Why are there more species in the tropics?
Noticed for two centuries, true for nearly every group of plants and animals counted, and explained by about thirty competing hypotheses.
The question
Walk one of the richest hectares of Amazon forest, near Iquitos, and you will pass about 300 species of tree at least 10 centimetres thick. The whole of Europe has 454 native trees.
The gradient is not subtle and it is not in doubt. What produces it has been argued since Darwin and Wallace were alive.
The pattern
Known. Strongly supported by evidence and broadly established.
Strongly supported by evidence and broadly established.
The pattern is nearly universal
Richness increases towards the equator for trees, birds, mammals and molluscs, on land and in the sea. A meta-analysis of nearly 600 published gradients found the decline holds across organisms, habitats and regions, more strongly at regional scales than at local ones. It reaches back before the dinosaurs, though perhaps not continuously: a 2014 review of the fossil record finds a tropical peak only in parts of the Palaeozoic and in the past 30 million years, with flatter gradients or temperate peaks in warmer times.
Exceptions exist, and they are informative. In the sea the gradient often looks bimodal, with a dip in richness at the equator and peaks to either side whose positions vary by group. How much of the dip is real is argued: a 2018 analysis traces most of it to how thinly tropical waters have been sampled. Fossil plankton, which preserve exceptionally well, show the dip opening as the ocean warmed after the last ice age, and the survey that described the bimodality suggests the equator may already be too hot for some species.
10 sources for this passage. Show them.
- Evolution and the latitudinal diversity gradient: speciation, extinction and biogeographyMittelbach and others, Ecology Letters · 2007
- On the generality of the latitudinal diversity gradientHillebrand, The American Naturalist · 2004
- Latitudinal gradients of biodiversity: pattern, process, scale, and synthesisWillig, Kaufman and Stevens, Annual Review of Ecology, Evolution, and Systematics · 2003
- Bimodality of latitudinal gradients in marine species richnessChaudhary, Saeedi and Costello, Trends in Ecology and Evolution · 2016
- Past and future decline of tropical pelagic biodiversityYasuhara and others, PNAS · 2020
- Tree species richness of upper Amazonian forestsGentry, PNAS · 1988
- European Red List of TreesRivers and others, IUCN · 2019
- The latitudinal biodiversity gradient through deep timeMannion, Upchurch, Benson and Goswami, Trends in Ecology and Evolution · January 2014
- Mapping knowledge gaps in marine diversity reveals a latitudinal gradient of missing species richnessMenegotto and Rangel, Nature Communications · November 2018
- Explaining global variation in the latitudinal diversity gradient: meta-analysis confirms known patterns and uncovers new onesKinlock and others, Global Ecology and Biogeography · 2018
Three kinds of explanation
A 2007 review sorted the proposals into three kinds: ecological limits on how many species a place can hold, differences in the rate at which lineages diversify, and differences in how long the tropics have been there to diversify in. Counted region by region, movement between regions is a fourth term, and leaving it out is how the same data has been read two ways.
The review found accumulating evidence for higher diversification in the tropics, from phylogenies and from fossils, and attributed it in part to faster tropical speciation. How extinction rates vary with latitude, it said, was little known. It also counted the greater age and former extent of tropical environments as part of the answer, an idea that had had far less attention than ecological ones.
Where the difference would have to come from
More are made
Uncertain. Meaningful scientific, historical or philosophical disagreement remains.
- Higher temperatures were proposed to raise evolutionary speed, and early evidence pointed to faster molecular evolution in some tropical taxa
- Narrower physiological tolerances and a larger tropical area were proposed to give more opportunity for populations to be isolated
- Phylogenies disagree: mammals form new species faster in the tropics, marine fishes fastest outside them, and a link from faster molecules to faster speciation had not been shown by 2007
Fewer are lost, and there has been longer
Uncertain. Meaningful scientific, historical or philosophical disagreement remains.
- Stable tropical climates were proposed to reduce the chance of extinction, by Darwin and by Wallace among others
- Pleistocene glacial cycles fell disproportionately on lineages living at high latitudes
- Tropical conditions have covered more of the planet for more of its history, and how extinction varies with latitude is notoriously hard to estimate
6 sources for this passage. Show them.
- Evolution and the latitudinal diversity gradient: speciation, extinction and biogeographyMittelbach and others, Ecology Letters · 2007
- Out of the Tropics: Evolutionary Dynamics of the Latitudinal Diversity GradientJablonski, Roy and Valentine, Science · October 2006
- Latitudinal gradients of biodiversity: pattern, process, scale, and synthesisWillig, Kaufman and Stevens, Annual Review of Ecology, Evolution, and Systematics · 2003
- Faster speciation and reduced extinction in the tropics contribute to the mammalian latitudinal diversity gradientRolland, Condamine, Jiguet and Morlon, PLOS Biology · January 2014
- An inverse latitudinal gradient in speciation rate for marine fishesRabosky and others, Nature · July 2018
- Speciation gradients and the distribution of biodiversitySchluter and Pennell, Nature · June 2017
Reading it out of the fossils
A 2006 study took the question to the marine fossil record, tracking genera and subgenera of bivalves through the past 11 million years and counting origination, extinction and immigration separately.
It found that taxa preferentially originate in the tropics and expand towards the poles without losing their tropical presence, a pattern the authors called out of the tropics.
That makes the tropics a cradle and a museum at once, against a dichotomy that had stood since 1974. A study of one genus of tropical American leaf beetles reached the same answer the same year, and the 2007 review took the pair as where the evidence stood.
3 sources for this passage. Show them.
- Out of the Tropics: Evolutionary Dynamics of the Latitudinal Diversity GradientJablonski, Roy and Valentine, Science · October 2006
- Evolution and the latitudinal diversity gradient: speciation, extinction and biogeographyMittelbach and others, Ecology Letters · 2007
- Tropical forests are both evolutionary cradles and museums of leaf beetle diversityMcKenna and Farrell, PNAS · July 2006
Speciation, extinction, time, and what moves in from somewhere else.
The explanations
Uncertain. Meaningful scientific, historical or philosophical disagreement remains.
Meaningful scientific, historical or philosophical disagreement remains.
Why the answer matters practically
If tropical richness reflects a slow accumulation over tens of millions of years, it cannot be rebuilt on any human timescale once lost.
If it reflects faster speciation, the loss is no less severe, because even fast speciation runs on a geological clock. The two stories still differ in what conservation is protecting: an archive in one, a source of new species in the other. The bivalve study put it flatly: a tropical diversity crisis would have evolutionary effects at every latitude.
2 sources for this passage. Show them.
Whether one explanation is required
Different groups may reach the same pattern by different routes, in which case looking for a single cause is a mistake in the framing of the question rather than a failure to find the answer.
A 2019 review says plainly that no consensus has been reached and that the observed patterns are often consistent with several explanations at once. That is less satisfying and better supported.
3 sources for this passage. Show them.
- Evolution and the latitudinal diversity gradient: speciation, extinction and biogeographyMittelbach and others, Ecology Letters · 2007
- The latitudinal diversity gradient: novel understanding through mechanistic eco-evolutionary modelsPontarp and others, Trends in Ecology and Evolution · 2019
- Faster speciation and reduced extinction in the tropics contribute to the mammalian latitudinal diversity gradientRolland, Condamine, Jiguet and Morlon, PLOS Biology · January 2014
The evidence
Primary research, scientific institutions and archives. Links open at the publisher.
- Evolution and the latitudinal diversity gradient: speciation, extinction and biogeographyMittelbach and others, Ecology Letters · 2007
- Out of the Tropics: Evolutionary Dynamics of the Latitudinal Diversity GradientJablonski, Roy and Valentine, Science · October 2006
- On the generality of the latitudinal diversity gradientHillebrand, The American Naturalist · 2004
- Latitudinal gradients of biodiversity: pattern, process, scale, and synthesisWillig, Kaufman and Stevens, Annual Review of Ecology, Evolution, and Systematics · 2003
- Bimodality of latitudinal gradients in marine species richnessChaudhary, Saeedi and Costello, Trends in Ecology and Evolution · 2016
- Past and future decline of tropical pelagic biodiversityYasuhara and others, PNAS · 2020
- Tree species richness of upper Amazonian forestsGentry, PNAS · 1988
- European Red List of TreesRivers and others, IUCN · 2019
- The latitudinal diversity gradient: novel understanding through mechanistic eco-evolutionary modelsPontarp and others, Trends in Ecology and Evolution · 2019
- Explaining global variation in the latitudinal diversity gradient: meta-analysis confirms known patterns and uncovers new onesKinlock and others, Global Ecology and Biogeography · 2018
- The latitudinal biodiversity gradient through deep timeMannion, Upchurch, Benson and Goswami, Trends in Ecology and Evolution · January 2014
- Mapping knowledge gaps in marine diversity reveals a latitudinal gradient of missing species richnessMenegotto and Rangel, Nature Communications · November 2018
- Faster speciation and reduced extinction in the tropics contribute to the mammalian latitudinal diversity gradientRolland, Condamine, Jiguet and Morlon, PLOS Biology · January 2014
- An inverse latitudinal gradient in speciation rate for marine fishesRabosky and others, Nature · July 2018
- Speciation gradients and the distribution of biodiversitySchluter and Pennell, Nature · June 2017
- Tropical forests are both evolutionary cradles and museums of leaf beetle diversityMcKenna and Farrell, PNAS · July 2006
Counting what is alive
How many kinds there are, why the counting is hard, and how fast they are going.
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