The biogeography of tropical reef fishes: endemism and provinciality through time

The largest marine biodiversity hotspot straddles the Indian and Pacific Oceans, driven by taxa associated with tropical coral reefs. Centred on the Indo‐Australian Archipelago (IAA), this biodiversity hotspot forms the ‘bullseye’ of a steep gradient in species richness from this centre to the periphery of the vast Indo‐Pacific region. Complex patterns of endemism, wide‐ranging species and assemblage differences have obscured our understanding of the genesis of this biodiversity pattern and its maintenance across two‐thirds of the world's oceans. But time‐calibrated molecular phylogenies coupled with ancestral biogeographic estimates have provided a valuable framework in which to examine the origins of coral reef fish biodiversity across the tropics. Herein, we examine phylogenetic and biogeographic data for coral reef fishes to highlight temporal patterns of marine endemism and tropical provinciality. The ages and distribution of endemic lineages have often been used to identify areas of species creation and demise in the marine tropics and discriminate among multiple hypotheses regarding the origins of biodiversity in the IAA. Despite a general under‐sampling of endemic fishes in phylogenetic studies, the majority of locations today contain a mixture of potential paleo‐ and neo‐endemic fishes, pointing to multiple historical processes involved in the origin and maintenance of the IAA biodiversity hotspot. Increased precision and sampling of geographic ranges for reef fishes has permitted the division of discrete realms, regions and provinces across the tropics. Yet, such metrics are only beginning to integrate phylogenetic relatedness and ancestral biogeography. Here, we integrate phylogenetic diversity with ancestral biogeographic estimation of lineages to show how assemblage structure and tropical provinciality has changed through time.     

Congruent spatial patterns of species richness and phylogenetic diversity in karst flora: Case study of Primulina (Gesneriaceae)

The karst landform in southern China is renowned for its high levels of species diversity and endemism. Globally, karst ecosystems are under threat from unsustainable anthropogenic disturbance and climate changes and are among the most threatened ecosystems worldwide. In this study, we used the typical karst endemic genus in southern China, Primulina Hance, as a model to identify areas within the karst landform with high diversity and to investigate congruence between phylogenetic and species‐based measures of diversity. Using phylogenetic information and species distribution data, we measured geographical patterns of diversity with four metrics: species richness (SR), corrected weighted endemism (CWE), phylogenetic diversity (PD), and phylogenetic endemism (PE). Our results revealed a high spatial congruence among SR, PD, and PE, with hotspot areas identified in the Nanling Mountains (i.e., north Guangdong and northeast Guangxi) and southeast Yungui Plateau (i.e., north and southwest Guangxi), whereas the hotspots of CWE are comparatively uniform throughout the geographic extent. The categorical analysis of neo‐ and paleoendemism identified a pattern of mixed neo‐ and paleoendemism in numerous grid cells, suggesting that karst areas in southern China have acted as both “museums” and “cradles” of plant evolution. Conservation gap analysis of hotspots revealed that the majority of prioritized hotspots (>90%) of the genus are outside of protected areas, therefore indicating the limited effectiveness of national nature reserves for the karst flora. Overall, our results suggest that the karst flora merits more conservation attention and SR can be an effective surrogate to capture PD in conservation planning.     

Spatial phylogenetics of the North American flora

North America is a large continent with extensive climatic, geological, soil, and biological diversity. As biota faces threat from habitat destruction and climate change, making a quantitative assessment of biodiversity becomes critically important. Rapid digitization of plant specimen records and accumulation of DNA sequence data enable a much‐needed broad synthesis of species occurrences with phylogenetic data. In this study, the first such synthesis of a flora from such a large and diverse part of the world is attempted, all seed plants from the North American continent (here defined to include Canada, United States, and Mexico), with a focus on examining phylogenetic diversity and endemism. We collected digitized plant specimen records and chose a coarse grain for analysis, recognizing that this grain is currently necessary for reasonable completeness per sampling unit. We found that raw richness and endemism patterns largely support previous hypotheses of biodiversity hotspots. The application of phylogenetic metrics and a randomization test revealed novel results, including a significant phylogenetic clustering across the continent, a striking east–west geographical difference in the distribution of branch lengths, and the discovery of centers of neo‐ and paleoendemism in Mexico, the southwestern USA, and the southeastern USA. Finally, our examination of phylogenetic beta diversity provides a new approach to compare centers of endemism. We discuss the empirical challenges of working at the continental scale and the need for more sampling across large parts of the continent, for both DNA data for terminal taxa and spatial data for poorly understood regions, to confirm and extend these results.     

Phylogenetic Composition of Angiosperm Diversity in the Cloud Forests of Mexico

Several members of the most ancient living lineages of flowering plants (angiosperms) inhabit humid, woody, mostly tropical habitats. Here we assess whether one of these forest types, the cloud forests of Mexico (CFM), contain a relatively higher proportion of phylogenetically early-diverging angiosperm lineages. The CFM houses an extraordinary plant species diversity, including members of earliest-diverging angiosperm lineages. The phylogenetic composition of CFM angiosperm diversity was evaluated through the relative representation of orders and families with respect to the global flora, and the predominance of phylogenetically early- or late-diverging lineages. Goodness-of-fit tests indicated significant differences in the proportional local and global representation of angiosperm clades. The net difference between the percentage represented by each order and family in the CFM and the global flora allowed identification of clades that are overrepresented and underrepresented in the CFM. Early-diverging angiosperm orders and families were found to be neither over- nor underrepresented in the CFM. A slight predominance of late-diverging phylogenetic levels among overrepresented clades, however, was encountered in the CFM. The resulting pattern suggests that cloud forests provide habitats where the most ancient angiosperm lineages have survived in the face of accumulating species diversity belonging to phylogenetically late-diverging lineages.     

Evolutionary heritage shapes tree distributions along an Amazon‐to‐Andes elevation gradient

Understanding how evolutionary constraints shape the elevational distributions of tree lineages provides valuable insight into the future of tropical montane forests under global change. With narrow elevational ranges, high taxonomic turnover, frequent habitat specialization, and exceptional levels of endemism, tropical montane forests and trees are predicted to be highly sensitive to environmental change. Using plot census data from a gradient traversing > 3,000 m in elevation on the Amazonian flank of the Peruvian Andes, we employ phylogenetic approaches to assess the influence of evolutionary heritage on distribution trends of trees at the genus‐level. We find that closely related lineages tend to occur at similar mean elevations, with sister genera pairs occurring a mean 254 m in elevation closer to each other than the mean elevational difference between non‐sister genera pairs. We also demonstrate phylogenetic clustering both above and below 1,750 m a.s.l, corresponding roughly to the cloud‐base ecotone. Belying these general trends, some lineages occur across many different elevations. However, these highly plastic lineages are not phylogenetically clustered. Overall, our findings suggest that tropical montane forests are home to unique tree lineage diversity, constrained by their evolutionary heritage and vulnerable to substantial losses under environmental changes, such as rising temperatures or an upward shift of the cloud‐base.     

Spatial phylogenetics of the native woody plant species in Hainan,China

To better identify biodiversity hotspots for conservation on Hainan Island, a tropical island in southern China, we assessed spatial variation in phylogenetic diversity and species richness using 18,976 georeferenced specimen records and a newly reconstructed molecular phylogeny of 957 native woody plants. Within this framework, we delineated bioregions based on vegetation composition and mapped areas of neoendemism and paleoendemism to identify areas of priority for conservation. Our results reveal that the southwest of Hainan is the most important hot spot for endemism and plant diversity followed by the southeast area. The distribution of endemic species showed a scattered, rather than clustered, pattern on the island. Based on phylogenetic range‐weighted turnover metrics, we delineated three major vegetational zones in Hainan. These largely correspond to natural secondary growth and managed forests (e.g., rubber and timber forests) in central Hainan, old‐growth forests and natural secondary growth forest at the margins of Hainan, and nature reserves on the island (e.g., Jianfeng and Diaoluo National Nature Reserves). Our study helps to elucidate potential botanical conservation priorities for Hainan within an evolutionary, phylogenetic framework.     

Phylogenetic endemism of the orchids of Megamexico reveals complementary areas for conservation

Orchid diversity provides a unique opportunity to further our understanding of biotic and abiotic factors linked to patterns of richness, endemism, and phylogenetic endemism in many regions. However, orchid diversity is consistently threatened by illegal trade and habitat transformation. Here, we identified areas critical for orchid conservation in the biogeographic province of Megamexico. For this purpose, we evaluated orchid endemism, phylogenetic diversity, and phylogenetic endemism within Megamexico and characterized orchid life forms. Our results indicate that the majority of the regions with the highest estimates of endemism and phylogenetic endemism are in southern Mexico and northern Central America, mostly located on the Pacific side of Megamexico. Among the most important orchid lineages, several belong to epiphytic lineages such as Pleurothallidinae, Laeliinae and Oncidiinae. We also found that species from diverse and distantly related lineages converge in montane forests where suitable substrates for epiphytes abound. Furthermore, the southernmost areas of phylogenetic diversity and endemism of Megamexico are in unprotected areas. Thus, we conclude that the most critical areas for orchid conservation in Megamexico are located in southern Mexico and northern Central America. We recommend that these areas should be given priority by the Mexican system of natural protected areas as complementary conservation areas.     

Among cradles and museums: seasonally dry forest promotes lineage exchanges between rain forest and savanna

ABSTRACT

Background: Cerrado comprises heterogeneous vegetation types. The flora of the vegetation types has distinct biogeographic origins and thus can be hypothesised to have distinct evolutionary imprints.

Aims: We hypothesised that the flora of riparian forest was phylogenetically overdispersed, whereas those of more open habitats were phylogenetically clustered.

Methods: We built a species-level phylogeny for 1,663 species of trees, shrubs and palms and analysed the pattern of lineage distribution, alpha and beta phylogenetic diversity among Cerrado vegetation types (riparian forest, rocky savana, savana and seasonally dry forest - SDTF).

Results: We found a gradient of high phylogenetic diversity and overdispersion from riparian forests towards phylogenetic clustering in vegetation in more open and drier habitats. Habitat shifts were common along the evolutionary history of all families analysed and most families showed a high frequency of shifts from SDTF to riparian forest and savanna.

Conclusions: SDTF seems to be a transitional habitat in evolutionary terms, promoting lineage exchanges between stands of riparian forest and savanna. While riparian forest can be seen as a ‘museum’, that harbours ancient lineages, savanna and rocky savanna are a ‘cradle’ of derived lineages. Habitat shifts are an important underlying drive of high present-day Cerrado flora diversity.     

Patterns in the diversity and endemism of extant Eocene age lineages across southern Africa

Southern Africa boasts a wealth of endemic fauna and flora, comprising both massive recent radiations such as those characteristic of the Cape flora, and solitary ancient species such as the peculiar desert gymnosperm Welwitschia. This study was undertaken to identify ancient biological lineages (tetrapod and vascular plant lineages of Eocene age or older) endemic to southern Africa, and to map their distribution across the region. Twenty‐seven (17 plant and ten animal) lineages were identified, and distribution maps were generated for each of them across 74 operational geographic units, which were then combined into total endemism and corrected weighted endemism per unit area. Total endemism peaked along South Africa's coast and Great Escarpment, but in the case of weighted endemism high values were also recorded along other portions of the Great Escarpment further north. A review of the lineages sister to southern African ancient endemic lineages showed that these are often globally widespread, and many of them differ substantially from the southern African ancient lineages in terms of morphology and ecology. The mechanisms of ancient lineage survival in the region are discussed, and their importance for conservation in southern Africa is emphasised.     

Contrasting patterns of diversification in a bird family (Aves: Gruiformes: Rallidae) are revealed by analysis of geospatial distribution of species and phylogenetic diversity

Geospatial patterns in the distribution of regional biodiversity reflect the composite processes that underpin evolution: speciation, dispersal and extinction. The spatial distribution and phylogeny of a globally widespread and species rich bird family (Rallidae) were used to help assess the role of large‐scale biogeographical processes in diversity and diversification. Here, we examine how different geostatistical diversity metrics enhance our understanding of species distribution by linking occurrence records of rail species to corresponding species level phylogeny. Tropical regions and temperate zones contained a large proportion of rail species richness and phylogenetic diversity whilst small islands in Australian, Oceanian and Oriental regions held the highest weighted and phylogenetic endemism. Our results suggest that habitat connectivity and dispersal were important ecological features in rail evolution and distribution. Spatial isolation was a significant driver of diversification where islands in Oceania were centres of neo‐endemism with recent multiple and independent speciation events and could be considered as nurseries of biodiversity. Palaeo‐endemism was mostly associated with older stable regions, so despite extensive long distance range shifting these areas retain their own ancient and distinct character. Madagascar was the major area of palaeo‐endemism associated with the oldest rail lineages and could be considered a museum of rail diversity. This implies a mixture of processes determine the current distribution and diversity of rail clades with some areas dominated by recent ‘in situ’ speciation while others harbour old diversity with ecological traits that have stood the test of time.     

Unravelling the evolutionary origins of biogeographic assemblages

Aim

Floristic and faunal diversity fall within species assemblages that can be grouped into distinct biomes or ecoregions. Understanding the origins of such biogeographic assemblages helps illuminate the processes shaping present‐day diversity patterns and identifies regions with unique or distinct histories. While the fossil record is often sparse, dated phylogenies can provide a window into the evolutionary past of these regions. Here, we present a novel phylogenetic approach to investigate the evolutionary origins of present‐day biogeographic assemblages and highlight their conservation value.

Location

Southern Africa.

Methods

We evaluate the evolutionary turnover separating species clusters in space at different time slices to determine the phylogenetic depth at which the signal for their present‐day structure emerges. We suggest present‐day assemblages with distinct evolutionary histories might represent important units for conservation. We apply our method to the vegetation of southern Africa using a dated phylogeny of the woody flora of the region and explore how the evolutionary history of vegetation types compares to common conservation currencies, including species richness, endemism and threat.

Results

We show the differentiation of most present‐day vegetation types can be traced back to evolutionary splits in the Miocene. The woody flora of the Fynbos is the most evolutionarily distinct, and thus has deeper evolutionary roots, whereas the Savanna and Miombo Woodland show close phylogenetic affinities and likely represent a more recent separation. However, evolutionarily distinct phyloregions do not necessarily capture the most unique phylogenetic diversity, nor are they the most species‐rich or threatened.

Main conclusions

Our approach complements analyses of the fossil record and serves as a link to the history of diversification, migration and extinction of lineages within biogeographic assemblages that is separate from patterns of species richness and endemism. Our analysis reveals how phyloregions capture conservation value not represented by traditional biodiversity metrics.

     

The Impact of Ecology and Biogeography on Legume Diversity,Endemism, and Phylogeny in the Caribbean Region: A New Direction in Historical Biogeography

The legume family is so well represented in the Caribbean that if a preserve was needed somewhere on earth to harbor all of the primary lineages in this family, the flora of just Cuba would suffice. Molecular phylogenetic, biogeographic, and evolutionary rates analysis all suggest that legume diversity and endemism in the Caribbean are mostly of recent origin and are likely a function of the abundance of seasonally dry tropical forests (SDTFs) throughout the neotropics. Legumes have a strong ecological affinity for SDTFs, and the Caribbean basin is well covered by this forest type. Rate-variable molecular clock analysis suggests that the majority of worldwide island lineages of legumes have ages of much less than 30 Ma. Singular historical events invoking land bridges or mobile continental plates are thus not needed to explain Caribbean legume diversity and endemism. The Greater Antilles are large islands located close to the American continent. They are therefore expected to fairly represent the diverse continental lineages of legumes. Yet, they are distant enough to be dispersal limited. As such, island lineages can speciate and diversify over evolutionary time unimpeded by high rates of immigration from the mainland. Vicariance and other standard phylogenetic methods of historical biogeography are likely to be replaced by those of ecological and island biogeography. This is because model selection approaches derived from the neutral concept of isolation by distance will be able to quantify patterns of alpha and beta diversity and detect niche assembly and phylogenetic niche conservatism within and among metacommunities that are hypothesized to constrain phylogeny.     

Relict floras of Atlantic islands: patterns assessed

St. Helena (South Atlantic Ocean) and Macaronesia have endemic plants that have been interpreted as relicts. This paper divides these relicts into three main types. It is suggested that type 1 relicts are of Miocene age. They are usually wet tropical forest trees, geographically and taxonomically highly disjunct from their nearest relatives. Type 2 relicts are of Late Miocene or Pliocene age, usually trees of tropical or subtropical seasonally dry vegetation, often with trans-African disjunction, and moderate taxonomic isolation. Type 3 relicts are of Late Pliocene or Pleistocene age, usually herbs of arid zone or temperate vegetation, with little taxonomic or geographical disjunction from their nearest relatives. Relict theory is defined, with discussion of its explanatory power to account for patterns of endemism, and more generally, taxonomic and geographical patterns of biodiversity.     

Phylogenetic assemblage structure of North American trees is more strongly shaped by glacial–interglacial climate variability in gymnosperms than in angiosperms

How fast does biodiversity respond to climate change? The relationship of past and current climate with phylogenetic assemblage structure helps us to understand this question. Studies of angiosperm tree diversity in North America have already suggested effects of current water–energy balance and tropical niche conservatism. However, the role of glacial–interglacial climate variability remains to be determined, and little is known about any of these relationships for gymnosperms. Moreover, phylogenetic endemism, the concentration of unique lineages in restricted ranges, may also be related to glacial–interglacial climate variability and needs more attention. We used a refined phylogeny of both angiosperms and gymnosperms to map phylogenetic diversity, clustering and endemism of North American trees in 100‐km grid cells, and climate change velocity since Last Glacial Maximum together with postglacial accessibility to recolonization to quantify glacial–interglacial climate variability. We found: (1) Current climate is the dominant factor explaining the overall patterns, with more clustered angiosperm assemblages toward lower temperature, consistent with tropical niche conservatism. (2) Long‐term climate stability is associated with higher angiosperm endemism, while higher postglacial accessibility is linked to to more phylogenetic clustering and endemism in gymnosperms. (3) Factors linked to glacial–interglacial climate change have stronger effects on gymnosperms than on angiosperms. These results suggest that paleoclimate legacies supplement current climate in shaping phylogenetic patterns in North American trees, and especially so for gymnosperms.     

The Azores diversity enigma: why are there so few Azorean endemic flowering plants and why are they so widespread?

Aim Endemism in the flora of the Azores is high (33%) but in other respects, notably the paucity of evolutionary radiations and the widespread distribution of most endemics, the flora differs markedly from the floras of the other Macaronesian archipelagos. We evaluate hypotheses to explain the distinctive patterns observed in the Azorean endemic flora, focusing particularly on comparisons with the Canary Islands. Location Azores archipelago. Methods Data on the distribution and ecology of Azorean endemic flowering plants are reviewed to ascertain the incidence of inter‐island allopatric speciation and adaptive, ecological speciation. These are contrasted with patterns for the Canary Islands. Patterns of endemism in the Azores and Canaries are further investigated in a phylogenetic context in relation to island age. beast was used to analyse a published molecular dataset for Pericallis (Asteraceae) and to investigate the relative ages of Azorean and Canarian lineages. Results There are few examples of inter‐island allopatric speciation in the Azorean flora, despite the considerable distances between islands and sub‐archipelagos. In contrast, inter‐island allopatric speciation has been an important process in the evolution of the Canary Islands flora. Phylogenetic data suggest that Azorean endemic lineages are not necessarily recent in origin. Furthermore, in Pericallis the divergence of the Azorean endemic lineage from its closest relative pre‐dates the radiation of a Canarian herbaceous clade by inter‐island allopatric speciation. Main conclusions The data presented do not support suggestions that hypotheses pertaining to island age, age of endemic lineages and ecological diversity considered individually explain the lack of radiations and the widespread distribution of Azorean endemics. We suggest that palaeoclimatic variation, a factor rarely considered in macroecological studies of island diversity patterns, may be an important factor. Palaeoclimatic data suggest frequent and abrupt transitions between humid and arid conditions in the Canaries during the late Quaternary, and such an unstable climate may have driven the recent diversification of the flora by inter‐island allopatric speciation, a process largely absent from the climatically more stable Azores. Further phylogenetic/phylogeographic analyses are necessary to determine the relative importance of palaeoclimate and other factors in generating the patterns observed.     

Fern and lycophyte diversity in the Pacific Northwest: Patterns and predictors

Recent floristic efforts in the Pacific Northwest (PNW) have now made it possible to characterize the broad‐scale patterns of fern and lycophyte diversity across this large and geologically‐complex region of western North America. The physiography of the PNW has been developing for over 200 million years, but Pleistocene glaciation‐induced migrations and recolonizations have strongly influenced the assembly of the flora. With the high dispersal potential of fern and lycophyte spores, the distribution patterns of pteridophytes may be representative of habitat suitability more than dispersal constraints. Our objective was to describe the biodiversity of pteridophytes in the PNW, determine the spatial distribution of that biodiversity in terms of phylogenetic diversity, identify centers of regional endemism, explore the correlations between biodiversity and environmental variables, and infer possible influences of past glaciation on the pteridophyte flora. We obtained presence‐only distribution data from two online databases. A phylogenetic tree was constructed using chloroplast DNA sequence data from GenBank. We used the Biodiverse software package to estimate and map phylogenetic diversity and phylogenetic endemism across the PNW, and to identify those regions of the PNW where diversity was higher or lower than expected in comparison to randomization models. Environmental correlates of diversity were identified using principal components analysis with bioclimatic data from WorldClim.org, and we used Maxent to predict habitat suitability for species under past and future climate conditions. We found evidence for the influence of past glaciations and glacial refugia on the patterns of pteridophyte diversity, that moisture availability and cold temperatures are strongly correlated with patterns of genus richness, phylogenetic diversity, and phylogenetic endemism. We infer that the topographic complexity of the region may be driving the assembly of the pteridophyte flora indirectly by influencing climate and precipitation patterns.     

Origins and patterns of endemic diversity in two specialized lizard lineages from the Australian Monsoonal Tropics (Oedura spp.)

Aim

Savanna biomes cover around 20% of land surfaces, yet the origins and processes that have shaped their biodiversity remain understudied. Here, we assess the timing of diversification and how patterns of genetic diversity vary along an aridity gradient in specialized saxicoline gecko clades (Oedura spp.) from the tropical savannas of northern Australia.

Location

Australian Monsoonal Tropics (AMT), Kimberley region (Western Australia).

Methods

We compiled mitochondrial and nuclear data for two Kimberley endemic lizard clades (Oedura filicipoda/murrumanu and O. gracilis), and allied non‐Kimberley taxa (O. marmorata complex). Species delimitation methods were used to identify evolutionary lineages, Maximum‐likelihood and Bayesian phylogenetic methods were employed to assess relationships and diversification timeframes, and rainfall data and range sizes were tested for correlations.

Results

Phylogenetic analyses of cryptic or recently discovered lineage diversity revealed late‐Miocene to early‐Pliocene crown ages. Microendemism and diversity were highest in high‐rainfall regions, while the most widespread lineages occurred in the central and south‐east Kimberley, and showed evidence of introgression with parapatric lineages.

Main conclusions

The initial diversification in both clades was broadly concordant with global climatic events linked to the expansion of savanna biomes in the lateMiocene. Higher endemism in mesic and refugial areas suggests long histories of localized persistence, while wider distributions and evidence of introgression suggest a dynamic history at the arid‐monsoonal interface.     

Spatial phylogenetics of the Chinese angiosperm flora provides insights into endemism and conservation

The flora of China is well known for its high diversity and endemism. Identifying centers of endemism and designating conservation priorities are essential goals for biodiversity studies.However, there is no comprehensive study from a rigorous phylogenetic perspective to understand patterns of diversity and endemism and to guide biodiversity conservation in China. We conducted a spatial phylogenetic analysis of the Chinese angiosperm flora at the generic level to identify centers of neo-and pale...     

Incorporating phylogenetic information for the definition of floristic districts in hyperdiverse Amazon forests: Implications for conservation

Using complementary metrics to evaluate phylogenetic diversity can facilitate the delimitation of floristic units and conservation priority areas. In this study, we describe the spatial patterns of phylogenetic alpha and beta diversity, phylogenetic endemism, and evolutionary distinctiveness of the hyperdiverse Ecuador Amazon forests and define priority areas for conservation. We established a network of 62 one‐hectare plots in terra firme forests of Ecuadorian Amazon. In these plots, we tagged, collected, and identified every single adult tree with dbh ≥10 cm. These data were combined with a regional community phylogenetic tree to calculate different phylogenetic diversity (PD) metrics in order to create spatial models. We used Loess regression to estimate the spatial variation of taxonomic and phylogenetic beta diversity as well as phylogenetic endemism and evolutionary distinctiveness. We found evidence for the definition of three floristic districts in the Ecuadorian Amazon, supported by both taxonomic and phylogenetic diversity data. Areas with high levels of phylogenetic endemism and evolutionary distinctiveness in Ecuadorian Amazon forests are unprotected. Furthermore, these areas are severely threatened by proposed plans of oil and mining extraction at large scales and should be prioritized in conservation planning for this region.     

Climate‐driven rampant speciation of the Cape flora

Aim To test whether the radiation of the extremely rich Cape flora is correlated with marine‐driven climate change. Location Middle to Late Miocene in the south‐east Atlantic and the Benguela Upwelling System (BUS) off the west coast of South Africa. Methods We studied the palynology of the thoroughly dated Middle to Late Miocene sediments of Ocean Drilling Program (ODP) Site 1085 retrieved from the Atlantic off the mouth of the Orange River. Both marine upwelling and terrestrial input are recorded at this site, which allows a direct correlation between changes in the terrestrial flora and the marine BUS in the south‐east Atlantic. Results Pollen types from plants of tropical affinity disappeared, and those from the Cape flora gradually increased, between 10 and 6 Ma. Our data corroborate the inferred dating of the diversification in Aizoaceae c. 8 Ma. Main conclusions Inferred vegetation changes for the Late Miocene south‐western African coast are the disappearance of Podocarpus‐dominated Afromontane forests, and a change in the vegetation of the coastal plain from tropical grassland and thicket to semi‐arid succulent vegetation. These changes are indicative of an increased summer drought, and are in step with the development of the southern BUS. They pre‐date the Pliocene uplift of the East African escarpment, suggesting that this did not play a role in stimulating vegetation change. Some Fynbos elements were present throughout the recorded period (from 11 Ma), suggesting that at least some elements of this vegetation were already in place during the onset of the BUS. This is consistent with a marine‐driven climate change in south‐western Africa triggering substantial radiation in the terrestrial flora, especially in the Aizoaceae.