Dr. L.

David Mech
Wolf Biologist

“A 50-year Career of Wolf Research”

Friday, April 17, 2009
4:00 pm
Marshall Auditorium
Reception to follow

Dr. Mech has studied wolves in Yellowstone Nat’l Park, Minnesota, Isle Royale, Alaska
and Canada's High Arctic (80°N). These photos are of High Arctic wolves.
 here Summary
• Habitat MAKES a difference!
• Non-lethal effects may be more important to
wildlife management than lethal ones.
• May be able to manage impact of predation
via habitat.
• Landscape of fear/opportunity may be the
most valuable management tool in
management AND conservation.
• Example?? Go to sheep presentation
 So what do we have?
• Habitat…Habitat….Habitat

• Brings us back to the beginning: wildlife

management is habitat management.
• Not just measure landscape of fear, modify
it!
• All we talked about, food, reproduction,
predation, population dynamics, energy
flow…. All rooted in habitat.
• Habitat is the medium in which energy is
captured…or not.

• So ecology is the play, habitat is the stage.

 Wildlife management
• Ok, covered the ecology part of the
course, lets look at the management part.

• First we will look at how we have been

managing wildlife

• Last we will look at how we should

manage wildlife!
 Chapters 13 and 14
• Deal with more practical aspects: counting
animals (13)
• Age structure (14)

• Mostly techniques stuff which game

agencies spend a lot of time doing!
• Annual herd counts, flock censuses, etc.
• If we have time, we will look into but…
 Goal of wildlife management
• Reasons for these is to aid in the REAL
goal of wildlife management: how many
can we kill! (wildlife harvesting, chap 19)
• The harvest
• The off take
• The whatever you want to call it,
• Human predation on selected wildlife
species.
 Second goal of wildlife
management
• Aside from trying to put more game in the
bag and more fish in the creel.
• Game agencies have a second goal of:
• Keeping other species from getting OUR
game!
• “wildlife control” (chap 20)
 Management in a nutshell
• These two goals: more for us, less for
them.

• Have and continue to be the center focus

of wildlife management in the U.S.

• So we need to look at them and end up by

asking if that is how it should be???
 But first
• Need to put current wildlife management
in its context.
• If we start from a point in time,
theoretically can go in an infinite number
of different directions.
• Once wildlife management got started,
why did it take the direction it did?
• A little bit of history
 A bit of history
• Where does the basis for wildlife
management come from?

• When Europeans arrived, came to a

continent “teeming” with wildlife.
• Deer, waterfowl, turkeys, abundant and
viewed as having no end.
 In less than 300 years
• First settlements in North America were
around the 1600’s
• In less than 300 years (late1800’s) our
ancestors had devastated these “endless”
wildlife populations.
• The list of overuse and abuse is long and
legendary, and in some cases is still
growing.
 Low points
• Eastern region had rid itself of all large
game except deer, and even these were
nearly eliminated from many areas
• The Midwest became the farm belt at the
expense of wildlife numbers rivaling the
plains of Africa
• The western regions reduced and eliminated
many wildlife populations: At one time in the
state of Colorado there were 0.00000 deer!
 But was this new??
• But even so, what continent had was a
shadow of what existed before.

• Mammoths, ground sloths, BIG animals

and their predators.
• One theory as to why disappeared was
when other group of humans came.
• Pleistocene overkill
 Imitating first immigrants
• The new immigrants were just doing what
the first ones had done, only more
effectively because of advanced
technology.
• Impact of earlier Americans limited by their
technology
• Now: guns vs arrows, horses vs feet
• Steel saws vs stone hatchets, and the
technology kept on growing!
 Compounding factors
• In Europe, socially, there existed few
landowners: kings, lords, etc.
• Owned the land AND the wildlife
• Wildlife in a large area were private property
of ONE person.
• Predation rate low (illegal killing severely
punished)
• Game keepers cared for the wildlife (that the
owner wanted).
 Majority of the people
• Had no use for wildlife because they could
not benefit from it
• Viewed as one of those privileges of the
rich.
• Now all of a sudden 100,000’s came to a
land were wildlife was free for the taking!
 A continental killing spree.
• European immigrants, raised in a social
environment where game were owned by
the large landowners, responded like
turning a wolf loose in a sheep pen!
• It became our “right” to use wildlife as we
saw fit.
• Commercial hunters had the same social
status as commercial fishermen still have
today.
 Double barreled
• So added to an individual’s right and need to
kill wildlife
• The commercialization of hunting added the
free enterprise aspect, except with a twist.
• If supply was low and demand high, kill more
because you will get more and they will then
be worth more.
• If supply was high and demand and price
low, kill more to make up for the low price
 And besides…
• Wildlife populations were “endless”!
 What was to be done?
• Some began to realize that the killing
could not continue. That some control had
to be exerted.
• Wildlife management was born as an
attempt to control the slaughter.
• Earliest “game laws” in the late 1600’s and
1700’s were attempts to limit our
effectiveness as a predator.
Was that wildlife management?
• One can argue that these early attempts at
controlling the kill were not actually wildlife
management as a collective effort to
“manage” wildlife and certainly not a
science!
• Many argue the science of wildlife
management got started in the 1920’s and
30’s
 Aldo Leopold
• argued that modern science and
technology could be used to restore and
improve wildlife habitat and thus produce
abundant "crops" of ducks, deer, and other
valued wild animals.
But what model should we use?
• If your going to start something like wildlife
management, usually creativity of ideas is
limited by historical and cultural confines.
• Trained in forestry (tree farming)
• So with this the mold was struck: Wildlife
were to be managed as livestock.
 Talk the talk
• The livestock model can be seen in most
aspects of how we talk about wildlife.
• Herd, flock
• Harvestable surplus: Harvest!!
• Sustainable yield
• Carrying capacity.
• Talking to a wildlife manager is like talking
to a cattle rancher.
 Set in motion
• Within this paradigm we began to try and
determine how many animals can we send
to slaughter each year and still maintain the
herd.
• Little more difficult because:
• 1) often don’t know how large the herd is.
• 2) have limited control over the harvest.
• Reason is because every hunter is a
rancher!
 But that is how it is, still…
• So, you as potential managers need to be
exposed to all this.

• Will go through chapter 19 and pull out the

information necessary to understand how
wildlife managers attempt to control the
harvest of wildlife species.
 Central theme
• Thru all of this runs one central theme
regarding wildlife harvesting
• Harvesting of wildlife must result in a
sustainable yield that can be taken year
after year without jeopardizing future
yields.
• Again, sound livestock ranching but true!
 Idea of a sustainable yield
• Strategy to achieve a sustainable yield is:
• To harvest the population at the same rate
as it can increase!
• Example: If population is increasing by 20%
per year, then you should be able to harvest
around 20%/year!
• This is a net rate of increase: after all
factors figured in (e.g.various other mortality
factors).
 How do we achieve it?
• Obviously sustainable yield levels are
affected by factors not under direct control
e.g. resource levels
• So most important aspect (controllable) is
the regulatory strategy (legal limits) used
to set harvest levels.
• Or basically, how do we actually control
the harvest?
 Fixed Quota harvest
• Obviously various techniques can be used.
• We will review them to see the philosophy
and logistics behind them.

• The first is the Fixed Quota Harvest

strategy.
• Will look at this one in detail to get feel for
how they are developed
 Fixed quota
• Most unharvested populations have a rate
of increase that averages zero!
• So what is the sustainable yield?
• The philosophy behind the fixed quota
harvest is:
• If you harvest a given number of individuals
(fixed number) then you stimulate that
population to increase by setting into
motion a chain of events.
 How is it done?
• Based on the idea that IF you increase
resource levels, a population will grow.
• More food = more animals
• The reason for this is because there are
more of the resource PER animal.
• So the key is increasing the amount of
the resource per animal.
 How is it done…cont.
• Ok, so if increasing the level of resource
increases per animal amount…..
• Keeping resource level the same but
DECREASING the number of animals will
give the same result!
• By removing individuals (the harvest), more
food to go around, more food, greater
fecundity/lower juvenile mortality and
population increases back toward original
level!!
 But how many should we
remove??
• In general the further the density is
reduced, the higher the yield as a
percentage of the population size.

• That is, If you remove 50 % of the

population and it “grows” back to original
level, your yield would be higher than if
you only removed 25%.
 But is there a limit?
• IF you do remove 50%, WILL it recover??
• Just how many can you remove?
• Based on intrinsic rate of increase of the
species.
• Based on maximum fecundity, maximum
survival, etc.
• This is based on maximum resource
levels!
here
Induced rate of increase
• So intrinsic rate of increase is based on
maximum resource levels, fecundity, etc.,
which only occurs when population is at a
minimum!
• Any N greater than that, rate of increase
decreases, eventually 0 at K
• So any reduction we do, will result in an
induced rate of increase < intrinsic
 Some declining function
• Induced rate of increase will be close to
intrinsic at a minimum of N but decline as
N increases.

• So higher N, lower induced rate of

increase.
• Based on that, we need to determine what
is the sustainable level we can harvest.
 How do we calculate it?
• First remember: we are attempting to
imitate how a population would respond
“naturally” without harvest
• The idea is that below K, there is a natural
induced rate of increase that will lead to a
net recruitment of new individuals as
population moves toward K.
 Harvest replaces recruitment
• So, IF population is reduced “naturally” to
some level, recruitment then replaces this
lose, if we harvest the same number of
animals, we reduce N and expect
recruitment to replace these animals.
• Trying to mimic recruitment under natural
conditions.
• Harvest can be what will be replaced by
this recruitment.
 But how much recruitment will
there be?
• Since induced rate of increase is density
dependent, the recruitment level will vary
across densities.
• At any given density it will be the result of
the combination of induced rate of
increase and density.
 Bell curve
• This can be envisioned as a bell shaped
curve over the range of density up to K
• At low N, low recruitment because of low
numbers of reproducers
• At high N, close to K so little recruitment
because of environmental resistance.
• In the middle somewhere it will peak.
Net recruitment

N K
 Now change recruitment to
harvest
• IF we harvest back the population to a
certain level, should expect a response
similar to what we would predict naturally.
• Sustainable yield any point along that line.

Net recruitment
or
harvest

N K
 Rule of thumb
• Best sustainable yields will be at intermediate
densities, highest recruitment rates.
• Example
• Small harvest of large population, induced rate
of increase small (large x small) 12 x 3 = 24
• Large harvest of population, induced rate of
increase large but remaining population small
(small x large) 3 x 12 =24
• Intermediate: 6 x 8 = 64
 So…How do we calculate it?
• Maximization problem:
• Trying to maximize the absolute yield
• Highest yield is at a density where the
induced rate of increase multiplied by the
density is maximum.
Looking for ideal sustained yield
• If we set harvest at a given level (20) note
that it intersects curve in two places.
• Called sustained-yield pair.
 So naturally
• Naturally, a recruitment rate of 20 would
be replaced either if the population had
been reduced to 80 OR 20.
• Means we could harvest back to 20 if we
want??
 Not necessarily
• To harvest these 20 is harder at lower
density
• Also it is unstable and any reduction below
the lower density would lead to
overharvest and population decline
because recruitment is declining.
• The upper population level is stable
because any decrease in density results in
increasing recruitement.
 But is this the maximum??
•How high should we push it?
•Maximum would be a little above 30.
•This is the maximum sustained yield.
•It is unstable because any reduction in N
will result in
declines.
Usually avoided.
 But….
• The major assumption of the fixed quota
harvest model is that there is no stochastic
variation in factors that influence net
recruitment.
• Means no uncontrollable changes in
population density due to weather, etc.
• Cows in a pen, which wildlife populations
are not!
 Results?
• Basically, can be shown that eventually,
you will overharvest the population and it
will decline, possibly to extinction!
• Gave you this model to show you the
rational used to manage wildlife!
 Better models?
• Fixed quota harvest is not good in most
cases, are there others?
• Fixed proportion harvest strategy
• How does it differ? Still fixed but now it
uses proportions.
• So number harvested changes depending
on density.
 Makes sense, kind of…
• Thus, if population declines for stochastic
reasons, percent taken is the same but the
absolute number is less.
• Result should be a changeable but
sustainable yield with a stable population
density even in the presence of stochastic
variation
• Sounds good!
 But…
• This requires managers to have perfect
information on abundance of the species to
set the quotas.
• Rarely do they have this information and
even if they do, usually not in time to set
quotas.
• Usually they base it on past success!
• Needless to say, they are often wrong and
overharvesting is possible.
 What else?
• Constant effort harvesting strategy
• Instead of controlling the harvest, this
attempts to control the harvesting effort.
• This is done via setting hunting seasons or
limiting the number of people harvesting
the population.
 How does this differ?
• So now instead of saying you can hunt
until you remove x or a proportion of N,
• You say you can only hunt x number of
days or only so many hunters can hunt the
area (usually also over a fixed time).
• This has a built in control mechanism in
that the number of animals killed over the
fixed time will be dependent on
abundance.
 High density/high success
• If population is low, hunter success will be
low and the total harvest will be low.
• If population is high, just the reverse.
• So it is somewhat self regulating and IF
you set the season length or the number
of hunters for the lower population levels,
should avoid overharvesting
 Others?
• Fixed escapement harvesting strategy
• This is based on the idea of maintaining a
population at a given level and only if it
goes over that level, can you harvest
individuals.
• Excess individuals above this target
threshold are termed escapement.
• Insures a minimum level of population
 Is it used?
• Fairly new and is not favored by managers
because of the variation in the harvest,
including no harvest in low years.
• But it begins to recognize that game
species are not just to shoot and actually
are “used” by others in non-consumptive
ways.
 Do they work?
• These are the harvest models and look
good on paper (maybe) but do they work
in the real world?
• Needless to say, it is hard to strictly apply
a mathematical model to real world, real
people situations.
• “Most harvesting of wildlife… has been
managed largely by trial and error”.
 Seat of the pants
• So taking in consideration all the fancy
models and math, we still control the
harvest based on “what we feel is right”!
• Usually managers have been conservative
so as to not overharvest but sometimes
mistakes are made, still can’t predict what
populations will be like
 Additional harvest technique
• Age or sex biased harvesting.
• So far have talked about removing
whatever animal from the population.
• This is common in species where sexes
and age are hard to determine.
• Grouse/woodcocks (sex nor age)
• Ducks (sex yes but age no)
 If you can tell the difference
• Most ungulate species can tell sex and
age (at least in males)
• When this is possible, many argue should
protect reproductive base (females) and
reproductive output (young).
• This leads to adult males only seasons.
• Is this desirable?
 Should we kill only the males?
• Population wise most expendable
• But… can lead to reduced hunting
opportunities.
• Example: start out with 100 animals
• 50 females/50 males.
• If we just kill of males, can lead to skewed
sex ratio where more of the population is
females and less room for males!
• IF area can only support given number of
animals AND you want to manage for
hunting opportunities male only harvest
can lead to reductions in these
opportunities.
• Strategy is to maintain females at an
adequate level for maximum production
but low enough so that all the energy is
not tied up in them.
 Having enough males
• Danger of producing skewed sex ratios is
not having enough males to go around!
 Within males
• In New York, recently have seen the use
of age harvest criteria within males (deer).
• Point limits, can only shoot bucks with
greater than x number of points.
• Idea is IF we consistently kill off the young
males (which often happens because they
are more numerous), they never get a
chance to grow into big ones!, which are
more desirable to many.
 Pros and cons
• Obviously aimed at “quality” of harvest.
• Makes sense IF this is your goal.
• Some argue that by doing so you are
killing off the good genes (tested big
bucks) and allowing inferior genes
(untested) ones replace them.
• Again cattle: do you send your biggest bull
to market??
 So arguments go on
• Sex and age biased harvests have always
been and will continue to be controversial.

• When I was in Michigan and they wanted

to start a doe season, accused of killing
bambi’s mom!
• Of course you could shoot good old dad!
Here Politics and harvest
management
• This brings us to probably the most
influential harvest strategy:
• Politics!
• As mentioned, game agencies may
manage wildlife but the public are the
owners!
• So a wildlife biologist may say all he wants
about what is the best way to manage the
harvest, but it is the public that decides!
 Political structure
• This public input is insured by the structure of
most game agencies.
• There can be an agency director but above
that person is the game commission.
• Commissions are noted for not being wildlife
people, except that sometimes they may be
hunters (but it is not necessary). They often
are political appointees.
 The process
• Field biologists make recommendations to
their superiors.
• The superiors get together to propose
policy.
• That policy is the adapted or not by the
commissioners, often after they get input
from Joe the hunter.
 Joe the hunter
• Joe the hunter bases his opinion on how
well he did last year!
• If he did good but the “experts” are
proposing decreases in the harvest, they
must be wrong, and visa versa!
• So we can have all the models we want but
when it comes to harvest strategies, the
final decisions are made by politicians who
know little or nothing about wildlife ecology.
 Summary
• How do we manage the harvest?
• Have models based on:
• 1) Agricultural heritage
• 2) mass action, no behavior of “prey”
• 3) biggest weakness, based on constant
populations.
 Summary
• Rarely work in real world so often
managers do what they think is best or
• What they are politically told to do.
• Last important aspect: They are by nature
single species concentrated.
• This we will address later.
 Next: wildlife control (Chap 20)
• So, managing how many we can kill is
difficult.
• BUT… if we can keep other species from
killing them, we should be able to kill more
our self…right??
 Manage vs control
• How do they differ?
• In control we desire to purposely REDUCE
a population to some lower level but not
for some sustainable harvest level.
• In fact, most controlled species are not
considered “harvestable”
• Harvestable meaning a species we kill for
its use.
 Control
• In control we are killing them because we
don’t want them!
• So they are rarely used and discarded.

• Laws against “wasting” game species

• But ok to discard 1,000s of kg of controlled
animals!
Why would we want to control?
• Before we go further, lets look at reasons
we would want to control a species.
• Basic reason is competition:
• Can be in two forms.
• 1) one species is competing for resources
of a desirable species.
• If we can control the first, leaves more
resources for the desirable one, more
harvest!
 Reasons for control
• 2) A species is competing directly for a
resource desirable by us.
• Can be plant foods: they are eating our
apples
• Can be for animal food; they are killing our
animals (domestic or wild).
 Need for control?
• So for one reason or another, WE decide
there are too many of a given species.
• Often we make that decision capriciously
without any sound basis, faith based!
• E.g. Eradication programs for predators
• How should we make those decisions?
 When is control appropriate?
• Some may argue never, we are imposing
our view on the natural systems!
• Some, who view the natural world as OUR
garden and anything we can’t use are
“weeds” needing to be controlled!
• Most see needs for control under certain
circumstances
• E.g. introduced species, cats on islands
(or is that the real case?)
 Circumstances for control
• 1) when benefits exceed the costs
• 2) when “pest” species is in fact the cause
of the perceived problem
• 3) when control has an acceptable impact
on non-target species.
 Cost/benefits
• A control action is undertaken to provide
some benefit (should be real and not
perceived!).
• Conducting the control action has costs
(monetary/environmental).
 Example: coyotes
• 2007 killed 90,326 coyotes
• Congress allocates $10M per year to kill
100,000 predators: mostly coyotes!
• Reasons: kill domestic stock, kill game
species.
• 270,000 sheep/year (4% of total sheep
population)
• ?? Game animals/year
 Perceived benefit
• Less coyotes/ more sheep

• Less coyotes/more game

 Is it worth it?
• It costs over a $100 for each animal that is
killed while the damages incurred by the
animal average $22.50
• Impact on game species debatable
• Other costs:
• 1 coyote eats 2,000 to 3,000 rodents/year
• 90,326 = 180 million/year!
• Rodents eat grass/plants that livestock
eat!
 Benefits of coyotes!
• Jackrabbits: 100/year/coyote
• 90,326 x 100 = 9 million jackrabbits!

• Arguments made both ways, not sure

which is the case because lack of studies!
 Is the coyote responsible?
• Livestock: 273,000 sheep but likely
inflated in that often, if a sheep does not
return, assumed it was killed by predator:
coyote!
• Again, still NO study showing definite
negative impact on wildlife populations.
 Impact on other species?
• Often use poisons to kill coyotes AND
other species.
• Coyotes are part of ecosystem. What
impacts are there upon its removal?
 Is it ever advisable?
• So when is control advisable?
• Exotics your best bet
• Natural systems???
 Why exotics?
• Often times easy to demonstrate they are
the cause of the problem: but how?
• Cats on islands:
• Introduced to control house mice from
ships (other exotic!)
• Killed mice but also ground nesting birds.
• How do we show it is the cause?
 Controls
• In any effort to demonstrate effect, need
controls.
• In this case, had islands where cats were
not present.
• Removed cats and saw increase breeding
success – 100% and decrease in chick
mortality – 0 %!
• Was it from cats? Comparison with cat free
island supported it.
 So it can be done
• But it is rarely so.
• Why?
• Objectives of control often poorly defined
or nonexistent!
• In any control effort, control in itself
should NOT be the objective!
• It is a management action to achieve the
objective
 Objectives of control
• The reason for the control action should be
clearly stated:
• E.g. increasing the density of a food plant
from one density to a higher one.
• Usually “objective” of control effort is
defined as trying to reduce the density of
the species being controlled.
• E.g. we need to reduce this species by x
percent.
• The management action then becomes
the goal!
• Example: never any clear objective and
eventually, just did it because….
• Some “objectives” shown not to be
accurate: deer had little effect on erosion.
 Back to coyotes
• Coyotes (and predators in general): we
know they kill sheep and game but have
very little data to demonstrate that
“controlling them” leads to x # less sheep
killed or y # more game in the field.
• In fact, we kill over 90,000/year and sheep
industry suffers the same level of loses/yr!
• Control becomes a means in itself!
 Length of control operation
• How long should you control a species?
• Important because once you start the
cost/benefit depends on how long you
need to commit resources to this.
• One year? two years? Forever??
• To understand what your getting into lets
look at the process of control.
 Process
• In any control operation, biologically we
are doing same as in harvesting:
• Lowering population via some form of
killing.
• In the harvest we WHAT the population to
rebound.
• In control we don’t want it to do so but…
 Same process
• Same process that we hope gives us
sustainable yields (higher recruitment rates
because of more resources/individual), also
applies to controlled populations.
• Population will spring back!
 Coyotes again!
• In control operations against coyotes, find
that remaining coyotes have increased
litter sizes!

• Wild horses/donkeys U.S.: same trends,

remove animals but population just comes
back!!
 So control is forever!
• Desert bighorn and cougars: attempting to
keep sheep populations going in the face
of other factors by reducing predation.
• Sheep will never get to a point where they
will be self sustainable WITHOUT cougar
control!
here
Conclusion
• Be sure you have clear justifiable
objectives for control….not just because
you think it will work!
• Be sure you have data on collateral effects
from control efforts
• Be aware that you are embarking on a
long term commitment.
• DON’T enter into it lightly!!
 Ok, it is justified
• How do you do it?
• First is by manipulating mortality: killing
them!
• Humane considerations need to be taken
into account (often difficult to do). Even the
most obviously deleterious species will
have defenders.
• The worse thing to do is make a sport of it!
 Your set to kill
• What should be the appropriate control
strategy?
• Return to harvest strategies: the one that
was most prone to overharvest?
• Fixed quota harvest!
• Saw earlier that even if you have a stable
population, can remove enough to cause
overharvesting.
 That is what you want!
• You want to exceed the maximum
sustained yield!
• Add to that the instability of populations, a
fixed quota harvest, set to overharvest
would provide the maximum control and
could even cause the population to go
extinct.
 Again, trick is how many
• Because you can overharvest anywhere
on the curve, setting the target kill then
affects the time it takes to reach
eradication.
 Examples
• Setting the kill rate of feral goats in aNew
Zealand park at 90%, each year would
lead to eradication in 12 years.
• If it was only 50% it would take 50 years!
 Other factors?
• Again, easy to say we will kill certain
number of animals/year but… as
mentioned, the less there are, the harder
they are to get!
• Adjust to hunting pressure
• Adjust habitat to hard to get to places,
• Adjust behavior: cows in Mexico.
• Judas goats
 Besides hunting them?
• Biological controls
 Is this the way it should be?
• If all we want is to “ranch” a single
species, ok.
• But is that all we want?
• Is that what is good for the ecosystems
that support these desired species?
• Sacrificing keystone species and
ecosystem health for one single focused
use of the ecosystem.