Allee effect | ConservationBytes.com

Perceptions on poverty: the rising Middle Class

16 03 2009

I’m being somewhat ‘lazy’ this week in that I have unfortunately less time to spend on pertinent blog posts than I’d like (lecturing, looming deadlines, that sort of thing). So, I start out this week’s posts with one of my favourite TED talks – Hans Rosling debunks myths about the developing world.

What’s the relevance to biodiversity conservation? I’ll admit, it may appear somewhat tangential, but there are a few important messages (both potentially good and bad):

1. POSSIBLE BENEFIT #1: The rising wealth in the developing world and associated reduction in family size may inevitably curb our human population growth rates;

2. POSSIBLE DISADVANTAGE #1: Rising wealth will necessarily mean more and more consumption, and as we know at least for tropical developing nations, resource consumption is killing biodiversity faster than anywhere else on the planet;

3. POSSIBLE DISADVANTAGE #2: As family wealth rises, so too do opportunities do opportunities for the Anthropogenic Allee effect (consuming rare species just because you can afford to do so);

4. POSSIBLE BENEFIT #2: Better health care associated with rising wealth and lower infant mortality might make education a higher priority, teaching more people about the necessity of safeguarding ecosystem services.

I’m not convinced the advantages will necessarily outweigh the disadvantages; regardless, Prof. Rosling’s amazing 20-minute presentation will both entertain and enlighten. I recommend it for a lunchtime sitting or that late-afternoon attention wain.

CJA Bradshaw

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Tags: biodiversity, conservation, development, Hans Rosling, poverty, science, TED
Categories : Allee effect, conservation, ecological literacy, environmental policy, habitat loss, harvest, human overpopulation, poverty, video

Rare just tastes better

11 02 2009

I had written this a while ago for publication, but my timing was out and no one had room to publish it. So, I’m reproducing it here as an extension to a previous post (That looks rare – I’ll kill that one).

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As the international market for luxury goods expands in value, extent and diversity of items (Nueno & Quelch 1998), the world’s burgeoning pool of already threatened species stands to worsen. Economic theory predicts that harvested species should eventually find refuge from over-exploitation because it simply becomes too costly to find the last remaining wild individuals (Koford & Tschoegl 1998). However, the self-reinforcing cycle of human greed (Brook & Sodhi 2006) can make rare species increasingly valuable to a few select consumers such that mounting financial incentives drive species to extinction (Courchamp et al. 2006). The economic and ecological arguments are compelling, but to date there has been little emphasis on how the phenomenon arises in the human thought process, nor how apparently irrational behaviour can persist. Gault and colleagues (2008) have addressed this gap in a paper published recently in Conservation Letters by examining consumer preferences for arguably one of the most stereotypical luxury food items, caviar from the 200-million-year-old sturgeon (Acipenser spp.).

Sturgeon (6 genera) populations worldwide are in trouble, with all but two of the 27 known species threatened with extinction (either Near Threatened, Vulnerable, Endangered or Critically Endangered) according to the International Union for Conservation of Nature and Natural Resources’ (IUCN) Red List of Threatened Species. Despite all 27 species also having strict international trade restrictions imposed by the Convention on International Trade in Endangered Species (CITES) (Gault et al. 2008), intense commercial pressure persists for 15 of these at an estimated global value exceeding US$200 million annually (Pikitch et al. 2005). The very existence of the industry itself and the luxury good it produces are therefore, at least for some regions, unlikely to endure over the next decade (Pala 2007). What drives such irrational behaviour and why can we not seem to prevent such coveted species from spiralling down the extinction vortex?

Gault and colleagues addressed this question specifically in an elegantly simple set of preference experiments targeting the very end-consumers of the caviar production line – French connoisseurs. Some particularly remarkable results were derived from presentations of identical caviar; 86 % of attendees of luxury receptions not only preferred falsely labelled ‘rarer’ Siberian caviar (A. baeri) after blind tasting experiments, they also scored what they believed was caviar from the rarer species as having a higher ‘gustative quality’. These high-brow results were compared to more modest consumers in French supermarkets, with similar conclusions. Not only were unsuspecting gourmands fooled into believing the experimental propaganda, subjects in both cases stated a preference for seemingly rarer caviar even prior to tasting.

The psycho-sociological implications of perceived rarity are disturbing themselves; but Gault and colleagues extended their results with a mathematical game theory model demonstrating how irrational choices drive just such a harvested species to extinction. The economic implications of attempting to curb exploitation as species become rarer when the irrationality of perceived rarity was taken into consideration were telling – there is no payoff in delaying exploitation as more and more consumers are capable of entering the market. In other words, the assumption that consumers apply a positive temporal discount rate to their payoff (Olson & Bailey 1981) is wrong, with the demographic corollary that total depletion of the resource ensues. The authors contend that such artificial value may drive the entire luxury goods market based mainly on the self-consciousness and social status of consumers able to afford these symbols of affluence.

The poor record of species over-exploitation by humans arising from the Tragedy of the Commons (Hardin 1968) is compounded by this new information. This anthropogenic Allee effect (Courchamp et al. 2006) provides a novel example mechanism for how small populations are driven ever-downward because low densities ensure declining fitness. Many species may follow the same general rules, from bluefin tuna, Napoleon wrasse lips and shark fins, to reptile skins and Tibetan antelope woollen shawls. Gault and colleagues warn that as the human population continues to expand and more people enter the luxury-goods market, more wildlife species will succumb to this Allee effect-driven extinction vortex.

The authors suggest that a combination of consumer education and the encouragement of farmed substitute caviar will be more effective than potentially counter-productive trading bans that ultimately encourage illegal trade. However, the preference results suggest that education might not promote positive action given that reluctance of affluent consumers to self-limit. I believe that the way forward instead requires a combination of international trade bans, certification schemes for ‘sustainable’ goods that flood markets to increase supply and reduce price, better controls on point-of-origin labelling, and even state-controlled ‘warning’ systems to alert prospective consumers that they are enhancing the extinction risk of the very products they enjoy. A better architecture for trading schemes and market systems that embrace long-term persistence can surely counteract the irrationality of the human-induced destruction of global ecosystem services. We just need to put our minds and pocketbooks to the task.

CJA Bradshaw

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Tags: biodiversity, conservation, science
Categories : Allee effect, bushmeat, captive breeding, caviar, conservation, conservation biology, cuisine, ecosystem services, environmental policy, exploitation, extinction, extinction vortex, fish, fisheries, harvest, Red List, sturgeon, threatened species

Just give them a lift

16 01 2009

One of the main problems in a rapidly changing world, whether that change arises from habitat loss, invasive species or climate change, is that often the pace of change is simply too fast for many species to keep up. History (both ‘deep-time’ and contemporary evidence) tells us this fact very clearly in the record of extinctions – species that have ‘slow’ life histories (i.e., those that mature late in their lives, produce few young and breed infrequently) are the most susceptible to extinction. More often than not, these tend to be the big organisms because the pace of life scales to body size nonlinearly (the so-called allometry of vital rates). The problem extends to evolution – when the pace of change happens faster than mutation and subsequent natural selection, you are unable to ‘evolve’ to the new environmental state fast enough. The end result – extinction.

So, can we help? Well, it’s fairly difficult to alter reproductive rates unless you do some assisted breeding programme (which generally don’t do much for the conservation status of a species) and you can’t really alter age at maturity or growth rates. You can stop or reverse habitat destruction, and you can translocate species in some circumstances.

So, in the case of climate change, if local conditions become too unbearable for a species (temperature, salinity, precipitation, etc.), just give them a lift to another spot where the new conditions suit! Sounds simple, but it could be rather difficult.

A relatively new Policy Forum piece in Science outlines how ‘assisted colonisation‘ could work for some species. The issues are many – most translocations fail for one reason or another (too few individuals moved, unforeseen predators or competitors, lack of appropriate habitat, etc.), but as we’ve seen the world over in the case of successful alien species, invasions can be remarkably successful (at least from the perspective of the invading species).

The key then is to think very carefully about which species to move and which to leave alone. Of course, generalist, highly adaptable and dispersed species probably don’t need the help, but restricted-range species or habitat specialists could really benefit from such action. You also run the risk of creating more problems than you solve (e.g., new invasive pests, disease introduction). However, a select group of species might just need this very assistance to persist given how much we’ve already change the biosphere, and how much more it will change due to shifting climate in the near future.

It’s controversial, but it could work in many circumstances. That’s why I’m adding this paper (Hoegh-Guldberg et al. – Assisted colonization and rapid climate change) to the Potential list.

CJA Bradshaw

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Tags: biodiversity, conservation, science
Categories : alien species, Allee effect, climate change, disease, invasive species, recovery, restoration

Classics: the Allee effect

22 12 2008

As humanity plunders its only home and continues destroying the very life that sustains our ‘success’, certain concepts in ecology, evolution and conservation biology are being examined in greater detail in an attempt to apply them to restoring at least some elements of our ravaged biodiversity.

One of these concepts has been largely overlooked in the last 30 years, but is making a conceptual comeback as the processes of extinction become better quantified. The so-called Allee effect can be broadly defined as a “…positive relationship between any component of individual fitness and either numbers or density of conspecifics” (Stephens et al. 1999, Oikos 87:185-190) and is attributed to Warder Clyde Allee, an American ecologist from the early half of the 20th century, although he himself did not coin the term. Odum referred to it as “Allee’s principle”, and over time, the concept morphed into what we now generally call ‘Allee effects’.

Nonetheless, I’m using Allee’s original 1931 book Animal Aggregations: A Study in General Sociology (University of Chicago Press) as the Classics citation here. In his book, Allee discussed the evidence for the effects of crowding on demographic and life history traits of populations, which he subsequently redefined as “inverse density dependence” (Allee 1941, American Naturalist 75:473-487).

What does all this have to do with conservation biology? Well, broadly speaking, when populations become small, many different processes may operate to make an individual’s average ‘fitness’ (measured in many ways, such as survival probability, reproductive rate, growth rate, et cetera) decline. The many and varied types of Allee effects can work together to drive populations even faster toward extinction than expected by chance alone because of self-reinforcing feedbacks (see also previous post on the small population paradigm). Thus, ignorance of potential Allee effects can bias everything from minimum viable population size estimates, restoration attempts and predictions of extinction risk.

A recent paper in the journal Trends in Ecology and Evolution by Berec and colleagues entitled Multiple Allee effects and population management gives a more specific breakdown of Allee effects in a series of definitions I reproduce here for your convenience:

Allee threshold: critical population size or density below which the per capita population growth rate becomes negative.

Anthropogenic Allee effect: mechanism relying on human activity, by which exploitation rates increase with decreasing population size or density: values associated with rarity of the exploited species exceed the costs of exploitation at small population sizes or low densities (see related post).

Component Allee effect: positive relationship between any measurable component of individual fitness and population size or density.

Demographic Allee effect: positive relationship between total individual fitness, usually quantified by the per capita population growth rate, and population size or density.

Dormant Allee effect: component Allee effect that either does not result in a demographic Allee effect or results in a weak Allee effect and which, if interacting with a strong Allee effect, causes the overall Allee threshold to be higher than the Allee threshold of the strong Allee effect alone.

Double dormancy: two component Allee effects, neither of which singly result in a demographic Allee effect, or result only in a weak Allee effect, which jointly produce an Allee threshold (i.e. the double Allee effect becomes strong).

Genetic Allee effect: genetic-level mechanism resulting in a positive relationship between any measurable fitness component and population size or density.

Human-induced Allee effect: any component Allee effect induced by a human activity.

Multiple Allee effects: any situation in which two or more component Allee effects work simultaneously in the same population.

Nonadditive Allee effects: multiple Allee effects that give rise to a demographic Allee effect with an Allee threshold greater or smaller than the algebraic sum of Allee thresholds owing to single Allee effects.

Predation-driven Allee effect: a general term for any component Allee effect in survival caused by one or multiple predators whereby the per capita predation-driven mortality rate of prey increases as prey numbers or density decline.

Strong Allee effect: demographic Allee effect with an Allee threshold.

Subadditive Allee effects: multiple Allee effects that give rise to a demographic Allee effect with an Allee threshold smaller than the algebraic sum of Allee thresholds owing to single Allee effects.

Superadditive Allee effects: multiple Allee effects that give rise to a demographic Allee effect with an Allee threshold greater than the algebraic sum of Allee thresholds owing to single Allee effects.

Weak Allee effect: demographic Allee effect without an Allee threshold.

For even more detail, I suggest you obtain the 2008 book by Courchamp and colleagues entitled Allee Effects in Ecology and Conservation (Oxford University Press).

CJA Bradshaw

(Many thanks to Salvador Herrando-Pérez for his insight on terminology)

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Tags: biodiversity, conservation, science
Categories : Allee effect, decline, ecology, extinction, extinction vortex, harvest, inbreeding depression, minimum viable population, population dynamics, population viability analysis, recovery

That looks rare – I’ll kill that one

12 12 2008

Here’s an interesting (and disturbing) one from Conservation Letters by Gault and colleagues entitled Consumers’ taste for rarity drives sturgeons to extinction.

I like caviar, I have to admit. I enjoy the salty fishy-ness and the contrast it makes with the appropriate selection of wine (bubbly or otherwise). I guess a lot of other people like it too, to the extent that worldwide sturgeon population’s have been hammered (all 27 species are listed in CITES Appendix I or II, and 15 species are still heavily exploited). Indeed, in the Caspian Sea from where 90 % of caviar comes, sturgeon populations have declined by 90 % since the late 1980s. Admittedly, I haven’t had sturgeon caviar very often, and I doubt I’ll ever eat it again.

Using a set of simple ‘preference’ experiments on epicurean (French) human subjects, Gault and colleagues found that when told that a particular type of caviar was rarer than the others (when in reality, they two choices were identical), these refined gourmets generally tended to claim that the rarer one tasted better.

This means that humans have a tendency to place exaggerated value on harvested species when they think they’re rare (in most instances, rarity is itself the result of over-exploitation by humans). This so-called ‘anthropogenic Allee effect‘ (see Courchamp et al. 2006) basically means that at least for the wildlife-based luxury market, there’s little chance that calls for reduced harvest will be heard because people continually adjust their willingness to pay more. This turns into a spiralling extinction vortex for the species concerned.

What to do? Ban all trade of caviar? This might do it, but with the reluctance to reduce highly profitable industries like this (see previous post on tuna over-exploitation here), there’s a strong incentive even for the harvesters to drive themselves out of a job. Consumer education (and a good dose of guilt) might help too, but I have my doubts.

CJA Bradshaw