What are we talking about?
Before
embarking on this exploratory journey in earnest, let’s first establish some
loose definitions for the temporal and descriptive characteristics that will be
referenced moving forward.
What: Fundamentally derived from the Greek megalo (meaning large), and the Latin fauna (meaning animal), megafauna are
simply defined as large animals. Defining ‘large’ in this context is then where
complexity arises. Delimitation of megafauna tends to be according to some
threshold mass of the adult animal, commonly >44 kg but varying greatly between
>5 kg and >1000 kg in different studies. Elsewhere, megafauna are defined
and stratified according to a trophic (relating to feeding and nutrition) herbivore-carnivore
cascade. Malhi et al. divide megafauna into megaherbivores (>= 1000 kg),
megacarnivores (>=100 kg), large herbivores (45-999 kg), and large
carnivores (21.5-999 kg) in order to incorporate intra-megafaunal trophic
interactions.
When: The timeframes to be dealt with are likely to
be variably and sometimes interchangeably referred to as the late Pleistocene
(from ~60,000 years ago to the beginning of the Holocene (~11,700 years ago)),
the late Quaternary (from ~60,000 years ago to the present), or in terms of
thousands of years before present. For the uninitiated (including myself until
very recently), the following table may be of some help:
Having, in
characteristically (and arguably unnecessarily) loquacious fashion, established
that megafauna are large animals and that we’re dealing with a timeframe of
~60,000 years to the present, let’s get stuck in.
History of Megafaunal Loss
Aside from
the Cretaceous-Tertiary mass extinction event, infamous for its termination of
the dinosaurs ~65 million years ago, megafauna experienced no significant
losses until ~1 million years ago in Early Pleistocene Africa, when, coincident
with the evolution of Homo Erectus into the carnivore niche space (increased
human consumption of meat, perhaps in relation to expanding brain size), several
carnivore lineages were locally cut short (some continued to flourish elsewhere
for thousands of years) and there was a substantial decrease in proboscidean
(order of mammals containing Elephants, Mammoths, similar) diversity.
The speed
and magnitude of these losses was to be dwarfed by the ~1 billion land
megafauna individuals lost globally in the late Pleistocene. ~50,000 years ago (50
kya) the earth hosted more than 150 genera of megafauna greater than or equal
to 44kg. By ~10 kya, at least 97 were
extinct. The figure below, from Barnosky et al summarises the number and suspected causes of megafaunal extinctions on each continent in the context of human arrival and climate change by genera:
In Eurasia
(northern Europe, Siberia and Alaska), megafaunal extinction occurred in 2
pulses:
- From 45-20 ky RCBP (thousands radio carbon years before present), coincident with decreasing temperatures and dispersion of Homo Sapiens
- From 12-9 ky RCBP, coincident with increasing temperatures and increasing Homo Sapiens populations
In North
America, megafaunal extinction coincided with climate change and the arrival of
advanced Homo Sapiens armed with a repertoire of advanced stone hunting tools between
11.5-10 ky RCBP, with more than 15 species becoming extinct during the Younger
Dryas (a shift from cold glacial to warmer interglacial state) between 11.4 and
10.8 ky RCBP.
In the southern
hemisphere (where a very significant proportion of the extinctions occurred), pending
further research, chronology is less certain. Current indications point to megafaunal
extinctions loosely around 45 kya in Australia, coincident with believed
timeframe of Homo Sapiens expansion in the region but interestingly predating
regional Last Glacial Maximum climate change. In South America, the current
consensus is that humans arrived between 12.5-12.9 ky BP and that megafaunal
extinction occurred sometime thereafter, probably coincident with changing climate.
Outside of these major regions, with the notable exception of Africa and Central Eurasia where hominids have been present for hundreds of thousands of years, megafaunal extinctions coincided with the global expansion of Homo Sapiens; ~30 kya in Japan, ~6 kya in the Caribbean, ~1-3 kya in the pacific islands, ~2 kya in Madagascar, ~0.7 kya in New Zealand.
Throughout
the world, megafauna that persisted into the mid-Holocene and beyond tended to
be found in areas where human populations never grew large – although some species,
such as horses and mammoths in mainland Alaska, became extinct without significant
human populations present, instead being the likely result of substantial
regional climate change.
Humans as
omnivorous, generalist super-predators capable of effecting and maintaining
predation pressure on even the very largest animals (who had been under very
little predation pressure before their arrival) had the ability to wreak havoc
on megafauna, particularly vulnerable to increases in predation pressure as a
result of their long lives and slow reproduction rates. Additionally, domestic
animals associated with humans may have introduced diseases into endemic
populations as well as competing with endemic carnivores for food. Resultant loss
of keystone (to be covered in detail later) megaherbivores, leading to changing vegetation and fire regimes and
loss of a food-base for megacarnivores, could then have compounded losses,
setting into motion subsequent cascades of extinctions. Yet in Africa, central
Asia and various other regions around the world, humans and megafauna coexisted without such
drastic megafaunal extinctions as elsewhere.
Climate
change has been proven to affect animals principally through the elicitation of
significant, abrupt vegetation change. However, while in some areas, the timing
of vegetation change coincided with various stages of extinctions, in others it
did not. Furthermore, climate shifts in
the late Pleistocene are believed not to have been unusual enough to effect considerable
ecological change, being neither faster nor of greater magnitude than other
such shifts in the past 700 kya that didn’t result in such megafaunal
devastation.
Neither
human nor climate effects appear to be sufficient in themselves, but the above
seems compelling evidence that a combination of human effects and climatic
change (likely leaning further towards humans than climate change in the majority of cases), interacting to apply theretofore unprecedented pressure, is the likely
culprit of the late Pleistocene megafaunal extinction event, characterised by
short term, regional pulses.
Much of the
information conveyed in this post was derived from a pair of fascinating
papers, Barnosky et al. 2004 and Malhi et al. 2015, both well worth a read in
their original entirety.
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