The Corridors Concept: learnings along the way
|A tiger crosses a river, beyond the designated Protected Area cover, in the central Indian wilderness, a rare parcel of land that is borderless, just a tad-bit careful about avoiding humans as they too amble along the same river.|
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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