Evolution and the latitudinal diversity gradient: speciation, extinction and biogeography

A latitudinal gradient in biodiversity has existed since before the time of the dinosaurs, yet how and why this gradient arose remains unresolved. Here we review two major hypotheses for the origin of the latitudinal diversity gradient. The time and area hypothesis holds that tropical climates are older and historically larger, allowing more opportunity for diversification. This hypothesis is supported by observations that temperate taxa are often younger than, and nested within, tropical taxa, and that diversity is positively correlated with the age and area of geographical regions. The diversification rate hypothesis holds that tropical regions diversify faster due to higher rates of speciation (caused by increased opportunities for the evolution of reproductive isolation, or faster molecular evolution, or the increased importance of biotic interactions), or due to lower extinction rates. There is phylogenetic evidence for higher rates of diversification in tropical clades, and palaeontological data demonstrate higher rates of origination for tropical taxa, but mixed evidence for latitudinal differences in extinction rates. Studies of latitudinal variation in incipient speciation also suggest faster speciation in the tropics. Distinguishing the roles of history, speciation and extinction in the origin of the latitudinal gradient represents a major challenge to future research.     

Botryosphaeriaceae on Syzygium cordatum across a latitudinal gradient in South Africa

The Botryosphaeriaceae is a family of endophytic fungi, many of which are latent pathogens of woody plants. Although extensively sampled in some parts of the world, little is known regarding their occurrence across different environmental conditions. This study considered the presence of the Botryosphaeriaceae on Syzygium cordatum trees across a latitudinal gradient. We examined the relative importance of different environmental factors on the presence of the Botryosphaeriaceae across this latitudinal gradient. Specifically, Botryosphaeriaceae community composition and species richness were analysed. The optimal growth temperature of the most common Botryosphaeriaceae isolates and its relation to isolate origin was also tested in culture. We identified 14 Botryosphaeriaceae species including seven each of Lasiodiplodia and Neofusicoccum species. The maximum historical temperature emerged as the environmental factor that best predicted the presence of Botryosphaeriaceae species in S. cordatum trees, specifically influencing Botryosphaeriaceae community composition. For all the Botryosphaeriaceae species studied in vitro, temperature strongly influenced mycelial growth and they all had an optimal growth temperature of 25 °C. Contrary to our hypothesis, the optimal growth temperature was not related to isolate origin. These results contribute to understanding the presence of the Botryosphaeriaceae in trees and our ability to detect these latent pathogens.     

Genetic differentiation across a latitudinal gradient in two co-occurring butterfly species: revealing population differences in a context of climate change

Genetic differentiation within a species' range is determined by natural selection, genetic drift, and gene flow. Selection and drift enhance genetic differences if populations are sufficiently isolated, while gene flow precludes differentiation and local adaptation. Over large geographical areas, these processes can create a variety of scenarios, ranging from admixture to a high degree of population differentiation. Genetic differences among populations may signal functional differences within a species' range, potentially leading to population or ecotype-specific responses to global change. We investigated differentiation within the geographical range of two butterfly species along a broad latitudinal gradient. This gradient is the primary axis of climatic variation, and many ecologists expect populations at the poleward edge of this gradient to expand under climate change. Our study species inhabit a shared ecosystem and differ in body size and resource specialization; both also find their poleward range limit on an island. We find evidence for divergence of peripheral populations from the core in both taxa, suggesting the potential for genetic distinctiveness at the leading edge of climate change. We also find differences between the species in the extent of peripheral differentiation with the smaller and more specialized species showing greater population divergence (microsatellites and mtDNA) and reduced gene flow (mtDNA). Finally, gene flow estimates in both species differed strongly between two marker types. These findings suggest caution in assuming that populations are invariant across latitude and thus will respond as a single ecotype to climatic change.     

Variation in behavioral traits across a broad latitudinal gradient in a livebearing fish

Evolutionary Ecology - Variation in animal behavior can have important consequences for ecological processes like mate choice, resource competition and invasive potential. Characterizing behavioral...     

Maintenance of forest species diversity and latitudinal gradient

The species diversity of trees maintained in tropical rain forests is much higher than in temperate, boreal, or seasonally dry tropical forests. Many hypotheses have been proposed for higher diversity in tropical rain forests, including: (i) higher specialization of resource use, (ii) different mode of disturbance, (iii) smaller opportunity for competition on oligotrophic soil, (iv) higher productivity, (v) more active specific herbivores and pathogens, (vi) evolutionary/ecological history. In this paper we report mathematical models for tree-by-tree replacement. First the analysis of random drift model shows that the effect of gap size to species diversity is not very strong. Second we study phenological segregation model, which has the following assumptions: Basic mechanism for many species to coexist in the community is assumed given by the storage effect of lottery model, as species differ in seasonality in peak fruit production and in the subsequent period of high regeneration ability. Gaps formed during unfavorable season accumulate and become available for regeneration in the beginning of the growing season. The resulting synchronization of regeneration opportunity jeopardizes the coexistence of many similar species in seasonal environments. Analysis of a mathematical model shows: (1) the existence of unfavorable season can greatly reduce the diversity of coexisting species. (2) Diversity in the equilibrium community can be high when niche width of each species is broad and resource use is strongly overlapped. (3) Equilibrium community may include several distinct groups of species differing in phenology of regeneration. Effect of unequal niche width and frequency dependent regeneration are also examined.     

Temporal population variability in local forest communities has mixed effects on tree species richness across a latitudinal gradient

Among the local processes that determine species diversity in ecological communities, fluctuation‐dependent mechanisms that are mediated by temporal variability in the abundances of species populations have received significant attention. Higher temporal variability in the abundances of species populations can increase the strength of temporal niche partitioning but can also increase the risk of species extinctions, such that the net effect on species coexistence is not clear. We quantified this temporal population variability for tree species in 21 large forest plots and found much greater variability for higher latitude plots with fewer tree species. A fitted mechanistic model showed that among the forest plots, the net effect of temporal population variability on tree species coexistence was usually negative, but sometimes positive or negligible. Therefore, our results suggest that temporal variability in the abundances of species populations has no clear negative or positive contribution to the latitudinal gradient in tree species richness.     

Night conditions affect morning incubation behaviour differently across a latitudinal gradient

Intermittently incubating birds alternate between sessions of egg warming and recesses for foraging during the day, but stay on the nest continuously at night. Hence, energy costs of nocturnal incubation (which increase during longer and colder nights) cannot be replenished until the next day. Night conditions might therefore be expected to affect morning incubation behaviour the day after. We tested this prediction by exploring latitudinal and seasonal trends in characteristics of the first recess in Eastern Bluebirds Sialia sialis over a 1400-km latitudinal gradient in the continental USA. The time from civil dawn to leaving the nest (latency) increased with latitude early in the breeding season but decreased with latitude late in the season. Birds breeding at higher latitudes also took longer first recesses throughout the season, which led to a larger drop in nest temperature. At the local scale, birds rose earlier after longer nights if the night was also cold, but night length did not predict latency following warm nights. The first recess was longer if the night was warmer, probably because birds could replenish reserves at lower risk of low egg temperature. Our study shows that characteristics of the night led to behavioural changes in features of early morning incubation that were evident at both continental and local scales. These responses also affected nest temperature. Hence, night conditions carry over to incubation behaviour the following morning, which in turn may impose thermal constraints on embryonic development.     

Effects of anthropogenic habitat disturbance on local pollinator diversity and species turnover across a precipitation gradient

Anthropogenic habitat disturbance can have profound effects on multiple components of forest biotas including pollinator assemblages. We assessed the effect of small-scale disturbance on local richness, abundance, diversity and evenness of insect pollinator fauna; and how habitat disturbance affected species turnover across the landscape and overall diversity along a precipitation gradient in NW Patagonia (Argentina). We evaluated the effect of disturbance on overall pollinator fauna and then separately for bees (i.e. Apoidea) and non-bee pollinators. Locally, disturbed habitats had significantly higher pollinator species richness and abundances than undisturbed habitats for the whole pollinator assemblage, but not for bees or non-bees separately. However, significant differences in species richness between habitats vanished after accounting for differences in abundance between habitat types. At a local scale Shannon–Weaver diversity and evenness did not vary with disturbance. A β diversity index indicated that, across forest types, species turnover was lower between disturbed habitats than between undisturbed habitats. In addition, rarefaction curves showed that disturbed habitats as a whole accumulated fewer species than undisturbed habitats at equivalent sample sizes. We concluded that small patches of disturbed habitat have a negligible effect on local pollinator diversity; however, habitat disturbance reduced β diversity through a homogenization of the pollinator fauna (in particular of bees) across the landscape.     

Areas, cradles and museums: the latitudinal gradient in species richness

Although numerous factors are postulated to be responsible for the gradient of increasing taxon richness towards lower latitudes, it has recently been suggested that the primary determinant is geographic area. This area model is appealing in its logic, but there is little empirical evidence to support it and several other mechanisms might also interact to obscure its effects. Nonetheless, the model has highlighted several fundamental issues concerning range size, speciation and extinction that, despite their considerable significance, remain poorly understood.     

Assessing recovery of biological soil crusts across a latitudinal gradient in Western Europe

Biological soil crusts are ecologically important communities in areas where vascular plant coverage is low, and their presence is often vital in prevention of soil erosion. Despite recurrent threats to biological soil crusts across different environments, their recovery after disturbance has been little studied. We therefore established experiments across a latitudinal gradient in Europe, from Öland, Sweden in the north, to Gössenheim, Germany and Hochtor, Austria, to Almeria, Spain in the south, spanning over 20° latitude and 2,300 m in altitude, and including natural and semi‐natural sites. At each site 10 (1 × 1 m) paired plots were constructed where the biological soil crusts were either completely removed, or left intact. Over a 2‐year period (2012–2014) the plots were regularly sampled to assess functional group recovery (cyanobacteria, algae, lichens, bryophytes, vascular plants), soil stability, and chlorophyll, carbon, and nutrient contents. Cyanobacterial assemblages were examined by denaturing gradient gel electrophoresis, a technique used to detect DNA in environmental samples. The results indicated that recovery was site dependent, suggesting that physical and climatic parameters play a major role in biological soil crust recovery. This conclusion is supported by the results of the soil properties, which were found to differ between sites, although they did not show meaningful recovery over the study period. Although 2 years was insufficient for pronounced biological soil crust recovery, this study documents changes over the initial recovery period, suggests management practices for future projects, and recommends proxies for measuring recovery over time.     

Bird species turnover and stochastic extinction in woodland fragments

Year-to-year turnover in bird species composition was recorded across, the whole size range (0 02-30 ha) of 146 woods studied The mean number of resident breeding species both lost and gained per wood between consecutive breeding seasons was 2 (range 0-8) No relationship was found between this absolute turnover rate and woodland area, or any other of 24 predictor variables (describing woodland structure, isolation, connectedness and surrounding land use) Extriction and colonisation rates (in terms of numbers of species lost and gained) were also unrelated to woodland area In all sizes of woods, the species most likely to show local extinctions and colonisations were those with small populations within those woods, but the identity of the species concerned changed as woodland area increased In the smallest woods, the majority of turnover involved common species, such as wren and dunnock, which occurred in only small numbers in these small woods As woodland area increased, these species attained sufficient numbers to usually avoid stochastic extinction The majority of turnover was then due to more specialist (and less numerous) woodland species, such as great-spotted woodpecker and marsh tit, which were usually lacking in small woods In Britain, much existing broadleaved woodland falls within the size range studied Thus the numbers of many bird species are liable to be small enough for yearly turnover in woodland bird communities to be appreciable, and for the long-term persistence of individual species in particular woods to depend on dispersal     

UV responses of Lolium perenne raised along a latitudinal gradient across Europe: a filtration study

Lolium perenne (cv. AberDart) was grown at 14 locations along a latitudinal gradient across Europe (37-68°N) to study the impact of ultraviolet radiation (UV) and climate on aboveground growth and foliar UV-B absorbing compounds. At each location, plants were grown outdoors for 5 weeks in a replicated UV-B filtration experiment consisting of open, UV-B transparent (cellulose diacetate) and UV-B opaque (polyester) environments. Fourier transform-infrared spectroscopy was used to compare plant metabolite profiles in relation to treatment and location. UV radiation and climatic parameters were determined for each location from online sources and the data were assessed using a combination of anova and multiple regression analyses. Most of the variation in growth between the locations was attributable to the combination of climatic parameters, with minimum temperature identified as an important growth constraint. However, no single environmental parameter could consistently account for the variability in plant growth. Concentrations of foliar UV-B absorbing compounds showed a positive trend with solar UV across the latitudinal gradient; however, this relationship was not consistent in all treatments. The most striking experimental outcome from this study was the effect of presence or absence of filtration frames on UV-absorbing compounds. Overall, the study demonstrates the value of an European approach in studying the impacts of natural UV across a large latitudinal gradient. We have shown the feasibility of coordinated UV filtration at multiple sites but have also highlighted the need for open controls and careful interpretation of plant responses.     

A 450 million years long latitudinal gradient in age‐dependent extinction

Leigh Van Valen famously stated that under constant conditions extinction probability is independent of species age. To test this 'law of constant extinction', we developed a new method using deep learning to infer age‐dependent extinction and analysed 450 myr of marine life across 21 invertebrate clades. We show that extinction rate significantly decreases with age in > 90% of the cases, indicating that most species died out soon after their appearance while those which survived experienced ever decreasing extinction risk. This age‐dependent extinction pattern is stronger towards the Equator and holds true when the potential effects of mass extinctions and taxonomic inflation are accounted for. These results suggest that the effect of biological interactions on age‐dependent extinction rate is more intense towards the tropics. We propose that the latitudinal diversity gradient and selection at the species level account for this exceptional, yet little recognised, macroevolutionary and macroecological pattern.     

Cactus species turnover and diversity along a latitudinal transect in the Chihuahuan Desert Region

A standardized sampling method was used to evaluate turnover (β diversity) among cactus species assemblages along a 798 km long latitudinal megatransect across the Chihuahuan Desert Region, from north-central Mexico to southern Texas. A total of 71 cactus species were found along the megatransect, 66.2% of which appeared at low frequencies, mostly as a consequence of their highly discontinuous distribution pattern. At the scale the study was conducted, there was always species turnover among cactus assemblages. The rate of turnover among contiguous sites primarily fluctuated from low to medium, but when all site combinations were considered (contiguous and non-contiguous), medium β diversity values were predominant (β = 0.331–0.66); however, 25.4% of the site pair combinations registered high values (β = 0.661–1.0). Our results showed that turnover among cactus species assemblages in the CDR does not consist for the most part of a process of species succession in the geographic space. Instead, we concluded that the continuous spatial changes in cactus species composition are primarily explained by the commonly intermittent distribution patterns of the species, by the presence in the megatransect of species at the margin of their distribution range, and, to a lesser extent, by the existence of narrowly endemic species.     

Effects of nutrients and fish on periphyton and plant biomass across a European latitudinal gradient

Replicated, factorial mesocosm experiments were conducted across Europe to study the effects of nutrient enrichment and fish density on macrophytes and on periphyton chlorophyll a (chl-a) with regard to latitude. Periphyton chl-a densities and plant decline were significantly related to nutrient loading in all countries. Fish effects were significant in a few sites only, mostly because of their contribution to the nutrient pool. A saturation-response type curve in periphyton chl-a with nutrients was found, and northern lakes achieved higher densities than southern lakes. Nutrient concentration and phytoplankton chl-a necessary for a 50% plant reduction followed a latitudinal gradient. Total phosphorus values for 50% plant disappearance were similar from Sweden (0.27 mg L⁻¹) to northern Spain (0.35 mg L⁻¹), but with a sharp increase in southern Spain (0.9 mg L⁻¹). Planktonic chl-a values for 50% plant reduction increased monotonically from Sweden (30 μg L⁻¹) to València (150 μg L⁻¹). Longer plant growing-season, higher light intensities and temperature, and strong water-level fluctuations characteristic of southern latitudes can lead to greater persistence of macrophyte biomass at higher turbidities and nutrient concentration than in northern lakes. Results support the evidence that latitudinal differences in the functioning of shallow lakes should be considered in lake management and conservation policies.     

Explaining the latitudinal gradient anomaly in ichneumonid species richness: evidence from butterflies

1. Despite the profusion of Lepidoptera and other potential hosts in the tropics, the Ichneumonidae are not more species-rich in the lowland tropics than in temperate communities. Alternate but partially complementary hypotheses emphasizing resource fragmentation, predation, or chemical defence have been proposed to explain this paradox.
2. Because different diversity gradients occur in the various subgroups of the Ichneumonidae, as well as among their hosts, general explanations are unlikely to account for this 'anomalous diversity.' Hypotheses of ecological causation are best tested at the taxonomic level where the pattern of interest applies.
3. Phylogenetic trends in the distribution, abundance, and palatability of butterflies (Papilionoidea), a group that increases dramatically in species richness towards the tropics, are presented as a case study for evaluating hypotheses of parasitoid species richness. Evidence is presented that supports the 'nasty host hypothesis' as an explanation for the lack of diversity of associated parasitoids.
4. An examination of hypotheses and the approaches used to test them suggests that the nasty-host hypothesis is best supported by the limited evidence available, while evidence in favour of other hypotheses is either lacking or ambiguous. In particular, use of the koinobiont/idiobiont dichotomy and parasitoid assemblage-size comparisons may result in contradictory conclusions.     

Comparative energetics and physiology of parental care in smallmouth bass Micropterus dolomieu across a latitudinal gradient

The energetic and physiological status of parental smallmouth bass Micropterus dolomieu was investigated across the majority of their latitudinal range at the onset and near the end of care. Variables such as tissue lipid stores, plasma indicators of nutritional status and chronic stress and white muscle were used to define energetic and physiological status. Results showed that northern males (48° N) were larger and heavier than mid-northern (44° N) and southern (36° N) latitude males. For a given body size, northern males had greater whole-body lipid across the parental care period and tended to feed more (based on gut contents) than mid-northern and southern latitude conspecifics. Indicators of nutritional status were also highest in northern males. Conversely, the southern males exhibited the greatest capacity for biosynthesis across the entire parental care period as indicated by the highest level of nucleoside diphosphate kinase activities. Collectively, these finding suggest that the energetic costs and physiological consequences of care vary across latitudes, providing some of the first mechanistic evidence of how environmental conditions can influence both the ecological and physiological costs of reproduction for wild animals during parental care. The data also suggest that lake-specific processes that can vary independently of latitude may be important, necessitating additional research on fish reproductive physiology across landscapes.