Biodiversity Gradient Hypothesis

Patterns in the past

The Latitudinal Diversity Gradient (LDG) is a noticeable pattern among modern organisms that
has been described qualitatively and quantitatively. It has been studied at various taxonomic
levels, through different time periods and across many geographic regions (Crame 2001). The
LDG has been observed to varying degrees in Earth's past, possibly due to differences in climate
during various phases of Earth's history. Some studies indicate that the LDG was strong,
particularity among marine taxa, while other studies of terrestrial taxa indicate the LDG had little
effect on the distribution of animals.

Hypotheses for pattern

Although many of the hypotheses exploring the latitudinal diversity gradient are closely related
and interdependent, most of the major hypotheses can be split into three general hypotheses.
1. Spatial/Area hypotheses
There are five major hypotheses that depend solely on the spatial and areal characteristics of the
tropics.
a. Mid-domain effect
Using computer simulations, Colwell and Hurtt (1994) and Willig and Lyons (1998) first pointed
out that if species’ latitudinal ranges were randomly shuffled within the geometric constraints of
a bounded biogeographical domain (e.g. the continents of the New World, for terrestrial species),
species' ranges would tend to overlap more toward the center of the domain than towards its
limits, forcing a mid-domain peak in species richness. Colwell and Lees (2000) called
this stochastic phenomenon the mid-domain effect (MDE), presented several alternative
analytical formulations for one-dimensional MDE (expanded by Connolly 2005), and suggested
the hypothesis that MDE might contribute to the latitudinal gradient in species richness, together
with other explanatory factors considered here, including climatic and historical ones. Because
"pure" mid-domain models attempt to exclude any direct environmental or evolutionary
influences on species richness, they have been claimed to be null models (Colwell et al. 2004,
2005). On this view, if latitudinal gradients of species richness were determined solely by MDE,
observed richness patterns at the biogeographic level would not be distinguishable from patterns
produced by random placement of observed ranges (Colwell and Lees 2000). Others object that
MDE models so far fail to exclude the role of environment at the population level and in setting
domain boundaries, and therefore cannot be considered null models (Hawkins and Diniz-Filho
2002; Hawkins et al. 2005; Zapata et al. 2003, 2005). Mid-domain effects have proven
controversial (e.g. Jetz and Rahbek 2001, Koleff and Gaston 2001, Lees and Colwell, 2007,
Romdal et al. 2005, Rahbek et al. 2007, Storch et al. 2006; Bokma and Monkkonen 2001, Diniz-
Filho et al. 2002, Hawkins and Diniz-Filho 2002, Kerr et al. 2006, Currie and Kerr, 2007). While
some studies have found evidence of a potential role for MDE in latitudinal gradients of species
richness, particularly for wide-ranging species (e.g. Jetz and Rahbek 2001, Koleff and Gaston
2001, Lees and Colwell, 2007, Romdal et al. 2005, Rahbek et al. 2007, Storch et al. 2006; Dunn
et al. 2007). Others report little correspondence between predicted and observed latitudinal
diversity patterns (Bokma and Monkkonen 2001, Currie and Kerr, 2007, Diniz-Filho et al. 2002,
Hawkins and Diniz-Filho 2002, Kerr et al. 2006).
b. Geographical area hypothesis
Another spatial hypothesis is the geographical area hypothesis (Terborgh 1973). It asserts that
the tropics are the largest biome and that large tropical areas can support more species. More
area in the tropics allows species to have larger ranges, and consequently larger population sizes.
Thus, species with larger ranges are likely to have lower extinction rates (Rosenzweig 2003).
Additionally, species with larger ranges may be more likely to undergo allopatric speciation,
which would increase rates of speciation (Rosenzweig 2003). The combination of lower
extinction rates and high rates of speciation leads to the high levels of species richness in the
tropics.
A critique of the geographical area hypothesis is that even if the tropics is the most extensive of
the biomes, successive biomes north of the tropics all have about the same area. Thus, if the
geographical area hypothesis is correct these regions should all have approximately the same
species richness, which is not true, as is referenced by the fact that Polar Regions contain fewer
species than temperate regions (Gaston and Blackburn 2000). To explain this, Rosenzweig
(1992) suggested that if species with partly tropical distributions were excluded, the richness
gradient north of the tropics should disappear. Blackburn and Gaston 1997 tested the effect of
removing tropical species on latitudinal patterns in avian species richness in the New World and
found there is indeed a relationship between the land area and the species richness of a biome
once predominantly tropical species are excluded. Perhaps a more serious flaw in this hypothesis
is some biogeographers suggest that the terrestrial tropics are not, in fact, the largest biome, and
thus this hypothesis is not a valid explanation for the latitudinal species diversity gradient (Rohde
1997, Hawkins and Porter 2001). In any event, it would be difficult to defend the tropics as a
"biome" rather than the geographically diverse and disjunct regions that they truly include.
The effect of area on biodiversity patterns has been shown to be scale dependent, having the
strongest effect among species with small geographical ranges compared to those species with
large ranges who are affected more so by other factors such as the mid-domain and/or
temperature.
 c. Species-energy hypothesis
The species energy hypothesis suggests the amount of available energy sets limits to the richness
of the system. Thus, increased solar energy (with an abundance of water) at low latitudes causes
increased net primary productivity (or photosynthesis). This hypothesis proposes the higher the
net primary productivity the more individuals can be supported, and the more species there will
be in an area. Put another way, this hypothesis suggests that extinction rates are reduced towards
the equator as a result of the higher populations sustainable by the greater amount of available
energy in the tropics. Lower extinction rates lead to more species in the tropics.
One critique of this hypothesis has been that increased species richness over broad spatial scales
is not necessarily linked to increased number of individuals, which in turn is not necessarily
related to increased productivity. Additionally, the observed changes in the number of
individuals in an area with latitude or productivity are either too small (or in the wrong direction)
to account for the observed changes in species richness. [6] The potential mechanisms underlying
the species-energy hypothesis, their unique predictions and empirical support have been assessed
in a major review by Currie et al. (2004).
The effect of energy has been supported by several studies in terrestrial and marine taxa.
d. Climate harshness hypothesis
Another climate-related hypothesis is the climate harshness hypothesis, which states the
latitudinal diversity gradient may exist simply because fewer species can physiologically tolerate
conditions at higher latitudes than at low latitudes because higher latitudes are often colder and
drier than tropical latitudes. Currie et al. (2004) found fault with this hypothesis by stating that,
although it is clear that climatic tolerance can limit species distributions, it appears that species
are often absent from areas whose climate they can tolerate.
e. Climate stability hypothesis
Similarly to the climate harshness hypothesis, climate stability is suggested to be the reason for
the latitudinal diversity gradient. The mechanism for this hypothesis is that while a fluctuating
environment may increase the extinction rate or preclude specialization, a constant environment
can allow species to specialize on predictable resources, allowing them to have narrower niches
and facilitating speciation. The fact that temperate regions are more variable both seasonally and
over geological timescales (discussed in more detail below) suggests that temperate regions are
thus expected to have less species diversity than the tropics.
Critiques for this hypothesis include the fact that there are many exceptions to the assumption
that climate stability means higher species diversity. For example, low species diversity is known
to occur often in stable environments such as tropical mountaintops. Additionally, many habitats
with high species diversity do experience seasonal climates, including many tropical regions that
have highly seasonal rainfall (Brown and Lomolino 1998).

2. Historical/Evolutionary hypotheses
There are three main hypotheses that are related to historical and evolutionary explanations for
the increase of species diversity towards the equator.
a. The historical perturbation hypothesis
The historical perturbation hypothesis proposes the low species richness of higher latitudes is a
consequence of an insufficient time period available for species to colonize or recolonize areas
because of historical perturbations such as glaciation (Brown and Lomolino 1998, Gaston and
Blackburn 2000). This hypothesis suggests that diversity in the temperate regions have not yet
reached equilibrium, and that the number of species in temperate areas will continue to increase
until saturated (Clarke and Crame 2003).
b. The evolutionary rate hypothesis
The evolutionary rate hypothesis argues higher evolutionary rates in the tropics have caused
higher speciation rates and thus increased diversity at low latitudes (Cardillo et al. 2005, Weir &
Schluter 2007, Rolland et al. 2014). Higher evolutionary rates in the tropics have been attributed
to higher ambient temperatures, higher mutation rates, shorter generation time and/or faster
physiological processes (Rohde 1992, Allen et al. 2006). Faster rates of microevolution in warm
climates (i.e. low latitudes and altitudes) have been shown for plants (Wright et al. 2006),
mammals (Gillman et al. 2009) and amphibians (Wright et al. 2010). Based on the expectation
that faster rates of microevolution result in faster rates of speciation, these results suggest that
faster evolutionary rates in warm climates almost certainly have a strong influence on the
latitudinal diversity gradient. More research needs to be done to determine whether or not
speciation rates actually are higher in the tropics. Understanding whether extinction rate varies
with latitude will also be important to whether or not this hypothesis is supported (Rolland et al.
2014).
c. The hypothesis of effective evolutionary time
This hypothesis assumes that diversity is determined by the evolutionary time under which
ecosystems have existed under relatively unchanged conditions, and by evolutionary speed
directly determined by effects of environmental energy (temperature) on mutation rates,
generation times, and speed of selection (Rohde 1992). It differs from most other hypotheses in
not postulating an upper limit to species richness set by various abiotic and biotic factors, i.e., it
is a nonequilibrium hypothesis assuming a largely non-saturated niche space. It does accept that
many other factors may play a role in causing latitudinal gradients in species richness as well.
The hypothesis is supported by much recent evidence, in particular the studies of Allen et al.
(2006) and Wright et al. (2006).

3. Biotic hypotheses
Biotic hypotheses claim ecological species interactions such as competition, predation,
mutualism, and parasitism are stronger in the tropics and these interactions promote species co-
existence and specialization of species, leading to greater speciation in the tropics. These
hypotheses are problematic because they cannot be the proximate cause of the latitudinal
diversity gradient as they fail to explain why species interactions might be stronger in the tropics.
An example of one such hypothesis is the greater intensity of predation and more specialized
predators in the tropics has contributed to the increase of diversity in the tropics (Pianka 1966).
This intense predation could reduce the importance of competition (see competitive exclusion)
and permit greater niche overlap and promote higher richness of prey. However, as discussed
above, even if predation is more intense in the tropics (which is not certain), as it cannot be the
ultimate cause of species diversity in the tropics because it fails to explain what gives rise to the
richness of the predators in the tropics.
Several recent studies have failed to observe consistent changes in ecological interactions with
latitude (Lambers et al. 2002, Hillebrand 2004). These studies suggest the intensity of species
interactions are not correlated with the change in species richness with latitude.