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• Overview of beech ecology• Role as wildlife food and habitat• Spatio-temporal cycles and masting

• Population demographics• Predator-prey relations• Human-wildlife conflict

• Beech Bark Disease impact on forest system• Conclusions & Implications• Current Projects

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Thomson et al. 2013

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Northern Research StationForest Inventory and Analysis

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American BeechBasal Area per Acre

Northern Research StationForest Inventory and Analysis

Sawtimber, paper, firewood, ornamental tree

Wildlife valuesFoodCavities for nestingDowned wood

Oft-maligned species

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• Model predicted 37% decline in nut production from BBD

• Significant and widespread impacts on wildlife and forest

Costello 1992

02468

10

1948 1989 2030

Beechnuts x100,000/ha

?

BBD

↓ Large northern hardwoods dominating canopy

↑ Beech saplings

↑ More, smaller beech logs

↓ Recent beech snags

↑ Advanced stages of BBD

↑ Shrub richness

↑ Beech nut production trendMcNulty and Masters 2004

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Component Non-thicket Thicket

Mean beech sapling density (stems/m2)

0.12 0.55

Mean litter depth (cm) 1.48 2.25

Plant species 34 22

Fern species 12 6

Mean shrub species 6.5 1

Cale et al. 2012

• Beech thicket• Light levels lower; competition higher?• Dense, lignin-rich beech litter makes herb establishment difficult –

phytotoxic?

• Higher diversity• Non-thicket had a third more species• Caveat: mechanism that caused a thicket not studied (soil,

drainage, land use history…?)

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• Vertebrates – runways, refugia• Fungi, insects – decomposition rate• Woodpeckers, chickadees – cavity nests in snags

• About 40 species• Deer eat nuts, but rarely browse leaves

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• Hard mast

• Produced at age 40-60• Ripe in late August -

November

• Beech Mast Cycle • 2-8 years• Northeast = 2 year cycle• Weather affected

Pre-denning period of hyperphagy

Stored Cache• Non-hibernators • Torpor-employers

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Component Beech nut Oak Herbs Corn

Protein 11% 6% 16% 11%

Fat 17 14 4 3

Carbohydrate 30 61 40 80

Fiber 27 18 26 2

Elowe and Dodge 1989Beech nuts:1.2 - 6 x more fat!

• Lignin – difficult to digest• Secondary plant compounds

• Polyphenolics• Allelopathic, impacts sugar maple seedling

development

• Structural • Tough husk, seed coat

• Beech nuts vs. oak acorns Hane 2003

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• Sow’s fat store winter influences number of cubs next spring

• Maine: 80% of sows reproduced after mast, 22% after poor nut year

• Beech mast density in the autumn before birth best explained annual Adirondack bear population size

Jakubas et al. 2005 LaMere et al. 2012

• Weasels, marten, fisher

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• Indirectly impacted by beech mast• Consume small mammals• Time lag for predators

• Owls• Raptors• Bobcats• Coyotes • Otters – direct!

Squirrels Deer Mice Jumping Mice Voles Shrews

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Blue Jays• Cache nuts up to 4 km away - can expand a tree's range

• Creates clusters of related beech• Chose 100% sound, green nuts vs. just

11% sound nuts hand-picked from tree

Ruffed Grouse• Overwinter survival

Turkeys• Increasing in northern regions

Johnson and Adkisson 1985

• Passenger pigeons ate beech nuts and acorns

Mott 1901

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• Unmanaged >300-year-oldnorthern hardwood forest

• Annual since 1988• Mid-November, 25 plots

• Five managed beech stands - 1989-present

0

50000

100000

150000

200000

250000

300000

350000

400000

450000

500000

1988 1990 1992 1994 1996 1998 2000 2002 2004 2006 2008

Beec

h M

ast P

rodu

ctio

n (s

ound

see

ds/h

a)

1-year lag (first order autocorrelation) = -0.544-year lag (fourth order autocorrelation) = 0.61

• Mast crop = 100x more nuts produced than during a mast failure

• Tree seed production synchronized (r = 0.48 – 0.64)

• Beech, sugar maple, mountain ash, conifers

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0

5

10

15

20

25

0

30

60

90

120

150

180

2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015

Mic

e pe

r 100

Tra

p N

ight

s

Bee

ch n

uts

colle

cted

Beech nuts Mice

Conflict during nut crop failure

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#

Adirondack Park

12

3

WildlifeManagementUnits

New York

Maine

New Brunswick

Consume • Small mammals• Nuts and berries• Birds, amphibians, etc.

• Impacted directlyand indirectly by beech masting

• Fur trappers’ data

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0

5

10

15

20

25

30

35

40

45

1988 1990 1992 1994 1996 1998 2000 2002 2004 2006 2008

Mea

n Sm

all M

amm

al C

atch

-per

-uni

t Eff

ort

(CPU

E; to

tal c

aptu

res/

100T

N)

~24,000 Trap-nights; 1992-2008Summer CPUE, all 8 species combined

-0.60

-0.40

-0.20

0.00

0.20

0.40

0.60

0.80

1.00

Cor

rela

tion

Coe

ffic

ent (

r)

American BeechSugar MapleConiferous Trees

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Marten/trapper Mean (SE)Mast Failure 1.82 (0.17)Intermediate Mast Crop 0.80 (0.14)Large Mast Crop 0.33 (0.04)

0.00

0.50

1.00

1.50

2.00

2.50

3.00

1988 1990 1992 1994 1996 1998 2000 2002 2004 2006 2008

Mar

ten

Har

vest

Rat

e (m

arte

ns tr

appe

d/m

arte

n tr

appe

r)

y = -0.000004x + 0.473r 2 = 0.84, P < 0.0001

-2.00

-1.50

-1.00

-0.50

0.00

0.50

1.00

1.50

0 50000 100000 150000 200000 250000 300000 350000 400000 450000 500000

American Beech Mast Production (sound seeds/ha)

Log(

Mar

ten

Harv

est R

ate)

(mar

tens

trap

ped/

mar

ten

trapp

er)

Harvest rates: strongly correlated with beech mast

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0.00

1.00

2.00

3.00

4.00

5.00

6.00

1988 1990 1992 1994 1996 1998 2000 2002 2004 2006 2008

Fish

er H

arve

st R

ate

(fish

ers

trapp

ed/s

ucce

ssfu

l fis

her t

rapp

er)

Central Adirondacks

Northern/SE NY

Harvest rates: weakly correlated with beech mast

y = -0.000003x + 4.194r 2 = 0.40, P = 0.008

0.00

1.00

2.00

3.00

4.00

5.00

6.00

0 50000 100000 150000 200000 250000 300000 350000 400000 450000 500000

American Beech Mast Production (sound seeds/ha)

Fish

er H

arve

st R

ate

(fish

ers

trapp

ed/s

ucce

ssfu

l fis

her t

rapp

er)

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0

500

1000

1500

2000

2500

3000

3500

1990 1992 1994 1996 1998 2000 2002 2004 2006

Tota

l Fis

her H

arve

st (M

aine

/New

Bru

nsw

ick)

0

100

200

300

400

500

600

700

800

Tota

l Fis

her H

arve

st

(New

Yor

k; C

entra

l Adi

rond

acks

)

Marten

0

500

1000

1500

2000

2500

3000

3500

1990 1992 1994 1996 1998 2000 2002 2004 2006

Tota

l Fis

her H

arve

st (M

aine

/New

Bru

nsw

ick)

0

100

200

300

400

500

600

700

800

Tota

l Fis

her H

arve

st

(New

Yor

k; C

entra

l Adi

rond

acks

)

Fisher

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• Cycles in beech mast production initiated immediate and time-lagged effects on mustelids, their small mammal prey, bear demography…

• Animals’ reliance on beech nuts affected by alternate food sources (also competition, habitat quality…)

• Synchronicity in space and time – environmental factors

• BBD/nut model: correct for single trees; wrong on landscape scale

• Forest continues to change with BBD

• The cascading effects of an invasive disease complex can negatively affect the diversity of non-host species

• Monitoring mast availability can be used to predict animal populations and periods of human-wildlife conflict and to inform harvest regulations

• Trophic interactions are affected by the beech mast cycle, e.g.• Herbivore-carnivore relationships• Interguild relationships: fewer beech nuts predation of bird nests by mammals

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• Dominant sugar maple, yellow birch, white ash (& beech) being replaced by a less-diverse forest of degraded beech

• Soils are high-quality (Skerry, Becket series)

• Is mechanical control possible and feasible?

• Can desirable species be retained and promoted?

• At what price?

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Commercial beech control

Study treatment impacts on • Forest structure and composition• Vascular plant/epiphyte diversity• Beech seed production by tree• Microclimate • Wildlife taxa

• Bats• Small mammals• Songbirds

Caroline Byrne

The project is designed to meet the challenge of retaining and encouraging a desirable mix of forest species and structure for both ecosystem function and economic value.

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Mammal GridsTreatment, 1971Treatment, 1952Uncut ControlALTEMP Control

• Beech growth, mortality and reproduction• Management ramifications

• Which beech can be retained to produce nuts/wildlife value

• How long might a beech tree survive given its BBD rating

• Integrate management objectives

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Paul Jensen, DEC Region 5Walter Jakubas, Maine Dept. Inland Fisheries and Wildlife

Charlotte Demers and Ray Masters, SUNY ESF Adirondack Ecological Center

Greg McGee, René Germain, Shannon Farrell, and Ralph Nyland, SUNY ESF

Paul Jensen, DEC Region 5Walter Jakubas, Maine Dept. Inland Fisheries and Wildlife

Charlotte Demers and Ray Masters, SUNY ESF Adirondack Ecological Center

Greg McGee, René Germain, Shannon Farrell, and Ralph Nyland, SUNY ESF

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Cale, J., S. McNulty, S. Teale and J. Castello. 2013. The impact of beech thickets on northern hardwood forest biodiversity. Biological Invasions 15:699-706.

Costello, C. 1992. Black bear habitat ecology in the central Adirondacks as related to food abundance and forest management. M.S. Thesis. State University of New York, College of Environmental Science and Forestry, Syracuse, 165 pp.

Elowe, K.D. and W.E. Dodge. 1989 Factors affecting black bear reproductive success and cub survival. Journal of Wildlife Management 53:962-968.

Hane, E.N. 2003. Indirect effects of beech bark disease on sugar maple seedling survival. Canadian Journal of Forest Research 33:807-813.Jakubas, W. J., C. R. McLaughlin, P. G. Jensen, and S. A. McNulty. 2005. Alternate year beechnut production and its influence on bear and

marten populations. Pages 79-87 in Evans, C.A., J.A. Lucas, and M.J. Twery, eds. Beech Bark Disease: Proceedings of the Beech Bark Disease Symposium. General Technical Report NE-331. Newtown Square, PA. USDA Forest Service, Northern Research Station. 149pp.

Jensen, P.G., C.L. Demers, S.A. McNulty, W. Jakubas, and M.M. Humphries. 2012. Marten and fisher responses to fluctuations in prey populations and mast crops in the northern hardwood forest. Journal of Wildlife Management 76:489-502. DOI: 10.1002/jwmg.322

LaMere, C. R., S. A. McNulty and J. E. Hurst. 2013. Human-black bear conflicts are related to mast production in the Adirondack Mountains of New York State. Proceedings of the Eastern Black Bear Workshop 2011.

McNulty, S. A. and R. D. Masters. 2005. Changes to the Adirondack forest: Implications of beech bark disease on forest structure and seed production. Pages 52-57 in Evans, C.A., J.A. Lucas, and M.J. Twery, eds. Beech Bark Disease: Proceedings of the Beech Bark Disease Symposium. General Technical Report NE-331. Newtown Square, PA. USDA Forest Service, Northern Research Station. 149pp.

Mott, E. 1901. The Extinct Passenger Pigeon.The New York Times Published August 18 1901.Thompson, J. R., D. N. Carpenter, C. V. Cogbill, and D. R. Foster. 2013. Four Centuries of Change in Northeastern United States Forests. B.

Bond-Lamberty, editor. PLoS ONE 8:e72540.

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