Edaphic Endemics Research Articles

The genetic basis of traits associated with the evolution of serpentine endemism in monkeyflowers.

The floras on chemically and physically challenging soils, such as gypsum, shale, and serpentine, are characterized by narrowly endemic species. The evolution of edaphic endemics may be facilitated or constrained by genetic correlations among traits contributing to adaptation and reproductive isolation across soil boundaries. The yellow monkeyflowers in the Mimulus guttatus species complex are an ideal system in which to examine these evolutionary patterns. To determine the genetic basis of adaptive and prezygotic isolating traits, we performed genetic mapping experiments with F2 hybrids derived from a cross between a serpentine endemic, M. nudatus, and its close relative M. guttatus. Few large effect and many small effect QTL contribute to interspecific divergence in life history, floral, and leaf traits, and a history of directional selection contributed to trait divergence. Loci contributing to adaptive traits and prezygotic reproductive isolation overlap, and their allelic effects are largely in the direction of species divergence. These loci contain promising candidate genes regulating flowering time and plant organ size. Together, our results suggest that genetic correlations among traits can facilitate the evolution of adaptation and speciation and may be a common feature of the genetic architecture of divergence between edaphic endemics and their widespread relatives.

Edaphic specialization onto bare, rocky outcrops as a factor in the evolution of desert angiosperms

Understanding the processes that enable organisms to shift into more arid environments as they emerge is critical for gauging resilience to climate change, yet these forces remain poorly known. In a comprehensive clade-based study, we investigate recent shifts into North American deserts in the rock daisies (tribe Perityleae), a diverse tribe of desert sunflowers (Compositae). We sample rock daisies across two separate contact zones between tropical deciduous forest and desert biomes in western North America and infer a time-calibrated phylogeny based on target capture sequence data. We infer biome shifts using Bayesian inference with paleobiome-informed models and find evidence for seven independent shifts into desert habitats since the onset of aridification in the late Miocene. The earliest shift occurred out of tropical deciduous forests and led to an extensive radiation throughout North American deserts that accounts for the majority of extant desert rock daisies. Estimates of life history and micro-habitat in the rock daisies reveal a correlation between a suffrutescent perennial life history and edaphic endemism onto rocky outcrops, an ecological specialization that evolved prior to establishment and diversification in deserts. That the insular radiation of desert rock daisies stemmed from ancestors preadapted for dry conditions as edaphic endemics in otherwise densely vegetated tropical deciduous forests in northwest Mexico underscores the crucial role of exaptation and dispersal for shifts into arid environments.

Limited seed dispersal distance in endemic species from tropical mountaintop grasslands may restrict upward migration in response to climate change

Understanding dispersal systems is a key step towards improving our ability to predict species migration trajectories in response to climate change. However, the abiotic seed dispersal systems of tropical plants remain largely overlooked, with studies inferring seed dispersal from fruit morphology than conducting empirical tests. Here, we determined seed dispersal season experimentally tested the seed dispersal mode, and estimated dispersal distance kernels for five edaphic specialists, mountaintop endemics from campo rupestre, a tropical, megadiverse and nutrient-impoverished grassland. We monitored fruiting phenology for 12 months and established field and laboratory experiments to assess the role of abiotic factors in releasing seeds from dry capsules and to measure seed dispersal distance. Contrary to previous studies that suggested abiotic dispersal, we found no evidence of water- or wind-operated primary seed dispersal mechanisms for all species. We show that fruiting peaks coincided with optimum conditions for seedling establishment, and that short seed dispersal distances are consistent with the idea of reduced dispersibility in edaphic specialists. Seed buoyancy experiments show a large potential for secondary seed dispersal by water run-off. Our data suggest that abiotic dispersal in the study species plays a limited role in displacing seeds away from parental plants, a pattern that markedly contrasts with that found in other vegetation types. Short-distance dispersal in campo rupestre is consistent with theory explaining the evolution of phylomatry, but it also represents reason for concern of species migration in face of global climate changes. The combination of edaphic endemism and limited dispersal in mountaintops species provides limited opportunity for migration and poses a significant threat to their long-term survival.

Testing the concept of edaphism for the quartz island flora of the Knersvlakte, South Africa

• Plant-functional traits are useful to study floras on unusual soils. • Quartz island flora, traits and environment differ from the surrounding. • Taxonomic, functional and edaphic diversity are high on quartz islands. • The flora of unusual edaphic habitats is worthy of particular protection. Edaphically unusual sites harbor a flora that is often unique in its taxonomic as well as functional composition and contain many endemic species. While this edaphic endemism, edaphism, has been discussed previously on various special soils, in this study we relate it for the first time to the floristic characteristics of so called “quartz islands”, edaphically driven special habitats in South Africa, and their surroundings (matrix habitat). We sampled the vegetation and soil for 245 vegetation plots in the Knersvlakte Nature Reserve and examined 336 plant species with respect to their evolutionary (plant family, habitat specificity), biogeographical (distributional range) as well as functional traits (life form, leaf succulence, flowering onset, dispersal mode). Based on the floristic uniqueness of the quartz islands, we expect them to host a highly specialized and locally endemic flora with smaller life forms, pronounced leaf succulence (both indicative of resource-conservative economics) and distinct reproductive traits than their surroundings. We find that, compared to the matrix habitat, quartz islands have significantly higher salinity and lower soil pH, and show higher proportion of leaf-succulent species of smaller life forms – some of which are restricted to quartz islands and have different timing of flowering onset, while dispersal via zoochory is less important. Like other special soils, quartz islands harbor unique plant species, which are distinguished by distinct trait patterns. We conclude that the concept of edaphism can be applied to quartz islands.

When disturbances favour species adapted to stressful soils: grazing may benefit soil specialists in gypsum plant communities

Background Herbivory and extreme soils are drivers of plant evolution. Adaptation to extreme soils often implies substrate-specific traits, and resistance to herbivory involves tolerance or avoidance mechanisms. However, little research has been done on the effect of grazing on plant communities rich in edaphic endemics growing on extreme soils. A widespread study case is gypsum drylands, where livestock grazing often prevails. Despite their limiting conditions, gypsum soils host a unique and highly specialised flora, identified as a conservation priority. Methods We evaluated the effect of different grazing intensities on the assembly of perennial plant communities growing on gypsum soils. We considered the contribution of species gypsum affinity and key functional traits of species such as traits related to gypsum specialisation (leaf S accumulation) or traits related to plant tolerance to herbivory such as leaf C and N concentrations. The effect of grazing intensity on plant community indices (i.e., richness, diversity, community weighted-means (CWM) and functional diversity (FD) indices for each trait) were modelled using Generalised Linear Mixed Models (GLMM). We analysed the relative contribution of interspecific trait variation and intraspecific trait variation (ITV) in shifts of community index values. Results Livestock grazing may benefit gypsum plant specialists during community assembly, as species with high gypsum affinity, and high leaf S contents, were more likely to assemble in the most grazed plots. Grazing also promoted species with traits related to herbivory tolerance, as species with a rapid-growth strategy (high leaf N, low leaf C) were promoted under high grazing conditions. Species that ultimately formed gypsum plant communities had sufficient functional variability among individuals to cope with different grazing intensities, as intraspecific variability was the main component of species assembly for CWM values. Conclusions The positive effects of grazing on plant communities in gypsum soils indicate that livestock may be a key tool for the conservation of these edaphic endemics.

Recent and ancient evolutionary events shaped plant elemental composition of edaphic endemics: a phylogeny-wide analysis of Iberian gypsum plants.

Summary The analysis of plant elemental composition and the underlying factors affecting its variation are a current hot topic in ecology. Ecological adaptation to atypical soils may shift plant elemental composition. However, no previous studies have evaluated its relevance against other factors such as phylogeny, climate or individual soil conditions.We evaluated the effect of the phylogeny, environment (climate, soil), and affinity to gypsum soils on the elemental composition of 83 taxa typical of Iberian gypsum ecosystems. We used a new statistical procedure (multiple phylogenetic variance decomposition, MPVD) to decompose total explained variance by different factors across all nodes in the phylogenetic tree of target species (covering 120 million years of Angiosperm evolution).Our results highlight the relevance of phylogeny on the elemental composition of plants both at early (with the development of key preadaptive traits) and recent divergence times (diversification of the Iberian gypsum flora concurrent with Iberian gypsum deposit accumulation). Despite the predominant phylogenetic effect, plant adaptation to gypsum soils had a strong impact on the elemental composition of plants, particularly on sulphur concentrations, while climate and soil effects were smaller.Accordingly, we detected a convergent evolution of gypsum specialists from different lineages on increased sulphur and magnesium foliar concentrations.

Target Enrichment and Extensive Population Sampling Help Untangle the Recent, Rapid Radiation of Oenothera Sect. Calylophus.

Oenothera sect. Calylophus is a North American group of 13 recognized taxa in the evening primrose family (Onagraceae) with an evolutionary history that may include independent origins of bee pollination, edaphic endemism, and permanent translocation heterozygosity. Like other groups that radiated relatively recently and rapidly, taxon boundaries within Oenothera sect. Calylophus have remained challenging to circumscribe. In this study, we used target enrichment, flanking noncoding regions, gene tree/species tree methods, tests for gene flow modified for target-enrichment data, and morphometric analysis to reconstruct phylogenetic hypotheses, evaluate current taxon circumscriptions, and examine character evolution in Oenothera sect. Calylophus. Because sect. Calylophus comprises a clade with a relatively restricted geographic range, we were able to extensively sample across the range of geographic, edaphic, and morphological diversity in the group. We found that the combination of exons and flanking noncoding regions led to improved support for species relationships. We reconstructed potential hybrid origins of some accessions and note that if processes such as hybridization are not taken into account, the number of inferred evolutionary transitions may be artificially inflated. We recovered strong evidence for multiple evolutionary origins of bee pollination from ancestral hawkmoth pollination, edaphic specialization on gypsum, and permanent translocation heterozygosity. This study applies newly emerging techniques alongside dense infraspecific sampling and morphological analyses to effectively reconstruct the recalcitrant history of a rapid radiation. [Gypsum endemism; Oenothera sect. Calylophus; Onagraceae; phylogenomics; pollinator shift; recent radiation; target enrichment.].

Seed Functional Traits Provide Support for Ecological Restoration and ex situ Conservation in the Threatened Amazon Ironstone Outcrop Flora.

Cangas (ironstone outcrops) host a specialized flora, characterized by high degree of edaphic endemism and an apparent lack of natural history knowledge of its flora. Due to intense pressure from iron ore mining this ecosystem is under threat and in need of restoration. We studied seed functional traits that are relevant for restoration, translocation and ex situ conservation in 48 species from cangas in eastern Amazon. Were determined the thermal niche breadth, classified seed dormancy and determined methods to overcome it, determined the effect of seed storage on germination, tested the association between germination traits and functional groups, and tested whether seed traits are phylogenetically conserved. We found a broad interspecific variation in most seed traits, except for seed water content. Large interspecific variation in the temperature niche breadth was found among the studied species, but only four species, showed optimum germination at high temperatures of 35–40°C, despite high temperatures under natural conditions. Only 35% of the studied species produced dormant seeds. Mechanical scarification was effective in overcoming physical dormancy and application of gibberellic acid was effective in overcoming physiological dormancy in five species. For the 29 species that seeds were stored for 24 months, 76% showed decreases in the germination percentage. The weak association between germination traits and life-history traits indicate that no particular plant functional type requires specific methods for seed-based translocations. Exceptions were the lianas which showed relatively larger seeds compared to the other growth-forms. Dormancy was the only trait strongly related to phylogeny, suggesting that phylogenetic relatedness may not be a good predictor of regeneration from seeds in cangas. Our study provides support to better manage seed sourcing, use, storage and enhancement techniques with expected reduced costs and increased seedling establishment success.

Iron islands in the Amazon: investigating plant beta diversity of canga outcrops.

The world’s largest mineral iron province, Serra dos Carajás, is home to an open vegetation known as canga, found on top of isolated outcrops rising out of the Amazon rainforest. Over one thousand vascular plants species have been recorded in these canga sites, including 38 edaphic endemics. A new survey adds to our investigation of biogeographic relationships between sixteen canga outcrops and the effect of the distance between site pairs on the number of shared species, regional species turnover and species distribution patterns. Plant collecting expeditions to the westernmost site, the Serra de Campos of São Félix do Xingu (SFX), were carried out followed by the identification of all collected specimens and the creation of a species database, built to perform biogeographical analyses. Floristic relationships among the sites were investigated regarding their similarity, using multivariate analyses. The correlation between canga areas and species richness was tested, as well as the geographical distance between pairs of outcrops and their shared species. Vascular plants at SFX total 254 species including 17 edaphic endemics. All canga sites are grouped with 25% of minimum similarity, and the SFX falls within a large subgroup of outcrops. The total species number shared between site pairs does not change significantly with geographical distance but is positively correlated with the area of each outcrop. Meanwhile, shared endemic species numbers between site pairs decline when geographical distance increases, possibly imposed by the barrier of the rainforest. Our data suggest higher shared similarity between the largest and species-richest sites as opposed to geographically nearby sites, and provide useful insight for drafting conservation and compensation measures for canga locations. The size of the canga outcrops is associated to higher floristic diversity but connectivity among islands also plays a role in their similarity.

Gypsum-exclusive plants accumulate more leaf S than non-exclusive species both in and off gypsum

Gypsum-exclusive species (gypsophiles), are restricted to gypseous soils in natural environments. However, it is unclear why gypsophiles display greater affinity to gyspeous soils than other soils. These plants are edaphic endemics, growing in alkaline soils with high Ca and S. Gypsophiles tend to show higher foliar Ca and S, lower K and, sometimes, higher Mg than non-exclusive gypsum species, named gypsovags. Our aim was to test if the unique leaf elemental signature of gypsophiles could be the result of special nutritional requirements linked to their specificity to gypseous soils. These nutritional requirements could hamper the completion of their life cycle and growth in other soil types. To test this hypothesis, we cultivated five gypsophiles and five gypsovags dominant in Spanish gypsum outcrops on gypseous and calcareous (non-gypseous) field soil for 29 months. We regularly measured growth and phenology, and differences in leaf traits, final biomass, individual seed mass, seed viability, photosynthetic assimilation and leaf elemental composition. We found all the gypsophiles studied were able to complete their life cycle in non-gypseous soil, producing viable seeds, attaining greater biomass and displaying higher photosynthetic assimilation rates than in gypseous soil. The leaf elemental composition of some species (both gypsophiles and gypsovags) shifted depending on soil, although none of them showed leaf deficiency symptoms. Regardless of soil type, gypsophiles had higher leaf S, Mg, Fe, Al, Na, Mn, Cr and lower K than gypsovags. Consequently, gypsophiles have a unique leaf chemical signature compared to gypsovags of the same family, particularly due to their high leaf S regardless of soil conditions. However, these nutrient requirements are not sufficient to explain why gypsophiles are restricted to gypsum soil in natural conditions.

Edaphic Endemism in the Amazon: Vascular Plants of the canga of Caraj\xe1s, Brazil

Amazonia is one of the most diverse biomes worldwide, and, as well as luxuriant forest, it includes mountain areas which, despite their small surface area, display fascinating endemism. In these regions, the specificity of edaphic factors is mirrored by a highly specialised, isolated flora adapted to survive adverse conditions. The Serra dos Carajás in the Brazilian state of Pará is one of world’s largest iron ore reserves. Known locally as canga, this ironstone formation occupies an area of 115.9 km2, and supports campo rupestre of canga vegetation on outcrops that are mostly in the Floresta Nacional de Carajás (FLONA of Carajás) and Parque Nacional dos Campos Ferruginosos (PNCF). The recent publication of the Flora of the cangas of Carajás lists 856 species of seed plants and 186 species of ferns and lycophytes. This project assessed the canga endemic species growing in the region, and further expeditions guided by SDM were carried out in order to ascertain their distribution outisde the area. Departing from an initial list of 58 putative endemics, the final list comprises 38 species of vascular plants (c. 4% of the local flora). These are distributed in 31 genera and 22 families, including three monotypic genera: Carajasia (Rubiaceae), Monogereion and Parapiqueria (Asteraceae). From these, 24 are classified as Rare Species for Brazil and seven as Highly Restricted Endemic (EEO < 100 km2). An illustrated account is provided, as well as further SDM to detect other possible areas of distribution based on the studied species. The knowledge generated is aimed at directing appropriate conservation plans for the area.

Speciation and subsequent secondary contact in two edaphic endemic primroses driven by Pleistocene climatic oscillation.

Climatic perturbation during the Pleistocene era has played a major role in plant evolutionary history by altering species distribution range. However, the relative roles of climatic and geographic factors in the distribution dynamics remain poorly understood; in particular, the edaphic endemics. In this paper, we examine the evolutionary history of two ultramafic primroses, Primula hidakana and Primula takedana. These species are ecologically and morphologically distinct with disjunct distributions on Hokkaido Island, Japan. Primula hidakana is found on various rocks in southern Hokkaido and P. takedana in serpentine areas in northern Hokkaido. We performed population genetics analyses on nuclear and chloroplast data sets and tested alternative phylogenetic models of divergence using approximate Bayesian computation (ABC) analyses. Nuclear microsatellite loci clearly distinguished the two sister taxa. In contrast, chloroplast sequence variations were shared between P. takedana and P. hidakana. ABC analyses based on nuclear data supported a secondary contact scenario involving asymmetrical gene flow from P. hidakana to P. takedana. Paleodistribution modeling also supported the divergence model, and predicted their latitudinal range shifts leading to past secondary contact. Our findings highlight the importance of the distribution dynamics during the Pleistocene climatic oscillations in the evolution of serpentine plants, and demonstrate that tight species cohesion between serpentine and nonserpentine sister taxa has been maintained despite past interspecific gene flow across soil boundaries.

Adaptation and divergence in edaphic specialists and generalists: serpentine soil endemics in the California flora occur in barer serpentine habitats with lower soil calcium levels than serpentine tolerators.

Adaptation to harsh edaphic substrates has repeatedly led to the evolution of edaphic specialists and generalists. Yet, it is unclear what factors promote specialization versus generalization. Here, we search for habitat use patterns associated with serpentine endemics (specialists) and serpentine tolerators (generalists) to indirectly test the hypothesis that trade-offs associated with serpentine adaptation promote specialization. We predict that (1) endemics have adapted to chemically harsher and more bare serpentine habitats than tolerators, and (2) edaphic endemics show more habitat divergence from their sister species than tolerators do among on- and off-serpentine populations. We selected 8 serpentine endemic and 9 serpentine tolerator species representing independent adaptation to serpentine. We characterized soil chemistry and microhabitat bareness from one serpentine taxon of each species and from a paired nonserpentine sister taxon, resulting in 8 endemic and 9 tolerator sister-taxa pairs. We find endemic serpentine taxa occur in serpentine habitats averaging twice as much bare ground as tolerator serpentine taxa and 25% less soil calcium, a limiting macronutrient in serpentine soils. We do not find strong evidence that habitat divergence between sister taxa of endemic pairs is greater than between sister taxa of tolerator pairs. These results suggest serpentine endemism is associated with adaptation to chemically harsher and more bare serpentine habitats. It may be that this adaptation trades off with competitive ability, which would support the longstanding, but rarely tested, competitive trade-off hypothesis.

Further Evidence of Edaphic Endemism of Lichens on Fox Hills Sandstone: Discovery and Description of Lecanora lendemerii as New to Science

Further Evidence of Edaphic Endemism of Lichens on Fox Hills Sandstone: Discovery and Description of Lecanora lendemerii as New to Science

Phylogeographic testing of alternative histories of single-origin versus parallel evolution of early flowering serpentine populations of Picris hieracioides L. (Asteraceae) in Japan

Serpentine, a chemically extreme substrate with a discontinuous distribution, supports a high proportion of endemic plant species. The evolutionary pathway of edaphic endemics in isolated areas may be of parallel origin (convergence), but this can be difficult to detect due to similar sets of phenotypic traits repeatedly selected for by the extreme conditions of serpentine environments. Testing the alternative histories of single origin vs. parallel evolution in serpentine plant populations provides insight into the ability of plants to adapt to extreme serpentine environments and to disperse between isolated serpentine areas. In this study, we focused on the herb Picris hieracioides L. (Asteraceae), which includes early-flowering serpentine populations surrounded by later flowering, non-serpentine populations on Hokkaido Island, Japan. We performed population genetic analysis based on genome-wide SNPs markers and quantified temporal overlap of flowering times under natural conditions. Phylogeographic analysis confirmed the occurrence of two genetic groups, corresponding to the serpentine and non-serpentine races. Although the two races were found to occur in close parapatry over multiple paired soil sites, little evidence for hybridization was detected. The flowering times of the two races showed significant divergence, with flowering peaks separated by more than 1 month. Overall, this study supported the single origin of serpentine Picris populations, and showed the role of divergent flowering time as a strong prezygotic barrier to maintain the edaphic races over short spatial distances.

The mid‐distance dispersal optimum, evidence from a mixed‐model climate vulnerability analysis of an edaphic endemic shrub

AbstractAimPredicting accurate species responses to climate change in fragmented landscapes is challenging in conservation biogeography. We assessed the role of dispersal, including long‐distance dispersal (LDD), for the long‐term persistence of a rare plant species (Vella pseudocytisus subsp. paui) under current climate conditions and four future climate scenarios; and analysed the effect of competition for its regional survival.LocationEster Iberian system, Aragón, Spain.MethodsWe used BioMove, a linked modelling platform that integrates demographic, dispersal and competition features with biogeographic predictions of range dynamics, and whose results can inform risk assessments and conservation planning. We linked Vella's population dynamics and habitat suitability models with its well‐documented life history traits and ecological characteristics to enhance our understanding of its inherent vulnerability to dispersal, competitive interactions and climate change.ResultsWe found thresholds in the effect of long dispersal distances on population persistence, suggesting a mid‐distance optimum, which reduces mortality risk associated with the increasing proportion of LDD seeds. Moreover, increasing the proportion of LDD propagules reduces Vella's ability to compete in currently occupied and nearby unoccupied habitats by reducing the number of local recruitments.Main conclusionsWhereas LDD ability is generally assumed to be beneficial for the long‐term persistence of plant species in fragmented landscapes, our results suggest that moderate to high distances between new colonization increase the species’ dispersal‐related mortality. Higher numbers of LDD events reduce the number of local propagules in established populations, reducing regional survival for Vella. Moreover, associated range shifts of potentially better climate‐adapted competitors could prevent Vella individuals from colonizing new climatically suitable habitats. These findings can support development of more efficient conservation management strategies for establishing new population patches to mitigate population losses due to climate change. Our findings may also apply as a framework for other narrowly distributed endemic shrub species, particularly edaphic endemics.

Habitat differentiation of obligate ultramafic Nepenthes endemic to Mount Kinabalu and Mount Tambuyukon (Sabah, Malaysia)

Localized ultramafic outcrops in Kinabalu Park (encompassing Mount Kinabalu and Mount Tambuyukon) in Sabah (Malaysia) are known for high levels of plant diversity and endemism, which have been stimulated by extreme soil chemistry and biogeographic factors, such as isolation. Characteristic of these outcrops are edaphic endemics of insectivorous pitcher plants from the Nepenthaceae-family including Nepenthes burbidgeae, N. edwardsiana, N. macrovulgaris, N. rajah and N. villosa, all of which (except N. macrovulgaris) are confined solely to the protected area of Kinabalu Park. Although the various aspects of plant taxonomy and nutrition of this genus have been extensively studied, the habitat of these rare species has to date not been studied in detail. It was hypothesized that while the Nepenthes of Mount Kinabalu and Mount Tambuyukon are excluders of excess trace elements, soil chemistry is nevertheless a major driver for creating the specific habitats in which ultramafic edaphic endemic Nepenthes occur, and that this is reflected in vegetation physiognomy and co-occurring species composition as well as in the foliar chemistry of Nepenthes. The results show that the characteristics of the physical environment and soil chemistry ‘induces’ a vegetation physiognomy that varies among sites and produces a series of different niches that are colonised by different Nepenthes-species. Although other plant species associated with these niches do not directly influence the distributions of the Nepenthes-species, some are characteristic. The unusual ultramafic soil chemistry is not reflected in the foliar chemistry of Nepenthes and they appear to be ‘Excluders’ of potentially toxic soil trace element concentrations. The populations of Nepenthes burbidgeae, N. edwardsiana, N, rajah, N. villosa are small (in particular, N. burbidgeae) and these species are thus likely to be vulnerable to the potential effects of climate change-induced drought and fire.

The evolution of halophytes, glycophytes and crops, and its implications for food security under saline conditions.

The effective development of salt tolerant crops requires an understanding that the evolution of halophytes, glycophytes and our major grain crops has involved significantly different processes. Halophytes (and other edaphic endemics) generally arose through colonization of habitats in severe disequilibrium by pre-adapted individuals, rather than by gradual adaptation from populations of 'glycophytes'. Glycophytes, by contrast, occur in low sodium ecosystems, where sodium was and is the major limiting nutrient in herbivore diets, suggesting that their evolution reflects the fact that low sodium individuals experienced lower herbivory and had higher fitness. For domestication/evolution of crop plants, the selective pressure was human imposed and involved humans co-opting functions of defense and reproductive security. Unintended consequences of this included loss of tolerance to various stresses and loss of the genetic variability needed to correct that. Understanding, combining and manipulating all three modes of evolution are now critical to the development of salt tolerant crops, particularly those that will offer food security in countries with few economic resources and limited infrastructure. Such efforts will require exploiting the genetic structures of recently evolved halophytes, the genetic variability of model plants, and endemic halophytes and 'minor' crops that already exist.

Multiple origins of serpentine-soil endemism explained by preexisting tolerance of open habitats

Plant specialization on soils derived from unusual parent materials is an important contributor to regional biodiversity. These stressful substrates include serpentine, gabbro, and other ultramafic rocks rich in heavy metals. The effect of substrate on plant diversity is illustrated by serpentine soils in California: they comprise less than 1% of the surface of the state (1), but serpentine endemics (species restricted to serpentine soils) make up about 10% of the flora (2). How such “edaphic endemics” (plants restricted to stressful soils) evolve is a long-standing question that remains largely unresolved. For example, plant tolerance of serpentine soils may often involve tradeoffs in competitive ability, and restriction to serpentine soil may reflect poor competitive ability on less stressful soils rather than obligate association (3⇓–5), although possible counter-examples exist (6). A common feature of plant communities on stressful soils is the wide spacing of plants and openness of the habitat. Openness may itself be stressful for a variety of reasons (detailed below) (7⇓–9). In PNAS, Cacho and Strauss (10) use a novel comparative experimental approach to explicitly assess the role of openness vs. soil chemistry as factors in the evolution of plant tolerance of, and endemism to, serpentine soils.

Conservation genetics of edaphic endemics in naturally isolated habitats: a case study withGeocarpon minimum(Caryophyllaceae)

Theory predicts that edaphic endemics should exhibit high levels of population differentiation due to restricted gene flow among patchily distributed habitats. Here, we tested this prediction with the federally threatened annual Geocarpon minimum, an edaphic endemic restricted to sandstone outcrops and slick spots associated with saline prairies in the Southeastern United States. We used AFLP data to quantify population genetic variation and structure in 13 G. minimum populations using a bulked sampling scheme. Modest but significant components of genetic variance are partitioned among populations (16%) and among regions (north, south; 3%). The maintenance of private bands within most G. minimum populations suggests limited gene flow among geographically isolated populations. However, a Mantel test failed to detect a pattern of isolation by distance. Estimates of within population variation in G. minimum are relatively low compared to other edaphic endemics with similar life-histories. Results suggest conservation efforts should focus on protecting genetically unique populations and ensuring that genetic variability is adequately captured in ex situ collections.