1

121 April 2015

2Charles McCole
3B.S. Wildlife and Fisheries Sciences
4Pennsylvania State University
5University Park, PA, USA
6Email: cvm5196@psu.edu
7
8RH: McCole et al. Ruffed Grouse Populations in Scotia Barrens
9Proposed Reversal of the Steady Decline in Ruffed Grouse
10Populations in the Scotia Barrens Region of Pennsylvania
11CHARLES MCCOLE, Department of Ecosystem Science and Management, Pennsylvania State
12

University, 416 Forest Resources Building, University Park, PA, USA

13MARK CHRONISTER, Department of Ecosystem Science and Management, Pennsylvania

14

State University, 416 Forest Resources Building, University Park, PA, USA

15SHARA HERMAN, Department of Ecosystem Science and Management, Pennsylvania State

16

University, 416 Forest Resources Building, University Park, PA, USA

17Corresponding author: cvm5196@psu.edu

18Present address: 416 Forest Resources Building, University Park, PA, USA
19ABSTRACT
20Ruffed grouse (Bonasa umbellus) are a species of concern for the state of Pennsylvania.
21Historical records compared to current population densities are revealing a sharp decline across
22the state at a rate of ~15% per year. Conservation efforts are needed to reverse the trend enabling
23the reestablishment of growing populations across the state. The Centre County population
24located on Pennsylvania Game Land #176, commonly called Scotia Barrens, is the basis for our
25study. We used an age-structured Leslie matrix model for our population projections. Models
26were ran which simulate the management actions that were considered: do nothing, increase
27suitable grouse habitat, and implement an open season on avian predators. Population data was

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28obtained from a previous long-term study done in the Scotia Barrens form 1976 to 1998 (Storm
292003). Data for the effects of habitat management and avian predator reduction on ruffed grouse
30populations was also obtained from the study. The current growth rate () of the grouse
31population that we studied is 0.84, which indicates a declining population. Our goal for the study
32was to determine if implementing the proposed management action could increase the -value
33greater than 1.0. After running the models, it was determined that our habitat management action
34would increase the by 0.19, which would result in a of 1.03. We also ran a model for our
35management action of reducing the avian predator population by opening a hunting season,
36removing a quarter of the predators. Our results showed that only increased by 0.05, up to 0.89,
37much lower than our target value. Although it is commonly known that ruffed grouse populations
38operate in a 10 year cycle, it is clear from our results that habitat plays a very large role in their
39success and managing habitat has the greatest effect on their numbers.
40Key Words: Ruffed Grouse, Bonasa Umbellus, Scotia, Habitat, Avian, Predation
41

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42

The Ruffed grouse (Bonasa umbellus) is the state bird of Pennsylvania. It has become a

43commonly hunted game species throughout the state. Ruffed grouse have a maximum lifespan in
44the wild of about 8 years. The females mate every year and on average lay 10 eggs per season.
45However, 63% of the first nesting events are unsuccessful, meaning that no eggs are hatched
46from the nest. In those cases, there is a 100% re-nesting rate, although only 33% of second nests
47are successful. Grouse chicks are precocial and will begin to walk on their own and forage upon
48hatching. In a week they are able to fly. Grouse mature at age 1 and have a constant fecundity
49rate once maturity is reached. Male grouse are territorial although grouse are not monogamous
50and both males and females can mate with multiple grouse. They thrive in early successional
51habitat which is growing scarce throughout the state of Pennsylvania, resulting in their declining
52numbers (Rusch 2000).
53

Pennsylvanias population of grouse is rapidly decreasing and if current rates of decline

54continue the population will be essentially extinct in 35 years. Specifically, in Scotia Barrens,
55abundance has declined 10-15% per year. Though the trend is thought to be primarily a result
56from habitat destruction, predation may also be a cause for concern. To reverse the decline of
57grouse, habitat must be analyzed to determine if a change in the landscape will contribute to the
58reversal in population trajectories (Storm 2003.).
59

The objective of our study is to increase the current rate of population growth () by a

60minimum 0.19, thereby increasing it to 1.03 and promote population increase. Current research
61indicated that grouse populations in PA are declining at a rate of 16% a year (=0.84). (Table 2,
623). To determine the success at meeting this objective, we looked at population abundance using
63flush counts and nest counts. Counting nests can give you the female population abundance as
64well as nest productivity numbers. Three actions were chosen and modeled to determine which

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65management practices would help us to achieve our objective. They are: do nothing, increase
66suitable grouse habitat, and implement an open season on avian predators.
67MANAGEMENT ACTIONS
68The first action that we are proposing is to do nothing. This is the easiest and most cost efficient
69action, as no extra effort will have to be made. For this action, no changes will be made to any of
70the current management plans concerning ruffed grouse and the populations will continue to be
71observed until quasi-extinction is reached.
72

Our second action is to increase the suitable habitat of the ruffed grouse in the study area

73by at least 50%. Suitable habitat is thought to be one of the biggest contributors to survivability
74of ruffed grouse. Grouse thrive in a very narrow age class of vegetation which makes it difficult
75to maintain suitable habitat for long periods of time. The constant creation of new habitat
76involves clear-cutting tracts of land to enable proper vegetation to grow. However, this process
77has the potential of being very costly if done over large tracts of land.
78

Our third action is to remove a specified number of avian predators from the study area.

79Avian predators account for 80% of the mortality of ruffed grouse. Northern goshawk, northern
80harriers, great horned owl, and barred owls are the top predators of grouse chicks. By reducing
81the number of predators in grouse habitat this would allow a higher rate of chicks to survive to
82maturity and a breeding age. By having more breeding adults in a population the abundance
83should increase dramatically over time.
84

To meet the objectives, full cooperation is required from all stakeholders that have an

85interest in the success of the species. Hunters provide a valuable source of income that aids in all
86management objectives implemented. A compromise needs to be made if the decline of a popular
87game species is to be averted. Both alternatives can be met with adjustments to the regular

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88hunting season and suspension of the birds of prey protection clause covered in the Migratory
89Bird Treaty Act. Allowing for the harvest of a species like the Red Tailed Hawk would provide a
90substitute to the temporary removal of Grouse as a game species. Additionally, the action would
91increase the survival of ruffed grouse by eliminating a direct prey species. This two-step solution
92has the ability to remove the additive effects of harvest and predation and could return the
93population to lambda levels greater than 1.0.
94STUDY AREA
95The Pennsylvania Game Commission maintains about 1.5 million acres of land across the state
96of Pennsylvania (PGC 2015). This land is open to the public and maintained by funds generated
97through license and firearms sales. Many sites are managed and manipulated to research various
98species of game animals to better the experience for hunters. Specific to this grouse project State
99Game Land (SGL) 176, also named Scotia Barrens, has been managed to improve and better
100understand the habits of ruffed grouse. SGL 176 characteristics are mixed hardwoods common
101to the Pennsylvania landscape. One major difference to the area is in the name. The barrens of
102the area are derived from the abundance of jack, pitch, and white pines growing in the acidic
103soils that are found at SGL176. Elevations ranged from 360 m to 450 m. Temperatures in the
104area range from -3.4 C to 21.9 C (National Oceanic and Atmospheric Administration 1985).
1051,120 ha of land with in SGL 176 (2500ha total) were used to establish a controlled area. This
106area has been selectively managed by the PGC since 1975 to establish ideal habitat for ruffed
107grouse and other various game species to prosper (Storm 2003).
108METHODS
109In our original population model, we used population abundance data obtained from a study done
110from 1976-1998 at our study site. Survival and fecundity rates were obtained from a grouse

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111demography report put out by the USGS as well as a breeding bird survey. An age-based Leslie
112matrix was used to estimate population abundance into the future, based on the current
113population, fecundity, and survivability numbers. The model showing the original data and
114population projection was used as our do nothing model.
115

In order to change the current population projection two methods of management were

116chosen; increasing favorable habitat and decreasing avian predation. The best grouse habitat is a
117mosaic of different age structures. They need early successional patches to nest, hide from
118predators, and soft mast crops. Mature growth habitat is needed for hard mast that will feed them
119through the winter.
120

In order to achieve this mosaic patchwork the Barrens were put on a clear-cut rotation.

121Every 10 years a 1-ha plot will be systematically chosen to be clear-cut and allowed to
122regenerate. Any harvestable trees will be sold while undesired trees will be cleared out of pocket.
123In particularly dense blocks an understory burn will take place before the cut in order to recycle
124nutrients back into the soil and open up the understory for easier navigation. Hard mast
125producing trees will be left in place to ensure that the regeneration is of the appropriate species.
126Based on the previous Scotia Barrens grouse study, grouse chick survivability increased
127proportionally with an increase in suitable grouse habitat, which is reflected in our tables and
128graphs (Storm 2003).
129

In order to decrease avian predation, there will be a reduction of avian predators in our

130study area. Over the entire study site, 2500 Ha, we determined that there were about 20 total
131individuals representing the four species that we targeted. The removal of 20% of all avian
132predators from the study area, a total of about five individuals over the site, is proposed. A
133temporary open season on avian predators will be enacted to allow for hunters to harvest the

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134allotted amount to achieve our management action. All falcons are protected under the birds of
135prey protection clause in the Migratory Bird Treaty Act. We chose 20% because many avian
136predators prey on other nuisance species as well. The changes in survivability and population
137abundance is shown in our tables and graphs.
138RESULTS
139Our do nothing model resulted in a grouse population declining at 16% per year, a of 0.84.
140was reached at the 7 year mark. At that rate, the ruffed grouse population at our study site would
141go extinct at the 35 year mark.
142

It was found that when suitable grouse habitat is increased by 50%, first year survival

143will increase by at least 50%. This will ensure that the chicks have a higher rate of survival
144during the first year which would bolster the upper age classes in the following years. This
145practice increased the lambda to 1.03 within 7 years, an increase of 0.19.
146

The removal of 20% of avian predators was not found to be as successful. At this rate the

147lambda only increased to .89, an increase of 0.05. In order to increase this rate by removal alone
148many more birds would need to be harvested which is not feasible.
149MANAGEMENT IMPLICATIONS
150At the current rate, the grouse population in Scotia Barrens of Pennsylvania will be extirpated
151within 35 years. Predation accounts for over 80% of all mortality. Avian predators account for
152most of the predation deaths. Management action is drastically needed to protect this vital game
153species. A couple management practices have been discussed in this paper such as increased
154habitat and an open season on avian predators.
155

Habitat has shown to be one of the most important determining factors in a successful

156ruffed grouse population. Habitat provides adequate cover and the proper food source for all age

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157classes. The data for our study was taken from a ruffed grouse study done at the Scotia Barrens
158in north central Pennsylvania. The study found that increasing suitable grouse habitat by fifty
159percent increases first year survival by at least 50%. For our study we manipulated the habitat
160cover to increase abundance. The result was that the lambda increased by 0.19. The resulting
161lambda was 1.03 which indicates an increasing population. If that data is accurate, this
162management practice would solve the current population decline problems. Additionally, it
163shows that even more habitat management could result in higher lambda values. This action has
164the potential to create more hunting opportunities and increase hunter satisfaction which would
165be a secondary benefit.
166

The second action that was researched was opening up a season on avian predators,

167mainly red-tailed hawks, northern harriers, great horned owls, and barred owls. The main goal
168was to increase the lambda above 1.0. After the removal of the allotted predators our lambda
169only increased by 0.05, from 0.84 to 0.89 which is far below the goal. However, this directive
170will extend the time before the population becomes extirpated from the region. An increase in
171predator harvest was considered but avian predators play other important roles in the
172environment and nearly all would need to be removed to have a dramatic impact on the growth
173rate. Overall, this action did not prove to be a viable option for our goals. For one, the likelihood
174that a season would be opened is very slim, as all of these species are protected under federal
175law. The second reason is that it isnt feasible to remove the number of individuals that are
176required to increase lambda by our desired amount.
177

It was found that it is possible to increase the growth rate of the ruffed grouse population

178in the region and to prevent future extirpation. The most ideal situation is to indirectly affect the

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179population by improving desirable habitat. The alterations to the environment have the
180possibility of benefitting other game species and increasing overall hunter satisfaction.
181REFERENCES
182Fearer, Todd M.; Stauffer, Dean F. 2004. Relationship of ruffed grouse Bonasa umbellus to
183

landscape characteristics in southwest Virginia, USA. Wildlife Biology. 10(2): 81-89.

184Hewitt, David G.; Keppie, Daniel M.; Stauffer, Dean F. 2001. Predation effects on forest grouse
185

recruitment. Wildlife Society Bulletin. 29(1): 16-23.

186Rusch, Donald H.; Destefano, Stephen; Reynolds, Michael C.; Lauten, David. 2000. Ruffed
187

grouse--Bonasa umbellus. In: Poole, A.; Gill, F., eds. The birds of North America. No.

188

515. Philadelphia, PA: Birds of North America. 28 p.

189Sauer, J. R., J. E. Hines, J. E. Fallon, K. L. Pardieck, D. J. Ziolkowski, Jr., and W. A. Link. 2014.
190

The North American Breeding Bird Survey, Results and Analysis 1966 - 2012. Version

191

02.19.2014 USGS Patuxent Wildlife Research Center, Laurel, MD

192Storm, G. L., W. L. Palmer, and D. R. Diefenbach. 2003. Ruffed grouse responses to

193

management of mixed oak and aspen communities in central Pennsylvania. Grouse

194

Research Bulletin No. 1, Pennsylvania Game Commission, Harrisburg, Pennsylvania,

195

USA

196National Oceanic and Atmospheric Administration. 1985. Climates of the states. II. Gale
197

Research Company, Detroit, MI. 1572pp.

198PGC, Pennsylvania Game Commission, 2015. Pennsylvania State Game Lands. April 24, 2015.
199

http://www.portal.state.pa.us/portal/server.pt/community/state_game_lands/11363

200

(accessed April 24, 2015).

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201Table 1. Survival and fecundity, presented in a Leslie matrix, for Ruffed Grouse in the Scotia
202Barren region of Pennsylvania adjusted for each of our actions. Survival rates were obtained
203from a USGS report on Ruffed Grouse. Fecundity rates and the initial population values were
204obtained from USGS Patuxent Wildlife Research Center, Laurel, MD North American Breeding
205Bird Survey. Fecundity rates reflect an initial nesting event and a secondary nesting event that
206occurs in the case of a failed first nest.

207

0
0.4447
0.069
0
0
0

1
2.5997
0
0.5
0
0

0
0.4447
0.0828
0
0
0

1
2.5997
0
0.5
0
0

0
0.4447
0.139
0
0
0

1
2.5997
0
0.5
0
0

Do Nothing
2
2.3497
0
0
0.45
0

3
1.8497
0
0
0
0.35

4+
1.3497
0
0
0
0.25

Predator Removal
2
3
2.3497
1.8497
0
0
0
0
0.45
0
0
0.35

4+
1.3497
0
0
0
0.25

Habitat Increase
2
2.3497
0
0
0.45
0

3
1.8497
0
0
0
0.35

4+
1.3497
0
0
0
0.25

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208Table 2. The proportion of the total population that is occupied in each age class at any given
209year once the asymptotic lambda is reached. All three management actions are represented in the
210table. Asymptotic lambda is reached in the 7th year of our projection.
Proportion of individuals in age class (Do Nothing)
0
0.853082666

1
0.069959518

2
0.041565822

3
0.02221529

4+
0.013176703

Year
7

Asym
Lambda
0.841399663

Year
7

Asym
Lambda
0.883262454

Year
7

Asym
Lambda
1.024524577

Proportion of individuals in age class (Predator Decrease)

0
0.841194106

1
0.078858414

2
0.044637943

3
0.022729369

4+
0.012580168

Proportion of individuals in age class (Habitat Increase)

0
0.803787567

211

1
0.109050549

2
0.053233548

3
0.023361108

4+
0.010567228

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Figures
Figure 1. Semi-log graph showing the population size projections for all age classes as well as
the total population, predicted 35 years into the future for the do nothing action. Age classes 1
through 4+ reach extinction between years 12 and 23. Age class 0, as well as the total population,
reaches extinction around year 35. The consistent population decline correlates with the
asymptotic lambda, 0.84.
Figure 2. Semi-log graph showing the population size projections for all age classes as well as
the total population, predicted 35 years into the future for the habitat increase action. An
asymptotic lambda of 1.024 is reached at year 7. Populations continue to increase throughout the
35 year projection.
Figure 3. Semi-log graph showing the population size projections for all age classes as well as
the total population, predicted 35 years into the future for the avian predator removal action.
Lambda was 0.89, which pushed extinction back 20 years.

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Figure 1.

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Figure 2.

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Figure 3.