradual increase in the change of abundance, but a small dip in lambda when there is 6,381 coyotes within the population. Lambda also depicts a leveling around 1. 1.05 1 0.95 0.9 0.85 0.8 0.75 0.7

Figure 5: This figure depicts a relationship between lambda and change in abundance within a 40% harvest rate. The lambda seems to have a gradual increase in the change of abundance, but a small dip in lambda when there is 5,617 coyotes within the population. Lambda also depicts a leveling around 1.

1 0.95 0.9 0.85 0.8 0.75 0.7

N Figure 6: This figure depicts a relationship between lambda and change in abundance within a 50% harvest rate. The lambda seems to have a gradual increase in the change of abundance, but a small dip and small increase in lambda when there is 5,000 to 2,000 coyotes within the population. Lambda also depicts a leveling around 0.95. 0.9 0.85 0.8 0.75 0.7 0.65 0.6

Figure 7: This figure depicts a relationship between lambda and change in abundance within a 50% harvest rate. The lambda seems to have a gradual increase in the change of abundance, but small peak (than decrease following) in lambda when there is 6,000 to 1,500 coyotes within the population. Lambda also depicts a leveling around 0.9.

Total Population in 2057

14000 12000 10000 8000 6000 4000 2000 0 Current Population 30% 40% 50% 60%

Harvest Rate (%)

Figure 8: This chart depicts the levels of total population in year 2057 within each type harvest rate (30%, 40%, 50%, and 60%). The current harvest rate is 20%. The chart depicts the current population is around 10,000 coyotes, the 30% harvest rate to increase the numbers of original coyotes (density-dependence indicator), decreasing in the 40% and 50%, but a possible extinction at 60% harvest rates.

1 0.98 0.96 0.94 0.92 0.9 0.88 0.86 0.84 0.82 0.8

ASymptotic Lambda

Current Population 30% 40% 50% 60%

Harvest Rate (%)

Figure 9: This chart depicts the levels of asymptotic lambda within each type harvest rate (30%, 40%, 50%, and 60%). The chart depicts the current population is around .96 lambda, the 30% harvest rate to increase the numbers of original coyotes (density-dependence indicator), decreasing in the 40%, 50%, and 60% harvest rates.

45000 40000 Popultion Abundance (N) 35000 30000 25000 20000 15000 10000 5000 0 30% 40% 50% Harvest Rate Scenario 60% currant year 10 yrs 20 yrs 30 yrs 40 yrs 50 yrs

Figure 10: This chart depicts the overall predicted change in population abundance (N) within each type of harvest rate scenario (HRS), as decades go by. Within 30% HRS, the current population is the highest and the following decades seem to decrease but come to a gradual level around 12,500 coyotes. Within 40% HRS, the current population is the highest and the following decades seem to decrease in intervals. Within 50% HRS, the current population is the highest and the following decades seem to decrease and cause population regeneration problems in the 40 and 50 decade. Within 60% HRS, the current population is the highest (same as 40% current population), but only can account for the 10th and 20th decade marker without causing the population to be unrecoverable.