‘Natural laboratories’ in perspective: a

review of literature on theories of

island biogeography
By Adrian Wang Xinting, University of Oxford

Abstract

Islands are diverse, well-defined, and relatively uncomplicated, facilitating significant

progress in the study of ecological and evolutionary processes. MacArthur and Wilson’s
(1963) Equilibrium Theory of Island Biogeography (ETIB) was a revolutionary framework
that facilitated the understanding of the biogeographical processes in insular ecosystems.
Over time, more sophisticated theoretical models have sought to advance the field of island
biogeography, with Whittaker et al.’s (2008) General Dynamic Model (GDM) being a
distinguished example, linking island age with species richness. However, there are several
factors that may influence the accuracy and reliability of island studies, such as palaeo-
configurations, anthropogenic activity, and evolutionary syndromes. This literature review
serves as a synthesis of critiques from published scientific journals on the use of models in
insular biogeography.

Glossary

Endemism: The state of a species being native to a single defined geographic location.

Insular biota: A term used to refer to animal, plant, and other organism life relating to,
dwelling, or situated on an island.

Phenotypic divergence: A term which describes variation within the population in terms of
the observable characteristics of an organism that result from the interaction of its genotype
with the environment.

Adaptive radiation: The process in which organisms undergo rapid diversification from a
single ancestral species into a range of new species that exhibit ecological traits specialised to
different niches.

Species-area relationship: A term that describes the general pattern of increase in species
richness with an increasing area of a habitat, or of part of the area of observation.

1. Introduction

Islands are the ‘logical laboratories of biogeography and evolution’, with each insular habitat
being ‘an experiment awaiting the analysis of evolution and ecology’.

Edward O. Wilson, cited in Losos et al., 2010, p.7.

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Diverse, well-defined, and relatively uncomplicated, islands are mediums through which one
can study the diversification of species as they adapt to different landscapes (Kohler 2002a,
cited in Hennessy, 2018). Moreover, as they vary in features such as area and degree of
isolation, islands serve as scientific experimental systems that allow for the study of
evolutionary responses of community properties (Lomolino et al., 2006). This essay will
explore the utility of islands as natural laboratories for the study of ecological, evolutionary,
and biogeographical processes, through models that predict and explain endemism.

Serving as a literature review, it will systematically examine the temporal development of

island biogeographical models and their prominent features and characteristics.
Simultaneously, it surfaces potential weaknesses of these models that may influence the
accuracy and reliability of these findings, such as not taking into account palaeo-
configurations, anthropogenic activity, and evolutionary syndromes.

Figure 1. MacArthur and Wilson’s equilibrium theory of island biogeography model (1963)
which demonstrates how the number of species on an island is determined by the rates of
colonization/immigration and extinction at equilibrium. Available
at http://californiachannelislands.blogspot.com/2012/04/macarthur-and-wilson-ii-
archipelagos.html

2. Equilibrium Theory of Island Biogeography

The Equilibrium Theory of Island Biogeography (ETIB) proposed by MacArthur & Wilson
(1963) posits that the species richness of an island represents a dynamic equilibrium between

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opposing rates of immigration and extinction (see Figure 1). The ETIB is grounded on three
fundamental premises of insular biotas: a positive correlation in the species-area relationship,
species number decreasing with increasing isolation, and there being a continual turnover of
species over time (Lomolino et al., 2010). Consequently, it is supposedly able to predict the
relative rates at which islands of ‘different sizes and degrees of isolation should return to
equilibrium, in the event that the biota is perturbed’ (ibid, p. 524). For instance, a near-source
island would return to equilibrium sooner as compared to a more distant island of the same
size, due to a disparity in immigration rate, assuming a constant extinction mechanism for
both islands (ibid). Furthermore, the ETIB hypothesises that the size and relative isolation of
the island affects local endemism. Larger islands, with lower extinction rates, permit the
accumulation of endemics, while more isolated islands, through the ‘radiation zone’ effect,
encourage greater proportions of endemism (Borregaard et al., 2016).

However, critics of the ETIB highlight how it does not account for several crucial factors that
determine the evolutionary processes of diversification and extinction within remote islands
(Borregaard et al., 2016). Firstly, it is argued that ETIB utilises island area as a sweeping
indicator of islands’ ability to sustain its species richness. As a result, the model disregards
the disparities in habitat prerequisites for different species of organisms, which are crucial in
the carrying capacity of similar islands (Lomolino et al., 2010). This is evident in the
presence of ‘specialist’ species, whose habitat specifications may only be fulfilled on sizeable
islands could further facilitate the diversification in richness (ibid). Moreover, the origins of
an island biota may be difficult to determine without examining the ‘systematics and
historical distribution of the species that are present’, with existing species on an insular biota
resulting from a range of factors, such as ‘over-water dispersal from continents and other
islands, historical links to other islands and in situ speciation’ (ibid, p. 527). Finally, the
ETIB omits the presence of disturbances, which contributes to setting the population off
equilibrium. In the occurrence of major environmental disturbances such as volcanic
eruptions and hurricanes, insular populations may never achieve equilibrium, or momentarily
at most (Heaney, 2000, cited in ibid).

3. General Dynamic Model

A comparatively more integrated theory based on oceanic island biogeography was

consequently found in the General Dynamic Model (GDM) proposed by Whittaker et al.
(2008). By incorporating the precepts of the ETIB within an evolutionary time-scale, it
highlights the increasingly influential roles of within-island speciation and geological
dynamics (Whittaker et al., 2008). The GDM posits that the physical geographical attributes
of oceanic islands demonstrate predictable dynamics over evolutionary time-scales, inducing
developments in the evolutionary and ecological adaptations of species in island biotas
(Borregaard et al., 2016). As the islands age, volcanic activity diminishes and they lose
elevation by erosion and subsidence. This leads to the local extinction of species adapted to
high-elevation habitats and lower nutrient availability in surfaces and soil (Price & Clague,
2002). Effectively, the GDM argues that the biotas of islands are determined by the relative
rates of immigration, local extinction and speciation, in accordance with the phases of
emergence, development, and submergence of island development (Whittaker et al., 2008).

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Figure 2. ATT2 model (log(Area) + Time + Time2) that predicts the species-area-time
relationship in islands as presented in the GDM (Whittaker et al., 2008). It demonstrates
a unimodal relationship with time and a linear rise with increasing area.

In contrast to the ETIB, the GDM suggests that islands’ distinct, dynamic geology have a
considerable impact on their habitat complexity and speciation over time (Borregaard et al.,
2017). Biodiversity indices, such as species richness and endemism, are influenced through
developments in the area, elevation, and environment of the island as it ages. This is
manifested in a unimodal/ hump-shaped trend in the carrying capacity of the island over its
life cycle (see Figure 2). Moreover, Borregaard et al. (2016) posits that the evolutionary rates
are augmented through two mechanisms – the area, elevation and topographic complexity of
the island determine the overall size of the resource base, while the topographical complexity
of the island influences the likelihood of reproductive isolation of species populations within
the island (p. 808). In terms of diversity-dependent dynamics, the GDM asserts that an
environmental carrying capacity restricts the species richness that can be supported on an
island at each stage (Borregaard et al., 2017). The initial periods of the island’s ontogeny
entails the resource base surpassing the realized species richness, due to there being
insufficient time for in situ speciation to mature. This untapped ecological potential
stimulates adaptive radiation, promoting productivity before species richness reaches a
plateau (ibid). Eventually, the resource base declines and the island founders, as speciation
rates decline and extinction rates climb (Whittaker et al., 2008).

In addition, Borregaard et al. (2016) found that low immigration rates on remote islands
meant that realised richness will invariably remain below resource supply levels for much of
the life cycle of the island. This is evidenced by the phenotypic divergence present on remote
islands – an instance of a diversity-dependent response to unoccupied niche space – as
mainland species adapted to particular ecological opportunities have yet to immigrate (Losos
& Ricklefs, 2009, cited in Borregaard et al., 2016). Consequently, it implies that over much
of an island’s lifespan, the system is arguably not in a dynamic equilibrium, contrary to the
assumption implied by the ETIB.

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4. Not necessarily an ideal laboratory

Yet, the results from experiments conducted on islands may not be widely applicable nor
transferable. These ‘natural laboratories’ influence their taxa via evolutionary syndromes, as
insular environments induce ‘fundamental transformations in the character of insular species’
(Lomolino, 2010: 986). The loss of dispersal prowess, a propensity towards flightlessness,
and developing naïveté toward predators are examples of converging traits of insularization
(Whittaker et al., 2017). For instance, flightlessness in birds is manifested through an
increased emphasis on forelimbs, as a ‘direct response to reduced predation pressure in the
absence of terrestrial mammals and specialist bird-hunting avian predators’ (ibid, p. 5).
Consequently, these reversals in natural selection left many insular species especially
vulnerable to introduced vertebrate predators, causing the former to be threatened with
extinction.

Moreover, islands may not be apt natural laboratories due to anthropogenic

influence. Through the means of transport, urbanization, and agriculture, human presence
has significantly modified the structure and ecology of local species in the forms of insular
colonization, habitat fragmentation, and dispersal of invasive species (van de Geer et al.,
2017). For example, the introduction of commensal mammals, such as pigs (Sus domesticus)
and goats (Capra hircus), has had a major negative effect on vegetation and has been
associated with the extinction of native species in Macaronesia (Nogué et al., 2017). These
anthropogenic disturbances have been immense, with Steadman (2006) estimating that of the
initial 500 to 1600 species of flightless rails that once inhabited the Pacific Islands, only 16
remain currently. Furthermore, Nogué et al. (2017) posits that human colonization and the
resulting cultural activity may have even influenced evolutionary processes through genetic
divergence, causing the evolution of unique island species and subspecies. The Canarian
Egyptian vulture (Neophron percnopterus) was found to have colonised the Canary Islands
under anthropogenic facilitation through the ‘introduction of domesticated livestock, as
suitable food sources were previously lacking’ (Agudo et al., 2010, cited in Nogué et al.,
2017). As a result, van de Geer et al. (2017) has found that ‘longstanding models of island
biogeography may prove inadequate unless their conceptual domains are expanded to include
… the impact of human activities on each of these processes’ (p. 1003).

Alternatively, instead of island models, palaeo-configurations may be a better method for the
study of biogeographical processes. Norder et al. (2018) found that extreme archipelago
configurations that have prevailed during the Last Glacial Maximum (LGM) are ‘not
sufficient to explain endemism patterns on volcanic oceanic islands’ (p. 192). Instead, palaeo-
configurations from the Pleistocene, observable at intermediate sea levels, left a superior
imprint on single-island endemic richness sequences. Interglacial ice-melt and sea-level rise –
induced by climatic fluctuations – caused the depletion of ecosystems at higher elevations,
impacting extinction, speciation and biota size (ibid). Thus, a more extensive temporal scale
may be more apt in understanding communities’ relationships.

5. Conclusions

In conclusion, islands are model ecosystems for biogeographic processes, such as trophic
relationships and species dynamics. Inferences from scientific studies could further
understandings of human-environment interactions, especially in the field of biodiversity
conservation, amidst increasing anthropogenic impact. However, as MacDonald et al. (2018)
posits, there is a substantial range of factors that ‘operate simultaneously to structure species

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diversity in insular and fragmented systems’ (p. 2741). Given the significance of evolutionary
syndromes, human influence, and the recency of island configurations on insular speciation,
islands may not necessarily serve as models for optimal natural laboratories. Moreover,
Hennessy (2018) argues that designating archipelagos as ‘natural laboratories’ may be
controversial, as it intrinsically encompasses the ‘conservationist territorial and biopolitical
management of nature’ (p. 500). Future research on islands could seek to further understand
the evolutionary dynamics of insular biotas across a range of dissimilar archipelagos under
contemporary developments, such as climate change and pollution. It could also seek to
develop enhanced models that take into account anomalies or extremities. An instance of this
is the Small Island Effect, which occurs when island area falls below a certain threshold. It
involves an extensive disruption of the species-area relationship, with species being
determined by habitat instead (Triantis et al., 2006).

Theory Key Contributions Critiques

Species richness of an island varies
with area and isolation – a dynamic
equilibrium between opposing rates
of immigration and extinction. Establishes key foundations of island
ETIB Species richness increases with an biogeography. However, overly
(MacArthur & increasing area of a habitat. Greater simplistic, omitting factors such as
Wilson, 1963) habitat heterogeneity induces differences in habitat conditions and the
reduced extinction rates. manifestation of disturbances.
Simultaneously, richness decreases
with rising island isolation, due to
diminishing immigration rates.
Positive relationship between
species richness and geological age
of island, together with area. Widely applicable.
GDM Species richness increases linearly Given the theory’s focus on oceanic
(Whittaker et with island area, while islands, a potential area of future research
al., 2008) demonstrating a a unimodal could involve specific niches in insular
relationship with time. habitats.
Environmental variables influence
level of endemism in islands.
Table 1. Summary of key theories

6. Acknowledgement

I am extremely thankful for the thorough feedback and support of Dr Marc Macias-Fauria
and the two anonymous reviewers of this essay.

7. References

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