V.1 Causes and Consequences of Species Extinctions Navjot S. Sodhi, Barry W. Brook, and Corey J. A. Bradshaw OUTLINE 4. Introduction 2. Extinction drivers 3. Extinction vulnerability 4..Consequences of extinctions 5. Conclusions The ive largest mass de-ffs in which 50-95% of species wee eliminated occurred during the Ordovicion {690-403 nilion years age yell, Devonian {617354 mya, Permian (39-280 myahTrissic (251-200 myal, and Cretoceous Ut mya periods, Most recent, human actions cope aly over the pat two centuries hove precipitated a globl exinton crisis o the “sith great extinction wave" come pantie tthe previous five: Increasing human populations Orr the last 50,000 years or so have tet measurable regatv ocprnts on bioiversit SLossaRY Allee effects. These factors cause a reduction in the rowth rate of small populations as they decline (cg, via reduced survival or ceproduetive success). ‘extinction. Extinction of one species triggers the loss of another species. : ‘tinction debt. This refers to the extinction of species ot populations long after habitat alteration. ‘tinction vertex. As populations decline, an insidious ‘mutual reinforcement occurs among biotic and abiotic processes driving population size downward toextinction, ‘tipation, This refers.to extinetion.of » population nailer than of an entie specs. insive species, These are nomindigends species in- troduced to areas outside of their natural range that have become established and have spread. megafauna. This refers to large-bodied (>44 kg) ani- ‘mals, commonly (but not exclusively) used to refer to the large mammal biota of the Pleistocene. minimum viable population. This is the number of in- dividuals in a population required to have a speci fied probability of persistence over a given period of ‘ime, 4, INTRODUCTION. J the Americas, charismatic large-bodied animals (megatasina) sch at sabertoothed css (Smifodon SPP) mammoth (Manmidhus spp) and gia proaed slots (Megalo jefferson) vanished fallowng hu san arsval rome 11,000-13,000 years apo. Stailas Tossa occurred’im Australia 43,000 years ago, ad in tany oceanic islands within few hundeed oars othe Srval of humans, Clasie examples of the lost of land endemice include the dodo (Rephus cuca) feom Maurtss, moat (e¢, Dinoris maximus) fron New Zealand and slephancbeds(Aepyoris moxie) from Madagascar. Megafaunal collage during de ny Pleistocene can largely be waved toa variety of epatve Inuman impacts, such ag ovetharvesting, balogiel ie vations, and habita tffiformation. - ‘The rate and extent #trempaf*mediated extinctions are debated, but theres genetal agreemen that fineion cates have soared over the pat few hasdeed year; largely as a result of aceleated habitat dee Trction following Enropean colonialism and the sb: Sequcm plobal expansion of the hutnan population during the eventeth century. Humans are ipliated director indvetly in the 400- to. 10,000 old ime eae the “natural” or “background” extncton fate thar normally occurs a a consequence of gaa Emviconmental change, newiy established compertiveCopyrighted Material interactions (by evolution or invasioy a i chance calamities such as fire storm oe ae Te current and future extinction rates are aie THE seca yer ae ined ug changes in the World Goiscémmrion Union's (Ce threat categories over time, Byssd on the globel Sesament ofall known species, some 31, 12, 0a NG, of known amphibian, bird, and mamnal specie spectively (by far the best-studied of al animal saan 3) we curently listed by the IUCN as under than just how many species are being lost each year i also hotly debated. Various estimate, mann ‘few thousand to more than 100,000 species being ex- tinguished every year, most without ever having been scientifically described. The mainly through the applicatioi relationships that vary substa ties and habitats. Despite sub: itis nevertheless certain that hi the structure and function of ravel, The past five great extinctions shared some inn. portant commonalities: (1) they caused a catastrophic loss of global biodiversity; (2) they unfolded rapidly (at least in the context of evolutionary and geological time); (3) taxonomically, their impact was not random. (that is, whole groups of related species were lost while other related groups remained largely unaffected); and (4) the survivors were often nor previously dominant evolutionary groups. All four of these features are rel- evant to the current biodiversity crisis. This sixth great extinction is likely to be most catastrophic in tropical regions given the high species diversity theré (more than two-thirds of all species) and the large, expanding human populations that threaten most species there as wel. The major “systematic drivers” of modern species loss are changes in land use (habitat loss degradation and fragmentation), overexploitation, invasive species, disease, climate change (global warming) connected to increasing concentration of atmospheric carbon di- oxide, and increases in nitrogen deposition. Mechan- isms for prehistoric (caused by humans >200 years ago) extinctions are likely to have been similar: over- hunting, introduced predators and diseases, and habi tat destruction when early people first arrived in virgin landscapes. nn Of various species~area intially among communi- stantial prediction error, uman actions are causing of natural systems to un- 2. EXTINCTION DRIVERS Some events can instantly eliminate all individuals of a particular species, such as an asteroid strike, a mas- sive volcanic eruption, or even a rapid loss of large areas of unique and critical habitat because of defor- estation, But ultimately, any phenomena that can cause large uncersainty comes - rently, and is projected to continue to be, * direct and indirect cause of reported extirpations) Fox Species Extinctions mortality rates t0 exc over a sustained period extinct. Such forces bs eed reproductive replacement can cause a species to become say act independently or syner. 6 difficult to identify a single Bistically, and it mayb species or disease age proving access to human hunters, bios ical conditions. As a result, any population to dwindle m: population to extinction. Evidence to date suggests that deforest jim- OF altering biophys- Y Process that causes a ay ultimately predispose that ation is cur- the prime example, it is predicted that up to 21% of Southeast Asian forest species will be lost by 2100 because of past and ongoing deforestation. Similar projections exist for biotas in other region: Overexploitation is also an important driver of ex tinctions among vertebrates and tends to operate syn- exgistically wich other drivers such as habitat loss, For example, roads and trails created to allow logging op- erations to penetrate into virgin forests make previ- ously remote areas more accessible to human hunters, who ean, in turn, cause the decline and eventual ex. tirpation of forest species. It is estimated that overex- ploitation is a major threat to at least one-thied of threatened birds and amphibians, with wildlife, cure rently extracted from tropical forests at approximately six times the sustainable rate. In other words, the quantity, and most likely the diversity, of human prey— both fisheries and “bush” (wild) meat—are rapidly diminishing. Megafauna—those species weighing in the tens to hhundzeds of kilograms—are among the most vulnera- ble to overexploitation. In genéral, a species? genera- + tion time (interval from birth to reproductive age) isa function of body mass (allometry), so larger, longer- lived, and slower-reproducing animal populations are generally unable to compensate for high rates of hat- vesting. Because slow-breeding large animals, such as apes, carnivores (c.g., the lion, Panthera leo), and Af- rican elephants (Loxodonta africana), are particularly vulnerable to hunting, the potential for population recovery in these animals over shore time scales is low. As an example supporting this generality, there is ev dence that 12 large vertebrate species have been trpated from Vietnam, primarily because of excessive hunting, within the past 40 years, The Steller’ sea cow (Hydrodamalis gigas), an aquatic herbivorous mam mal that inhabited the Asian coast of the Bering St® is the quintessential example of the rapid demise of #Copyrighted Material st Conservation Biology species as a result of overexploitation, Discovered in 1741, it became extinct by 1768 because of overhunt- ing by sailors, seal hunters, and fur traders, This species was hunted for food, its skin for making boats, and its subcutaneous fat for use in oil lamps. ‘The ccosystem and biological community changes precipitated by invasive species represent another Teading cause of biodiversity loss. Of 170 extinct spe cies for which causes have been identified reliably, invasive species contributed directly to the demise of 91 (54%). In particular, the rates of extinctions oc- curring on islands have been greatly elevated by the introduction of novel predators. Several ecological and life-history attributes of island species, such as their naturally constrained geogeaphic range, small popula tion sizes, and particular traits (e4g., lack of flight in birds or lack of thorns in plants) make island biotas vulnerable to predation from invading species. For example, the introduction of the brown tree snake (Boiga irregularis) shortly after World War Il wreaked havoc on the biodiversity of the island of Guam in the South Pacific, In all likelihood, tree snakes were dic rectly responsible for the loss of 12 of 18 native bird species, and they also reduced the populations of other vertebrates such as flying foxes (Menopysurariannus), mainly Because of the inability $4 the island's native species to recognize the novel predatot“as a threat Despite an annual expenditure of USS44.6 million for the management of this problem, tree snakes on Guam are still not under control, largely because of their ability to penetrate artificial snake barriers such as fences. The mosquito Culex quinquefasciatus was inad- vertently introduced to Hawaii in 1826, and the disease-causing, parasite (Plasmodiun relictum) it cat sries arrived soon after. Since then, ayian malaria’ (in conjunction with other threats) has been responsible for the decline and extinction of some 60 species of endemic forest birds on the Hawaiian Islands. Having evolved in the absence of the disease, Hawaiian bird species were generally unable to cope with the debili- tating effects of the novel parasite. However, more than 100 years after the establishment of the disease, some native thrushes (Myadestes spp.) are now show- ing resistance to the disease. Sadly, many of the re- maining species, especially forest birds in the family Drepanididae, are still vulnerable and are now re- stricted to altitudes where temperatures are below the therinal tolerance limits of the mosquito vector. Global warming is predicted to increase the altitudinal distri- bution of the mosquito, thus spelling doom for disease~ susceptible birds as mosquito-free habitats disappear. The most feasible method of reducing transmission of malaria is to. reduce or eliminate vector mosquito populations through chemical treatments and the elim- ination of larval habitats. __ Perhaps one of the most infamous examples of an invasion catastrophe occurred in the world’s largest freshwater lake—Lake Victoria in tropical East Africa, Celebrated for its amazing collection of over 600 endemic haplochromine (i.e., formerly of the genus Haplochromis) cichlid fishes (Family Cichlidae), the Lake Victoria cichlid community is perhaps one of the most rapid, extensive, and recent vertebrate radiations known. There is also a rich community of endemic noncichlid fish that inhabit the Lake, In addition to the threats posed to this unique biota by a rapid rise in fisheries exploitation, human density, deforestation, and agriculture during the past century, without doubt the most devastating effect was the introduction of the predatory Nile perch (Lates niloticus) in the 1950s. This voracious predator, which ca 2m in length, w 18 grow to more than, introduced from Lakes Albert and Turkana (Uganda and Kenya, respectively) to com- pensate for depleting commercial fisheries in Lake Victoria. Although the Nile perch population remained relatively low for several decades after its introduc- tion, an eventual population explosion in the 1980s caused the devastating direct or indirect extinction of 200-400 cichlid species endemic to the Lake as well as the extinction of several noncichlid fish species. Al- though many other threats likely contributed to the observed extinctions, including direct overexploitation and eutrophication from agriculture and deforestation leading to a change in the algal plankton community, there are few other contemporary examples of such a rapid and massive extinction event involving a single group of closely related species. Human-mediated climate change represents a po- tentially disastrous sleeping giant in terms of future biodiversity losses. Climate warming can affect species, in five principal ways: (1) alterations of species densi- ties (including altered community composition and structure); (2) range shifts, either poleward or upward in elevation; (3) behavioral changes, such as the phe- nology (seasonal timing of life cycle events) of migea- tion, breeding, and flowering; (4) changes in mor- phology, such as body size; and (5) reduction in genetic diversity that leads to inbreeding depression. A related threat for island and coastal biotas is the predicted loss of habitat via inundation by rising sea levels. Although, large fluctuations in climate have occurred regularly throughout Earth’s history, the implications of an- thropogenic global warming for contemporary biodi- versity are particularly pessimistic because of the rate of change and previous heavy modification of land- scapes by humans. Good empirical evidence for some of these effects is rare, and speculations abound, butthere are already many local or regional exam, and model-based predictions that seprer: ales that rapid climate change, acting in conccrtwnng Soe divers of species loss and habitat degradation sae cone of the most pressing conservation ise, global biodiversity faces over the coming concurs 7 ssible future crisis comes af ‘Monterverde (Costa Rica), 3 frog and toad species dis. appeared following synchronous popalesorenieee 1987, with most crashes linked to a rapid promentin warming and drying of the local climate. The local endemic golden toad 2 (Bufo periglenes) was one of high-profile casualties in this aren, fe hen oe ae gested that climate warming resulted in a retreat clouds and a drying of the mountain habitats, wealing amphibians more susceptible to fungal and’ parashe outbreaks. Indeed, the pathogenic chytrid fungus Bat trachochytrium dendrobatidis, which grows on aime phibian skin and increases mortality rates, hes been" implicated in the loss of harlequin frogs (Atelopus spp.) in Central and South America and reductions in other amphibian populations elsewhere. It is hypoth- esized that warm and dry conditions may stress am- phibians and make them more vulnerable to the fungal infection. where 40% (20 of 50) oF Irrespective of the reason for a population's decline from a large to small population size, unusual (and often random and detrimental) events assume promi- nence at low abundances. For instance, although competition among individuals is reduced at low den- sities and can induce a population rebound, a coun- tervailing phenomenon known as the “Allee effect” can act to draw populations toward extinction by (for in- stance) disrupting behavioral patterns that depend on numbers (e.g., herd defense against predators) or by genetic threats such as inbreeding depression. Small populations, dominated by chance events and Allee effects, are often considered to have dipped below their “minimum viable population” size. Thus, once a major population decline has occurred (from habitat loss, overexploitation, or in response to many other possible stressors), an “extinction vortex” of positive feed- back loops can doom species to extinction, even if the original threats have been alleviated. Further, many species may take decades to perish following habitat degradation. Although some species may withstand the initial shock of land clearing, factors such as the lack of food resources, breeding sites, and dispersers may make populations unviable, and they eventually succumb to extinction, This phenomenon evokes the Concept of living-dead” species, or those “commited to extinction.” The eygtualaless of such species is * referred to as the “extinction ,Jebt” caused by past Copyrighted Material | Species Extinctions 517 habitat loss. For example, even if net deforestation rates can be reduced or even halted, the extinction debt of remnant and secondary forest patches will see the extinction of countless remaining species over this interval. 3, EXTINCTION VULNERABILITY Certain life-history, behavioral, morphological, dnd physiological characteristics appear to make some spe- cies more susceptible than others to the extinction drivers described above. In general, large-sized species with a restricted distribution that demonstrate habitat specialization tend to be at greater risk of extinction from human agency than others within theit respective taxa (e.g., Javan rhinoceros, Rhinoceros sondaicius), especially to processes such as rapid habitat loss. Because of their high habitat specificity and/or low . population densities, rare spéties may'be mare.prone to extinction tian common species. The size of a species’ range is also a major determinant of its extinction proneness. Small ranges may make species more vul- nerable to stochastic perturbations, even if local abun- dance is high; for example, proportionally more passerines (perching birds) with relatively small geo- graphic ranges in the Americas are at risk of extinction than their more widely distributed counterparts, Such trends are worrisome because those species with shrinking ranges as @résult of adverse human activities become particularly »etmecglffe to other drivers such a¢ climate change. Habitat loss also reduces the patch sizes necessary for species requiring large home ranges, making them vulnerable to extinction from a loss of subpopulation connectedness, reduced dispersal ica- pacity, and the ensuing lower population viability. Larger-bodied vertebrates are considered to be more extinetion-prone than smaller-bodied ones when the threatening process unfolds rapidly or intensely. In- deed, threatened mammals are an order of magnitude heavier than nonthreatened ones. A common expla- nation for this trend is that body size is inversely cor- related with population size, making large-bodied an- imals less abundant and more vulnerable to chronic environmental perturbations (while, being buffered against short-term environmental fluctuations). The extinction proneness of large-bodied animals to human activities is further enhanced because of other corte lated traits, such as their requirement of large aft greater food intake, high habitat specificity, and lower reproductive rate. Large species can also be more vulnerable to bums" persecution such as hunting, whereas smaller species arc generally more vulnerable to habitat loss. It is i=” portant, however, to be cautious when constructiasCopyrighted Material ai Conserval Nn Biology. generalized rules regarding the role of bad size in the Extinction process, Because they have a slower repro- ductive rate, larger parrots are more vulnerable to overexploitation than smaller finches, despite fewer ‘numbers of the former being captured for the pet trade. However, some smaller species (e.g, white-eyes, Zos: terop: spp.) with small population sizes are also vul- nerable to extinction becatise of heavy harvest rates for the pet trade, suggesting that only when the threatening processes are approximately equivalent will the larger df two species being compared demonstrate a hightt risk of extinction. In addition to body size, other morphological characteristics affect extinction prone- ness. For instance, large investment in secondary sexual characteristics may render highly dimorphic species less adaptable in a changing environment or more at. tractive to specimen or pet-trade collectors. ‘When an environment is altered abruptly or sys- tematically ata rate above normal background change, or beyond the capacity of adaptation via natural se. Ietion, specialist species wth narrow ecologies niches often bear the brunt of progressively unfavorable conditions such habitat loss and degradation. For in. stance, highly specialized forestdependent taxa are acutely vulnerable to extinction following deforesta- tion and forest fragmentation. Possible mechanisms include reductions in breeding and feeding sites, in- creased predation, clevated soil erosion and nutrient loss, dispersal limitation, enhanced edge effects, and other stressors. Conversely, non-forest-dependent spe- cies or those that prefer open habitats are often better able to persist in disturbed landscapes and may even be favored by having fewer competitors or expanded ranges following deforestation, It is important to be aware that in relatively stable systems, evolution en- spenders the speciation of taxa that occupy all available niches so both specialist and generalist species can co- exist, As a result, the rapid pace of habitat and climate change renders specialization a modern “curse” in evolutionary terms, Foraging specialization is one mechanism that can compromise a species’ ability to persist in altered habitats. Many studies have shown that frugivorous and insectivorous birds are more extinction-prone than other avian feeding guilds, with the lack of year-round ‘access to fruiting plants in fragmented forests being the culprit for the former. A number of hypotheses have been proposed to explain the disappearance of insec- tivorous birds from deforested or fragmented areas. First, deforestation may impoverish the insect fauna and reduce selected insectivore microhabitats (eg-, dead leaves). Second, insectivores may be poor dis- ersers and have near-ground nesting habits, the latter ‘wait making them more vulnerable to nest predators Penetrating smaller forest fragments. Absence of some insectivorous bird species from small fragments may not be related to food scarcity; rather, it may result from their poorer dispersal abilities. The ability to disperse in birds and insects depends on morphological characteristics such as wing loading, and physiological restrictions such as intolerance to sunlight when mov- ing within the nonforested matrix landscape separat- ing fragments. As a result, poor dispersal ability may make certain species vulnéfable to extinction because they cannot.readily supplerient sink habitats (habi- tats in which populations cannot replace themselves), supporting otherwise unviable subpopulations, of colonize new areas. Because of poor dispersal abil. ity, patchy distributions, and generally low popula. tion densities, the genctic diversity of species in fragmented landscapes may be difficult to maintain, with the resulting inbreeding depression further re. ducing population size toward extinction. However, clear and quantitative dethonstrations of the role of life-history teaits in the exsuetionrocess of biotas are still rare, 4, CONSEQUENCES OF EXTINCTIONS ‘The extinction of certain species such as large preda- tors and pollinators may have more devastating eco- logical consequences than the extinction of others, Ironically, avian vulnerability to predation is often exacerbated when certain large predatory species be- come rarer in tropical communities. For example, al though large cats such as jaguars (Panthera onica) do not prey on small birds directly, they exert a limiting force on smaller predators such as medium-sized and small mammals (mesopredators), which become more abundant with the former species’ decline. The cor- ollary is that abundant mesopredators inflict an above- average predation rate on the eggs and nestlings of” small birds. Although this “mesopredator-release” hypothesis has been applied largely to mammals (e.g Australian dingoes, Canis lupus, suppressing foxes and cats; coyotes in California controlling cat abundance), the loss of large predatory birds such as the harpy eagle (Harpia harpyja) may have similar ecosystem effects. Similar mesopredator release has been demonstrated for the frst time in the marine environment, where the overexploitation of large pelagic sharks resulted in an increase in rays and skates that eventually suppressed commercially important scallop populations. Likewise, does the disappearance of a competitor result in the niche expansion and higher densities of subordinate species? This phenomenon has been observed between unrelated taxa—the extinction of insectivorous birds from scrub forests of West Indian islands correlatedCopyrighted Material with’ the subsequent higher biomass of competing Anolis lizards. Conservation biologists have traditionally focused on the study of the independent declines, extirpations, or extinctions of individual species while paying rela- tively less attention to the possible cascading effects of species coextinctions (e.g., hosts and their parasites). However, it is likely that many coextinctions between interdependent taxa have occurred, but most have gone ‘unnoticed in these relatively understudied systems. For example, an extinct feather louse (Columbicola ex- tinctus) was discovered in 1937, 23 years after likely coextinction with its host passenger pigeon (Ectopistes migratorius). Ecological processes disrupted by ex- tinction or species decline may also lead to cascading and catastrophic coextinctions. Frugivorous animals and fruiting plants on which they depend have a key interaction linking plant reproduction and dispersal with ‘animal nutrition. Thus, the two interdependent taxa are placed in jeopardy by habitat degradation. ‘Many trees produce large, lipid-rich fruits adapted for animal dispersal, so the demise of avian frugivores may have serious consequences for forest regeneration, even if the initial drivers of habitat loss and degradation are annulled. Essential ecosystem functions provided by forest invertebrates are also highly susceptible when species are lost after habitat loss and degradation. Acting as keystone species in Southeast Asian rainforests, figs rely on tiny (1-2 mm) species-specific wasps for their pollination. Some fig wasps may have limited dispersal ability, suggesting that forest disturbance can reduce wasp densities and, by proxy, the figs that they polli nate. Similarly, dung beetles are essential components of ecosystem function because they contribute heavily to nutrient-recycling processes, seed dispersal, and the reduction of disease Gsif"axsgeiated with dung accu- mulation. In Venezudla, heayjer dung beetles were more extinction-prone than lighter species on artifi- cially created forested islands, which predicts particu- larly dire ecosystem functional loss given the former group's greater capacity to dispose of dung. ‘Almost all flowering plants in tropical rainforests are pollinated by animals, and an estimated one-third of the human diet in tropical countries is derived from insect-pollinated plants. Therefore, a decline of forest- dwelling pollinators impedes plint reproduction not _ only in*forests but also in neighboring agricultural *. areas visited by'these species. Lowland coffee (Coffea canephora) is an important tropical cash crop, and it depends on bees for cross-pollination. A study in Costa Rica found that forest bees increased coffee yield by 20% in fields within 1 km of the forest edge. Between 2000 and 2003, the pollination services provided by Species Extinction 4 forest bees were worth US$60,000 to a 1100-ha ¢ : A forest patch as small as 20 ha located neae am can increase coffee yield and thus bring lant nomic benefits to the farmers. Such findings iff feo, the imperative of preserving native forest ney forestry systems to facilitate the travel by tes dependent pollinating insects. tty. 5. CONCLUSIONS | Although extinctions are a normal part of human modifications to the planet in the lag centuries, and pezhaps even millennia, have gy accelerated the rate at which extinctions ocaus fe? loss remains the main driver of extinctions, ti act synergistically with other drivers such as on! harvesting and pollution, and, in the future, clinsy change. Large-bodied species, rare species, and hie specialists are particularly prone to extinction asa. sult of rapid human modifications of the plane, tinctions can disrupt vital ecological processes sik pollination and seed dispersal, leading to cascads losses, ecosystem collapse, and a higher extinction rat overall. | evo FURTHER READING | Brook, Barry W., Navjot S. Sodhi, and Peter K. L. Ne! 2003 Catastrophic extinctions follow deforestation in Sng pore, Nature 424; 420-423. This is one of few pope reporting broad-scale extinctions driven by tropical de forestation, | Clavero, Miguel, and Emili Garcia-Berthou. 2005. Invasrt species are a leading cause of animal extinctions. Trend. Ecology and Evolution 20: 110. The article highlights! invasive species represent one of the primary thteds biodiversity, Dirzo, Rudolfo, and Peter J. Raven, 2003. Global ste biodiversity and loss. Annual Review of Environment Resources 28: 137-167. The article constitutes 4 msi review of the state of the modern global biodiverst its associated losses, Fegan, William F., and E, E, Holmes, 2006, Quang only anon Vortex. Ecology Letters 9: 51-60. Thi 8 only study yet to quantify the final phases of extn 1 vertebrates for which date of extinction was known, GN Red List of threatened species, Downlosd oa HF teow iucnredlist.org, This presents sm spt fee Of and reasons for a listed species’ conse™ Koh, Lian P,, Robert R, nat K. Colwell soit 2009. le Heather C. Pro: 5 1 Science 305: 1632-1634, This mo oh Science 305 . This models how loss Fes indirectly result in the extinction of 1Copyrighted Material 0 Conservation Biology Pim, Start L, and Peter Raven. 2000, Extinction by rombers. Nature 403: 843-845. The article summrices the ikly extent of biodiversity losis asa result of aman cites. Pounds, J. Alan, Marin R. Bustamante, Luis A. Coloma, ‘Jem A. Consuegra, Michael P.L. ogden, Pou. Foster, Enrique La Marca, Karen L, Masters, Andres Merino. ‘Viren, Robert Puschendost, Santiago R. Ron, G. Arturo Sunchez-Azofeifa, Christopher J. Stil, and Brace E. ‘Young. 2006. Widespread amphibian extineions from cpdemic disease driven by global warming. Nature 439, U6I-16T, The article provides evidence on the role of climate change in recent amphibian extinctions, Rickets, Taylor H, Gretchen C. Daly, Patl Rc Ehalch, and ‘C.D. Michener 2004. Economic value of topical fost wet to coffee production, Proceedings of the National Acad: femy of Sciences US.A. 34; 12579-12582. This work shows how the lose of ecosystem services can affect pol- ination of commercial ereps Rosser, Alisoa M., and Sue A. Manika: 2002, Over ‘exploitation and species extinctions, Conservation Biol: 8y 16: 584-586. This work provides a quantitating Overview of the extent of threat faced By birds and mammals from direct exploitation by people, Sckercioglu, Gagan H,, Gretchen C. Daly, and Paul R. Ehr- Hich. 2004. Ecosystem consequences of bied declines. ‘Proceedings ofthe National Atademy of Sciences USA, 301: 18042-18047, This article provides a framework for Z di ENVIRONMENTAL (NATURAL HABITATS) Stalus of taxa range easual Figure 1. Schematic representation of the phases between the introduction of an organism to a locality through human action, and its establishment and proliferation in n: ‘nvironments. The ability of a ; a he new environment 'us as an alien. This scheme objectively defines "casual", "naturalize 7.W i and "invasive" species [adapted cons from Richardson et al. (2002a)]. futu ‘There are two main groups of definitions of ‘Invasive’; This (|. inter 1. Those based on ‘biological/ecological’ principles a jective | exe fi ind more or less objectivé and measurable criteria - invasive species are Buber Fe ie species | ioe that produce reproductive offspring, ofte; nin very large numbers, are able |disperse considerable distances from parent populations, and thus have the potential to spread over a large area (Richardson et al., 2000b; Pysek et al., 2004a), The advantage of this definition Is that “invasive’ taxa can be defined, Using reasonably objective criteria, according to their position along the ‘naturalization-Invasion’ continuum (Richardson et al., 2000b; see Figure 1). Importantly, the definition infers no connotation of impact. The second category of definitians may be lumped under the Héading ~*~ + me ‘anthropocentric’ invasive species are thosé that are alien to the ecosystem under question, and whose introduction causes, or is likely to cause, economic or environmental harm to human health. This definition has been widely adopted in policies, including such influential ones as the Executive Order 13112 of 1999 signed by President Clinton in the USA (btta://wens,invasivespecies.uov/laws/execorter.html). The rationale for such a definition is usually stated as being the need to focus the attention of policy makers on the biggest problems. A substantial problem with this definition is that many impacts are not readilyemeasurable or, in the absence of objective criteria for comparing impacts of difgerent®pecies in an different ecosystems, assigned meaningful currency. At what’stage in an Invasion can an alien organism be considered to be causing ‘harm’? And, given the problem inherent in predicting which alien species are likely to invade (i.e. spread), how can we decide which species are likely to cause harm? Nonetheless, the definition adopted by GISP reads “Invasive slien species are non-native organisms that cause, or have the potential to cause, harm to the environment, economies, or human health", A most readable review of the debate in this regard, dealing with issues such as whether it is practical or desirable to include impact as an integral part of the definition of ‘invasive’ or not, and discrepancies between scientific and popular usage of the term is provided by Carlton (2002), The most intense and challenging debate on defining ‘alien’ and ‘invasive’ species hhas focused on plants. This is probably because there is generally much more precise geographical data available for plant species than for most other types of organisms. ° This is partly because most plants stand still and wait to be counted, allowing their Populations to be mapped accurately. They do not migrate. Evaluating geographical Fanges is generally much more difficult for animals, even for large, conspicuous species. Although the term ‘invasive’ has lost its precise meaning through convoluted usage, itis likely to remain in widespread use, especially in socio-political circles. Some prominent ecologists are abandoning the term ‘invasive’ as a scientific concept. Back ic Top i i ers, 7. What terminology should research scientists, policy makers, conservationists and quarantine staff use for practical applications in the future? i in the relevant forums and This question has yet to be debated with sufficient vigour in the re International conventions, Urgent attention Is required to harmonize concepts and develop international guidelines for practical application by polley makers, researchers, phytosanitary organizations, conservationists, etc. WhateverPractical use of concepts such ag ‘potentially invasive’ and ‘potential inannGently required. Thirdly, if impact is used os criterion for la lly causing f, invasive, objective parameters that define eng impact should be belling speci Used. "9 SPecieg 3" 8. References Abbott RJ, 1992. Plant i invasions, interspecific hybridisation and the evolution of ney Plant taxa. Trends in Ec ‘ology and Evolution, 7:401-405. Carlton JT, 1996, Biological invasions and cryptogenic species, Ecology, 77:1653. 1655, Carlton JT, 2002. Bioinvasion ecolo. 7 Leppakoski E, Gollasch S, Olenin Ss, Distribution, Impacts, and Management. Dordrecht, The Netherlands: Kluwer Academic Publishers, 7-19, Chittka L, Schiirk ens S, 2001. Successful invasion of a floral market. Nature, 411:653, Colautti IR, McIsaac HJ, 2004. a neutral terminology to define ‘invasive’ species. Diversity and Distributions, 10:135-141 Daehler CC, 2001. Two Ways to be an invader, but one Is more suitable for ecology: Bulletin of the Ecological Society of America, 82: 206, Daehler CC, Carino D, 2001. Hybri consequences. In: Lockwood 'y M, eds, Biotic Homogenization. The | Netherlands: Kluwer/Plenum Publishers,.B1:102.._ es | Daehler CC, Strong DR, 1994, Variable neeedustive output among clones of Spartin? alt emiflora (Poaceae) including San Francisco Bay, California: the influence ofIndicators of Global Climate Change b; ry Prof A. Balasubramanian Centre for Advanced Studies in Earth Science University of Mysore Mysore-6 Introduction: Global Climate Change refers to a change in the state of the elim: in the mean and/or the variability of its properties and that persists bserved since about 1950, Warming of the climate system is unequivocal, and since the 1950s, many ofthe observed chenrs sro unprecedented over decades to millennia. ‘The atmosphere and ocean have warmed, the amounts of snow and i mini r 1 , unts of snow and ice have diminished, sea tic ane connate ose pe a eave in ta been successively warmer at the Barthys surface than any preceding decade since 1850. Global rrecipitation change average i i ; Precipitation change averaged over global land areas since 1901 is low prior to 1951 and medium Ocean warming dominates the increase in energy stored in the elimate system, accounting for m than 90% ofthe energy accumulated between 1971 and 2010, Ooesn hen corneas oe on mi . increased more slowly during 2003 to 2010, Regions of high salinity where evaporation dominates have become more saline. Over the last two decades, the Greenland and Antarcsie iee shee have been losing mass, glaciers have continued to shrink almost worldwide, and Arctic ses ies and Northen Hemisphere spring snow cover have continued to decrease in extent. ‘The rate of sea level rise since the mid-19th century has been larger than the mean rate during the previous two millennia, Over the period 1901 to 2010, global mean sea level rose by 0.19 (0.17 to: 0.21] m. The atmospheric concentrations of carbon dioxide, methane, and nitrous oxide have increased to levels unprecedented in at least the last 800,000 years. Carbon dioxide concentrations have increased by 40% since pre-industrial times, primarily from fossil fuel emissions and secondarily from net land use change emissions. ‘The ocean has absorbed about 30% of the emitted anthropogenic carbon dioxide, causing ocean acidification. ‘What made to think? jodi anging, The global temperatures are rising, snow and rainfall stag act ot shite. We ety ee aisrp end more exteme climate events—like heavy rainstorms and unevenly-high temnperatGres. These are already taking place and we could ‘witness these within our life span. Scientists are highly confident that many ofthese observed, changes can be linked to the levels of carbon dioxide and other greenhouse g : qvhich have increased ytadeeof human activi ies. = How is the Climate Changing? ‘The Earth's al patterns are showing a lot of shifts. inthe 1¢ world population have added a gan ine pre This happened largely due to burning of After the Industrial Revolution which bei Igs—as well as by clearing into the atmosp! significant it of greenhouse gases into di .d power vehicl fossil fuels to generate electricity, heat and cool buildings, and pow AVE! forests for urbanisation. 7i ! - \ \ded to the atmosphere are carbon dioxide, mo dae aeaoes are emitted into the atmosphere, mn thn) ide to thousands of years, Per MANY eng : j as I The major greenhdus¥ gases that people nitrous oxide, and fluorinated gases. When thes there for long time periods, ranging from a deca i Such emissions seen in the past have affect our atmosphefe in the present day. hence, the cama future emissions will continue to increase the levels of these gases in our atmosphere for the foreseeable future. These are called as “Greenhouse gases” because they trap heat (energy) jp, | greenhouse in the lower part of the atmosphere. As more of these gases are added to thie atmosphere, more heat is trapped. This. extra heat leads g her air temperatures near the Earth’s surface, alters weather patterns, ant ae f fe temperature i the oceans. These observed changes affect people and the environment in imp tn ways, é ‘What is Climate Change? . a Climate change refers to any substantial change in measures of climate (such as temperature or precipitation) lasting for an extended period (decades or longer), Natural factors have caused the climate to change during the earlier periods of the Earth's geology history. But now, due to human activities the present day’ changes are being observe: ‘The observed facts are the following: » 1. Global Surface Temperatures have increased over the 20th Century 2. The Decline of Winter- Snow Cover and Ice extent has decreased 3. Global Average Sea level has risen and Ocean Heat content has increased 4. More Frequent and Intense extreme weather events 5. Evidence in Small Island Developing States 6. Other noteworthy changes in aspects of climate- precipitation (rainfall) 7. Climate induced changes: changes in at least 420 physical processes and biological specis, communities, 8. Anthropogenic issues compounding the climate change crisi: There are a number of human activities and practices (besides the burning of fossil fuels) are exacerbating the crisis posed by climate change. These include deforestation; pollution; unplanned development; social pressures, and unsustainable farming practices, Why do we bother about climate change Indicators? Indicators are observations or calculations that can be used to track conditions and trends. : One important way to track and communicate the causes and effects of climate change is through tt use of indicators. An indicator represents the state or trend of certain environmental or societal conditions over a given area and for a specified period of time. Change indicators related to climete which may be physical, ecological, or societal, can be used to understand how environmental | conditions are changing, assess risks and vulnerabilities, and help inform resiliency and planning climate impacts. and by various countries. Drivers of Climate Change: Radiating Forces(RF):le, in ang Sor at es Radiative forcing by a climate variable is a change in Earth’s ener reen incomit ins te nd cine ra tas ECO ery se ren nin sl other factors are held constant. Radiative Forcing (RF) is the measurement of the capacity of a gas other foreing agents to affect that energy balance, thereby contributing te elimmns change. To define it more simply, RF expresses the change in energy inthe atmosphere dus fo OWI ater i Radiative forcing is also called as climate forcing. It is the difference of insolatic i absorbed by the Earth and energy radiated back to space. Natural parr obama eee and processes that alter the Barth’s energy budget are drivers of climate changes Ratinsne teen y continues to be a useful tool to estimate, to a first order, the relative climate impacts (viz. relative slobal mean surface temperature responses) due to radiatively induced pertateeneae 1. Changes in Radiation Budgets: ‘The radiation budget of the Earth is a central element of the climate system. On average, radiative processes warm the surface and cool the atmosphere, which is balanced by the byérological cycle and sensible heating, Spatial and temporal energy imbalances due to radiates oo {stent heating produce the general circulation of the atmosphere and oceans, Anthropogenie itlinrce ‘on climate occurs primarily through perturbations of the components of the Earth radiator budget. te radiation budget at the Top Of The Atmosphere (TOA) includes the absorption of solar radiation Earth, determined gs the difference between the incident and reflected solar radiation atthe TOA, as well as the thaymatUtifgoing radiation emitted to space. The surface radiation budget takes ‘The impact analysis includes the following: 1. Radiative Forcing from Greenhouse Gases, 2. Anthropogenic Aerosols . 3. Radiative Forcing from Land Surface Changes Anthropogenic Aerosols: ~ The indirect effect of aerosols (a decrease in the precipitation efficiency, increase in cloud water content and cloud lifetime) is another potentially important mechanism for climate change. The Radiation Forcing(RF) due to aerosol-radiation interaction is scattering and absorption of shortwave and longwave radiation by atmospheric aerosols. Several different aerosol types from various sources are present in the atmosphere (see Section 8.2). Most of the aerosols primarily scatter solar radiation, but some components absorb solar radiation to various extents with BC as the most absorbing component. RF of aerosols in the troposphere is often calculated at the TOA because it is similar to tropopause values. An increase in the hygroscopic aerosol abundance may enhance the concentration of cloud condensation nuclei (CCN). Oe 2. Changes in Surface Radiation Budget: Surface Solar Radiation: a e Actosols ean directly attenuate SSR by scattering and absorbing solar radiation, or indirectly, trough their ability to act as cloud condensation nuclei, thereby changing cloud reflectivity and lifetime. Surface Thermal and Net Radiation: é Thermal radiation, also known as longwave, terrestrial or far-IR ra atmospheric GHGs, temperature and humidity. n is sensitive to changes in Changes in Temperature : -. Land Surface Air Temperature: : ee | Global land-surface air temperature (LSAT) had increased over the instrumental period Of econ | with the warming rate approximately double that reported over the oceans since 1979, Diurnal Temperature Range: i 1. Land Use Change and Urban Heat Island Effects! | 2. Upper Air Temperature. . { Zhe term "land-use change” refers to a change in the use or management of land, Such change mt ‘eu from various human activities such as changes in agriculture and irrigation, deforestation” reforestation and afforestation, but also from urbanisation or traffic. Land-use change results in changing the physical and biological properties of the land surface and thus the climate system, Urbanisation is another kind of land-use change. This may affect the local wind climate through its» influence on the surface roughness. 3. Changes in Hydrological Cycle: | ~The main aspects of the hydrological cycle, including large-scale average precipitation, stream fibw and runoff, soil moisture, atmospheric water vapour, and clouds, Precipitation and Global Land Areas: Precipitation has generally increased over land north of 30°N over the period 1900-2005. | Streamflow and Runoff: The runoff and river discharge generally increased at high latitudes, with some exceptions. No consistent long-term trend in discharge was reported for the world’s major rivers on global scale. River discharge is unique among water cycle components in that it bath spatially antd ‘temporally | integrates surplus waters upstream within acatchment, ° - ‘ Evapotranspiration Including Pan Evaporation: Decreasing trends were found in records of pan evaporation over recent decades over the USA, India, Australia, New Zealand, China and Thailand and speculated on the causes including decreased | surface solar radiation, sunshine duration, increased specific humidity and increased clouds, Surface Humidity: o Widespread increases in surface air moisture content since 1976, alongéetth,sar-constant relative humidity over large seales though with some significant changes specific to region, time of day or season. In summary, itis very likely that global near surface air specific humidity has increased since the 1970s. Tropospheric Humidity: Observations from rdiofotide and GPS measurements over land, and satellite measurements over ocean indicate increases in'tropospheric water vapour at near-global spatial scales which are consistent with the observed increase in atmospheric temperature over the last several decades. Tropospheric water vapour plays an important role in regulating the energy balance of the surface, 4. Changes in Snowfall: | Statistically significant increases were found in most of Canada, parts of northern Europe and Russia. 4ow 5, Changes in Ocean Temperature and Heat Content: Effects of Sampling on Ocean Heat Content Estimates includes: 1. Upper Ocean Temperature 2. Upper Ocean Heat Content 3. Deep Ocean Temperature and Heat Content 4, Sea Surface Temperature 5. Marine Air Temperature 6. Global Combined Land and Sea Surface Temperature, 6. Changes in Salinity and Freshwater Content : 1. Global to Basin-Scale Trends 2. Regional Changes in Upper Ocean Salinity 3.” Evidence for Change of the Hydrological Cycle from Salinity Changes 7. Changes in Ocean Surface Fluxes: Air-Sea Heat Fluxes ede eur I i ‘Turbulent Heat Fluxés and Evaporation . Surface Fluxes of Shortwave and Longwave Radiation ‘Net Heat Flux and Ocean Heat Storage Constraints Ocean Precipitation and Freshwater Flux. Wind Stress. SPaYNe 8. Changes in Water-Mass Properties & Changes in Surface Waves: 1. Intermediate Waters 2. Deep and Bottom Waters 9. Changes in Ocean Circulation: 1. Global Observations of Ocean Circulation Variability 2. Wind-Driven Circulation Variability in the Pacific Ocean 3. The Atlantic Meridional Overturning Circulation 4. The Antarctic Meridional Overturning Circulation S. Water Exchange Between Ocean Basins 10. Sea Level Change, Including Extremes 1. Trends in Global Mean Sea Level and Components 2. Regional Distribution of Sea Level Change | 3. Assessment of Evidence for Accelerations in Sea Level Rise 4, Changes in Extreme Sea Level. 11. Ocean Biogeochemical Changes: 1. Carbon - : 2. Anthropogenic Ocean Acidification 3. Oxygen 4. Nutrients.12. Changes in Extremes : Hl 1, Temperature Extremes, 2. Extremes of the Hydrological Cycle. 1 3. Precipitation Extremes, 4, Floods, ' 5. Droughts. i 6 Severs Local Weather Events-severe thunderstorms or hailstorms. I 7, Tropical Storms, 8. Extratropical Storms. | 13. Overall Climate Change Indicators: Climate change can lead to other effects on the Earth’s physical system that are also indicators of climate change. Such integrative indicators include changes in sea level (ocean warming + land melt), in ocean acidification (ocean uptake of CO2) and in the amount of ice on ocean and lang (temperature and hydrological changes). | There are many indicators of climate change. These include physical responses such as changes in the following: 14, Ozone: = ae Stratospheric Ozone/TropsSpheric Ozon * Tropospheric ozone is a short-lived trace gas that either originates in the stratosphere or is produced situ by precursor gases and sunlight. Tropospheric ozone also impacts human health and vegetation at the surface. Its average atmospher lifetime of a few weeks produces a global distribution highly variable by season, altitude and lovati Ozone-Depleting Substances (Chlorofluorocarbons, Chlorinated Solvents, and Hydrochlorofluorocarbons): : a5 CFC atmospheric abundances are decreasing “becalise of the successful reduction in emissions resulting from the Montreal Protocol. 15. Changes in Atmospheric Composition: Greenhouse Gas Concentrations: Gases of most concern are, CO2, CH, and N;O. Well-Mixed Greenhouse Gases: «Increasing atmospheric burdens of well-mixed GHGs resulted in a 9% increase from 1998 to 2005. Since 2005, the atmospheric abundances of many well-mixed GHG increased further. Emissions are predominantly from surface sources, for CO», CH,, and N30. Kyoto Protocol Gases (Carbon Dioxide, Methane, Nitrous Oxide, Hydrofluorocarbons, Perfluorocarbons and Sulphur Hexafluoride). The main contributors to increasing atmospheric CO, abundance are fossil fuel combustion and and use change. i From 1980 to 2011, the average annual increase in globally averaged,CO; (ftom 1 January in one year to 1 January in the next year) was 1.7 ppm yr-ls.’ ‘The CO; growth rate varies from year to Year, since 1980 the range in annual increase is 0.7 + += 0.1 ppm in 1992 to2.9* +- Q.1 ppm in 1998.Methane: . Globally averaged CH4 in 1750 was 722 + 25 ppb. In 2011, the global annual mean was 1803 +- 2 ppb. Nitrous Oxide: Globally averaged N;O in 2011 was 324.2 ppb, an increase of 5.0 ppb over the value reported for 2005. Measurements of NzO and its isotopic composition in firn air suggest the increase, at least since the early 1950s, is dominated by emissions from soils treated with synthetic and organic (manure) nitrogen fertilizer. Hydrofluorocarbons, Perfluorocarbons, Sulphur Hexafluoride and Nitrogen Trifluorid Currently, the largest emissions of HFC-23 are from East Asia. CF, and CaF (PFCs) have lifetimes of $0 kyr and 10 kyr, respectively, and they are emitted as by- products of aluminium production and used in plasma etching of electronics. CF has a natural lithospheric source. Carbon Monoxide, Non-Methane Volatile Organic Compounds and Nitrogen Dioxide: Emissions of carbon monoxide (CO), non-methane volatile organic compounds (NMVOCS) and NOx (NO+NOz) do not have a direct effect. Z 16. Aerosols: The presence of dense aerosol changes the visibility of sky and promote for global dimming and brightening. Aerosol from anthropogenic sources (j.e., fossil and biofuel burning) are confined mainly to populated regions in the Northern Hemisphere, whereas aerosol from natural sources, such as desert dust, sea salt, volcanoes and the biosphere, are important in both hemispheres and likely dependent on climate and land use change. 17. Surface Temperature : Global and Regional Surface Temperatures 18. Atmospheric Water Vapour: Stratospheric Water Vapour: Stratospheric HO vapour has an important role in the Earth’s radiative balance and in stratospheric chemistry. Increased stratospheric HO vapour causes the troposphere to warm and the stratosphere to cool , and also causes increased rates of stratospheric Os loss. 19. Precipitation: Farge-Scale Changes yi Pemepitation: . Bete : Stributi cipitation i ani or indicator. ‘Variation in frequency and distribution of precipitation is a major 20. Severe Events + ‘The least likely events in a statistical sense are called "extreme events" Extremo weather in one i be normal in another. In Fee ae wave) pd over longer periods, but much les fo extemes, Impacts of extreme il be fe ‘and society and may be destructive. events will be felt strongly by ecosystems y and m cae L