Climatic zonation drives latitudinal variation in speciation mechanisms

Many groups of organisms show greater species richness in the tropics than in the temperate zone, particularly in tropical montane regions. Forty years ago, Janzen suggested that more limited temperature seasonality in the tropics leads to greater climatic zonation and more climatic barriers to organismal dispersal along elevational gradients in the tropics relative to temperate regions. These factors could lead to differences in how species arise in tropical versus temperate regions and possibly contribute to greater tropical diversity. However, no studies have compared the relationships among climate, elevational distribution and speciation in a group inhabiting both tropical and temperate regions. Here, we compare elevational and climatic divergence among 30 sister-species pairs (14 tropical, 16 temperate) within a single family of salamanders (Plethodontidae) that reaches its greatest species richness in montane Mesoamerica. In support of Janzen's hypothesis, we find that sister species are more elevationally and climatically divergent in the tropics than in the temperate zone. This pattern seemingly reflects regional variation in the role of climate in speciation, with niche conservatism predominating in the temperate zone and niche divergence in the tropics. Our study demonstrates how latitudinal differences in elevational climatic zonation may increase opportunities for geographical isolation, speciation and the associated build-up of species diversity in the tropics relative to the temperate zone.     

Latitude, elevational climatic zonation and speciation in New World vertebrates

Many biodiversity hotspots are located in montane regions, especially in the tropics. A possible explanation for this pattern is that the narrow thermal tolerances of tropical species and greater climatic stratification of tropical mountains create more opportunities for climate-associated parapatric or allopatric speciation in the tropics relative to the temperate zone. However, it is unclear whether a general relationship exists among latitude, climatic zonation and the ecology of speciation. Recent taxon-specific studies obtained different results regarding the role of climate in speciation in tropical versus temperate areas. Here, we quantify overlap in the climatic distributions of 93 pairs of sister species of mammals, birds, amphibians and reptiles restricted to either the New World tropics or to the Northern temperate zone. We show that elevational ranges of tropical- and temperate-zone species do not differ from one another, yet the temperature range experienced by species in the temperate zone is greater than for those in the tropics. Moreover, tropical sister species tend to exhibit greater similarity in their climatic distributions than temperate sister species. This pattern suggests that evolutionary conservatism in the thermal niches of tropical taxa, coupled with the greater thermal zonation of tropical mountains, may result in increased opportunities for allopatric isolation, speciation and the accumulation of species in tropical montane regions. Our study exemplifies the power of combining phylogenetic and spatial datasets of global climatic variation to explore evolutionary (rather than purely ecological) explanations for the high biodiversity of tropical montane regions.     

Minimal effects of latitude on present-day speciation rates in New World birds

The tropics contain far greater numbers of species than temperate regions, suggesting that rates of species formation might differ systematically between tropical and non-tropical areas. We tested this hypothesis by reconstructing the history of speciation in New World (NW) land birds using BAMM, a Bayesian framework for modelling complex evolutionary dynamics on phylogenetic trees. We estimated marginal distributions of present-day speciation rates for each of 2571 species of birds. The present-day rate of speciation varies approximately 30-fold across NW birds, but there is no difference in the rate distributions for tropical and temperate taxa. Using macroevolutionary cohort analysis, we demonstrate that clades with high tropical membership do not produce species more rapidly than temperate clades. For nearly any value of present-day speciation rate, there are far more species in the tropics than the temperate zone. Any effects of latitude on speciation rate are marginal in comparison to the dramatic variation in rates among clades.     

甘肃鸡类物种多样性研究

甘肃鸡类有19种,随纬度变化形成了物种多样性梯度。本文通过气候(年无霜天数、元月均温、年均温、7月均温和年降水量),海拔高差,纬度和植被类型多样性与各县鸡类多样性相关统计分析,揭示了7月均温,年均降水量,海拔高差和纬度是控制甘肃鸡类多样性的主要因素,而甘肃鸡类多样性与植被类型多样性无关。在降水量400—650mm和海拔高差1500—3500m的地带种类最多。一个地区物种的多少不仅取决于其离物种库的远近,还取决于这一地区环境空间异质性程度。     

Large-scale phylogenetic analyses reveal the causes of high tropical amphibian diversity

Many groups show higher species richness in tropical regions but the underlying causes remain unclear. Despite many competing hypotheses to explain latitudinal diversity gradients, only three processes can directly change species richness across regions: speciation, extinction and dispersal. These processes can be addressed most powerfully using large-scale phylogenetic approaches, but most previous studies have focused on small groups and recent time scales, or did not separate speciation and extinction rates. We investigate the origins of high tropical diversity in amphibians, applying new phylogenetic comparative methods to a tree of 2871 species. Our results show that high tropical diversity is explained by higher speciation in the tropics, higher extinction in temperate regions and limited dispersal out of the tropics compared with colonization of the tropics from temperate regions. These patterns are strongly associated with climate-related variables such as temperature, precipitation and ecosystem energy. Results from models of diversity dependence in speciation rate suggest that temperate clades may have lower carrying capacities and may be more saturated (closer to carrying capacity) than tropical clades. Furthermore, we estimate strikingly low tropical extinction rates over geological time scales, in stark contrast to the dramatic losses of diversity occurring in tropical regions presently.     

THE OCCURRENCE AND DISTRIBUTION OF ACRASIEAE IN FORESTS OF SUBTROPICAL AND TROPICAL AMERICA

Forests of the subtropical and tropical regions of North America harbor cellular slime molds not found in the soils of temperate deciduous forests investigated previously. However, most species found in the temperate forest are common in the tropics. Although the diversity of forms is greater in the soils of tropical forests the numbers of Acrasieae per unit of soil are comparable. Characteristic of tropical and subtropical forest soils are Acrasieae bearing crampon bases, of which four new species of Dictyostelium are presently known. Also present, but less frequently isolated, are two other new species of the genus Dictyostelium and two still undescribed species of the Guttulinaceae. Occasional isolates of D. purpureum and D. discoideum were found that produce macrocysts, which seem, also, to be confined to tropical and subtropical areas. Macro-cysts were previously known only in D. mucoroides and D. minutum isolated from temperate forest soils. The occurrence and distribution of Acrasieae in warm temperate desert and mesquite-scrub, in subtropical hammock, and in tropical thorn, deciduous, seasonal evergreen, rain, and cloud forests were investigated. Acrasieae were well represented in all of these forests except desert. The number of species and the total populations were largest in seasonal evergreen forests. The composition of the cellular slime mold populations and the dominant species within these populations could be related to the soil environment as expressed by the dominant vegetation.     

Seaweeds, latitudinal diversity patterns, and Rapoport's Rule

We analysed the geographic distribution of 645 species of marine benthic algae along the Atlantic coast of Europe and Pacific coast of temperate South America to test for the existence of an association between geographic range size and latitude (Rapoport's Rule) and for three key components of the explanations offered for it. We found that species in high diversity areas are characterized by small geographic ranges and by low specific growth rates as compared to species with large geographic ranges, thus supporting the Rapoport-rescue hypothesis. However, the pattern is not related to species' tolerances, to abiotic conditions or to climatic variability. Further, the inverse latitudinal diversity pattern shown by the marine algal flora of temperate Pacific South America, and the opposite patterns shown by tropical and subantarctic species within this flora, stressed that small geographic ranges are linked to high diversity areas in general, and not only in relation to the pole to tropic species diversity gradient.     

Geographic patterns in fruit colour diversity: do leaves constrain the colour of fleshy fruits?

We tested for geographic patterns in fruit colour diversity. Fruit colours are thought to promote detection by seed dispersers. Because seed dispersers differ in their spectral sensitivities, we predicted that fruit colour diversity would be higher in regions with higher seed disperser diversity (i.e. the tropics). We collected reflectance data on 232 fruiting plant species and their natural backgrounds in seven localities in Europe, North and South America, and analysed fruit colour diversity according to the visual system of birds—the primary consumer types of these fruits. We found no evidence that fruit colours are either more conspicuous or more diverse in tropical areas characterised by higher seed disperser diversity. Instead, fruit colour diversity was lowest in central Brazil, suggesting that fruit colours may be more diverse in temperate regions. Although we found little evidence for geographic variation in fruit hues, the spectral properties of fruits were positively associated with the spectral properties of backgrounds. This result implies that fruit colours may be influenced by selection on the reflectance properties of leaves, thus constraining the evolution of fruit colour. Overall, the results suggest that fruit colours in the tropics are neither more diverse nor more conspicuous than temperate fruits, and that fruit colours may be influenced by correlated selection on leaf reflectance properties. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Niche-tracking migrants and niche-switching residents: evolution of climatic niches in New World warblers (Parulidae)

Differences in life-history traits between tropical and temperate lineages are often attributed to differences in their climatic niche dynamics. For example, the more frequent appearance of migratory behaviour in temperate-breeding species than in species originally breeding in the tropics is believed to have resulted partly from tropical climatic stability and niche conservatism constraining tropical species from shifting their ranges. However, little is known about the patterns and processes underlying climatic niche evolution in migrant and resident animals. We evaluated the evolution of overlap in climatic niches between seasons and its relationship to migratory behaviour in the Parulidae, a family of New World passerine birds. We used ordination methods to measure seasonal niche overlap and niche breadth of 54 resident and 49 migrant species and used phylogenetic comparative methods to assess patterns of climatic niche evolution. We found that despite travelling thousands of kilometres, migrants tracked climatic conditions across the year to a greater extent than tropical residents. Migrant species had wider niches than resident species, although residents as a group occupied a wider climatic space and niches of migrants and residents overlapped extensively. Neither breeding latitude nor migratory distance explained variation among species in climatic niche overlap between seasons. Our findings support the notion that tropical species have narrower niches than temperate-breeders, but does not necessarily constrain their ability to shift or expand their geographical ranges and become migratory. Overall, the tropics may have been historically less likely to experience the suite of components that generate strong selection pressures for the evolution of migratory behaviour.     

Parasite loads are higher in the tropics: temperate to tropical variation in a single host-parasite system

Parasites are important selective forces upon the evolutionary ecology of their hosts. At least one hypothesis suggests that high species diversity in the tropics is associated with higher parasite abundance in tropical climates. Few studies, however, have directly assessed whether parasite abundance is higher in the tropics. To address this question, it is ideal, although seldom achievable, to compare parasite abundance in a single species that occurs over a geographical area including both temperate and tropical regions. We examined variation in blood parasite abundance in seven populations of a single lizard host species ( Eulamprus quoyii ) using a transect that spans temperate and tropical climates. Parasite prevalence (proportion of the host population infected) showed no geographical pattern. Interestingly though, parasite load was higher in lizard populations in the tropics, and was related to mean annual temperature, but not to rainfall. We speculate that in this system the relationship between latitude and parasite load is most likely due to variation in host life history over their geographic range.     

Environmental filters shaping angiosperm tree assembly along climatic and geographic gradients

Question

Global‐scale forest censuses provide an opportunity to understand diversification processes in woody plant communities. Based on the climatic or geographic filtering hypotheses associated with tropical niche conservatism and dispersal limitation, we analysed phylogenetic community structures across a wide range of biomes and evaluated to what extent region‐specific processes have influenced large‐scale diversity patterns of tree species communities across latitude or continent.

Location

Global.

Methods

We generated a data set of species abundances for 21,379 angiosperm woody plants in 843 plots worldwide. We calculated net relatedness index (NRI) for each plot, based on a single global species pool and regional species pools, and phylogenetic β‐diversity (PBD) between plots. Then, we explored the correlations of NRI with climatic and geographic variables, and clarified phylogenetic dissimilarity along geographic and climatic differences. We also compared these patterns for South America, Africa, the Indo‐Pacific, Australia, the Nearctic, Western Palearctic and Eastern Palearctic.

Results

NRI based on a global‐scale species pool was negatively associated with precipitation and positively associated with Quaternary temperature change. PBD was positively associated with geographic distance and precipitation difference between plots across tropical and extratropical biomes. Moreover, phylogenetic dissimilarity was smaller in extratropical regions than in regions including the tropics, although temperate forests of the Eastern Palearctic showed a greater dissimilarity within extratropical regions.

Conclusions

Our findings support predictions of the climatic and geographic filtering hypotheses. Climatic filtering (climatic harshness and paleoclimatic change) relative to tropical niche conservatism played a role in sorting species from the global species pool and shaped the large‐scale diversity patterns, such as the latitudinal gradient observed across continents. Geographic filtering associated with dispersal limitation substantially contributed to regional divergence of tropical/extratropical biomes among continents. Old, long‐standing geographic barriers and recent climatic events differently influenced evolutionary diversification of angiosperm tree communities in tropical and extratropical biomes.     

Life-history strategies of Australian lizards: a comparison between the tropics and the temperate zone

Summary Previous studies have suggested that tropical and temperate-zone lizards may differ fundamentally in life histories. We tested the applicability of this idea to Australian species by comparing temperate-zone species of agamid and scincid lizards with their congeners from the seasonal tropics. Data were derived from dissection of 1,941 specimens and from published information. Clutch size and egg size were positively correlated with mean maternal body size in most lizard species from both climatic zones. Mean body size of the lizards studies did not differ between the tropics and the temperate zone, nor did egg or hatchling size. However, tropical skinks showed considerably (approximately 20%) lower clutch size and relative clutch mass than did temperate-zone skinks. This difference was partly due to the higher incidence of species with low, invariant clutch size in the tropical lizard fauna (as seen in other continents as well), but primarily due to a trend for lineages (especially genera) with relatively high fecundity to be more common in the temperate zone than in the tropics. In contrast to studies on African lizards, our data suggested that modification of clutch size between areas has not occurred within genera: congeneric species from the tropics and temperate zone did not differ in clutch size. Production of more than one clutch per annum by individual females was common in both climatic zones. Tropical lizards may differ from temperate-zone species in showing higher reproductive frequencies, more rapid growth and earlier maturation. However, most of these effects may be due to phenotypic responses to environmental conditions (especially longer annual activity season), rather than to genetically based lifehistory adaptations.     

Life history strategies of cladocerans: comparisons of tropical and temperate taxa

We review recent works on different life history variables of cladoceran taxa in tropical and temperate freshwater bodies, comparing the strategies that cladocerans have evolved to adapt to contrasting environmental conditions in the two geographical regions. These life-history parameters relate to age and size at maturity, survival, fecundity, life-expectancy at birth, lifespan, gross, and net reproductive rates, generation time, the rate of population increase, peak population density and day of peak abundance. We also discuss the role of photoperiod and temperature on some of these life history parameters. We found a general paucity of experimental work and field data in tropics on cladocerans. There is very limited information on the few Daphnia species found in the tropics. The misconception of low species diversity of cladocerans in the tropics arose due to several reasons including lack of extensive and intensive field collections. Higher water temperatures apparently promote permanent infestation of tropical waters with toxic cyanobacteria, which reduce the zooplankton diversity. In addition to higher temperatures in the tropics, the year-round high predation pressure of planktivorous fish probably causes the tropical species, particularly in pelagic habitats, to reach maturity earlier (< 3 days) than in temperate regions. Species of Daphnia in temperate regions are particularly adapted to living at food concentrations that are much lower and seasonably more variable than those for tropical genera such as Diaphanosoma. This is further corroborated by the more than an order of magnitude higher threshold food concentration (TFC) for tropical Cladocera than for their temperate counterparts. Fecundity patterns differ between tropical and temperate cladoceran taxa: cultured under optimal temperature regimes, tropical taxa have fewer eggs than temperate species of a comparable body size. Predation pressure may act differently depending on the size of the cladoceran neonates and thus on their population size structure. Global warming and climate changes seem to affect the behaviour (migration), distribution, and abundance of cladocerans. Apparently, in direct response to these changes, the possibility of encountering the tropical cladocerans in the northern, temperate hemisphere (bioinvasions) is on the rise.     

Phytoplankton and its dynamics in two tropical lakes: a tropical and temperate zone comparison

Temporal patterns of phytoplankton biomass and community structure are described for two Kenyan lakes and subsequently compared with patterns reported in other tropical and temperate lakes. Lake Naivasha had a lower and more seasonally variable (10×) biomass, with a seasonal shift between diatoms and blue-greens, while the L. Oloidien biomass was less variable (3.7×) and dominated by blue-greens. Biomass and chlorophyll a were strongly correlated and in turn were coupled to the level of total phosphorus. A total of 143 and 94 taxa were described for L. Naivasha and L. Oloidien, respectively.The comparative analysis showed: a) a paucity of exclusively tropical species; b) that more than 30 percent of the species in two highly saline Kenyan lakes were also present in the two freshwater lakes; c) no evidence for a postulated decline of phytoplankton species abundance with latitude from the temperate zone to the tropics; d) that the low fraction of chrysophyte biomass in tropical lakes is a function of trophy rather than of latitude; e) that the fraction of chlorophyte biomass in tropical lakes is generally higher than in temperate lakes; f) that the proportion of nannoplankton in the two Kenyan freshwater lakes is not different from that in temperate lakes of the same trophy; g) that seasonal or annual biomass oscillations in the tropics are not systematically lower than in the temperate zone; h) evidence for large inter-year difference in the max.:min. biomass ratio in the only tropical lake (L. Naivasha) for which such data are available; i) that an average biomass ratio appears predictable for tropical lakes from the proportion of the sediment surface in contact with epilimnetic water. Overall, no evidence was found that the freshwater tropical phytoplankton composition or dynamics differ in any fundamental fashion from that observed in the temperate lakes during the summer.Contribution number 147 of the Limnology Research Centre, McGill University.     

Faster Speciation and Reduced Extinction in the Tropics Contribute to the Mammalian Latitudinal Diversity Gradient

The increase in species richness from the poles to the tropics, referred to as the latitudinal diversity gradient, is one of the most ubiquitous biodiversity patterns in the natural world. Although understanding how rates of speciation and extinction vary with latitude is central to explaining this pattern, such analyses have been impeded by the difficulty of estimating diversification rates associated with specific geographic locations. Here, we use a powerful phylogenetic approach and a nearly complete phylogeny of mammals to estimate speciation, extinction, and dispersal rates associated with the tropical and temperate biomes. Overall, speciation rates are higher, and extinction rates lower, in the tropics than in temperate regions. The diversity of the eight most species-rich mammalian orders (covering 92% of all mammals) peaks in the tropics, except that of the Lagomorpha (hares, rabbits, and pikas) reaching a maxima in northern-temperate regions. Latitudinal patterns in diversification rates are strikingly consistent with these diversity patterns, with peaks in species richness associated with low extinction rates (Primates and Lagomorpha), high speciation rates (Diprotodontia, Artiodactyla, and Soricomorpha), or both (Chiroptera and Rodentia). Rates of range expansion were typically higher from the tropics to the temperate regions than in the other direction, supporting the “out of the tropics” hypothesis whereby species originate in the tropics and disperse into higher latitudes. Overall, these results suggest that differences in diversification rates have played a major role in shaping the modern latitudinal diversity gradient in mammals, and illustrate the usefulness of recently developed phylogenetic approaches for understanding this famous yet mysterious pattern.     

Potential effects of elevated CO2 and changes in temperature on tropical plants

Abstract. Very little attention has been directed at the responses of tropical plants to increases in global atmospheric CO₂ concentrations and the potential climatic changes. The available data, from greenhouse and laboratory studies, indicate that the photosynthesis, growth and water use efficiency of tropical plants can increase at higher CO₂ concentrations. However, under field conditions abiotic (light, water or nutrients) or biotic (competition or herbivory) factors might limit these responses. In general, elevated atmospheric CO₂ concentrations seem to increase plant tolerance to stress, including low water availability, high or low temperature, and photoinhibition. Thus, some species may be able to extend their ranges into physically less favourable sites, and biological interactions may become relatively more important in determining the distribution and abundance of species. Tropical plants may be more narrowly adapted to prevailing temperature regimes than are temperate plants, so expected changes in temperature might be relatively more important in the tropics. Reduced transpiration due to decreased stomatal conductance could modify the effects of water stress as a cue for vegetative or reproductive phenology of plants of seasonal tropical areas. The available information suggests that changes in atmospheric CO₂ concentrations could affect processes as varied as plant/herbivore interactions, decomposition and nutrient cycling, local and geographic distributions of species and community types, and ecosystem productivity. However, data on tropical plants are few, and there seem to be no published tropical studies carried out in the field. Immediate steps should be undertaken to reduce our ignorance of this critical area.     

Trichomycete gut fungi from tropical regions of the world

Fifty-nine species of gut fungi in the orders Harpellales and Asellariales, Class Trichomycetes, have been collected in eight tropical regions of the world, some species occurring in more than one geographic region. Regarding the Harpellales, the rather low number of taxa, compared to reports in more temperate localities, is due primarily to relatively few collections in the tropics, as well as the usually warmer waters found in tropical regions that often have a lower species richness of potential immature insect hosts. Asellariales in freshwater, marine, and terrestrial habitats, likewise, have been seldom inventoried in the tropics. Nonetheless, it is clear that the tropics are fertile grounds for discovering new genera and species, and future investigations will undoubtedly reveal many new taxa that will lead to a better understanding of the evolution and biogeography of Trichomycetes.     

Lichens as rapid bioindicators of pollution and habitat disturbances in the tropics

Lichens have value as bioindicators of environmental pollution, climate change, and ecological continuity. Extensive work has been undertaken in temperate areas, but in only few cases have the techniques been applied in the tropics. Most tropical studies to date are in relation to air pollution and forest disturbance, but these are scattered geographically and remain to be undertaken in most tropical regions. The potential of lichens as rapid bioindicators in the tropics can start to be realized even where the species described are not fully identified as they are perennial and separable by eye or hand lens, and a lack of training is identified as the main constraint. An extensive bibliography is included.     

EVOLUTIONARY AND BIOGEOGRAPHIC ORIGINS OF HIGH TROPICAL DIVERSITY IN OLD WORLD FROGS (RANIDAE)

Differences in species richness between regions are ultimately explained by patterns of speciation, extinction, and biogeographic dispersal. Yet, few studies have considered the role of all three processes in generating the high biodiversity of tropical regions. A recent study of a speciose group of predominately New World frogs (Hylidae) showed that their low diversity in temperate regions was associated with relatively recent colonization of these regions, rather than latitudinal differences in diversification rates (rates of speciation–extinction). Here, we perform parallel analyses on the most species-rich group of Old World frogs (Ranidae; ∼1300 species) to determine if similar processes drive the latitudinal diversity gradient. We estimate a time-calibrated phylogeny for 390 ranid species and use this phylogeny to analyze patterns of biogeography and diversification rates. As in hylids, we find a strong relationship between the timing of colonization of each region and its current diversity, with recent colonization of temperate regions from tropical regions. Diversification rates are similar in tropical and temperate clades, suggesting that neither accelerated tropical speciation rates nor greater temperate extinction rates explain high tropical diversity in this group. Instead, these results show the importance of historical biogeography in explaining high species richness in both the New World and Old World tropics.