Some ecosystems are transient in nature—their character is temporary and will change because of disturbance and succession. Consequently, species that occupy those transient habitats are bound to be naturally extirpated at one time or another. Consider, for example, a small population of wildflowers occurring in a river’s floodplain; at some stage, there is going to be a flood that will wash away those flowers. But the flooding also disperses seeds downstream, allowing for new wildflower populations to establish in suitable habitat elsewhere. These shifting populations linked by movements between them are better characterized as a metapopulation (a “population of populations”) (Figure 11.8) consisting of several subpopulations. For some metapopulations, every subpopulation is transient: their distribution changes dramatically with each generation. Other metapopulations involve relatively permanent subpopulations with only a few individuals dispersing each generation. Some metapopulations consist of one or more source populations whose sizes are stable or increasing, and several sink populations whose sizes fluctuate depending on environmental conditions. Some sink subpopulations may undergo such dramatic fluctuations that they would be extirpated in unfavourable years were it not for population rescue by immigrants from source populations.
A metapopulation (a “population of populations”) consists of several subpopulations linked by movements of individuals between them.
Habitat fragmentation threatens metapopulation dynamics by reducing opportunities for dispersal across the landscape (Chapter 5). When there is too little movement of individuals between habitat fragments, the dwindling subpopulations within those fragments are at risk of extirpation or even extinction (Section 8.7). In contrast, well-connected subpopulations maintain themselves by colonising empty niches, exchanging genetic material, and adapting to changing environments. Dispersal also maintains critical ecosystem processes, such as pollination and seed dispersal (Section 4.2.5). Consequently, conservation biologists have invested significant resources in recent years to maintain and restore wildlife movements within fragmented ecosystems.
Connectivity in terrestrial ecosystems
Maintaining and restoring ecosystem connectivity—the ability of ecosystems to facilitate the dispersal of individuals between different areas—involves maintaining and restoring wildlife movements that are (at risk of being) impeded by human activities. The most popular method to maintain (or restore) connectivity in a fragmented landscape is to maintain (or restore) habitat linkages, also called wildlife corridors, habitat corridors, dispersal corridors, or movement corridors. All these terms refer to continuous tracts of suitable habitat with little to no dispersal barriers that connect otherwise isolated habitat patches and populations.
Maintaining and restoring ecosystem connectivity is an important strategy for conserving wildlife whose movements are impeded by human activities.
Some of the most prominent efforts to restore habitat linkages involve habitat restoration. For example, plans are currently underway to use forest regeneration to reconnect nine forest fragments in Tanzania’s East Usambara Mountain; if successful, this project would establish the largest contiguous forest block (over 3,000 km2) in the Eastern Arc Mountain Biodiversity Hotspot (Newmark, 2008). The positive impact of this project is expected to be immense. It has been estimated that the restoration of just 80 km2 of forest would stave off the first fragmentation-induced extinctions by over 2,000 years, compared to an estimated seven years until the first extinction if these forest fragments were to remain unconnected (Newmark et al., 2017).
Connectivity is important in every ecosystem on Earth. However, given the linear characteristic of riparian zones along rivers and stream—and hence a larger proportional impact of edge effects (Section 5.1.2)—we might consider connectivity in these spatially restricted systems to be particularly important (Figure 11.9). Protecting and restoring riparian zones as habitat linkages resonates with a variety of people because these areas provide a range of important ecosystem services, including flood control and water purification (Section 4.2.4). Conservationists can tap into this energy by lobbying for laws that prohibit activities such as logging, housing, and industrial developments within a certain distance from a river or stream. By protecting ecosystem services associated with riparian zones, these laws simultaneously also maintain wildlife refuges (Monadjem and Reside, 2008), source populations (Vosse et al., 2008), and habitat linkages (Bentrup et al., 2012; McLennan and Plumptre, 2012). In contrast, inadequate protection of riparian ecosystems not only compromises connectivity, but also negatively affect species not overtly dependent on these buffer areas. For example, research from Southeast Asia has shown that losing riparian ecosystems in an otherwise palm oil dominated landscape reduced stream quality, which in turn reduced local fish diversity by up to 36% (Giam et al., 2015). In contrast, protecting riparian zones were found to increase palm oil yields (Horton et al., 2018). With so many riparian areas currently being degraded and destroyed, there is an urgent need for stronger riparian protection laws (Chapter 12), and for more effective enforcement of those laws.
Restoring connectivity may also involve removing or otherwise mitigating human constructs that block wildlife dispersal. This is a major aim of TFCAs, which aim to restore dispersal between protected areas (Jones et al., 2012) by removing fences and other human constructs while still maintaining sustainable land tenures (Andersson et al., 2013). These efforts, accomplished through partnerships with local communities, are re-establishing historical mass migration routes, which in turn will hopefully also boost those areas’ ecotourism potential (Box 11.3). Efforts to revive extinct mass migrations also seem to be paying off! For example, in Botswana, the removal of veterinary fences—meant to prevent spread of diseases from wildlife to livestock, but also cutting off the world’s second largest wildebeest migration—have seen several hundred plains zebras (Equus quagga, NT) returning to old migration routes within four years (Bartlam-Brooks et al., 2011).
Simon M. Munthali
Kavango-Zambezi Transfrontier Conservation Area Programme,
TFCAs are components of a larger ecosystems that straddles the border between two or more countries, encompassing one or more protected areas as well as multiple-resource areas used by communities and private landholders. They are also managed for sustainable use of natural resources (Singh, 1998). The concept recognizes that borders are political rather than ecological (Dallimer and Strange, 2015), and aims to ensure that key ecological processes continue to function where political borders have divided ecosystems, river basins, or wildlife corridors (Cumming, 1999).
TFCAs are widely being established in Africa. One of these is the 520,000 km2 Kavango-Zambezi TFCA (KAZA)—a conservation and development initiative of Angola, Botswana, Namibia, Zambia, and Zimbabwe.
The benefits of the KAZA include:
- Re-establishment of the seasonal wildlife migration routes and connectivity among the many protected areas (national parks, community conservancies, and wildlife and forest reserves) within the region (Figure 11.C). The primary wildlife focus is the savannah elephant (Loxodonta africana), whose population of about 250,000 is predominantly concentrated in Chobe National Park (Botswana), Hwange National Park (Zimbabwe), and Bwabwata National Park (Namibia). Elephants need unimpeded movement to protected areas where population densities are much lower, such as Luengue-Luiana and Mavinga National Parks (Angola), and Sioma Ngwezi and Kafue National Parks (Zambia). This movement would reduce pressure on the ecosystems that are currently overpopulated and enable elephants and other species to better coexist—especially grazing herbivores that depend on the same habitats as the elephant.
- Expanding the wildlife-based economy, primarily ecotourism, into agricultural marginal areas (with predominantly Kalahari sand soils), through community-private partnerships. Through these partnerships, local communities would benefit from employment and business opportunities in ecotourism activities.
- Opportunities for local communities to participate in decision-making, and influencing policies and legislation related to natural management such as coordination of the fishing closed season between Namibia and Zambia during the fish breeding season (December–March) in the Zambezi River.
- Formation of alliances among different stakeholders (governments, private sector, NGOs, and local communities) to maximise skills and resources in promoting sustainable land use, conserving biodiversity and alleviating poverty.
Despite these benefits, there are obstacles to progress in attaining the benefits of the KAZA. Notable among these are social and political factors, such as increasing human population density, increasing cultivation of land, and expanding human settlements in wildlife corridors. Many of these factors trigger human-wildlife conflicts and poaching both for local consumption of bushmeat and for the illegal sale of elephant ivory. To mitigate these threats, the following strategies are being implemented:
- A Master Integrated Development Plan for the KAZA has been developed, which provides initial zoning. Its key feature is spatially allocating land into various uses (human settlement, agriculture, and protected wildlife areas, including wildlife dispersal corridors). The Master Integrated Development Plan also assists in creating awareness about the value of the wildlife corridors, which traverse communal areas.
- Promotion of conservation agriculture as a tool for improving land stewardship, intensification of agriculture, and improving crop yields per unit area of land, and therefore decreasing the likelihood of cutting down forested areas in and around wildlife corridors to plant new agricultural fields. Currently, within the KAZA, conservation agriculture is being piloted in Angola, Namibia, and Zambia. Conservation agriculture is crop production that strives to achieve acceptable profits together with high and sustained production levels while concurrently conserving the environment.
- Promotion of community-private partnerships in ecotourism development. Over the past four years, Ngoma safari lodge (Botswana), and Machenje sport fishing lodge (Zambia) have been developed specifically in support of securing wildlife corridors. They also provide incentives to the local communities for adopting wildlife conservation as a supplement to their land use practices. These lodges are in addition to the numerous existing tourist resorts in the KAZA.
- A law enforcement and anti-poaching strategy for the KAZA is being developed to coordinate transboundary law enforcement surveillance and fines to prevent poaching of protected wildlife. In addition, KAZA partner countries are integrating other security agencies, such as the military, police, immigration, and customs officials, to prevent the illegal export of wildlife products such as elephant ivory and bushmeat out of the KAZA.
- Reducing human-wildlife conflicts (Section 14.4) through improved land use planning, solar-powered electrified fencing encircling clusters of village fields and facilities and use of chilli-pepper-based olfactory repellents to deter elephants from entering crop fields.
The KAZA has made considerable progress to date in coordinating conservation efforts among the wildlife agencies and national parks across five countries in Southern Africa. The principal success has been measures to allow the continued migration of elephants along existing migration routes across international borders. The challenges ahead—from inadequate funding for wildlife patrolling and anti-poaching activities to increasing populations of rural people outside the protected areas and across migration routes—remain significant.
Section 5.1.1 discussed how inconsiderate fence placements threaten wildlife, while the paragraph above explained how removing fences can improve connectivity. Ironically, and illustrating the difficulties conservationists face when dealing with conflicting demands, strategically placed fences can sometimes also be used as a conservation tool. For example, researchers working on a fragmented lion population in Botswana found that the most effective way to improve this population’s viability was through strategic placement of fences to direct dispersal between protected areas (Cushman et al., 2016). Strategically placed predator-proof fences may at times also be required to avoid human-wildlife conflict (Packer et al., 2013, but see Creel et al., 2013), and to facilitate the recovery of threatened species, as is the case for Africa’s rarest antelope, the hirola (Beatragus hunter CR) (Ng’weno et al., 2017). The final word here is that management must remain responsive to both positive and negative impacts of tools, such as fences, rather than relegating them to bins, such as good or bad. (See also Dupuis-Desormeaux et al.,  for the use of fence-gaps and exclusionary fences to mitigate some negative fence impacts.)
Protecting and restoring stepping stone habitats can maintain connectivity in areas where it is impractical to establish or restore continuous habitat linkages.
At times, when it is impractical to establish or restore continuous habitat linkages, biologists may opt to protect and restore stepping stone habitats (Figure 11.10). As the name implies, stepping stone habitats are a special type of habitat linkage that facilitate dispersal along a patchwork of isolated habitat patches within a matrix of unsuitable or inhospitable habitat. Stepping stones thereby divide long dispersal events through a long stretch of inhospitable terrain up into shorter, and thus more manageable, sections. Stepping stone habitats are particularly important for migratory species that rest and refuel at stop-over sites between the end-points of their migratory route (Runge et al., 2015)—each stop-over site can be viewed as a stepping stone habitat. Prominent examples of stepping stone habitats that deserve protection include sacred forests which can act as stop-over sites for migratory forest birds; wetlands and estuaries (see Box 5.3), which can act as stop-over sites by migratory waterbirds; and small forest reserves, which can act as stepping stones between a network of other protected areas (Riggio and Caro, 2017).
Connectivity in freshwater ecosystems
Dams have always played an important role in hydropower generation and securing a year-round supply of water for farms, industries, and cities. Unfortunately, recent evidence suggests that reservoirs may create more problems than they solve (Section 5.3.2). Of concern is their contribution to greenhouse gases (Deemer et al., 2016), as well as their role in blocking dispersal of aquatic organisms. To counter these negative impacts, governments across the world are decommissioning and removing dams and other types of artificial water impoundments. For instance, over the past 30 years more than 1,174 dams were removed in the USA; the 72 dams removed in 2016 alone restored more than 3,000 km of streams (Thomas-Blate, 2016). Similar efforts are also underway in Europe (http://www.ecrr.org), where river restoration efforts have been initiated at over 1,100 locations across 31 countries. Unfortunately, not only are efforts to restore freshwater connectivity lagging across Africa; in many cases, even more rivers are currently being dammed (Winemiller et al., 2016).
While dams play an important role in hydropower generation and securing a year-round supply of water, recent evidence suggests that they create many environmental problems, including blocking species dispersal.
Connectivity in marine ecosystems
Ecosystem connectivity is also important in marine ecosystems. Many marine organisms, including economically important species, breed and feed in different areas at different times of the year, and use established dispersal routes to move between those areas. It is thus important to protect these dispersal routes so we can maintain these marine ecosystems and ecosystem services.
Maintaining movement dynamics in marine seascapes involves protecting and restoring marine corridors, estuarine linkages, and coastal habitat linkages.
There are three main strategies to maintain and restore movement dynamics of marine seascapes. First, marine corridors—zones used by whales and other marine species to move between feeding and breeding grounds—should be protected. Marine biologists in several countries successfully reduced collisions between whales and ocean-faring vessels with minor adjustments to shipping lanes that previously crossed marine corridors (Silber et al., 2012). Second, estuarine linkages should be protected, and restored where needed. For example, biologists in South Africa restored the natural flow regime of the St Lucia Estuary, Africa’s largest estuarine lake, by removing dredge spoil in the estuary mouth (Nunes et al., 2018). Third, coastal habitat linkages—beaches and littoral shallows used by wildlife for dispersal, breeding, and feeding—need to be maintained. Studies from South Africa have highlighted how poor protection of connectivity pathways between coastal habitats can compromise these areas’ high levels of species richness and endemism (von der Heyden, 2009; Harris et al., 2014).
In the absence of habitat linkages, wildlife managers may be able to mimic dispersal dynamics by sporadically translocating a few individuals between subpopulations. Managing populations in this way may be a good alternative in cases where areas earmarked for translocations are too small to sustain a single viable population. Such is the case in South Africa, where conservation biologists occasionally move threatened predators within a small and fragmented protected areas network, where none of the areas are large enough to host a viable population on their own (see Box 8.3). Managing isolated and small populations so intensively nearly always requires sound underlying principles and extensive quantitative analyses (Chapter 9) for guidance.
Management considerations in connectivity conservation
While intuitively appealing, there are a few potential drawbacks to connectivity that conservation planners should consider when planning to establish new habitat linkages (reviewed in Haddad et al., 2014). Prominently, connecting historically isolated populations may lead to outbreeding depression, for example when populations with different local adaptations are connected. Habitat linkages may also act as bottlenecks that expose dispersing animals to greater risks of predation and enable pests and diseases to spread easier. Care must be taken to ensure that wildlife do indeed perceive the landscape “connected”; a habitat linkage that may look good to the human eye may in fact be perceived as inhospitable habitat to wildlife (Newmark, 2008). A recent study from the Americas has shown that the habitat quality of a single stepping stone habitat can determine whether a migration is successful or not (Gómez et al., 2017).
Although the benefits for reconnecting fragmented landscapes generally outweigh the drawbacks, it is important to carefully plan to avoid those drawbacks.
Although the benefits for connecting landscapes for conservation generally outweigh the drawbacks (Haddad et al., 2014), it is important to carefully plan to avoid those drawbacks. Genetic studies can be useful in both determining connectivity among populations (von der Heyden, 2009; Godley et al., 2010) and help researchers detecting potential deleterious factors, such as outbreeding depression (Figure 11.11, see also Frankham et al., 2011; Ralls et al., 2018). Modelling approaches that combine a target species’ movement limitations with radio tracking technologies (e.g. Godley et al., 2010) or remotely sensed environmental variables (e.g. Wegmann et al., 2014) could help to estimate whether a landscape is indeed connected. Much effort has also been invested in finding the optimal width of habitat corridors. For example, one study in lowland forests suggested that corridors that are 30–40 m wide might be adequate for migration of most species while corridors that are 200 m wide will be adequate for all species (Laurance and Laurance, 1999). This is useful guidance, but ecosystems vary, as do target species (Wilson et al., 2010; Pryke and Samways, 2012) and, thus, some corridors may need to be even wider.