The Corridors Concept: learnings along the way
The process of population isolation, driven by habitat loss and fragmentation, leads to population extinctions and reduction in biological diversity (Rosenberg, Noon & Meslow, 1997). That isolated populations are significantly more prone to extinction with increasing interpopulation distance has been observed in various taxa, including insects (Saccheri et al., 1998), fishes (Magnuson et al., 1998), frogs (Sjögren, 1991), snakes (Webb, Brook & Shine, 2002), and mammals – from the small island marsupials (see Miller et al., 2011) to large carnivores such as tigers (see Sagar et al., 2021), as has been theoretically put forth by Wright (1943) in the iconic ‘isolation by distance’, and later demonstrated by MacArthur and Wilson (1967) in their treatise ‘The Theory of Island Biogeography’. Human-mediated habitat loss and fragmentation are considered to be the greatest threat to biodiversity, particularly terrestrial mammals, with studies predicting on an average ten ecoregional mammal extinctions due to human land use change (Kuipers et al., 2021), and with global threat of climate change, it is likely to exacerbate threat to over 54% of biodiversity in 18.5% of the ecoregions (Segan, Murray & Watson, 2016).
The concept of
corridors came to the forefront nearly two centuries after the earliest
Protected Areas were established. As protected areas transformed into islands amidst
mosaic human-dominated landscapes, the concept of biological-, wildlife-, and
habitat-corridors was advocated in early 1980s ‘to increase the connectivity of
otherwise isolated patches’ (Rosenberg, Noon & Meslow, 1997; Beier &
Noss, 1998), with cues from island biogeography theory to show how isolation
influences extinction and immigration regulates balance. Like life on islands,
this connectivity relies on several dynamic features a population is subject to;
spatial, such as distance to the nearest habitat patch of any size, the nearest
large patch, and the nearest occupied source patch (Prugh et al., 2008), life-history
requirements such as food availability and territory (example Harihar &
Pandav, 2012; Chanchani & Gerber, 2018), and behaviour, including movement
patterns, often on the limit of the realised niche of that species, such as in-and-around
human-inhabited areas for many of the large terrestrial mammals (Habib et al.,
2020; Barber-Meyer et al., 2012; Harihar & Pandav, 2012).
Corridors were considered
a paradigm in conservation of large terrestrial mammals, enabling interchange
of individuals from isolated populations which would increase local and
regional population persistence, reduce extinction rates, and increase
colonization rates (Rosenberg, Noon & Meslow, 1997). Studies that looked at
extinction vis-à-vis isolation increasingly affirmed that corridor function was
paramount to conservation, but how and in what ways it would manifest was known
in bits-and-pieces across taxa, distances, ecosystems, and increasingly complex
landscapes. A study underscoring the threat of extinction due to isolation focused
on a butterfly species, Melitaea cinxia in Finland, which showed
decreasing heterozygosity in isolated metapopulations, indicated by adversely
affected larval survival, adult longevity, and egg-hatching rate, leading to
extinctions in several isolated populations (Saccheri et al., 1998). Inbreeding as one of the direct effects of
isolation is also seen in large carnivores such as tigers. The rather-common
occurrence of pseudomelanistic tigers in Similipal Tiger Reserve in India,
characterized by broad, darker and merged stripes, was considered an anomalous
phenotype in natural populations associated with loss of genetic diversity in
bottlenecked or inbred populations, with the results suggesting that genetic
rescue to increase heterozygosity would likely decrease inbreeding depression (Sagar
et al., 2021). Saccheri et al. (1998) mention that although heterozygosity was
a significant explanatory variable for the Melitaea cinxia butterfly,
demographic and environmental factors are also known to significantly affect
extinction risk, concluding that, even as ‘demographic and environmental
factors are likely to be the primary determinants of extinction risk, the
contribution of inbreeding should not be underestimated, especially in species
with a highly fragmented population structure.’ Sagar et al. (2021) also mention
that the associated high frequency of pseudomelanistic phenotype is associated
with the reserve’s small and isolated population (eight tigers at 0.39 per 100
sq km density), driven by recent bottlenecks, adding that the apparent absence
of this trait elsewhere ‘suggests strong stochastic effects and inbreeding
operating locally in this population.’
Maintaining – or
managing – immigration is significant for increasing genetic diversity. While genetic
markers are an indicator of a weakening link driven by reproductive isolation, often
these are noticeable as mutations begin to pop up when a population has been in
isolation for several generations. The effects are also directly visible in
terms of declining populations, decreased breeding potentials, biased sex
ratios, and declining survival rate. In most cases, the pervasive drivers of
extinction are environmental and demographic stochastic factors. The former
signifies ‘unpredictable spatiotemporal fluctuations in environmental
conditions’ (Fujiwara & Takada, 2017) and the latter as variation among
individuals within populations caused by random variations in survival and
reproduction (Lee, Saether & Engen, 2021). While demographic stochasticity
has a limited to no effect on large populations – allowing limited to no
genetic drift, smaller populations tend to show a substantial effect on
population dynamics (ibid), and environmental factors may span from freak
storms in the deserts, rivers changing paths, droughts, or now increasingly more
human-mediated changes such as roads fragmenting a contiguous habitat overnight
and the long-term climate-change-related effects.
The best way to
understand isolation vs extinction is to look at landlocked waterbodies, or
waterlocked lands. Magnuson et al. (1998) studied what-drives-what in fish diversity
and assembly of small temperate woodland freshwater lakes in USA – the remnants
of molten glaciers from ages ago – extinction or isolation. Features such as
high acidity and seasonally low oxygen, or both, including predation and
invasion – the environmental and demographic stochastic factors – frequently
led to extinction. The authors found a strong relation between fish species
richness and lake area, similar to, the authors mention, ‘the steeper slopes
observed for more distant island archipelagos, more isolated islands, less
vagil species,’ however, the authors mention that “recruitment in lakes may be
best predicted by extinction models whereas recruitment in oceans may be best
predicted by colonization models,” because, in the lakes, high frequencies of
extinction occur only when invasions continue to populate the lakes: ‘to go
extinct, a species must first have arrived at the lake or returned following a
previous local extinction.’ In simpler words, the authors conclude, ‘the
greater the isolation is among insular sites, the more important extinction
will be in determining patterns of richness and assembly.’ This was also seen
in another group of animals. A study of multiple metapopulations of pool frogs in
Sweden showed that inbreeding depression among populations within two kilometres
or less from the neighbouring population was not a determining factor as much
as combined environmental and demographic stochasticity in isolated populations
which brought about a reduction or absence of egg-carrying females in some
years. In addition, predation also stunted population growth. With increased
isolation, the likelihood of populations facing extinction increased (Sjögren,
1991). Often and more increasingly, human-mediated factors accelerate
population declines. The direct mediators of extinction, such as removing
individuals through their habitat, has been observed in several taxa, from the
endangered broad-headed snake in Australia driven to local extinctions due to
the illegal pet trade (Webb, Brook & Shine, 2002), to the Malayan tiger,
facing an intermediate population crash with only 200 individuals remaining in
isolated rainforests of Malaysia, with poaching, human-tiger conflicts,
decreasing habitat quality, and infectious diseases accelerating the threat of
extinction (Ten et al., 2021).
A corridor, from a
stream to a forested patch, allows for immigration, colonization, and reduces
isolation, but there is a difference: it is influenced by the landscape it is
embedded in. Prugh et al. (2008) remark that unlike landlocked waters and
waterlocked lands, habitat patches are not islands, the surroundings provide
sufficient benign conditions which may serve as areas where the niche may extend
into, with area sensitivity higher in human-dominated matrix than in natural
matrix, such as agricultural fields and green cover along streams in
non-forested areas. For the core concept of terrestrial corridors for
terrestrial mammals, the deterministic factors of the functionality of the
corridor can be broadly classified into two physical components, the distance
and shape of the corridor, and the composition of the corridor. A study of four
carnivore species – three large and one small – in central India showed that
genetic connectivity is influenced by land-use and land-cover for the large
carnivores, with dispersal ability differing significantly between the four
species based on body size and trophic level occupied by the species (Thatte et
al., 2019). The composition of the corridor incorporates the habitat type and
how conducive it is to the species using the area. Large, wide-ranging mammals
such as the African Elephant showed spatiotemporal changes in use of corridors,
influenced by vegetation cover, human disturbance, but also the social and
resource needs of individual elephants (Green et al., 2018).
Given the fact that corridors
are often linear (with length greater than width) unlike chunks of protected
areas, and ecologically different from the matrix on the either size (Rosenberg,
Noon & Meslow, 1997), their role in a species’ life-history requirements
are different – from mere dispersal to being a part of a territory. In central
India, tiger populations connected by forest corridors showed highest rates of
contemporary gene flow than those that have lost a considerable forest cover
and hence connectivity (Sharma et al., 2013). In spite of their structure, corridors
as habitats and not merely dispersal routes are also increasingly being
considered. In addition to movement facilitation, some species meet some
life-history requirements beyond dispersal. For a small carnivore such as the
Jungle Cat, the maximum resistance the central Indian population counters is
density of linear features, but given that it occurs at higher densities than
other large carnivores and with median dispersal distance at 8 km (Thatte et
al., 2019), it is likely occupying the habitat within the corridor, with
increased densities of linear features serving as barriers threatening
connectivity for this small cat. Corridors with larger spaces which encompass
small meta-populations serve as stepping-stones or even a hotspot of genetic
admixtures, as evidenced in some forests of central India, where a 963 sq km
chunk of forested habitat harbours a population of roughly 9 out of an
estimated 30-40 tigers connecting at least four tiger reserves – Kanha, Pench
Madhya Pradesh, Pench Maharashtra, and Navegaon Nagzira (Talegaonkar et al.,
2020). Even as they occupy this land, their behaviour is quite different. Distinct
behavioural patterns have been observed between individual tigers with home
ranges inside a protected area and outside; those outside of protected areas
showed significant displacement in the night than in the day – indicating
human-avoidance behaviour, although both showed relatively low difference in
total hourly displacement rate within and outside protected areas (Habib et
al., 2020). In case of the African Elephant, herds were shown to prefer
corridor sites with lower disturbance during the day and moved closer to roads
at night to traverse the corridor (Green et al., 2018).
A study that aims to
understand habitat use and habitat connectivity has to acknowledge a corridor
as a state-space which is a crucial part of a species’ life-history requirement
not limited to dispersal. In this context, understanding extinction and
isolation gradients for a species requires a holistic approach. How resistant
is the matrix within or beyond the designated corridor for a species is often
influenced by environmental and human-influenced factors, for instance, how
humans and wildlife interact in such shared-space also needs a broad
understanding. While surface resistance such as high human density and built-up
areas, linear infrastructure, and dams and extractive industries, as well as
rivers and valleys, mountains, and other large natural features, are identified
as physical barriers to animal movement, human presence and behaviour as a
factor affecting corridor functionality is an important aspect. Dubbed
‘anthropogenic resistance,’ Ghoddousi et al. (2020) define it as impacts of
human behaviours on species’ movement, including psychological (individual),
social (group), and policy decisions. Factors such as risk to wellbeing and
property also influence connectivity (ibid), an interaction that often
leads to retaliation in the form of hunting corridor animals through illegal
means such as poisoning, trapping, and actively shooting. On the other hand,
that wild animals and humans cooccur and coexist in parts of the world such as
in central India in spite of centuries of destruction of wild animals for sport,
stands the test of time that large carnivores and humans can and do share space
in the 21st century, calling it ‘resistance’ therefore is covering
only one dimension of a cultural landscape; an apt term would be ‘anthropogenic
fluidity’, where resistance but also coocurrence, and rarely but not
exceptionally, coexistence, play important parts in connectivity, for the
animals and for the people.
A study of how local
communities navigate spaces in two central Indian tiger reserves for their
daily livelihood revealed that behavioural responses varied with the type of
risk. For movement through a wilderness area, the speed and directedness of
movement changed if people perceived presence of a wild animal, especially a
large carnivore, in the area (Read et al., 2021). There is a reason for what
the authors call ‘landscape of fear’ in the context of human inhabitants: sudden
encounters with wild animals or damage to property are not isolated events even
in corridors. Several countries provide compensation for the loss of life or
property by a wild animal. In the context of ‘landscape of fear’ for the
animals, how these are managed is a key determinant of whether a corridor
serves its purpose or becomes a death-trap. In India, a majority of states (27
out of 29 states for at least one or more policy) provide compensation
irrespective of where the incident took place, a protected area or outside of
it (Karanth, Gupta & Vanamamalai, 2018). While compensation benefitting
people and wildlife remains to be tallied and audited (Nyhus et al., 2003; Karanth,
Gupta & Vanamamalai, 2018), in view of negative interactions and mitigation
programmes with an objective to reduce retaliatory killings of wild animals
while compensating for the loss, the question whether corridors are places of
coexistence or cooccurrence, or, in other words, favouring wild animals for
humans, is a burning issue. In the 21st century, as the world stares
at two extremes of local-to-global awareness of the natural world and the global-to-local
effects of manmade climate-change, a wildlife-, biological-, and
habitat-corridor remains an important area for conservation interventions made
possible so long as the integrity and functionality of these spaces are
collectively conserved: that a habitat we dub a corridor cannot exist without
animals – or humans.
In our quest to allow
wild populations to interact, we are perhaps missing the forest for the trees,
that corridors, irrespective of their state-space, are shared habitats. In
fact, I will go on a limb to say that a corridor is only one dimension of a
complex land history. Often in a state of gradually reducing or deteriorating
shape and habitat quality compared to the protected areas around, as indicated
by a India-wide study which showed that along with the expansion of croplands,
industrial development and mining, excessive economic dependence on forest
resources is also one of the causes of forest loss (Meiyappan et al., 2017), they
still serve as habitats for a number of flora and fauna, for natural springs,
streams, and origins of rivers; such spaces remain because humans need that
space, intrinsically and extrinsically. To look at a parcel of land in one
dimension, therefore, is missing the big picture. Corridor conservation usually
encompasses conservation of such habitats, but the methods of conservation need
to be integrative as opposed to the protected-area-like exclusionist, fortress-models.
A comprehensive assessment of habitats traversed by elephants, detailed in the
report ‘Right of Passage’ (Menon et al., 2017), lists voluntary relocation and
suitable compensation as an important part of the conservation plan for the
identified elephant corridors, with suggestions for rights to NTFP collection
and felling revoked and restrictions on livestock grazing and fodder collection
being enforced in certain places. Corridors such as these have been established
giving exclusive rights to elephants with little or no resistance today. On the
other hand, such corridor conservation practice is also displacing, albeit
voluntarily, a community that doesn’t merely derive sustenance or livelihood
from the land but considers it a part of themselves as documented by Shaji (2021).
In such practices, whose right of way and whose landscape of fears is a
contentious issue, and enforcing protected area-like management practices as
‘corridors’ which by concept are porous passages of varying sensitivities, requires
a relook at the meaning of corridors. For example, replace the right of passage
for elephants with large carnivores such as tigers, and a corridor becomes a
tiger reserve, exclusive to the large carnivore and its prey-base; the
resulting spill-over requiring ‘creation’ of new corridors for their dispersal.
Comparing tiger with a megaherbivore that does not breed like a cat, but requires large
areas, is not justifiable, but it is justifiable when it comes to
human-wildlife conflicts: if the problem was conflict all along, there are ways
to address the issue without removing the victims of conservation (communities
of that region) or the victims of their habits (elephants of that region). While
exclusionist corridor conservation provides the voluntarily relocated
communities with support to generate new livelihoods, such as in the form of
fishery ponds “to feed” relocated farmers in Assam, in addition to developing
entrepreneurship among women and youth groups, livelihood programmes to reduce
dependence on forests and erecting solar-powered fences, among other activities
(World Lands Trust, 2021), how much this helps with socio-economic upliftment
is strictly a case-by-case, region-specific outcome that is never a win-win model,
even as elephant populations nation-wide fall due to electrocution, train
collisions, poaching, and poisoning – deaths not related to the right of
passage. Furthermore, while elephants make their own corridors, often amounting
to conflict in areas of high human densities and low tolerance, how
exclusionist corridor conservation helps elephants maintain genetic diversity
at the same time reducing conflict with humans, also remains to be seen.
That humans are a part
of a corridor, not merely using it, needs more than just an acknowledgment. The
difference lies in the thin line that separates cooccurrence and coexistence. In
India, the central India and Eastern Ghats, covering eight states and with over
24 Tiger Reserves, comprise roughly a third of India’s tiger population (about
1,033 tigers out of 2,967, Jhala et al., 2020). Of this, over half
(approximately 526 tigers) are present in the state of Madhya Pradesh, of which
14% live outside of protected areas, most of them (est. 57 tigers) in four
corridor areas. The central Indian tiger corridors are well studied for their
connectivity particularly for large carnivores (Rathore et al., 2012; Borah et
al., 2015; Sharma et al., 2013), mapping of habitat connectivity (Dutta et al.,
2015), modelling threat of extinction (Thatte et al., 2018), land-use and
land-cover change (Banerjee, Kauranne & Mikkila, 2020) as well as for ecosystem
restoration (Dutta, Sharma & DeFries, 2018), providing a comprehensive
understanding of the corridor- and landscape-specific ecological and
anthropogenic influences in this tiger stronghold. Factors which influence
habitat-use, such as peoples’ reliance on forest-based produces, scale of
human-wildlife conflicts, pervasiveness of wildlife crime, and frequency of
forest fires, are elements that also affect connectivity laterally – that is,
it is often underrepresented (eg. human-wildlife conflict), its threat underestimated
(eg. forest fires), or its significance not known because it occurs under the
radar (eg. wildlife crime). These problems, as also faced by elephants, cannot
be solved with exclusionist corridor conservation methods. A method that
remains to be seen in India but can be witnessed at smaller spatial scales in
community conservation areas, in areas with community rights, and in
loosely-held concept of corridor conservation between some protected areas
where ad-hoc interventions exist, is that of corridors with the rights to local
communities, especially in the face of industrial and large-scale land
conversions which are a bigger threat to the flow of wild populations. This
idea is where we’re split into two schools of thoughts: if not unique, it is
dystopian for some and utopian for some. As we’re seeing exclusionist corridor
conservation, community corridor conservation is also in the works, if not
under that title, and that might be a bigger gamechanger.
--
A lot can be opined,
discussed, and debated about wildlife corridors. This piece is, as I hope you
can tell from these loosely collected thoughts, a self-study on understanding
the basic concept of corridors, some key studies on why we need this concept,
on why it is so important in some contexts especially pertaining to large
mammal conservation, and how we’re still, in practice, experimenting with ways
in which to conserve such areas even as this concept is evolving, and my final
realisation as I pieced this together, that the corridor concept is more than
just about a passage in space and time: in this day and age, it is more than
the meaning of the word itself.
(This is an adapted
and long-form version of the original written as an introduction to a report on
the habitat use and habitat connectivity of a central Indian corridor, which I
hope to talk more about soon.)
--
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This year of Sahyadrica was the year of opinions – disgorged into the vastness of this space, but more rightly directed in my own mind – particularly of inclusive models of conservation. I’ve set my goal on only a few objectives for the coming year, and as I work towards it, Sahyadrica may go through yet-another period of dormancy. I take this opportunity as we close 2021 to wish you the happiest for what’s to come in 2022, may the world return to normalcy again, and the lot of us back on track.
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