Frequent and parallel habitat transitions as driver of unbounded radiations in the Cape flora

The enormous species richness in the Cape Floristic Region (CFR) of Southern Africa is the result of numerous radiations, but the temporal progression and possible mechanisms of these radiations are still poorly understood. Here, we explore the macroevolutionary dynamics of the Restionaceae, which include 340 species that are found in all vegetation types in the Cape flora and are ecologically dominant in fynbos. Using an almost complete (i.e., 98%) species‐level time calibrated phylogeny and models of diversification dynamics, we show that species diversification is constant through the Cenozoic, with no evidence of an acceleration with the onset of the modern winter‐wet climate, or a recent density‐dependent slowdown. Contrary to expectation, species inhabiting the oldest (montane) and most extensive (drylands) habitats did not undergo higher diversification rates than species in the younger (lowlands) and more restricted (wetland) habitats. We show that the rate of habitat transitions is more closely related to the speciation rate than to time, and that more than a quarter of all speciation events are associated with habitat transitions. This suggests that the unbounded Restionaceae diversification resulted from numerous, parallel, habitat shifts, rather than persistence in a habitat stimulating speciation. We speculate that this could be one of the mechanisms resulting in the hyperdiverse Cape flora.     

Fire and other variables as determinants of forest/fynbos boundaries in the Cape Province

Abstract. Forests in the southwestern Cape are restricted to sheltered stream banks and patches of rock scree owing to protection from recurrent fire and the more favourable soil moisture conditions in these habitats. The effects of a prescribed burn on 17 March 1987 on the forest / fynbos boundaries in the Swartboskloof catchment are described. No evidence of erosion of the forest edge was recorded, although this could be ascribed to the low intensity of the fire. Soil properties, primarily pH and aluminium and available phosphorus levels, differed markedly between forest and fynbos. It is argued, however, that this is a result, not a cause of vegetation change. Light extinction levels and soil moisture availability after the fire were lower in the fynbos than in the forest, and are considered to preclude incursions of forest species into fynbos of a low post-fire age.     

POSTFIRE RESPONSE AND GENETIC DIVERSITY IN ERICA COCCINEA: CONNECTING POPULATION DYNAMICS AND DIVERSIFICATION IN A BIODIVERSITY HOTSPOT

Understanding the processes of biological diversification is a central topic in evolutionary biology. The South African Cape fynbos, one of the major plant biodiversity hotspots out of the tropics, has prompted several hypotheses about the causes of generation and maintenance of biodiversity. Fire has been traditionally invoked as a key element to explain high levels of biodiversity in highly speciose fynbos taxa, such as the genus Erica. In this study, we have implemented a microevolutionary approach to elucidate how plant‐response to fire may contribute to explain high levels of diversification in Erica. By using microsatellite markers, we investigated the genetic background of seeder (fire‐sensitive) and resprouter (fire‐resistant) populations of the fynbos species Erica coccinea. We found higher within‐population genetic diversity and higher among‐population differentiation in seeder populations and interpreted these higher levels of genetic diversification as a consequence of the comparatively shorter generation times and faster population turnover in the seeder form of this species. Considering that genetic divergence among populations may be seen as the initial step to speciation, the parallelism between these results and the pattern of biodiversity at the genus level offers stimulating insights into understanding causes of speciation of the genus Erica in the Cape fynbos.     

Diversity versus disparity and the radiation of modern cetaceans

Modern whales are frequently described as an adaptive radiation spurred by either the evolution of various key innovations (such as baleen or echolocation) or ecological opportunity following the demise of archaic whales. Recent analyses of diversification rate shifts on molecular phylogenies raise doubts about this interpretation since they find no evidence of increased speciation rates during the early evolution of modern taxa. However, one of the central predictions of ecological adaptive radiation is rapid phenotypic diversification, and the tempo of phenotypic evolution has yet to be quantified in cetaceans. Using a time-calibrated molecular phylogeny of extant cetaceans and a morphological dataset on size, we find evidence that cetacean lineages partitioned size niches early in the evolutionary history of neocetes and that changes in cetacean size are consistent with shifts in dietary strategy. We conclude that the signature of adaptive radiations may be retained within morphological traits even after equilibrium diversity has been reached and high extinction or fluctuations in net diversification have erased any signature of an early burst of diversification in the structure of the phylogeny.     

Ecological opportunity alters the timing and shape of adaptive radiation

The uneven distribution of diversity is a conspicuous phenomenon across the tree of life. Ecological opportunity is a prominent catalyst of adaptive radiation and therefore may alter patterns of diversification. We evaluated the distribution of shifts in diversification rates across the cichlid phylogeny and the distribution of major clades across phylogenetic space. We also tested if ecological opportunity influenced these patterns. Colonization‐associated ecological opportunity altered the tempo and mode of diversification during the adaptive radiation of cichlid fishes. Clades that arose following colonization events diversified faster than other clades. Speciation rate shifts were nonrandomly distributed across the phylogeny such that they were disproportionally concentrated around nodes that corresponded with colonization events (i.e., of continents, river basins, or lakes). Young clades tend to expand faster than older clades; however, colonization‐associated ecological opportunity accentuated this pattern. There was an interaction between clade age and ecological opportunity that explained the trajectory of clades through phylogenetic space over time. Our results indicate that ecological opportunities afforded by continental and ecosystem‐scale colonization events explain the dramatic speciation rate heterogeneity and phylogenetic imbalance that arose during the evolutionary history of cichlid fishes.     

火烧强度对兴安落叶松群落叶片功能性状及功能多样性的影响

林火是北方针叶林的重要生态因子,直接影响火烧迹地物种多样性及功能多样性,进而影响森林群落的演替.以牙克石地区火后自然恢复12年的兴安落叶松群落为研究对象,在群落尺度上分析火烧迹地土壤养分含量、叶片功能性状、物种多样性和功能多样性在不同火烧强度(轻、中和重度火烧)下的变化规律.结果表明: 火烧显著降低了土壤全氮含量,对土壤全磷含量无显著影响.轻、中度火烧有助于维持群落较高的物种多样性和功能多样性;中度火烧显著增加了群落的物种多样性,物种丰富度指数、Shannon指数、Simpson指数和Pielou均匀度指数均在中度火烧时有最大值;火烧降低了群落的功能丰富度和功能离散度指数,二者分别在中、轻度火烧时最大(除未过火外),而群落的功能均匀度和二次熵指数在林火干扰后增加,轻度火烧后最大.随火烧强度增加,叶干物质含量、叶组织密度和叶磷含量总体呈显著增大趋势,而比叶面积、叶含水量、叶氮含量和叶N∶P则表现出未过火>中度火烧>轻度火烧>重度火烧的规律,叶片厚度呈先增大后减小的变化规律.火烧强度对森林群落的叶片功能性状和功能多样性均有显著影响,且适度的火干扰对森林群落的恢复具有促进作用.     

A conceptual and statistical framework for adaptive radiations with a key role for diversity dependence

Abstract In this article we propose a new framework for studying adaptive radiations in the context of diversity-dependent diversification. Diversity dependence causes diversification to decelerate at the end of an adaptive radiation but also plays a key role in the initial pulse of diversification. In particular, key innovations (which in our definition include novel traits as well as new environments) may cause decoupling of the diversity-dependent dynamics of the innovative clade from the diversity-dependent dynamics of its ancestral clade. We present a likelihood-based inference method to test for decoupling of diversity dependence using molecular phylogenies. The method, which can handle incomplete phylogenies, identifies when the decoupling took place and which diversification parameters are affected. We illustrate our approach by applying it to the molecular phylogeny of the North American clade of the legume tribe Psoraleeae (47 extant species, of which 4 are missing). Two diversification rate shifts were previously identified for this clade; our analysis shows that the first, positive shift can be associated with decoupling of two Pediomelum subgenera from the other Psoraleeae lineages, while we argue that the second, negative shift can be attributed to speciation being protracted. The latter explanation yields nonzero extinction rates, in contrast to previous findings. Our framework offers a new perspective on macroevolution: new environments and novel traits (ecological opportunity) and diversity dependence (ecological limits) cannot be considered separately.     

不同火烈度火烧迹地内油松叶功能性状的变化

叶功能性状对林火的响应是林火生态领域的研究热点之一,研究火后油松叶功能性状变化能够揭示油松为适应火环境形成的生长策略,为促进油松火后恢复提供参考。以山西省沁源县火烧迹地内油松为研究对象,选择当年生叶片分析叶功能性状在不同火烈度(未过火、轻度火烧、中度火烧)火烧迹地间的变化规律,并研究不同火烧迹地内叶经济谱的变化特征。结果表明: 除氮磷比外,叶功能性状在不同火烈度的火烧迹地间存在显著差异,其中,叶面积的差异最为明显,是最敏感的性状。随火烧迹地内火烈度的增加,叶面积、叶厚度、叶干物质含量、叶氮含量和叶磷含量升高,比叶面积、叶有机碳含量降低。部分叶功能性状间存在显著的相关关系,但其相关性在不同火烈度的火烧迹地间存在差异。叶经济谱沿着“未过火-轻度火烧-中度火烧”的火烧迹地环境总体向“快速投资-收益型”的资源权衡策略移动,低烈度火烧迹地内油松的生长恢复会加快。     

Contrasting mechanisms of resilience at mesic and semi-arid boundaries of fynbos,a mega-diverse heathland of South Africa

Biome boundaries are expected to be sensitive to changes in climate and disturbance, because it is here that ecological communities are at environmental, ecological or disturbance limits. Using palaeoecology to study ecosystem dynamics at biome boundaries provides opportunities for understanding ecosystem resilience or sensitivity at ecologically meaningful timescales, and under varying climatic and disturbance conditions.The fynbos biome is a megadiverse Mediterranean type shrubland, found only in South Africa, that is threatened by climate change, land-use change and invasion by alien species. We used palaeoecological records from the semi-arid and mesic boundaries of the fynbos biome to test hypotheses regarding ecosystem resilience over timescales of centuries to millennia. We hypothesised that fynbos would expand at its mesic boundary at the expense of afrotemperate forest under drier and / or more fire prone conditions. In contrast, we hypothesised that at the semi-arid boundary, fynbos would expand at the expense of succulent karoo under wetter and cooler and / or more fire-prone conditions. Contrary to our expectations, the fossil pollen record at both biome boundaries showed remarkable stability at centennial - millennial timescales. To explain our results, we generated new hypotheses exploring possible mechanisms that might confer resilience.At the mesic (temperate) boundary, we suggest that decreased seasonality of rainfall during drier phases favoured fire and fynbos persistence, while in wetter periods, increased seasonality of rainfall resulted in enhanced summer drought stress, inhibiting forest expansion. At this boundary, internal reorganisation from grassy to proteoid fynbos states conferred resilience through resistance. At the succulent karoo boundary, we suggest that increased aridity was offset by less seasonality of rainfall, which enhanced biomass and allowed fire to persist, favouring persistence of fynbos. At this boundary, fynbos sensu stricto retreated during arid phases but recovered during climate amelioration, consistent with resilience through recovery. In both cases, this mega-diverse, disturbance-adapted flora provided a range of traits that enabled fynbos to persist despite environmental perturbation. Our findings agree with general observations that for ecosystems in regions of ample resource availability (i.e. at the mesic boundary), biotic interactions and disturbance tend to become more important in ecosystem dynamics, whereas in regions of scarce resources (in this case water scarcity at the semi-arid boundary) abiotic stress is more important. Our findings contribute to debates over the mechanisms that confer resistance and resilience to environmental change. Understanding and conserving the processes and mechanisms underpinning its resilience will be critical to effective conservation planning.     

ECOLOGICAL OPPORTUNITY AND THE RATE OF MORPHOLOGICAL EVOLUTION IN THE DIVERSIFICATION OF GREATER ANTILLEAN ANOLES

The pace of phenotypic diversification during adaptive radiation should decrease as ecological opportunity declines. We test this prediction using phylogenetic comparative analyses of a wide range of morphological traits in Greater Antillean Anolis lizards. We find that the rate of diversification along two important axes of Anolis radiation—body size and limb dimensions—decreased as opportunity declined, with opportunity quantified either as time elapsed in the radiation or as the diversity of competing anole lineages inferred to have been present on an island at different times in the past. Most previous studies of the ecological opportunity hypothesis have focused on the rate of species diversification; our results provide a complementary perspective, indicating that the rate of phenotypic diversification declines with decreasing opportunity in an adaptive radiation.     

Pollination ecotypes of Satyrium hallackii (Orchidaceae) in South Africa

Intraspecific variation in floral traits may reflect adaptive shifts in the pollination system of a plant. This idea was tested by examining the pollination biology of Satyrium hallackii H. Bolus., an orchid which has spurs varying from 8 to 36 mm among populations in southern Africa. Field observations showed that the short-spurred form ( S. hallackii subsp. hallackii ) in coastal fynbos habitats is pollinated by bees, while the long-spurred form ( S. hallackii subsp. ocellatum ) in grassland habitats is pollinated primarily by hawkmoths and, secondarily, by long-tongued flies. The shift between hawkmoths and bees as pollinators may have been promoted by an ecological gradient in South Africa: carpenter bees are common in coastal fynbos habitats, but rare in grassland habitats where there are few available nesting sites. On the other hand, hawkmoths are common in grassland habitats, but rare in fynbos where there are few palatable host-plants. The formation of pollination ecotypes across pollinator gradients has probably been a major factor in the diversification of South African plants with specialized pollination systems.     

Effects of alien pine plantations on small mammal community structure in a southern African biodiversity hotspot

Commercial plantations and alien tree invasions often have substantial negative impacts on local biodiversity. The effect of plantations on faunal communities in the fire‐adapted fynbos vegetation of the Cape Floristic Region biodiversity hotspot is not yet well quantified. We studied small mammal community structure in alien Pinus radiata plantations and adjacent fynbos regenerating after clear‐felling of plantations on the Cape Peninsula, South Africa. Small mammal sampling over 1,800 trap‐nights resulted in 480 captures of 345 individuals (excluding recaptures) representing six species. Significantly more species, individuals (12 X) and biomasses (29 X) of small mammals occurred on recovering fynbos sites compared to plantations. This was commensurate with a higher diversity of plant growth forms, vegetation densities and live vegetation biomass. Only one small mammal species, the pygmy mouse (Mus minutoides), was consistently trapped within plantations. Fynbos sites were dominated by three small mammal species that are ecological generalists and early successional pioneer species, rendering the recovering fynbos slightly depauperate in terms of species richness and evenness relative to other studies done in pristine fynbos. We make three recommendations for forestry that would facilitate the restoration of more diverse natural plant communities and progressively more diverse and dynamic small mammal assemblages in a key biodiversity hotspot.     

Do habitat shifts drive diversification in teleost fishes? An example from the pufferfishes (Tetraodontidae)

Habitat shifts are implicated as the cause of many vertebrate radiations, yet relatively few empirical studies quantify patterns of diversification following colonization of new habitats in fishes. The pufferfishes (family Tetraodon‐tidae) occur in several habitats, including coral reefs and freshwater, which are thought to provide ecological opportunity for adaptive radiation, and thus provide a unique system for testing the hypothesis that shifts to new habitats alter diversification rates. To test this hypothesis, we sequenced eight genes for 96 species of pufferfishes and closely related porcupine fishes, and added 19 species from sequences available in GenBank. We time‐calibrated the molecular phylogeny using three fossils, and performed several comparative analyses to test whether colonization of novel habitats led to shifts in the rate of speciation and body size evolution, central predictions of clades experiencing ecological adaptive radiation. Colonization of freshwater is associated with lower rates of cladogenesis in pufferfishes, although these lineages also exhibit accelerated rates of body size evolution. Increased rates of cladogenesis are associated with transitions to coral reefs, but reef lineages surprisingly exhibit significantly lower rates of body size evolution. These results suggest that ecological opportunity afforded by novel habitats may be limited for pufferfishes due to competition with other species, constraints relating to pufferfish life history and trophic ecology, and other factors.     

A comparison of fynbos and non-fynbos coenoclines in the lower Gamtoos River Valley,southeastern Cape,South Africa

Patterns in the relative importance of structural attributes and growth forms along fynbos and non-fynbos coenoclines were studied to test the hypothesis that there would be less structural variation in the former because the overriding influence of low levels of soil nutrients would be manifest in a great deal of structural convergence in fynbos. The coenoclines were ranged along identical environmental gradients of increasing altitude, rainfall and soil moisture and decreasing climatic variability. Results showed that along the entire fynbos coenocline vegetation was structurally a small-leaved sclerophyllous shrubland with a graminoid understorey and, usually, a large-leaved (proteoid) shrub overstorey. Fynbos structure was interpreted largely as a response to low levels of soil nutrients. Non-fynbos vegetation ranged from mixed succulent-sclerophyllous and spiny large-leaved thicket at lower altitudes to tall mesic forest at the upper end of the gradient. Non-fynbos structure was explained in terms of variations in soil moisture and climate. An analysis of the biogeographical affinities of sample floras at sites along the coenoclines showed that fynbos vegetation was dominated by taxa endemic to the Cape phytochorion, although phytochorological mixing was pronounced at the lower altitude sites. The level of local endemism in the fynbos coenocline was relatively high; nearly all endemics were Cape fynbos taxa and their incidence increased with increasing altitude. These data indicate that fynbos vegetation has had a lengthy history in the southeastern Cape and that high altitude sites would have comprised a refuge for Cape taxa during unfavourable climatic periods. Non-fynbos vegetation ranged from dry subtropical Tongaland-Pondoland thicket with a strong Karoo-Namib component to temperate Afromontane forest. Levels of endemism were lower than the fynbos coenocline and decreased with increasing altitude. The high number of karroid endemics found in both coenoclines at low altitudes suggests that karroid vegetation would have been more widespread in the past, probably during the last glacial which was considerably drier than the present Holocene interglacial.     

Morphological innovation,ecological opportunity,and the radiation of a major vascular epiphyte lineage

The emergence of angiosperm‐dominated tropical forests in the Cretaceous led to major shifts in the composition of biodiversity on Earth. Among these was the rise to prominence of epiphytic plant lineages, which today comprise an estimated one‐quarter of tropical vascular plant diversity. Among the most successful epiphytic groups is the Polypodiaceae, which comprises an estimated 1500 species and displays a remarkable breadth of morphological and ecological diversity. Using a time‐calibrated phylogeny for 417 species, we characterized macroevolutionary patterns in the family, identified shifts in diversification rate, and identified traits that are potential drivers of diversification. We find high diversification rates throughout the family, evidence for a radiation in a large clade of Paleotropical species, and support for increased rates of diversification associated with traits including chlorophyllous spores and noncordiform gametophytes. Contrary to previous hypotheses, our results indicate epiphytic species and groups with humus‐collecting leaves diversify at lower rates than the family as a whole. We find that diversification rates in the Polypodiaceae are positively correlated with changes in elevation. Repeated successful exploration of novel habitat types, rather than morphological innovation, appears to be the primary driver of diversification in this group.     

PLANT RESPONSES TO CLIMATE IN THE CAPE FLORISTIC REGION OF SOUTH AFRICA: EVIDENCE FOR ADAPTIVE DIFFERENTIATION IN THE PROTEACEAE

Local adaptation along environmental gradients may drive plant species radiation within the Cape Floristic Region (CFR), yet few studies examine the role of ecologically based divergent selection within CFR clades. In this study, we ask whether populations within the monophyletic white protea clade (Protea section Exsertae, Proteaceae) differ in key functional traits along environmental gradients and whether differences are consistent with local adaptation. Using seven taxa, we measured trait–environment associations and selection gradients across 35 populations of wild adults and their offspring grown in two common gardens. Focal traits were leaf size and shape, specific leaf area (SLA), stomatal density, growth, and photosynthetic rate. Analyses on wild and common garden plants revealed heritable trait differences that were associated with gradients in rainfall seasonality, drought stress, cold stress, and less frequently, soil fertility. Divergent selection between gardens generally matched trait–environment correlations and literature‐based predictions, yet variation in selection regimes among wild populations generally did not. Thus, selection via seedling survival may promote gradient‐wide differences in SLA and leaf area more than does selection via adult fecundity. By focusing on the traits, life stages, and environmental clines that drive divergent selection, our study uniquely demonstrates adaptive differentiation among plant populations in the CFR.     

Concentration of ultraviolet-B radiation absorbing compounds in leaves of a range of fynbos species

Leaves from 38 species representing 17 families were collected from contrasting elevations (100 to 824 m above sea level) in sclerophyllous mountain fynbos vegetation near Hermanus, South Africa, and the leaf percentage dry mass (PDM), specific leaf mass (SLM) and ultraviolet-B (UV-B, 280 to 320 nm) absorbance properties determined. Leaf PDM, SLM and UV-B absorbance were generally high compared to mesophyllous plants. Leaves collected at high elevation exhibited higher absorbances per unit dry mass at both 280 and 320 nm than those from the same species at low elevation. No differences in absorbance per unit leaf area were observed. UV-B absorbance properties differed between families, with high values obtained for the Apiaceae, Asteraceae, Cyperaceae, Ericaceae, Penaeaceae and Proteaceae, but lower values for the Anacardiaceae, Fabaceae and Geraniaceae. A positive correlation was found between absorbance at 280 nm per unit leaf area and SLM. It was concluded that most fynbos species, on account of their highly sclerophyllous leaves and large accumulation of UV-B absorbing compounds (flavonoids and related phenolics) may be well protected against future increases in UV-B radiation.     

Trophic novelty is linked to exceptional rates of morphological diversification in two adaptive radiations of Cyprinodon pupfish

Adaptive radiations are known for rapid morphological and species diversification in response to ecological opportunity, but it remains unclear if distinct mechanisms drive this pattern. Here, we show that rapid rates of morphological diversification are linked to the evolution of novel ecological niches in two independent Cyprinodon radiations nested within a wide-ranging group repeatedly isolated in extreme environments. We constructed a molecular phylogeny for the Cyprinodontidae, measured 16 functional traits across this group, and compared the likelihoods of single or multiple rates of morphological diversification. We found that rates of morphological diversification within two sympatric Cyprinodon clades containing unique trophic specialists are not part of an adaptive continuum with other clades, but are instead extreme outliers with rates up to 131 times faster than other Cyprinodontidae. High rates were not explained by clade age, but were instead linked to unique trophic niches within Cyprinodon, including scale-eating, zooplanktivory, and piscivory. Furthermore, although both radiations occur in similar environments and have similar sister species, they each evolved unique trophic specialists and high rates of morphological diversification in different sets of traits. We propose that the invasion of novel ecological niches may be a key mechanism driving many classic examples of adaptive radiation.     

Ecomorphological diversification following continental colonization in muroid rodents (Rodentia: Muroidea)

The emergence of exceptionally diverse clades is often attributed to ecological opportunity. For example, the exceptional diversity in the most diverse superfamily of mammals, muroid rodents, has been explained in terms of multiple independent adaptive radiations. If multiple ecological opportunity events are responsible for generating muroid diversity, we expect to find evidence of these lineages ecologically diversifying following dispersal into new biogeographical areas. In the present study, we tested the trait‐based predictions of ecological opportunity using data on body size, appendages, and elevation in combination with previously published data on biogeographical transitions and a time‐calibrated molecular phylogeny. We identified weak to no support of early ecological diversification following the initial colonizations of all continental regions, based on multiple tests, including node height tests, disparity through time plots, evolutionary model comparison, and Bayesian analysis of macroevolutionary mixtures. Clades identified with increased diversification rates, not associated with geographical transitions, also did not show patterns of phenotypic divergence predicted by ecological opportunity, which suggests that phylogenetic diversity and phenotypic disparity may be decoupled in muroids. These results indicate that shifts in diversification rates and biogeographically‐mediated ecological opportunity are poor predictors of phenotypic diversity patterns in muroids.