Beaver as a modifier of

boreal riparian
ecosystems
Tartu 21.11.2022

Petri Nummi
Department of Forest Sciences
University of Helsinki

http://wetlandecologygroup.it.helsinki.fi/
 Beaver concepts
◼ Keystone species
◼ Ecosystem engineer
◼ Facilitator
◼ Disturbance agent
◼ Herbivore
Kolkku
 I Beaver – ecosystem engineer,
keystone species & facilitator
◼ keystone species strongly affect other
species
◼ ecosystem engineers modify the pathways
of organic matter and energy
◼ facilitators benefit other species
II Beaver – disturbance agent &
herbivore

◼ Disturbance agents affect the succession

of e.g. forests
◼ As a herbivore beaver peculiar since it can
cut large trees (elephants!)
 Keystone species

◼ ”All
animals are equal but some
are more equal than the others”
(Orwell 1945)
 Vääriselupaik (võtmebiotoop, key biotope)

Võtmebiotoop on paik kaitse-või tulundusmetsas, kus

tõenäosus ohustatud, ohualdiste või haruldaste liikide
esinemiseks on suur nagu väikeste veekogude ja allikate
lähiümbrus. Väikesed lodud, põlendikud ja soosaared, liigirikkad
metsalagendikud, metsa kasvanud kunagised aiad,
metsaservad, astangud
A key biotope is a location in a protection or
commercial forest where the probability of
occurrence of endangered, vulnerable or rare species
is high, such as in the immediate vicinity of small
water bodies and springs. Small marshes, burned
areas and bog islands, species-rich open areas in the
forest, former gardens grown into forest, forest
edges, terraces
 Vääriselupaik
(võtmebiotoop)
ALAD, MILLE ELUSTIK JA
KESKKONNATINGIMUSED OMAVAD
SUURT MÕJU ÜMBRITSEVA, HÄIRITUD
KOOSLUSE TAASTUMISEKS

Areas of which biological and environmental

conditions have a significant impact on the recovery
of the surrounding, disturbed community
 Metsade bioloogiline mitmekesisus

◼ Maal elab hinnanguliselt üle 13 miljoni liigi

◼ Eestis teatakse elavat umbes 23 000 liiki (tegelik 40 000)
◼ Eesti metsades ligikaudu 20 000 hulkrakset (multicellular)
liiki
◼ selgrootuid (invertebrates) üle 10 000 (valdavalt
putukad)
◼ seeni 2500 liiki

◼ samblikke 500

◼ soontaimi üle 450 (vascular plants)

◼ selgroogsed loomad 150 liiki

 Keystone species

◼ ”All
animals are equal but some
are more equal than the others”
(Orwell 1945)
 Keystone species
◼ ”A species whose loss is likely to trigger a
siginificant number of secondary
extinctions”.
◼ Ebenman & Jonsson 2005. Using community viability
analysis to identify fragile systems and keystone species.
TREE 20: 568-575.

◼ Keystone species not easily identified

 Kinds of keystone species?

◼ Top predators (sea otter, Paine 1974)

◼ Predator removal is mediated downwards in
the food chain (top down regulation):
◼ Sea urchins increased so that algae and
associated species disappear
◼ Keystone species also among other
ecological groups: prey, pollinators,
ecological engineers…
◼ Keystone species
- extirpation may affect
many parts of the
food chain:
- predators
- consumers
producers

Ebenman & Jonsson 2005 TREE

-

◼ Important to idenfy
in conservation

(a) Loss of a top predator (collapse of kelp forest communities following the loss of sea
otters). (b) Loss of a consumer leading to the breakdown of consumer-mediated coexistence
among prey (extinctions of algal species in marine intertidal communities following the loss of the
herbivorous periwinkle snail). (c) Loss of a primary producer triggering a bottom-up extinction
cascade.
 Ecosystem engineers

◼ …organisms that directly or indirectly

modulate the availability of resources to
other species,
◼ by causing physical state changes in biotic
or abiotic materials.
◼ (Jones et al. 1994)
Autogenic
engineers
change environment
via their own
physical stuctures
= e.g. trees (Case 3)

Allogenic
engineers
change environment
by transforming
material to another
physical state
= beaver (Case 4)

(Jones et al. 1994)

Engineering: material flow and heat transfer
Gutierrrez & Jones 2006
 Ecological facilitation

◼ For long, ecologists have worked on

negative interactions: competition,
predation…
◼ Recent research indicates that positive
interactions are at least as important in
shaping communities
 The realized niche (green
circle) often can be smaller
than the spatial range
predicted by the fundamental
niche (dashed line) (ai).

Incorporating facilitation into

the niche concept (aii)
recognizes processes that
can expand the amount of
space that meets the
requirements of the
fundamental niche and can
mitigate the effects of niche-
shrinking factors.

Bruno et al. 2003 TREE

2006
Models of alternate system states. Current state (red circles),
desired state (green squares). Green area = conditions that
lead to the restoration goal.

(bi) No ecosystem engineer added to

the system

(bii) restoration-facilitating ecosystem

engineer added to the system

(biii) restoration-inhibiting ecosystem

engineer added to the system.

Solid arrows = human intervention on biotic and/or abiotic properties,

Dashed arrows = post-intervention, natural readjustment of the system
 Models of alternate system states. Current state (red circles), desired state
(green squares). Green area = conditions that leads to the restoration goal.

Restoration target is achieved with:

(bi) considerable intervention with no

ecosystem engineer;

(bii) the least human effort with a desirable

ecosystem engineer;

(biii) not achieved for the illustrated effort

with an undesirable ecosystem engineer
present.

Solid arrows = human intervention on biotic and/or abiotic properties,

Dashed arrows = post-intervention, natural readjustment of the system
Beaver in restoration
 Beaver effects

◼ beavers have been absent from their

areas for many centuries
◼ wetlands have been studied in an
”unnatural” state
 Beaver distribution in Europe

- Majavien reliktipopulaatioita A, B, C ja Dneprin ja Donin alueella, vain 1000!

- nykyään taas vähintään 600 000
 Beaver effects
◼ beaver changes the structure of the
shore zone
◼ beaver returns forest succession to
an early stage ->
◼ disturbance
 Beaver disturbance

◼ resembles forest fire and storm – but

more predictable
◼ beaver flooding turns a shore meadow to
an aquatic ecosystem
◼ after beaver abandonment, a terrestrial
succession begins
Nummi & Kuuluvainen 2013 BER
Nummi & Kuuluvainen 2013 BER
 Beaver effects
◼ beaver affects habitats at the level of:
◼ 1) patches and

◼ 2) landscape
A three-dimensional beaver patch

Johnston & Naiman 1987

 Changes in element flow

◼ 10 m3 dam holds 5000 m3 sediment ->

◼ Within 320 km of rivers, 3.2 milj. m3 of
sediment
◼ amount of e.g. carbon 12 kg/m2 in beaver
ponds, 4 kg/m2 in the riffle
◼ Methane emission
 Beaver and carbon

◼ Short and long term processes

◼ DOC (dissolved organic carbon)
◼ CH4 (methane)
◼ Organic sediments
Beaver flood and DOC in lakes

◼ Carbon firstly
increases in the water
of the flooded pond
◼ Later on decreases
◼ But not in
downstream ponds
◼ Where does it go?

Vehkaoja et al. Submitted

 Beaver and greenhouse gases
”Normal boreal
ponds:

~ 5 mmol m2/day

New beaver pond

Lazar et al.2014 JEQ

 Long-term beaver deposits

Persico & Meyer 2013

Older deposits partly removed by erosion (Yellowstone
 Beaver herbivory

◼ beaver browsing modifies forest structure

even above the flood line
◼ a beaver family may cut over 1000 kg
trees per hectare
◼ a biomass effect comparable of that of a
antilope herd in Serengeti savanna
Beaver flooding and succession Knudsen 1962
Succession paths in beaver patches
Evo experimental flowage before
flood, 1984
Evo flowage, 1986
Evo flowage, 2002
 Tree survival

◼ Salix survives best

◼ Adventitious roots

Nummi1989
◼ More biomass in
leaf litter
◼ Effect on litter
based production
 Plant succession in
beaver ponds
30
Emergent
Floating and submerged
25 Trees and bushes

Number of species
20

-vegetation lines 15

10

0
1985 1990 1995
Year

Emergent
Floating and submerged
Trees and bushes

100

Proportion (%) of total abundance

80

60

40

20

0
1985 1989 1995

Hyvönen & Nummi 2011, Return of the beaver Year

 Beaver flooding and trees

Havupuut CONIFEROUS
Lehtipuut DECIDUOUS
100

80
Importance value (%)

60

40

20

0
D L D L D L D L D L D L
Pond I Pond II Pond III Pond IV Pond V Pond VI

Importance values of dead (D) and living (L)

coniferous and deciduous trees in Evo Hyvönen & Nummi 2008 WB
Tree density and
height after beaver
- more saplings of
deciduous trees

- deciduous trees taller

2 years old beaver meadow
5 years old beaver meadow

Terho Hyvönen
Beaver dynamics in Evo
Beaver landscape in Evo in 1998
◼ Shannon Diversity Index
(flood history diversity)

◼ Nearest Neighbour Index

(clustering, random,
dispersion)

◼ Connectivity index
(exponential of distances
between beaver sites)

◼ Site abundance within

different buffer zones
(500, 1000, 1500 and
2000m wide)
 BEAVER
LANDSCAPE
- flowages in
Kabetogama-
peninsula in 1940
& 1986

Naiman et al 1988 BioScience

 Beaver in Kabetogama
◼ 1940:
◼ 71 dams
◼ less than 1% of the peninsula flooded

◼ 1986:
◼ 835 dams (2.5/creek km)
◼ 13 % of the peninsula flooded
◼ another 13 % changed by browsing
◼ water habitats increased by 440 %
Beaver and biodiversity

Sakari
 Effect of beaver on plant species
number
◼ on patch level no
difference between
beaver and other
riparian zones
◼ on a landscape level
beaver increases
species number of
plants

Wright ym. 2002

 Effects of beaver flooding on
animal life 1/3
◼ flooding affects many animals since:
◼ nutrients are released -> more food
◼ shallow shores are created -> amelioration
of habitat structure

◼ after beaver herbivores benefit from the

bushy beaver meadow
Nummi & Kattainen 2006, Suomen Riista 52
 Teals in boreal lakes

Nummi et al. 2005 SR 51

 6

5 Teal - Beaver pond

4
3

Teal pairs
1

◼
Change in pair numbers

increase after
-1

-2
1 2 3 4 5 6 7 8 9 10 11 12 13

beaver
-3

6 damming
5 Teal - control
4

1
0

-1

-2
-3 1 2 3 4 5 6 7 8 9 10 11 12 13

Nummi & Pöysä 1997, Ecography

 Beaver activity and ice melting:

- open water earlier in

active beaver ponds

- open water present

adjacent to the lodge
entrance and over the
food cache

Bromley & Hood 2013 Mamm Biol

 Beaver and geese
Access to open water earlier in the season might allow for:
- earlier nesting times
- earlier access to aquatic vegetation and invertebrates
- longer maturation times

Percent of active and inactive beaver lodges with breeding Canada geese
◼ Teal broods increase
after damming

Nummi & Hahtola ,

2008, Ecography
Broods in beaver flowages
% 70

40
60
Observed and expected use

50 16 242

45
I
40
II
III
30
20 20 Juv
74
10
20 65 143

185 43
10

0
Mallard Teal Goldeneye

Nummi & Pöysä 1995

 Relative influence of habitat, pair and weather covariates in
explaining number of Teal age class II broods during 1989-2008.

Holopainen et al. 2014 Freshwater Biol Beaver flood

Pictures by Petri Timonen
 Invertebrates caught
in activity traps

Pictures by Petri Timonen

 Invertebrates in beaver ponds
400

350

300
n = 12
Invertebrate index

250

200

150

100

n = 28
50 Mann-Whitney U-test,
0
p < 0.000
Beaver flowages Undisturbed ponds

Nummi & Hahtola 2008

 Teal brood density
and invertebrates

1,20

1,00
Broods / shoreline km

0,80

0,60

0,40

0,20

0,00
black = beaver
0 100 200 300 400 500 600 700 800 900 grey = other
Invertebrate abundance
Habitat amelioration
 Duckling survival in beaver ponds

0,04 - mortality lower

in beaver ponds
0,03 (Mann-Whitney, P 0.001)
Mortality rate

0,02

0,01

0
Beaver Non-beaver

Nummi & Hahtola 2008

 Bats and beaver

http://www.medicine.uiowa.edu/CIgW/image_cartoon/beaver.gif
 Bat food

◼ Bat food consists mainly of insects

◼ Insects emerging from the water abundant:
◼ chironomids, trichopterans

Henry Ekholm
 Bat habitats

◼ Two third of Europes wetlands lost during

the last 100 years
◼ In England only 1 % of bat habitats good
wetlands
◼ How to get more wetlands ->
◼ Beaver!?
 Bats in beaver ponds

Number of detections

80
70
beaverpond
60
Batobservations

control
50
40
30
20
10
0
1 2 3 4 5 6 7 8 9 10 11 12 13

Nummi et al. 2011, Biodiv & Cons.

Bats and emerging insects

Nummi ym. 2011, Biodiv Cons

Whole community facilitation

Risto Salovaara/Yle
 Frog community and beaver

Bufo bufo
Rana temporaria
Rana arvalis
Croak index

Beaver ponds Lakes Seasonal ponds Vehkaoja & Nummi, Herpetozoa 2015
Engineering and species richness
 Beaver and waterbird richness

◼ Species number increases

during the first two years
after flooding

◼ Stays on a higher level

continuously
Beaver and waterbird abundance
Beaver ecosystem services
Beaver and waterflow

Buckley et al. 2011

◼ Beaver dams can hold up to 30-60 % of
the base flow and discharge it later (Kay
1994)
 Thank you!

http://wetlandecologygroup.it.helsinki.fi/