Population “Climb-and-Collapse”

Outcomes in “Vast open-space”
( 2/1000ths of 1% occupied // 99.998% unoccupied ) Conditions
The classical climb-and-collapse
collapse population graph shown
left (after Scheffer, 1951) tracks a population of reindeer
on an Alaskan island (1911-1950
1950). Following the introduction
tion of 25 reindeer to the 106 km2 island in 1911, the
population experienced
ed a period of rapid growth (left
side of graph). This
his initial period of growth, however,
was subsequently followed
wed by a calamitous
calamitou 99%-plus
Note that the graph of this classical study does not exex
hibit an s-shaped (“sigmoid”
“sigmoid” or logistic)
logistic curve. Instead it
exemplifies a J-curve followed by a second real-world
population outcome known as climb-and-collapse.
gap in the graph reflects data that were unable to be
collected during World War II.)

collapse in Population Systems
This article addresses a potentially-calamitous
potentially calamitous outcome of rapid population growth in natural
systems which is known as “climb
climb-and-collapse.” In these patterns, a rapidly-growing
population fails to stabilize its numbers within the ""carrying capacity"" limits of its environment and
instead continues its growth. Then, by exceeding one or more environmental limits, the population enters a perilous condition known as “overshoot .”
This article outlines both the dynamics and real
real-world examples of such population
climb-andcollapse outcomes and die-offs
offs. The climb-and-collapse population outcomes outlined here
differ strikingly from other models that diagram, imagine, or postulate “density
density-dependent” or
“logistic curve” population outcomes.
Other broad patterns of population growth (such as exponential patterns,
logistic patterns, and cyclic patterns) are addressed in other articles.

“Climb-and-collapse” population
opulation patterns
patterns earn their name because graphs of their early stages
commonly exhibit a sharply-rising
rising, exponential, or J-curve increase in numbers.
(Notice that the left-hand portion
on of the above reindeer-herd
graph shows a steep increase in the herd’s population.)

In climb-and-collapse
collapse trajectories, however, such populations do not stabilize their numbers or
enter sustainable equilibrium with their environments.
environments Instead, they overshoot such limits and
enter a dramatic “die-off”” or collapse in numbers. In the above reindeer study (and other realreal
world examples), such collapses can exhibit 99%-plus
offs or even worse mass mortalities.

How and why (and by what mechanisms) do such climb-and-collapse outcomes come about?
One trigger that can initiate a sudden burst of rapid population growth is known as “ecological
release.” Such ‘release’ can occur when an important competitor, pathogen, predator, or limitation is removed from a system. When such removals occur, they can suddenly free a population from natural pressures and mortalities that previously held its numbers in check.
As one real-world example of ecological release, sea otters (Enhydra lutris) on the California
coast feed on sea urchins on the sea floor. If too many sea otters are killed, however (through
hunting, for example), the sea urchins can undergo a population explosion arising from the
reduced sea otter feeding pressures.
Environmentally, the explosive increases in the numbers of grazing sea urchins began to damage underwater kelp forests on which the urchins feed (e.g., Estes and Duggins, 1995). Thus,
by eating sea urchins, sea otters help suppress or stabilize sea urchin numbers and help allow
kelp algal forests to persist.
As a second example of ecological release, wild coyote populations (e.g., Canis latrans) feed
on, among other things, jackrabbits (e.g., (Lepus californicus). What if large numbers of coyotes are removed from such a system by local ranchers or by dust-storm events such as those
of the 1930s? The resulting population explosion of jackrabbits reflects another classical example of population explosions resulting from ecological release.
Additional examples of ecological release can be seen when new species (invasives) accidentally arrive (or are introduced into) a new environment. If native species have no experience
with the new invasives, the new arrivals may suddenly explode in numbers and begin to overwhelm, alter, or damage the new environment in which they find themselves. In this case, native
species can be adversely impacted as they suddenly confront competition, predation, or diseases that they have never encountered before.
In other cases, some populations may undergo a sudden explosion in numbers (ecological release) if unexpected bursts of nutrients become suddenly-available. As examples, river flooding
or seasonal upwellings of deep-sea nutrients may suddenly deliver added nutrients to marine
or aquatic habitats. In these cases, the sudden bursts of explosive population growth can result
in oxygen depletion (eutrophication), red-tides, or other harms.

Overshoot and collapse phases
During its overshoot and collapse phases, a population undergoing a climb-and-collapse dynamic may begin to encounter and transgress ecological thresholds and limits.
Examples of limits and limiting factors that instinctively come to mind include instinctive
necessities, of course, such as food and water. Despite such "running-out-of" suppositions,
however, there are other decisive limiting factors which are less-instinctive.
As a population grows ever-larger, for instance, it may begin to damage, reduce, or eradicate
crucial environmental life-support systems. This may occur by release of wastes (as in red
tides in marine systems) or through widening damage, impacts, and eradications of support

machinery itself (think of photosynthesis each day producing the molecular O2 that we breathe
every few seconds).
Cumulative effects of such impacts may damage, undermine, or collapse system capacities for
essential functions such as ecological services, self-maintenance, self-perpetuation, or self-repair.
Thus, J-curve graphs together with growing, endless, and/or ever-accumulating impacts, eradications, wastes, and/or various resource limitations may signal serious limits, tipping points,
and/or threshold transgressions. If such signals and feedbacks are ignored, support-systems
damage and failing functional capacities may lead to die-offs or mass mortalities.

Three classical examples of “Climb-and-collapse” outcomes
This section outlines three classical examples of climb-and-collapse/mass-mortality outcomes in
real-world population systems.
The first of these was described in a classical paper by V.B. Scheffer in 1951. Scheffer’s study
followed a small population of 25 reindeer (Rangifer tarandus) that were introduced to a 106 km2
island off the coast of Alaska in 1911. Within just 27 years the originally-small population of 25
increased in numbers roughly eighty times to become a herd of more than 2000 individuals.
(Notice that the left-hand portion of above graph of Scheffer’s data shows the herd’s numbers
climbing steeply.) Following achievement of its peak numbers, however, the herd underwent a
nearly-annihilating 99%-plus die-off leaving just eight survivors by 1950.
In a second such study, 29 reindeer were introduced to another Alaskan island in 1944 (Klein,
1968). In the initial stages of its climb-and-collapse graph, this herd increased to more than
6000 individuals within just 19 years. Following that peak, however, this second herd also saw
a 99%-plus die-off over the next three years, closing the study in 1966.
A third instance of climb-and-collapse resulting in far-reaching mass-mortalities is exemplified
by outbreaks of red-tide in marine systems. One famous form of red-tide mortalities results from
population explosions of one-celled marine dinoflagellates known as Karenia brevis. Their
outbreaks of red-tide can sometimes cover hundreds of square kilometers and can kill millions
of tons of fish and even large organisms such as manatees.
It is worth noting that K. brevis red-tide calamities arise from wastes called brevetoxins that the
populations of the dinoflagellate cells release into their surroundings. Thus, during population
explosions of K. brevis, vastly-increasing populations release vastly-increasing quantities of
wastes into their surroundings. As these toxins accumulate they begin to affect the wider systems in which the dinoflagellates reside, as well as many other species beyond the dinoflagellates themselves.
What implications might such dinoflagellate examples have for humankind’s own population
futures and trajectories? As one cautionary implication, consider that the cited K. brevis dynamics result from cellular and metabolic wastes that the dinoflagellates release into their surroundings.

At the same time, however, dinoflagellate populations have no capacity to supplement their
cellular and metabolic wastes with additional quantities of societal and industrial wastes.
In addition, no dinoflagellate outbreaks have ever occurred or dispensed their wastes on a
worldwide, decades long, and accumulating scale. Nor have dinoflagellates devised thousands of mechanized means - (such as, for example, chain saws, Earth moving tractors, or
machine guns) - to eradicate their surroundings or other species by physical, mechanical, or
technological means.
Such matters may have highly dangerous implications, however, if we contemplate another species that appears to exhibit exactly such patterns of behavior.

The examples cited in this pdf also offer still other powerful insights into whole-systems aspects
of population-environment dynamics. Its examples underscore, for instance, the role played by
natural "top-down" pressures, mortalities, and interactions that help maintain population and
systems stability. They also show that when such controlling pressures are removed, a population explosion arising from ecological release can result. In that case, an ecologically-released
population may begin to overwhelm or displace other species or begin to undermine or overwhelm functional system stability itself.
Note that in an ecological release a key interplay emerges between fertility rates and unexpected reductions in external mortality pressures. The response and speed of response (or nonresponse) of fertility rates of the species experiencing ecological release becomes decisive.
If guided, for instance, by tradition, habits, or instinct, a population may continue to reproduce at
previously-existing (or even greater) fertility rates. Imagine, therefore, a combination of external
mortality pressures being reduced, together with continued (or even enhanced) levels of ongoing fertility. Combining the two, a rapidly-growing population can find itself suddenly approaching and then “overshooting” various carrying capacity limitations of its environment.

Civilizational implications
Considerations of ecological release/population explosion dynamics may have implications for
civilization’s population-environment impacts today. For example, desirable advances in health
and medicine have thankfully suppressed many deadly pathogens that once held our numbers
in check.
A potentially-crucial aspect of this, however, may be that humankind’s ecological release may
constitute (or reflect) a repetitive process rather than a one-time event. Thus, humankind’s
soaring J-curve graph of worldwide population growth may reflect not one, but multiple, sudden,
and unexpected cases ecological releases one after another.
Might our own species be currently exhibiting something like an ecological release on steroids?
If one ecological release can raise the possibility of climb-and-collapse outcomes, what might be
expected of multiple such releases in quick succession?

It seems worth noting, for instance, that a graph of humankind’s
d’s worldwide population growth (shown left) over
the 10,000 years of civilization is quite-noticeably
extremely pronounced J-curve.
The above becomes, if anything, even more sobering if
we bear in mind that: (a) As J-curve
curve progressions skyrocket upward along their y-axes,
axes, they have a rather de
decided tendency to flatten everything around themselves
in every direction, (b) That up untilil now the two most famous J-curves
curves in all of human history have both been
atomic detonations, and (c) That the J-curve
shown left
is not a localized phenomenon but is a worldwide phephe
nomenon that is taking place in the middle of the only
planetary life-support
support machinery so far known to exist
anywhere in the universe.
In addition, it shows world population totals increasing by
more than five billions just since 1930. Likewise, recent
U.N. medium and high-fertility
fertility projections show that world
population may be headed toward 10,
1 11, 12, 13, 14, or
15.8 billion by 2100.

A sentient species in seemingly 'empty' surroundings?
Somewhat surprisingly, real-world
world population die-offs
and collapse outcomes can classically
occur in seemingly ‘vast open-space’
conditions and surroundings that can visually appear to
remain almost entirely ‘empty.’
For example, all three populations outlined in this briefing peaked and then began their collapse in surroundings that remained roughly 99.998% unoccupied. At their peak populations,
for instance, both reindeer-herds
herds discussed in this article lived on islands that remained roughly
99.998% unoccupied. Likewise, deadly red-tides
red tides can occur even as the combined K. brevis
cells occupy roughly 2/1000ths of 1% of the water samples in which they live.
None of the three populations cited in this article consisted, of course, of sentient species. Even
if they were sentient, however, it is interesting to ask whether their
ir population collapse, thresholds, extent, and proximity would seem imminent?
Given vast-open-space (99.998% unoccupied) surroundings, would even their brightest
scholars and most conscientious leaders appreciate the nearness and degree of danger?
Could they have readily perceived or even imagined both the proximity and degree of the approaching thresholds their populations faced? And could they have even begun to fully-apprecifully
ate the possible degree (99%-plus
die-offs) of the approaching emergencies that their populations faced? Given that their populations seemed to be surrounded by such seemingly ‘vast
open-space’ conditions,, would they assess their
the population trajectories as an emergency matter
demanding immediate attention?

Lastly, red-tide or harmful
ul algal bloom (HAB) impacts may also suggest potential implications
for humankind. For example, the adverse impacts of red-tides
red tides arise from their release of cellular
and metabolic wastes into their surroundings. On the other hand, dinoflagellate populatio
populations exhibit no capacity to supplement such wastes with additional quantities of industrial and societal
wastes. Nor have they devised any mechanized
mechanized means (such as millions of chain saws, E
rs, machine guns, and shopping malls, etc.)
etc. to eradicate their
heir surroundings or other
species by physical, mechanical, or technological means.
Thus, we know of no dinoflagellate species that possesses physical or engineering capacities
to amplify the degree, extent, speed, and efficiency of non-stop eradications
eradication on an endless,
ever-widening ,and ever-accumulating
accumulating worldwide scale. In contrast, of course, we DO know of
another species, perhaps, that DOES exhibit such capacities
ies and which appears to employ
them on a global scale?

All three classical climb-and-collapse
collapse outcomes cited in this briefing
took place when the physical bodies (or cells) of each population
cited physically-occupied
occupied roughly 2/1000ths of 1% of environmental
surroundings that visually-speaking,
speaking, appeared to remain “almost
entirely empty.”
(To visualize 2/1000ths of 1% in more familiar terms, envision a circle roughly twice
the diameter of a baseball on an otherwise empty basketball court.)
In all three classical cases referenced, the cited collapses or calamities either
eith began
or were already well-underway
underway in “vast open space” conditions that remained roughly
99.998% unoccupied.
A question suggested by these assessments?
Suppose, as a thought experiment, that any or all of the three classical populations
cited were sentient species. How many of even their brightest scholars and most
conscientious leaders of even a sentient and educated species could have imagined
that the vast open space conditions depicted here and referenced in this
thi article
already constituted “too-late” / “they waited too-long”
long” population conditions?

This article has shown that climb-and-collapse
can arise from ecological release and population explosions that result when controlling top-down
top down pressures and mortalities
are removed.
In turn, an exploding population of such ecologically-released
ecologically released organisms can begin to exert adverse and cascading
g impacts on other species and upon the functional life
life-support systems upon which they themselves depend.
Scheffer, V.B., 1951. The rise and fall of a
reindeer herd. Scientific Monthly 73:356-362
downloadable JSTOR PDF of this paper is
accessible at

Klein, D.R., 1968. The introduction, increase,
and crash of reindeer on St. Matthew Island.
Journal of Wildlife Management 32: 350
350-367. A
web copy of this paper is accessible
at http://dieoff.org/page80.htm

Bushaw-Newton, K.L. and Sellner, K.G. 1999.
Harmful Algal Blooms IN: NOAA’s State of the
Coast Report, Silver Spring, Md. NOAA.
Accessible at
(And see also
s/sotc_pdf/hab.pdf )
Estes, J.A. and D.O. Duggins. 1995. Sea otters
and kelp forests in Alaska: generality and
variation in a community ecological paradigm.
Ecological Monographs 65:75-100.
A downloadable JSTOR PDF of this paper is
accessible at

Knowlton, F. F., and L. C. Stoddart. 1992. Some
observations from two coyote-prey studies.

Pages 101 – 121 in A. Boer, editor. Ecology and
management of the eastern coyote.
Wildlife Research Unit, University of New
Brunswick, Fredericton, New Brunswick,
Bartel, R. A., Knowlton, F. F., and Stoddart,
L.C., 2005. Mammal abundance indices in the
northern portion of the Great Basin, 1962–1993.
Ecology 86:3130.
Flewelling, L. J.; et al. (2005). "Red tides and
marine mammal mortalities". Nature 435(7043):
755–756. doi:10.1038/nature435755a. PMC
2659475. PMID 15944690.
WDFW [Washington Department of Fish and
Wildlife]. Annual report, 2012.

Further Reading
Meadows, D., Randers, J. and D, Meadows,
2004. Limits to Growth: The 30-Year Update.
Chelsea Green Publishing, White River
Junction, Vermont.
Cohen, J.E., 2003. Human population: The
next half-century. Science 302:1172-1175.
Press, F., Atiyah, M., and Raven, P., et al., 1992.

Population growth, resource consumption,
and a sustainable world. U.S. National
Academy of Sciences and Royal Society of
148&page=5 and
Kendall, H., et al., 1992. An urgent warning to
humanity. Union of Concerned Scientists,
Cambridge, Massachusetts.

Actual text with list of signatories: http://wwwformal.stanford.edu/jmc/progress/ucs-

Catton, W.R., 2009. Bottleneck: Humanity's
Impending Impasse. Xlibris Corporation, 2009.
290 pp. ISBN 978-1-4415-2241-2
Myers, N. and R.A. Mittermeier, et al., 2000.
Biodiversity hotspots for conservation priorities.
Nature 403:853-858.
Catton, W.R., 1980. Overshoot: The Ecological
Basis of Revolutionary Change. Urbana:
University of Illinois Press, 1980. 298 pp. ISBN
Encyclopedia of Life: Red-tide / Karenia brevis
Femmer, R. 2013 Six PDF executive summaries
for policymakers and academia. Accessible at

Climb-and-collapse in natural systems

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