cervids density and the ecological integrity of forest

1
Cervids density and the ecological integrity of forest ecosystems: are deer impacts directly proportional to their density? NSERC - Produits forestiers Anticosti Industrial Research Chair 1 . Cervids as global disturbance agents Cervids are key components of forest ecosystems, yet, as their density increases, they also compromise forest integrity (Côté et al. 2004). Depending on the level of alteration imposed by cervids, forests may lose their capacity to absorb disturbance and to maintain their composition, structure and functions. This is a concern in ecosystems where cervids populations are uncoupled from their natural regulating factors such as density-dependent predation or competition for resources and now act as global agents of disturbance (Tremblay et al. Invited revision). Understanding the functional relationships between forest ecosystems integrity and deer density is essential for ecological-based management of deer-forest systems. In our effort to achieve this, we: Investigated the relationships between the integrity of the forest ground layer and deer density Assessed white-tailed deer (Odocoileus virginianus) densities compatible with the conservation of the integrity of eastern balsam fir (Abies balsamea) – white birch (Betula papyrifera) forests of Anticosti Island, Québec, Canada. On Anticosti, a large introduced deer population has been exerting dramatic impacts on native plant communities which culminated with the conversion of balsam fir stands to white spruce (Picea glauca; Potvin et al. 2003). 0 10 20 30 40 50 Time Deer density (deer/km 2 ) low Potential recovery paths High integrity dynamic regime Low integrity dynamic regime maintained by browse tolerant/resistant species Irreversible threshold - + high Level of alteration ��������Grasses Deer density (deer/km 2 ) Aboveground biomass (g/m 2 ) 0 0 10 20 30 40 50 100 200 300 cutover 2004 forest understory 2004 ��������Abies balsamea Deer density (deer/km 2 ) Aboveground biomass (g/m 2 ) 0 2 4 6 ±1 se max 2002 2003 2004 cutvover 2004 cutover 2003 Epilobium angustifolium 0 0 10 20 30 40 50 0 10 20 30 40 50 10 20 30 40 cutover 2002 cutvover 2004 cutvover 2003 cutover st understory fores Deer density Plant response The classical linear succession hypothesis Nonlinear, potentially multiple equilibriums, hypotheses x 3 replicated blocks x 3 years 10 ha 0 deer/km 2 40 ha x 3 deer 7.5 deer/km 2 20 ha x 3 deer 15 deer/km 2 In situ density (unfenced) 27 to 56 deer/km 2 cutover forest understory 6 . Concluding remarks: an ecosystem-based management model inspired from the catastrophe theory (Scheffer et al. 2001) 2 . Three types of functional relationships proposed as research hypotheses 3 . A controlled browsing experiment designed to monitor functional relationships between forest integrity and deer density 4 . Results Exponential decay functions link biomass of preferred seedlings and forbs to deer density in cutover Exponential growth of grasses biomass as deer density increases in cutover Negative sigmoid functions in the morphological responses of Maianthemum canadense to deer density in cutover 5 . Summing up: several functional relationships within a simple deer-forest system Exponential decay functions dominate in cutover after 2-3 years Usually no response thresholds or thresholds <7.5 deer/km 2 Impacts of deer on forest integrity at low density levels Maximum impact reached at 15 deer/km 2 Positive response of browse tolerant (e.g. grasses) or resistant (e.g. white spruce) species No short term response in forest understory 7 . References cited Côté, S. D., T. P. Rooney, J.-P. Tremblay, C. Dussault, and D. M. Waller. 2004. Ecological impacts of deer overabundance. Annual Review of Ecology, Evolution, and Systematics 35: 113-147. Potvin, F., P. Beaupré, and G. Laprise. 2003. The eradication of balsam fir stands by white-tailed deer on Anticosti Island, Québec: a 150-year process. Ecoscience 10: 487-495. Scheffer, M., S. R. Carpenter, J. A. Foley, C. Folke, and B. Walker. 2001. Catastrophic shift in ecosystems. Nature 413: 591-596. Tremblay, J.-P., I. Thibault, C. Dussault, J. Huot, and S. Côté. 2005. Long-term changes in the availability of white-tailed deer (Odocoileus virginianus) winter forage in the boreal forest and the potential of litterfall as an alternate food source. Canadian Journal of Zoology: Invited revision. The model implies that: Deer act as a global disturbance slowly undermining forest resilience and may force regime shifts Browse tolerant or resistant plants gain a competitive advantage over preferred species and contribute to maintain the system into an alternate dynamic forest regime through positive feedbacks Reduction of deer density >3 years after a canopy disturbance may not be sufficient to restore forest integrity Restoration of altered deer-forest systems may require other direct intervention such as scarification The perspective of catastrophic regime shifts call for a precautionary approach in the management of deer-forest systems, an ecosystem-based perspective and better link between science and management (adaptive management). Rapid inhibition of sexual reproduction as deer density increases in cutover [email protected] Jean-Pierre Tremblay 1,2,3 , Jean Huot 1,2,3 and François Potvin 1,2,4 1 NSERC – Produits forestiers Anticosti Industrial Research Chair, 2 Département de biologie, Université Laval, 3 Centre d’études nordiques, 4 Ministère des Ressources naturelles et de la faune

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Page 1: Cervids density and the ecological integrity of forest

Cervids density and the ecological integrity of forest ecosystems: are deer impacts directly proportional to their density?

NSERC - Produitsforestiers AnticostiIndustrial ResearchChair

1. Cervids as global disturbance agents

Cervids are key components of forest ecosystems, yet, as their density increases, they also compromise forest integrity (Côté et al. 2004). Depending on the level of alteration imposed by cervids, forests may lose their capacity to absorb disturbance and to maintain their composition, structure and functions. This is a concern in ecosystems where cervids populations are uncoupled from their natural regulating factors such as density-dependent predation or competition for resources and now act as global agents of disturbance (Tremblay et al. Invited revision).

Understanding the functional relationships between forest ecosystems integrity and deer density is essential for ecological-based management of deer-forest systems. In our effort to achieve this, we:

• Investigated the relationships between the integrity of the forest ground layer and deer density

• Assessed white-tailed deer (Odocoileus virginianus) densities compatible with the conservation of the integrity of eastern balsam fi r (Abies balsamea) – white birch (Betula papyrifera) forests of Anticosti Island, Québec, Canada.

On Anticosti, a large introduced deer population has been exerting dramatic impacts on native plant communities which culminated with the conversion of balsam fi r stands to white spruce (Picea glauca; Potvin et al. 2003).

0 10 2030

4050

Time

Deer density(deer/km2)

low

Potentialrecovery paths

High integrity dynamic regime

Low integrity dynamic regimemaintained by browse tolerant/resistant species

Irreversiblethreshold

-

+

high

Leve

l of a

ltera

tion

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Grasses

Deer density (deer/km2)

Abo

vegr

ound

bio

mas

s (g

/m2 )

00 10 20 30 40 50

100

200

300

cutover 2004

forest understory 2004

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Abies balsamea

Deer density (deer/km2)

Abo

vegr

ound

bio

mas

s (g

/m2 )

0

2

4

6±1 se max

2002 2003 2004

cutvover 2004

cutover 2003

Epilobium angustifolium

00 10 20 30 40 500 10 20 30 40 50

10

20

30

40

cutover 2002

cutvover 2004

cutvover 2003

cutover forest understoryforest understory

Deer density

Pla

nt r

espo

nse

The classical linearsuccession hypothesis

Nonlinear, potentiallymultiple equilibriums,

hypotheses

x 3 replicated blocks x 3 years

10 ha0 deer/km2

40 ha x 3 deer7.5 deer/km2

20 ha x 3 deer15 deer/km2

In situ density (unfenced)27 to 56 deer/km2

cutover forest understory

6. Concluding remarks: an ecosystem-based management model inspired from the catastrophe theory (Scheffer et al. 2001)

2. Three types of functional relationships proposed as research hypotheses

3. A controlled browsing experiment designed to monitor functional

relationships between forest integrity and deer density

4. Results

Exponential decay functions link biomass of preferred seedlings and forbs to deer density in cutover

Exponential growth of grasses biomass as deer density increases in cutover

Negative sigmoid functions in the morphological responses of Maianthemum canadense to deer density in cutover

5. Summing up: several functional relationships within a simple deer-forest system

• Exponential decay functions dominate in cutover after 2-3 years• Usually no response thresholds or thresholds <7.5 deer/km2

• Impacts of deer on forest integrity at low density levels• Maximum impact reached at ≥ 15 deer/km2

• Positive response of browse tolerant (e.g. grasses) or resistant (e.g. white spruce) species• No short term response in forest understory

7. References cited

Côté, S. D., T. P. Rooney, J.-P. Tremblay, C. Dussault, and D. M. Waller. 2004. Ecological impacts of deer overabundance. Annual Review of Ecology, Evolution, and Systematics 35: 113-147.Potvin, F., P. Beaupré, and G. Laprise. 2003. The eradication of balsam fi r stands by white-tailed deer on Anticosti Island, Québec: a 150-year process. Ecoscience 10: 487-495.Scheffer, M., S. R. Carpenter, J. A. Foley, C. Folke, and B. Walker. 2001. Catastrophic shift in ecosystems. Nature 413: 591-596.Tremblay, J.-P., I. Thibault, C. Dussault, J. Huot, and S. Côté. 2005. Long-term changes in the availability of white-tailed deer (Odocoileus virginianus) winter forage in the boreal forest and the potential of litterfall as an alternate food source. Canadian Journal of Zoology: Invited revision.

The model implies that:• Deer act as a global disturbance slowly undermining forest resilience and may force

regime shifts• Browse tolerant or resistant plants gain a competitive advantage over preferred species

and contribute to maintain the system into an alternate dynamic forest regime through positive feedbacks

• Reduction of deer density >3 years after a canopy disturbance may not be suffi cient to restore forest integrity

• Restoration of altered deer-forest systems may require other direct intervention such as scarifi cation

The perspective of catastrophic regime shifts call for a precautionary approach in the management of deer-forest systems, an ecosystem-based perspective and better link between science and management (adaptive management).

Rapid inhibition of sexual reproduction as deer density increases in cutover

[email protected] Tremblay1,2,3, Jean Huot1,2,3 and François Potvin1,2,4

1NSERC – Produits forestiers Anticosti Industrial Research Chair, 2Département de biologie, Université Laval, 3Centre d’études nordiques, 4Ministère des Ressources naturelles et de la faune