Ecological Release and Climb-and-collapse

Outcomes in real-world Population Systems

This article addresses one primary category of classical outcomes of rapid population growth in
natural systems which is known as “climb-and-collapse” in which a rapidly-growing population
fails to stabilize its numbers within the "carrying capacity" limits of its environment and instead
continues its growth, progressing into a condition of “overshoot” which transgresses or exceeds at
least one or more environmental limits, thresholds, capacities, or limitations [1].

The purpose of this article is to address such real-world climb-and-collapse outcomes because they
stand in stark contrast to widespread but theoretical suppositions and expectations (sometimes true
and sometimes not) that underlie mathematical models that envision “demographic transition” out-
comes or appear to mistakenly-assume that "logistic curve" population outcomes must supposedly
constitute some universal automatic rule or inviolate law of nature. (Such s-curve expectations,
however, are .not. universal outcomes, nor inviolate, nor an automatic outcome or law of nature.)

The classical “climb-and-collapse” dynamics outlined here earn their name because graphs of such
populations over time tend to exhibit an early growing phase (which often rises sharply in an expo-
nential or “J-curve” pattern as in the left-hand portion of reindeer graph below), exhibiting a pat-
tern of population growth that does not stabilize or achieve a sustainable equilibrium with its en-
vironmental systems and limits, but instead overshoots such limits and subsequently exhibits a
calamitous "die-off" or collapse in numbers as seen in the right-hand portion of the depicted graph.

If we assess classical real-world studies (such as Scheffer, 1951[2] and Klein, 1968 [3]) populations
that exhibit such climb-and-collapse trajectories can undergo rapid and nearly-annihilating 99%-
plus die-offs (and/or even-worse mass mortalities as in Bushaw-Newton and Sellner, 1999) [4].

The classical climb-and-collapse graph shown left reflects
data after Scheffer (1951) which tracked a population of rein-
deer on St. Paul Island, Alaska (1911-1950). Note that the
data from this classical study, when graphed, do not produce
a logistic or sigmoid-curve as often envisioned by “demogra-
phic-transition” models, suppositions, and theorists, but in-
stead exemplifies a second real-world population outcome
known as climb-and-collapse. (The gap in the graph reflects
data that were unable to be collected during World War II.)

This graph depicts the population growth and eventual collapse of a reindeer herd introduced to an
Alaskan island in 1911, leading to overshoot and a subsequent and nearly-annihilating 99%-plus
die-off. The graph reflects data from a classical climb-and-collapse population study reported by
V.B. Scheffer in 1951 (The rise and fall of a reindeer herd. Scientific Monthly 73:356-362).
Mechanisms –
Including ‘Ecological release’

How and why (and by what mechanisms) do such classical climb-and-collapse outcomes come
about? One trigger that can lead to such a pattern is known as “ecological release” in which a key
competitor, pathogen, predator, or limitation is unexpectedly removed from a functioning system,
which suddenly frees a population from natural mortalities or pressures that previously held its
numbers in check.

Such limitations might include, for example, instinctively-appreciated matters such as limited sup-
plies of food or water or other ‘resource’ limitations. Other barriers, obstacles, or suppressing fac-
tors however, can be equally powerful and important (but may be somewhat less-instinctive), and
can include the suppressing effects of pathogens and disease, competition with one or more other
species, and/or the feeding pressure or predation by other species.

As one example of ecological release, sea otters (Enhydra lutris) along the Alaskan and California
coasts feed on sea urchins, so that if sea otter numbers are reduced (through hunting, for example),
a resulting population explosion of sea urchins can ensue (ecological release), which, in this case,
can begin to damage large areas of the underwater kelp forest upon which the urchins feed (e.g.,
Estes and Duggins, 1995).[5] By their predation, therefore, sea otters help suppress and control sea
urchin numbers. At the same time, however, any weakening or removal of this controlling external
pressure results in a large, sudden, or dramatic increase in sea urchin numbers with potential dam-
age to the kelp-beds together with systems-wide and cascading impacts on multiple other species.

As a second example of ecological release, wild coyote populations (e.g., Canis latrans) feed on,
among other things, jackrabbits (e.g., Lepus californicus). If large numbers of coyotes are re-
moved from a system (by experiment, for example, or by hunting, or by dust-storm events such as
those of the 1930s), the jackrabbit populations upon which they feed can suddenly explode in
numbers (e.g., Knowlton and Stoddart, 1992) [6]. We thus see illustrative examples of natural "top-
down" regulatory mechanisms wherein mortality or feeding pressures exert a check on the size of
a given population. Thus, as the above examples show, if such checks or limiting-factors or limit-
ing-influences are removed (“ecological release”), a prey population explosion can result, with
multiple and potentially cascading impacts on other species or even with systems-persistence im-
plications and effects.
As the above examples thus illustrate, when natural populations undergo ecological release and
find themselves suddenly freed from one or more negative external pressures that would normally
hold their numbers in check, they may simply continue to reproduce at previously-existing (or ev-
en enhanced) fertility rates and a sudden and unexpected population explosion ensues.

The combination, therefore, of outside mortality pressures being reduced, together
with previously-routine, instinctive, or enhanced levels of ongoing fertility,
results in a population with rapidly growing numbers that can find itself suddenly cascading
toward and “over-shooting” various carrying capacity and environmental systems limitations.
There are thus implications here for humankind and our own current, gigantic, dangerous, non-stop, on-
going, and worldwide population explosion. The two quick examples of population explosions that we
have just examined, for example (jackrabbits and sea urchins), each occurred in response to an ecological
release (and our next sections will explore three calamitous examples of the catastrophic climb-and-collapse
die-offs that can follow).

In what ways, therefore might our own planet-wide population explosion be similar to and dissimilar to the
sea urchins and jackrabbits population explosions just cited? First, our own species has clearly undergone
an ecological-release – and not just locally, but on a global scale. One of the most critical factors triggering
our release have been our advances in medicine, allowing more and more individuals of each generation to
live longer, so that death rates around the world have fallen (and we all like that). What else has happened,
however, is that all around the world (and in some places more than others) despite falling death rates, too
often our societies and families have continued with their previously-routine, instinctive, or traditional lev-
els of ongoing fertility.

As a result on a worldwide scale, birth rates on average now exceed death rates (and the mathematics of
that disparity has launched us upon – like the sea urchins and the jackrabbits in our example - a gigantic
population explosion). And the calamitous mass mortalities observed in three classical real-world examples
of such climb-and-collapse outcomes will be outlined in the sections which follow. First, however, one
more thing about humankind. Yes, somewhat like the sea urchins and jackrabbit populations in our initial
examples, we humans have undergone an ecological release. But our dynamics is actually dissimilar in
exceedingly dangerous ways:

[1] Our population explosion is taking place on a worldwide scale and is damaging, eradicating, and oblit-
erating vast portions of the only planetary life-support machinery so far known to exist anywhere in the
universe; and

[2] Our species has not undergone ONE ecological release, but multiple ecological releases, one after ano-
ther and another and another. Each new advance, for example, in medicine, antibiotics, and vaccines has
happily and thankfully helped reduce death rates. Each such advance, however, commonly arrives as a
welcome - but unexpected – surprise. Thus, death rates as a result can begin to fall immediately, but in
each case there is a decided delay or a lag-time before families and societies and traditions begin to reduce
or adjust fertility accordingly (or to the same extent, or at all), with the result that humankind’s worldwide
numbers continue to explode upwards.

Other ecological releases in humankind result from advances in agriculture, industry, and fishing technologies, etc., which have
allowed us to overcome past difficulties and negative external pressures that once held our numbers in check. Thus, unlike the
cited ecological release / population explosion examples already cited above, humankind has undergone one ecological release
after another and another and another, so falling fertility is never able to catch-up and our worldwide population explosion does
not stabilize.

Overshoot and collapse phases .
During the overshoot and collapse phases of such population histories, such a population may not
only exceed or exhaust critical resources such as food and water, etc., but may simultaneously be-
gin to damage, reduce, and/or degrade its surroundings and environmental support systems with
wastes and/or eradications, and/or to damage, diminish, degrade, or collapse the capacities of such
systems for self-maintenance, self-perpetuation, ecological services, and/or self-repair. (Damage,
wastes, environmental impacts, J-curve graphs, and/or various resource shortages or limitations
can be signals of potentially-serious population, ecological, and systems risks, while the “collapse”
phase of such populations can reflect assorted combinations of wastes, impacts, damages, eradi-
cations, diminished resources and/or damaged or failed functional capacities that result in die-offs
and/or mass mortalities.)
Examples of real-world “climb-and-collapse” die-offs and outcomes
This section outlines three classical real-world examples of actual population climb-and-collapse
/mass-mortality outcomes. The first of these, described in a paper by V.B. Scheffer in 1951 [7],
began with a small population of 25 reindeer (Rangifer tarandus) introduced to a 106 km2 island
off the coast of Alaska in 1911. Within just 27 years the originally-small herd of 25 increased in
numbers to more than 2000 individuals (roughly 80x larger; and notice the roughly exponential
increase seen in this article’s opening graph). Following achievement of its peak numbers, how-
ever, the herd underwent a nearly-annihilating 99%-plus decline (die-off) leaving just eight sur-
vivors by the close of the study in 1950.

In a second such study (1944-1966), 29 reindeer were introduced to another Alaskan island in 1944
(Klein, 1968). [2] This herd increased its numbers to more than 6000 individuals within just nine-
teen years. That peak, however, was immediately followed by a nearly-annihilating die-off over
the next three years with a loss of 99%-plus of the herd by the close of the study in 1966. [Notice
therefore a second calamitous-scale extent of the mass mortalities.]

A third classical example of real-world mass-mortality calamities arising from explosive popula-
tion growth is exemplified by population explosions of one-celled marine dinoflagellates such as
Karenia brevis which induce the marine environmental calamities known as “red-tides” which
can sometimes cover hundreds of square kilometers and can kill millions of tons of fish and even
other species such as manatees (see, for example, harmful algal blooms, red-tides, and Bushaw-
Newton and Sellner, 1999 [8]). It is interesting that such red-tide calamities arise from wastes (such
as brevetoxins, for example) that the dinoflagellate populations release into their surroundings, so
that their increasing numbers result in increasing wastes that end up affecting both the wider sys-
tems in which they reside as well as many other species beyond the dinoflagellates themselves (do
we know of any other species that appears to exhibit a somewhat similar behavior?).

Die-offs in seemingly 'vast-open-space' conditions

Recent mathematical analyses of the three real-world population climb-and-collapse calamities ci-
ted above [22] have shown that such calamities can “actually unfold in ‘vast-open-space’ surround-
ings which, visually-speaking, appear to remain almost entirely ‘empty.’ [ibid]

It is often surprising, therefore, to realize that real-world population climb-and-collapse calamities
can actually unfold in seemingly ‘vast open-space’ surroundings which, visually-speaking, appear
to remain almost entirely ‘empty’ (e.g. roughly .99.998%. .unoccupied. by the explosively-growing
population itself). For instance, in all three of the classical ‘climb-and-collapse’ die-offs/mass mor-
talities already described in this article (two mammalian populations and one population of marine
dinoflagellates; Scheffer, Klein, and Bushaw-Newton), in all three cases the collapse and calami-
tous die-off outcomes were either about to begin or were already well-underway when the com-
bined bodies (or combined cells) of each of the populations physically-occupied roughly 2/1000ths
of 1% of their environments. (In other words, all three of the populations in the quintessential
climb-and-collapse calamities already described had already peaked or had already begun to enter
their calamitous phases while occupying surroundings that were roughly 2/1000ths of 1% occupied
and which remained roughly 99.998% unoccupied.

(See supporting graphs and mathematics link in article addenda, together with external links to related executive-brief-
ing and ocw pdfs reviewing real-world collapse outcomes in seemingly ‘vast open-space’ surroundings).

In the case of the dinoflagellate red-tides just discussed the mass mortalities and obliterating out-
comes of such outbreaks can begin (or may already be well-underway) when the dinoflagellate
cells responsible for the outbreak reach, approach, or exceed densities of approximately 1,000,000
cells per liter.[22] However, when the actual physical size of all 1,000,000 such cells combined is
compared to the physical volume of the one liter sample in which they reside, the total cells per
liter exhibit a roughly 2/1000ths of 1% occupancy while most of the sampled liter remains roughly
99.998% unoccupied [22] so that it would visually appear to remain almost entirely ‘empty.’

Cognitions? Capacities? Lag-times? Non-responses?

The observation has recently been made [22] that none of the three populations cited in this article
consisted, of course, of sentient species. ”Even if they were sentient, however, it is provocative to
contemplate that, given such seemingly vast open-space (roughly 99.998% unoccupied) surround-
ings, whether even their brightest scholars and most conscientious leaders, could have perceived
or imagined: (a) The proximity of potential climb-and-collapse thresholds their populations faced
and/or (b) The possible degree of the potential population-environment emergency conditions,
especially when such ‘vast open-space’ surroundings appeared to seemingly prevail (recall the
99%-plus die-offs and/or mass mortality calamities in all three examples cited in this article).
Would they assess the condition as a worry worthy of immediate attention?” [22]

Recent observations have also suggested that “Such ecological release / population explosion
events may be relevant to humankind’s own population-environment conditions today because in
effect, our own population explosion today appears to reflect a classical instance of “ecological
release” (or multiple and repetitive such releases, one after another and another and another) in
which desirable advances in fields such as health and medicine have thankfully, greatly, and re-
peatedly lessened the effects of deadly pathogens that once held our numbers in check.” [17]

A resulting side-effect of such desirable advances in agriculture, health, and medicine, of course,
has also been a sudden environmental imbalance as worldwide death rates have fallen (with the
help of science and medicine), even as worldwide birth rates have not fallen to an equal extent.

The result of this imbalance has been a sudden ecological release with the sudden, rapid, and
dramatic increase in humankind's worldwide numbers and our planetary-scale impacts. The
results of our non-stop and ever-growing worldwide numbers, together with growing, ever-
widening and ever-accumulating impacts, wastes, damage, and eradications to the only which so
much has been written in recent decades (such as, for example, formal population environment
warnings by 99 Nobel Laureates and hundreds of top scientists [9], Officers of the U.S. National
Academy of Sciences and the Royal Society of London (1992) [10], and multiple books[11]).

Possible implications

Might examples such as those cited above have any implications for humankind’s own population
futures and trajectories? Consider, for example, that dinoflagellate impacts such as those cited
above result from cellular and metabolic wastes that dinoflagellates release into their local sur-
roundings over a relatively short span of time.

Meanwhile our own species: (a) Not only exhibits a similar pattern of behavior, but does so on an
endless, growing, non-stop, worldwide, and ever-accumulating scale, and, in addition, (b) Our own
species also supplements its cellular and metabolic wastes with an endless, non-stop, on-going,
daily, planet-wide, ever-growing, and ever-accumulating avalanche of societal and industrial

Suppose that we try to reassure ourselves, however, that we humans are smarter than a population
of mindless, one-celled dinoflagellates. (And actually, of course, when it comes to libraries, space
travel, communications, symphonies, and great literature, our species is clearly much smarter than
dinoflagellates. Dinoflagellates, for example, cannot memorize Hamlet, write computer code, or
manufacture a refrigerator. Nor can they launch a satellite, write their own history, or write a book.)

(On the other hand, mindless one-celled dinoflagellates have not devised things such as bulldozers,
chain saws, wood-chippers, Earth-movers, and other machinery to quickly and efficiently eradicate
entire forests, nor have they built and dispatched industrial-scale fishing fleets that employ long-
lines, drift-nets, radar, and GPS to pursue and catch entire schools of fish. Nor have they invented
and manufactured millions of trucks, buses, automobiles, and interstate highways, together with
coal mines, oil wells, and power plants to pump greenhouse gases into the atmosphere, nor ways
to flood forests, produce nuclear weapons, shoot elephants, rhinos, endangered species and each
other, while also draining aquifers, diverting rivers, injecting fracking fluids, and eradicating water
bodies like the Aral Sea; etc.)

The above might give us pause, then, since:
(1) We are the only animals on Earth that do this,
(2) We are the only animals on Earth that have EVER done this, and

(3) We are doing so on a global scale throughout the entire world,
all at the same time, in less than a single human lifetime.
(Which clearly shows that we are smarter, right?)
Thus, if populations of red-tide dinoflagellates were required to take an exam testing their know-
ledge of biospheric life-support machinery, this pdf, whole-systems ecology, literacy involving
exponential mathematics, J-curves, the enormity of the difference between a million and a billion,
ecological release, demographics, population explosions, Earth’s razor-thin surface films of atmo-
sphere, oceans, soils, and seas, planetary carrying capacity, limits, limiting factors, real-world pop-
ulation climb-and-collapse outcomes in ‘vast-open-space’ conditions (e.g. 99.998% unoccupied as
‘too-late’ / ‘waited-too-long’ conditions), and the only planetary life-support machinery so far
known to exist anywhere in the universe, our dinoflagellate colleagues would be (a) virtually illit-
erate, and (b) completely clueless.
But unfortunately, for the most part, of course,
so would we – (including most of our leaders).

Lastly, note that in this graph of worldwide
human population growth over the last ten
thousand years (which exhibits an extreme

That humankind's worldwide population
increased by roughly FIVE additional billions
in less than a single human lifetime
between 1930 and 2011,

(And according to recent U.N.
medium and high-fertility projections
may be headed toward 11, 12, 13,
14, 15, or 16.6 billion by 2100).

This article is an open-courseware document
and is entirely free for use by scientists,
students, and educators anywhere in the world.

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