Latitudinal gradients in species diversity

From Wikipedia, the free encyclopedia

Jump to navigationJump to search

Map latitudinal gradient of living terrestrial vertebrate species richness (Mannion

2014)
Species richness, or biodiversity, increases from the poles to the tropics for a
wide variety of terrestrial and marine organisms, often referred to as the
latitudinal diversity gradient (LDG).[1] The LDG is one of the most widely
recognized patterns in ecology.[1] The LDG has been observed to varying degrees in
Earth's past.[2] A parallel trend has been found with elevation (elevational
diversity gradient),[3] though this is less well-studied.[4]

Explaining the latitudinal diversity gradient has been called one of the great
contemporary challenges of biogeography and macroecology (Willig et al. 2003, Pimm
and Brown 2004, Cardillo et al. 2005).[5] The question "What determines patterns of
species diversity?" was among the 25 key research themes for the future identified
in 125th Anniversary issue of Science (July 2005). There is a lack of consensus
among ecologists about the mechanisms underlying the pattern, and many hypotheses
have been proposed and debated. A recent review [6] noted that among the many
conundrums associated with the LDG (or LBG, Latitudinal Biodiversity Gradient) the
causal relationship between rates of molecular evolution and speciation has yet to
be demonstrated.

Understanding the global distribution of biodiversity is one of the most

significant objectives for ecologists and biogeographers. Beyond purely scientific
goals and satisfying curiosity, this understanding is essential for applied issues
of major concern to humankind, such as the spread of invasive species, the control
of diseases and their vectors, and the likely effects of global climate change on
the maintenance of biodiversity (Gaston 2000). Tropical areas play prominent roles
in the understanding of the distribution of biodiversity, as their rates of habitat
degradation and biodiversity loss are exceptionally high.[7]

Contents
1 Patterns in the past
2 Hypotheses for pattern
2.1 Spatial/Area hypotheses
2.1.1 Mid-domain effect
2.1.2 Geographical area hypothesis
2.1.3 Species-energy hypothesis
2.1.4 Climate harshness hypothesis
2.1.5 Climate stability hypothesis
2.2 Historical/Evolutionary hypotheses
2.2.1 The historical perturbation hypothesis
2.2.2 The evolutionary speed hypothesis
2.2.3 The hypothesis of effective evolutionary time
2.2.4 The integrated evolutionary speed hypothesis
2.3 Biotic hypotheses
3 Synthesis and conclusions
3.1 The generality of the latitudinal diversity gradient
3.2 Data robustness
3.3 Conclusion
4 See also
5 References
Patterns in the past
The 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, particularly among marine taxa, while other
studies of terrestrial taxa indicate the LDG had little effect on the distribution
of animals.[2]

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.

Spatial/Area hypotheses
There are five major hypotheses that depend solely on the spatial and areal
characteristics of the tropics.

Mid-domain effect
Using computer simulations, Cowell and Hurt (1994) and Willing 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 (Cowell 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 called dinosures(Colwell and Lees 2000). Others object
that MDE models so far fail to exclude the role of the 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)[5][8] 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).

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.[5]

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 an increased number of individuals,
which in turn is not necessarily related to increased productivity.[9]
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.[9] 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).[10]

The effect of energy has been supported by several studies in terrestrial and
marine taxa.[7]

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)[10] 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.

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).

Historical/Evolutionary hypotheses
There are four main hypotheses that are related to historical and evolutionary
explanations for the increase of species diversity towards the equator.

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 has not yet reached equilibrium and that the
number of species in temperate areas will continue to increase until saturated
(Clarke and Crame 2003). However, in the marine environment, where there is also a
latitudinal diversity gradient, there is no evidence of a latitudinal gradient in
perturbation.

The evolutionary speed hypothesis

The evolutionary speed hypothesis[11] argues higher evolutionary rates due to
shorter generation times in the tropics have caused higher speciation rates and
thus increased diversity at low latitudes.[12] Higher evolutionary rates in the
tropics have been attributed to higher ambient temperatures, higher mutation rates,
shorter generation time and/or faster physiological processes,[13][14] and
increased selection pressure from other species that are themselves evolving.[15]
Faster rates of microevolution in warm climates (i.e. low latitudes and altitudes)
have been shown for plants,[16] mammals,[17] birds,[18] fish[19] and amphibians.
[20] Bumblebee species inhabiting lower, warmer elevations have faster rates of
both nuclear and mitochondrial genome-wide evolution.[21] 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.[22]

The hypothesis of effective evolutionary time

The hypothesis of effective evolutionary time 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.[14] 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.[13] and Wright et al.[23]

The integrated evolutionary speed hypothesis

The integrated evolutionary speed hypothesis argues that species diversity
increases due to faster rates of genetic evolution and speciation at lower
latitudes where ecosystem productivity is generally greater.[24] It differs from
the effective evolutionary time hypothesis by recognizing that species richness
generally increases with increasing ecosystem productivity[25][26][27] and declines
where high environmental energy (temperature) causes water deficits.[28] It also
proposes that evolutionary rate increases with population size, abiotic
environmental heterogeneity, environmental change and via positive feedback with
biotic heterogeneity. There is considerable support for faster rates of genetic
evolution in warmer environments,[24] some support for a slower rate among plant
species where water availability is limited [29] and for a slower rate among bird
species with small population sizes.[30] Many aspects of the hypothesis, however,
remain untested.

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 coexistence and specialization of species, leading to
greater speciation in the tropics. These hypotheses are problematic because they
cannot be the ultimate 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. Some recent large-scale experiments suggest
predation may indeed be more intense in the tropics,[31][32] although this cannot
be the ultimate cause of high tropical diversity because it fails to explain what
gives rise to the richness of the predators in the tropics. Interestingly, the
largest test of whether biotic interactions are strongest in the tropics, which
focused on predation exerted by large fish predators in the world's open oceans,
found predation to peak at mid-latitudes. Moreover, this test further revealed a
negative association of predation intensity and species richness, thus contrasting
the idea that strong predation near the equator drives or maintains high diversity.
[33] Other studies have failed to observe consistent changes in ecological
interactions with latitude altogether (Lambers et al. 2002),[1] suggesting that the
intensity of species interactions is not correlated with the change in species
richness with latitude. Overall, these results highlight the need for more studies
on the importance of species interactions in driving global patterns of diversity.

Synthesis and conclusions

There are many other hypotheses related to the latitudinal diversity gradient, but
the above hypotheses are a good overview of the major ones still cited today. It is
important to note that many of these hypotheses are similar to and dependent on one
another. For example, the evolutionary hypotheses are closely dependent on the
historical climate characteristics of the tropics.

The generality of the latitudinal diversity gradient

An extensive meta-analysis of nearly 600 latitudinal gradients from published
literature tested the generality of the latitudinal diversity gradient across
different organismal, habitat and regional characteristics.[1] The results showed
that the latitudinal gradient occurs in marine, terrestrial, and freshwater
ecosystems, in both hemispheres. The gradient is steeper and more pronounced in
richer taxa (i.e. taxa with more species), larger organisms, in marine and
terrestrial versus freshwater ecosystems, and at regional versus local scales. The
gradient steepness (the amount of change in species richness with latitude) is not
influenced by dispersal, animal physiology (homeothermic or ectothermic) trophic
level, hemisphere, or the latitudinal range of study. The study could not directly
falsify or support any of the above hypotheses, however, results do suggest a
combination of energy/climate and area processes likely contribute to the
latitudinal species gradient. Notable exceptions to the trend include the
ichneumonidae, shorebirds, penguins, and freshwater zooplankton.

Data robustness
One of the main assumptions about LDGs and patterns in species richness is that the
underlying data (i.e. the lists of species at specific locations) are complete.
However, this assumption is not met in most cases. For instance, diversity patterns
for blood parasites of birds suggest higher diversity in tropical regions, however,
the data may be skewed by undersampling in rich faunal areas such as Southeast Asia
and South America.[34] For marine fishes, which are among the most studied
taxonomic groups, current lists of species are considerably incomplete for most of
the world's oceans. At a 3° (about 350 
km2) spatial resolution, less than 1.8% of
the world's oceans have above 80% of their fish fauna currently described.[35]

Conclusion
The fundamental macroecological question that the latitudinal diversity gradient
depends on is "What causes patterns in species richness?". Species richness
ultimately depends on whatever proximate factors are found to affect processes of
speciation, extinction, immigration, and emigration. While some ecologists continue
to search for the ultimate primary mechanism that causes the latitudinal richness
gradient, many ecologists suggest instead this ecological pattern is likely to be
generated by several contributory mechanisms (Gaston and Blackburn 2000, Willig et
al. 2003, Rahbek et al. 2007). For now, the debate over the cause of the
latitudinal diversity gradient will continue until a groundbreaking study provides
conclusive evidence, or there is general consensus that multiple factors contribute
to the pattern.

See also
Asynchrony of Seasons Hypothesis
Biodiversity
Biomes
Effective evolutionary time
Evolution
Extinction
Latitude
Speciation
Species richness
Tropics
References
References
Hillebrand, H (February 2004). "On the generality of the latitudinal diversity
gradient" (PDF). The American Naturalist. 163 (2): 192–211. doi:10.1086/381004.
PMID 14970922. S2CID 9886026.
Sahney, Sarda; Benton, Michael J (7 April 2008). "Recovery from the most profound
mass extinction of all time". Proceedings of the Royal Society B: Biological
Sciences. 275 (1636): 759–765. doi:10.1098/rspb.2007.1370. PMC 2596898. PMID
18198148.
McCain, Christy M. (February 2005). "Elevational Gradients in Diversity of Small
Mammals" (PDF). Ecology. 86 (2): 366–372. doi:10.1890/03-3147. S2CID 37759984.
Archived from the original (PDF) on 2020-02-08.
Rahbek, Carsten (June 1995). "The elevational gradient of species richness: a
uniform pattern?". Ecography. 18 (2): 200–205. doi:10.1111/j.1600-
0587.1995.tb00341.x.
Mora, Camilo; Robertson, D. Ross (July 2005). "Causes of latitudinal gradients in
species richness: a test with fishes of the Tropical Eastern Pacific" (PDF).
Ecology. 86 (7): 1771–1782. doi:10.1890/04-0883. S2CID 35273509. Archived from the
original (PDF) on 2019-03-07.
Dowle, E. J.; Morgan-Richards, M.; Trewick, S. A. (2013). "Molecular evolution and
the latitudinal biodiversity gradient". Heredity. 110 (6): 501–510.
doi:10.1038/hdy.2013.4. PMC 3656639. PMID 23486082.
Tittensor, Derek P.; Mora, Camilo; Jetz, Walter; Lotze, Heike K.; Ricard, Daniel;
Berghe, Edward Vanden; Worm, Boris (August 2010). "Global patterns and predictors
of marine biodiversity across taxa". Nature. 466 (7310): 1098–1101.
Bibcode:2010Natur.466.1098T. doi:10.1038/nature09329. PMID 20668450. S2CID 4424240.
Mora, Camilo; Chittaro, Paul M.; Sale, Peter F.; Kritzer, Jacob P.; Ludsin, Stuart
A. (February 2003). "Patterns and processes in reef fish diversity". Nature. 421
(6926): 933–936. Bibcode:2003Natur.421..933M. doi:10.1038/nature01393. PMID
12606998. S2CID 4311686.
Cardillo, M.; Orme, C. D. L.; Owens, I. P. F. (2005). "Testing for latitudinal
bias in diversification rates: An example using New World birds". Ecology. 86 (9):
2278–2287. doi:10.1890/05-0112.
Currie, D. J.; Mittelbach, G. G.; Cornell, H. V.; Kaufman, D. M.; Kerr, J. T.;
Oberdorff, T. (2004). "Predictions and tests of climate-based hypotheses of broad-
scale variation in taxonomic richness". Ecology Letters. 7 (11): 1121–1134.
doi:10.1111/j.1461-0248.2004.00671.x.
Rensch, Bernhard (1959-03-02). Evolution Above the Species Level. Columbia
University Press. doi:10.7312/rens91062. ISBN 978-0-231-88186-9.
Rohde, Klaus (1992). "Latitudinal Gradients in Species Diversity: The Search for
the Primary Cause". Oikos. 65 (3): 514–527. doi:10.2307/3545569. ISSN 0030-1299.
JSTOR 3545569.
Allen, Andrew P.; Gillooly, James F. (August 2006). "Assessing latitudinal
gradients in speciation rates and biodiversity at the global scale". Ecology
Letters. 9 (8): 947–954. doi:10.1111/j.1461-0248.2006.00946.x. ISSN 1461-023X. PMID
16913938.
Rohde, Klaus (1992). "Latitudinal Gradients in Species Diversity: The Search for
the Primary Cause". Oikos. 65 (3): 514–527. doi:10.2307/3545569. ISSN 0030-1299.
JSTOR 3545569.
Schemske, Douglas W.; Mittelbach, Gary G.; Cornell, Howard V.; Sobel, James M.;
Roy, Kaustuv (December 2009). "Is There a Latitudinal Gradient in the Importance of
Biotic Interactions?" (PDF). Annual Review of Ecology, Evolution, and Systematics.
40 (1): 245–269. doi:10.1146/annurev.ecolsys.39.110707.173430. S2CID 53470632.
Archived from the original (PDF) on 2020-10-30.
Wright, Shane D.; Keeling, Jeannette; Gillman, Len N. (May 2006). "The road from
Santa Rosalia: a faster tempo of evolution in tropical climates". Proceedings of
the National Academy of Sciences. 103 (20): 7718–7722. doi:10.1073/pnas.0510383103.
PMC 1472511. PMID 16672371.
Gillman, Len N.; Keeling, D. Jeanette; Ross, Howard A.; Wright, Shane D. (2009-09-
22). "Latitude, elevation and the tempo of molecular evolution in mammals".
Proceedings of the Royal Society B: Biological Sciences. 276 (1671): 3353–3359.
doi:10.1098/rspb.2009.0674. PMC 2817169. PMID 19556254.
Gillman, Len N.; McCowan, Luke S. C.; Wright, Shane D. (September 2012). "The
tempo of genetic evolution in birds: body mass and climate effects: The tempo of
evolution in birds". Journal of Biogeography. 39 (9): 1567–1572.
doi:10.1111/j.1365-2699.2012.02730.x. S2CID 83114083.
Wright, Shane D.; Ross, Howard A.; Jeanette Keeling, D.; McBride, Paul; Gillman,
Len N. (2011-03-01). "Thermal energy and the rate of genetic evolution in marine
fishes". Evolutionary Ecology. 25 (2): 525–530. doi:10.1007/s10682-010-9416-z. ISSN
1573-8477. S2CID 38535658.
Wright, Shane D.; Gillman, Len N.; Ross, Howard A.; Keeling, D. Jeanette (June
2010). "Energy and the tempo of evolution in amphibians: Energy and the tempo of
evolution in amphibians". Global Ecology and Biogeography: no. doi:10.1111/j.1466-
8238.2010.00549.x.
Lin, Gonghua; Huang, Zuhao; Wang, Lei; Chen, Zhenhua; Zhang, Tongzuo; Gillman,
Lennard N; Zhao, Fang (2019-06-01). "Evolutionary Rates of Bumblebee Genomes Are
Faster at Lower Elevations". Molecular Biology and Evolution. 36 (6): 1215–1219.
doi:10.1093/molbev/msz057. ISSN 0737-4038. PMC 6526908. PMID 30865278.
Rolland, Jonathan; Condamine, Fabien L.; Jiguet, Frederic; Morlon, Hélène (2014-
01-28). "Faster Speciation and Reduced Extinction in the Tropics Contribute to the
Mammalian Latitudinal Diversity Gradient". PLOS Biology. 12 (1): e1001775.
doi:10.1371/journal.pbio.1001775. ISSN 1545-7885. PMC 3904837. PMID 24492316.
Wright, Shane; Keeling, Jeannette; Gillman, Len (2006-05-16). "The road from Santa
Rosalia: A faster tempo of evolution in tropical climates". Proceedings of the
National Academy of Sciences. 103 (20): 7718–7722. doi:10.1073/pnas.0510383103.
ISSN 0027-8424. PMC 1472511. PMID 16672371.
Gillman, Len N.; Wright, Shane D. (January 2014). Ladle, Richard (ed.). "Species
richness and evolutionary speed: the influence of temperature, water and area".
Journal of Biogeography. 41 (1): 39–51. doi:10.1111/jbi.12173. ISSN 0305-0270.
S2CID 84888131.
Gillman, Len N.; Wright, Shane D. (May 2006). "The Influence of Productivity on
the Species Richness of Plants: A Critical Assessment". Ecology. 87 (5): 1234–1243.
doi:10.1890/0012-9658(2006)87[1234:TIOPOT]2.0.CO;2. ISSN 0012-9658. PMID 16761602.
Cusens, Jarrod; Wright, Shane D.; McBride, Paul D.; Gillman, Len N. (October
2012). "What is the form of the productivity–animal-species-richness relationship?
A critical review and meta-analysis". Ecology. 93 (10): 2241–2252. doi:10.1890/11-
1861.1. ISSN 0012-9658. PMID 23185885.
Gillman, Len N.; Wright, Shane D.; Cusens, Jarrod; McBride, Paul D.; Malhi,
Yadvinder; Whittaker, Robert J. (January 2015). "Latitude, productivity and species
richness". Global Ecology and Biogeography. 24 (1): 107–117. doi:10.1111/geb.12245.
ISSN 1466-822X.
Hawkins, Bradford A.; Field, Richard; Cornell, Howard V.; Currie, David J.;
Guégan, Jean-François; Kaufman, Dawn M.; Kerr, Jeremy T.; Mittelbach, Gary G.;
Oberdorff, Thierry; O'Brien, Eileen M.; Porter, Eric E. (December 2003). "Energy,
Water, and Broad-Scale Geographic Patterns of Species Richness". Ecology. 84 (12):
3105–3117. doi:10.1890/03-8006. ISSN 0012-9658.
Goldie, Xavier; Gillman, Len; Crisp, Mike; Wright, Shane (2010-09-07).
"Evolutionary speed limited by water in arid Australia". Proceedings of the Royal
Society B: Biological Sciences. 277 (1694): 2645–2653. doi:10.1098/rspb.2010.0439.
PMC 2982047. PMID 20410038.
Wright, Shane D.; Gillman, Len N.; Ross, Howard A.; Keeling, D. Jeanette
(September 2009). "Slower Tempo of Microevolution in Island Birds: Implications for
Conservation Biology". Evolution. 63 (9): 2275–2287. doi:10.1111/j.1558-
5646.2009.00717.x. PMID 19473390. S2CID 23035757.
Roslin, Tomas; Hardwick, Bess; Novotny, Vojtech; Petry, William K.; Andrew, Nigel
R.; Asmus, Ashley; Barrio, Isabel C.; Basset, Yves; Boesing, Andrea Larissa;
Bonebrake, Timothy C.; Cameron, Erin K.; Dáttilo, Wesley; Donoso, David A.; Drozd,
Pavel; Gray, Claudia L.; Hik, David S.; Hill, Sarah J.; Hopkins, Tapani; Huang,
Shuyin; Koane, Bonny; Laird-Hopkins, Benita; Laukkanen, Liisa; Lewis, Owen T.;
Milne, Sol; Mwesige, Isaiah; Nakamura, Akihiro; Nell, Colleen S.; Nichols,
Elizabeth; Prokurat, Alena; Sam, Katerina; Schmidt, Niels M.; Slade, Alison; Slade,
Victor; Suchanková, Alžběta; Teder, Tiit; Nouhuys, Saskya van; Vandvik, Vigdis;
Weissflog, Anita; Zhukovich, Vital; Slade, Eleanor M. (19 May 2017). "Higher
predation risk for insect prey at low latitudes and elevations". Science. 356
(6339): 742–744. Bibcode:2017Sci...356..742R. doi:10.1126/science.aaj1631. PMID
28522532. S2CID 206653702.
Hargreaves, A. L.; Suárez, Esteban; Mehltreter, Klaus; Myers-Smith, Isla;
Vanderplank, Sula E.; Slinn, Heather L.; Vargas-Rodriguez, Yalma L.; Haeussler,
Sybille; David, Santiago; Muñoz, Jenny; Almazán-Núñez, R. Carlos; Loughnan,
Deirdre; Benning, John W.; Moeller, David A.; Brodie, Jedediah F.; Thomas, Haydn J.
D.; M, P. A. Morales (1 February 2019). "Seed predation increases from the Arctic
to the Equator and from high to low elevations". Science Advances. 5 (2): eaau4403.
Bibcode:2019SciA....5.4403H. doi:10.1126/sciadv.aau4403. PMC 6382403. PMID
30801010.
Roesti, Marius; Anstett, Daniel N.; Freeman, Benjamin G.; Lee-Yaw, Julie A.;
Schluter, Dolph; Chavarie, Louise; Rolland, Jonathan; Holzman, Roi (31 March 2020).
"Pelagic fish predation is stronger at temperate latitudes than near the equator".
Nature Communications. 11 (1): 1527. Bibcode:2020NatCo..11.1527R.
doi:10.1038/s41467-020-15335-4. PMC 7109113. PMID 32235853.
 Clark, Nicholas; Clegg, S.; Lima, M. (2014). "A review of global diversity in
avian haemosporidians (Plasmodium and Haemoproteus: Haemosporida): new insights
from molecular data". International Journal for Parasitology. 44 (5): 329–338.
doi:10.1016/j.ijpara.2014.01.004. hdl:10072/61114. PMID 24556563.
Mora, Camilo; Tittensor, Derek P; Myers, Ransom A (22 January 2008). "The
completeness of taxonomic inventories for describing the global diversity and
distribution of marine fishes". Proceedings of the Royal Society B: Biological
Sciences. 275 (1631): 149–155. doi:10.1098/rspb.2007.1315. PMC 2596190. PMID
17999950.
Bibliography
Allen, A. P.; Gillooly, J. F.; Savage, V. M.; Brown, J. H. (2006). "Kinetic effects
of temperature on rates of genetic divergence and speciation". PNAS. 103 (24):
9130–9135. Bibcode:2006PNAS..103.9130A. doi:10.1073/pnas.0603587103. PMC 1474011.
PMID 16754845.
Blackburn, T. M.; Gaston, K. J. (1997). "The relationship between geographic area
and the latitudinal gradient in species richness in New World birds". Evolutionary
Ecology. 11 (2): 195–204. doi:10.1023/A:1018451916734. S2CID 42509143.
Bokma, F. J.; Monkkonen, M.; Mönkkönen, Mikko (2001). "Random processes and
geographic species richness patterns: why so few species in the north?". Ecography.
24: 43–49. doi:10.1034/j.1600-0587.2001.240106.x.
Brown, J. H., and M. V. Lomolino. 1998. Biogeography. Sinauer Associates,
Sunderland.
Cardillo, M.; Orme, C. D. L.; Owens, I. P. F. (2005). "Testing for latitudinal bias
in diversification rates: An example using New World birds". Ecology. 86 (9): 2278–
2287. doi:10.1890/05-0112.
Clarke, A., and J. A. Crame. 2003. The importance of historical processes in global
patterns of diversity. Pages 130-151 in T. M. Blackburn and K. J. Gaston, editors.
Macroecology Concepts and Consequences. Blackwell Scientific, Oxford.
Colwell, R. K.; Hurtt, G. C. (1994). "Nonbiological gradients in species richness
and a spurious Rapoport effect". American Naturalist. 144 (4): 570–595.
doi:10.1086/285695. S2CID 84542533.
Colwell, R. K.; Lees, D. C. (2000). "The mid-domain effect: geometric constraints
on the geography of species richness". Trends in Ecology & Evolution. 15 (2): 70–
76. doi:10.1016/s0169-5347(99)01767-x. PMID 10652559.
Colwell, R. K.; Rahbek, C.; Gotelli, N. (2004). "The mid-domain effect and species
richness patterns: what have we learned so far?" (PDF). American Naturalist. 163
(3): E1–E23. doi:10.1086/382056. PMID 15026983. S2CID 10498450. Archived from the
original (PDF) on 2020-02-27.
Colwell, Robert K.; Rahbek, Carsten; Gotelli, Nicholas J. (1 November 2005). "The
Mid‐Domain Effect: There's a Baby in the Bathwater". The American Naturalist. 166
(5): E149–E154. doi:10.1086/491689. S2CID 28510753.
Connolly, Sean R. (July 2005). "Process‐Based Models of Species Distributions and
the Mid‐Domain Effect" (PDF). The American Naturalist. 166 (1): 1–11.
doi:10.1086/430638. PMID 15937785. S2CID 37200186.
Crame, J. A. (2001). "Taxonomic diversity gradients through geological time".
Diversity and Distributions. 7 (4): 175–189. doi:10.1046/j.1472-4642.2001.00106.x.
Currie, D. J.; Kerr, J. T. (2007). "Testing, as opposed to supporting, the Mid-
domain Hypothesis: a response to Lees and Colwell (2007)". Ecology Letters. 10 (9):
E9–E10. doi:10.1111/j.1461-0248.2007.01074.x.
Currie, D. J.; Mittelbach, G. G.; Cornell, H. V.; Kaufman, D. M.; Kerr, J. T.;
Oberdorff, T. (2004). "Predictions and tests of climate-based hypotheses of broad-
scale variation in taxonomic richness". Ecology Letters. 7 (12): 1121–1134.
doi:10.1111/j.1461-0248.2004.00671.x.
Diniz-Filho, J. A.; Rangel, T. F. L. V. B.; De Souza, Marcia Christianne; Rangel,
Thiago F. L. V. B. (2002). "Null models and spatial patterns of species richness in
South American birds of prey" (PDF). Ecology Letters. 5: 47–55. doi:10.1046/j.1461-
0248.2002.00289.x. S2CID 86814324. Archived from the original (PDF) on 2020-10-31.
Dunn, R. R.; McCain, C. M.; Sanders, N. (2007). "When does diversity fit null model
predictions? Scale and range size mediate the mid-domain effect". Global Ecology
and Biogeography. 16 (3): 305–312. doi:10.1111/j.1466-8238.2006.00284.x.
Gaston, K. J. (2000). "Global patterns in biodiversity". Nature. 405 (6783): 220–
227. doi:10.1038/35012228. PMID 10821282. S2CID 4337597.
Gaston, K. J., and T. M. Blackburn. 2000. Pattern and processes in macroecology.
Blackwell Scientific, Oxford.
Hawkins, B. A.; Diniz-Filho, J. A. F. (2002). "The mid-domain effect cannot explain
the diversity gradient of Nearctic birds". Global Ecology and Biogeography. 11 (5):
419–426. doi:10.1046/j.1466-822x.2002.00299.x.
Gillman, L. N.; Keeling, D. J.; Ross, H. A.; Wright, S. D. (2009). "Latitude,
elevation and the tempo of molecular evolution in mammals". Proceedings of the
Royal Society B: Biological Sciences. 276 (1671): 3353–3359.
doi:10.1098/rspb.2009.0674. PMC 2817169. PMID 19556254.
Hawkins, B. A.; Diniz-Filho, J. A. F.; Weis, A. E. (2005). "The mid-domain effect
and diversity gradients: is there anything to learn". American Naturalist. 166 (5):
E140–E143. doi:10.1086/491686. PMID 16224716. S2CID 33984562.
Hawkins, B. A.; Porter, E. E. (2001). "Area and the latitudinal diversity gradient
for terrestrial birds". Ecology Letters. 4 (6): 595–601. doi:10.1046/j.1461-
0248.2001.00271.x.
Hillebrand, Helmut (1 February 2004). "On the Generality of the Latitudinal
Diversity Gradient" (PDF). The American Naturalist. 163 (2): 192–211.
doi:10.1086/381004. PMID 14970922. S2CID 9886026.
Jetz, W.; Rahbek, C. (2001). "Geometric constraints explain much of the species
richness pattern in African birds". Proceedings of the National Academy of Sciences
of the USA. 98 (10): 5661–5666. Bibcode:2001PNAS...98.5661J.
doi:10.1073/pnas.091100998. PMC 33269. PMID 11344307.
Kerr, J. T.; Perring, M.; Currie, D. J. (2006). "The missing Madagascan mid-domain
effect". Ecology Letters. 9 (2): 149–159. doi:10.1111/j.1461-0248.2005.00860.x.
PMID 16958880.
Koleff, P.; Gaston, K. J. (2001). "Latitudinal gradients in diversity: real
patterns and random models". Ecography. 24 (3): 341–351. doi:10.1111/j.1600-
0587.2001.tb00207.x.
Lambers, J. H. R.; Clark, J. S.; Beckage, B. (2002). "Density-dependent mortality
and the latitudinal gradient in species diversity". Nature. 417 (6890): 732–735.
Bibcode:2002Natur.417..732H. doi:10.1038/nature00809. PMID 12066182. S2CID 4307545.
Lees, D. C.; Colwell, R. K. (2007). "A strong Madagascan rainforest MDE and no
equatorward increase in species richness: Re-analysis of 'The missing Madagascan
mid-domain effect', by Kerr J.T., Perring M. & Currie D.J (Ecology Letters 9:149-
159, 2006)". Ecology Letters. 10 (9): E4–E8. doi:10.1111/j.1461-0248.2007.01040.x.
PMID 17663706.
Pianka, E. R. (1966). "Latitudinal gradients in species diversity: a review of
concepts". American Naturalist. 100 (910): 33–46. doi:10.1086/282398. S2CID
84244127.
Pimm, S.L.; Brown, J.H. (2004). "Domains of diversity". Science. 304 (5672): 831–
833. doi:10.1126/science.1095332. PMID 15131295. S2CID 128866488.
Rahbek, C.; Gotelli, N.; Colwell, R.K.; Entsminger, G.L.; Rangel, T.F.L.V.B.;
Graves, G.R. (2007). "Predicting continental-scale patterns of bird species
richness with spatially explicit models". Proceedings of the Royal Society of
London B. 274 (1607): 165–174. doi:10.1098/rspb.2006.3700. PMC 1685854. PMID
17148246.
Rohde, K. (1992). "Latitudinal gradients in species diversity: the search for the
primary cause" (PDF). Oikos. 65 (3): 514–527. doi:10.2307/3545569. JSTOR 3545569.
S2CID 40145348. Archived from the original (PDF) on 2019-03-03.
Rohde, K. (1997). "The larger area of the tropics does not explain latitudinal
gradients in species diversity". Oikos. 79 (1): 169–172. doi:10.2307/3546102. JSTOR
3546102.
Rolland, J.; Condamine, F.L.; Jiguet, F.; Morlon, H. (2014). "Faster speciation and
reduced extinction in the tropics contribute to the mammalian latitudinal diversity
gradient". PLOS Biology. 12 (1): e1001775. doi:10.1371/journal.pbio.1001775. PMC
3904837. PMID 24492316.
Romdal, T. S.; Colwell, R. K.; Rahbek, C. (2005). "The influence of band sum area,
domain extent and range sizes on the latitudinal mid-domain effect" (PDF). Ecology.
86: 235–244. doi:10.1890/04-0096. S2CID 55114736. Archived from the original (PDF)
on 2020-10-30.
Rosenzweig, M. L. (1992). "Species diversity gradients: we know more and less than
we thought". Journal of Mammalogy. 73 (4): 715–730. doi:10.2307/1382191. JSTOR
1382191.
Rosenzweig, M. L. 2003. How to reject the area hypothesis of latitudinal gradients.
Pages 87–106 in T. M. Blackburn and K. J. Gaston, editors. Macroecology: Concepts
and Consequences. Blackwell Publishing, Oxford.
Storch, D.; Davies, R.G.; Zajicek, S.; et al. (2006). "Energy, range dynamics and
global species richness patterns: reconciling mid-domain effects and environmental
determinants of avian diversity". Ecology Letters. 9 (12): 1308–1320.
doi:10.1111/j.1461-0248.2006.00984.x. PMID 17118005.
Terborgh, J. (1973). "On the notion of favorableness in plant ecology". American
Naturalist. 107 (956): 481–501. doi:10.1086/282852. S2CID 84577772.
Weir, J.T.; Schluter, D. (2007). "The latitudinal gradient in recent speciation and
extinction rates of birds and mammals". Science. 315 (5818): 1574–1576.
Bibcode:2007Sci...315.1574W. doi:10.1126/science.1135590. PMID 17363673. S2CID
46640620.
Willig, M. R.; Lyons, S. K. (1998). "An analytical model of latitudinal gradients
of species richness with an empirical test for marsupials and bats in the New
World" (PDF). Oikos. 81 (1): 93–98. doi:10.2307/3546471. JSTOR 3546471. S2CID
54953465. Archived from the original (PDF) on 2019-03-08.
Willig, M. R.; Kaufmann, D. M.; Stevens, R. D. (2003). "Latitudinal gradients of
biodiversity: pattern, process, scale and synthesis". Annu. Rev. Ecol. Syst. 34:
273–309. doi:10.1146/annurev.ecolsys.34.012103.144032.
Wright, S.; Keeling, J.; Gillman, L. (2006). "The road from Santa Rosalia: a faster
tempo of evolution in tropical climates". Proceedings of the National Academy of
Sciences. 103 (20): 7718–7722. Bibcode:2006PNAS..103.7718W.
doi:10.1073/pnas.0510383103. PMC 1472511. PMID 16672371.
Zapata, F. A.; Gaston, K. J.; Chown, S. L. (2003). "Mid-domain models of species
richness gradients: assumptions, methods and evidence". Journal of Animal Ecology.
72 (4): 677–690. doi:10.1046/j.1365-2656.2003.00741.x. PMID 30893962.
Zapata, F. A.; Gaston, K. J.; Chown, S. L. (2005). "The mid-domain effect
revisited". American Naturalist. 166 (5): E144–E148. doi:10.1086/491685.
hdl:10019.1/119950. PMID 16224717. S2CID 45289227.
Categories: BiodiversityEcology
Navigation menu
Not logged in
Talk
Contributions
Create account
Log in
ArticleTalk
ReadEditView history
Search
Search Wikipedia
Main page
Contents
Current events
Random article
About Wikipedia
Contact us
Donate
Contribute
Help
Learn to edit
Community portal
Recent changes
Upload file
Tools
What links here
Related changes
Special pages
Permanent link
Page information
Cite this page
Wikidata item
Print/export
Download as PDF
Printable version

Languages
Deutsch
Español
Français
Magyar
Edit links
This page was last edited on 29 June 2022, at 23:47 (UTC).
Text is available under the Creative Commons Attribution-ShareAlike License 3.0;
additional terms may apply. By using this site, you agree to the Terms of Use and
Privacy Policy. Wikipedia® is a registered trademark of the Wikimedia Foundation,
Inc., a non-p