Nitrogen deposition suppresses ephemeral post-fire plant diversity.

Fire is a dominant force shaping patterns of plant diversity in Mediterranean-type ecosystems. In these biodiversity hotspots, including California's endangered coastal scrub, many species remain hidden belowground as seeds and bulbs, only to emerge and flower when sufficient rainfall occurs after wildfire. The unique adaptations possessed by these species enable survival during prolonged periods of unfavorable conditions, but their continued persistence could be threatened by nonnative plant invasion and environmental change. Furthermore, their fleeting presence aboveground makes evaluating these threats in situ a challenge. For example, nitrogen (N) deposition resulting from air pollution is a well-recognized threat to plant diversity worldwide but impacts on fire-following species are not well understood. We experimentally evaluated the impact of N deposition on post-fire vegetation cover and richness for three years in stands of coastal sage scrub that had recently burned in a large wildfire in southern California. We installed plots receiving four levels of N addition that corresponded to the range of N deposition rates in the region. We assessed the impact of pre-fire invasion status on vegetation dynamics by including plots in areas that had previously been invaded by nonnative grasses, as well as adjacent uninvaded areas. We found that N addition reduced native forb cover in the second year post-fire while increasing the abundance of nonnative forbs. As is typical in fire-prone ecosystems, species richness declined over the three years of the study. However, N addition hastened this process, and native forb richness was severely reduced under high N availability, especially in previously invaded shrublands. An indicator species analysis also revealed that six functionally and taxonomically diverse forb species were especially sensitive to N addition. Our results highlight a new potential mechanism for the depletion of native species through the suppression of ephemeral post-fire bloom events.

A relic of the past: current genetic patterns of the palaeoendemic tree Nothofagus macrocarpa were shaped by climatic oscillations in central Chile.

BACKGROUND AND AIMS: The Mediterranean-type forest of central Chile is considered a 'biodiversity hotspot' and a relic of a wider ancient distribution produced by past climatic oscillations. Nothofagus macrocarpa, commonly known as 'roble de Santiago', is a threatened palaeoendemic of this forest, poorly represented in the protected area system. This tree has been repeatedly misidentified as the sister species N. obliqua, which has affected its recognition and protection. Only a few populations of N. macrocarpa remain within a matrix of intensive land use that has been affected by recent forest fires. We tested the hypothesis that current populations of N. macrocarpa are a relic state of a previously widespread range, with the aim of contributing to its identification, its biogeographical history and the design of conservation measures using genetic information. METHODS: We analysed remnant N. macrocarpa forests using nuclear (nDNA) and chloroplast DNA (cpDNA) sequences, conducted phylogenetic and phylogeographical analyses to reconstruct its biogeographical history, and assessed microsatellites [simple sequence repeats (SSRs)] to determine contemporary patters of diversity within and among all remnant populations. We also examined the degree of past, current and potential future isolation of N. macrocarpa populations using ecological niche models (ENMs). KEY RESULTS: The species N. macrocarpa was confirmed by nDNA sequences, as previously suggested by chromosomal analysis. Small isolated populations of N. macrocarpa exhibited moderate to high genetic diversity according to SSRs. cpDNA analysis revealed a marked past latitudinal geographical structure, whereas analysis of SSRs did not find such current structure. ENM analyses revealed local expansion-contraction of the N. macrocarpa range during warmer periods, particularly in the northern and central ranges where basal-most cpDNA haplotypes were detected, and recent expansion to the south of the distribution. CONCLUSIONS: Genetic patterns confirm that N. macrocarpa is a distinct species and suggest a marked latitudinal relic structure in at least two evolutionarily significant units, despite contemporary among-population gene flow. This information must be considered when choosing individuals (seeds and/or propagules) for restoration purposes, to avoid the admixture of divergent genetic stocks.

Invasive annuals respond more negatively to drought than native species.

In his foundational list of 'ideal weed' characteristics, Baker (1965) proposed that weedy plants maximize reproductive output under high resource availability. Since then, the idea that invasive plant species are more responsive to fluctuating resources compared with native or noninvasive species has gained considerable traction, although few studies extend this hypothesis to include reproductive output. We revisit Baker's hypothesis in the context of invasion and drought in California grasslands, exploring whether invasives show greater growth and reproductive responses to water availability compared with the native wildflowers they displace. In an outdoor potted study, we grew eight native and eight invasive species of annuals commonly found in southern California grasslands to reproductive maturity under both well-watered and drought conditions. While drought negatively impacted plant performance overall, invasives showed more negative responses for growth and reproductive traits. Invasives also grew larger than native species, especially under well-watered conditions, and produced seed with higher rates of germination. Invasives may be more negatively impacted by drought compared with natives, but they are also able to capitalize on high resource conditions and greatly increase reproductive output. Such opportunistic responses exhibited by invasives might explain previously observed fluctuations in their abundance under variable precipitation.

Can we condition native plants to increase drought tolerance and improve restoration success?

A common method in ecological restoration is the transplanting of nursery-grown seedlings to the field and, with proper resources, this technique can be highly successful. However, stressors such as drought may negatively impact plant performance and restoration success, especially in dryland ecosystems. Furthermore, increasing environmental change may hamper the ability of practitioners to restore native vegetation. A growing body of research suggests that exposing plants to a stressor may improve tolerance to subsequent stress events later in life. We sought to understand if such a phenomenon could be exploited in order to improve plant drought-tolerance and aid native plant restoration in southern California. In a multi-phase experiment, we first exposed seedlings of native perennials to episodic drought and then later compared the response of these plants to a second drought event to that of well-watered controls. We also transplanted replicates of both treatments to a restoration site in the field to test whether exposure to drought as a seedling could improve plant performance. Plant species responded to drought differently, with species exhibiting the full range of positive, neutral, and negative responses to temporal variability in water stress. However, some species appeared to benefit from drought preconditioning, exhibiting greater growth and increased water-use efficiency compared to well-watered plants. This suggests that simple applications of stress treatments could improve plant growth and stress tolerance, but the success of this method is likely very species specific. Restoration practitioners should consider conducting pilot studies with target plant species to better understand if this technique could assist in achieving restoration goals.

Precipitation and topography modulate vegetation greenness in the mountains of Baja California, México.

Vegetation greenness (normalized difference vegetation index, NDVI) showed significant temporal and spatial correlations with precipitation and topography-derived features within the context of slope aspect (South- (SFS) and North-facing slopes (NFS) and an intermountain valley (IMV)) in a semi-arid Mediterranean-climate watershed in northwestern Baja California, México. Rank correlation with annual precipitation (1986-2016) showed a strong positive relationship with wet season NDVI at SFS (Rs = 0.82), IMV (Rs = 0.79), and NFS (Rs = 0.65) but moderate relation and only on hillslopes in the dry season (SFS, Rs = 0.47; NFS, Rs = 0.39). Thus, the vegetation on the more xeric SFS sites was more responsive to intra-annual and inter-annual precipitation than on either IMV or NFS. The correlation of NDVI with six topography-derived environmental attributes (elevation, slope gradient, curvature, drainage density, topographic wetness index, solar radiation) was weak to moderate, varied in degree and significance between years with exceptionally high or low NDVI, and often differed in sign between SFS, NFS, and IMV. Results showed that precipitation controlled vegetation greenness, under the three aspect conditions, more closely than did the other topography-derived features, and the sparse deciduous vegetation of SFS showed stronger associations with precipitation than IMV or NFS. The measurement of these relationships should be continued and complemented by other studies to improve the overall model, because they are important to modeling ecohydrology and productivity, and may be of use for projecting and hindcasting vegetation dynamics.

Assessing the Importance of Native Mycorrhizal Fungi to Improve Tree Establishment after Wildfires.

The Chilean matorral is a heavily threatened Mediterranean-type ecosystem due to human-related activities such as anthropogenic fires. Mycorrhizal fungi may be the key microorganisms to help plants cope with environmental stress and improve the restoration of degraded ecosystems. However, the application of mycorrhizal fungi in the restoration of the Chilean matorral is limited because of insufficient local information. Consequently, we assessed the effect of mycorrhizal inoculation on the survival and photosynthesis at set intervals for two years after a fire event in four native woody plant species, namely: Peumus boldus, Quillaja saponaria, Cryptocarya alba, and Kageneckia oblonga, all dominant species of the matorral. Additionally, we assessed the enzymatic activity of three enzymes and macronutrient in the soil in mycorrhizal and non-mycorrhizal plants. The results showed that mycorrhizal inoculation increased survival in all studied species after a fire and increased photosynthesis in all, but not in P. boldus. Additionally, the soil associated with mycorrhizal plants had higher enzymatic activity and macronutrient levels in all species except in Q. saponaria, in which there was no significant mycorrhization effect. The results suggest that mycorrhizal fungi could increase the fitness of plants used in restoration initiatives after severe disturbances such as fires and, consequently, should be considered for restoration programs of native species in threatened Mediterranean ecosystems.

Shoot and root biomass production in semi-arid shrublands exposed to long-term experimental N input.

Anthropogenic nitrogen (N) deposition has affected the primary production of terrestrial ecosystems worldwide; however, ecosystem responses often vary over time because of transient responses, interactions between N, precipitation, and/or other nutrients, and changes in plant species composition. Here we report N-induced changes in above- and below-ground standing crop and production over an 11-year period for two semi-arid shrublands, chaparral and coastal sage scrub (CSS), of Southern California. Shrubs were exposed to 50 kgN ha-1 in the fall of each year to simulate the accumulation of dry N deposition, and shoot and root biomass and leaf area index (LAI) were measured every 3 months to assess how biomass production responded to chronic, dry N inputs. N inputs significantly altered above- and below-ground standing crop, production, and LAI; however, N impacts varied over time. For chaparral, N inputs initially increased root production but suppressed shoot production; however, over time biomass partitioning reversed and plants exposed to N had significantly more shoot biomass. In CSS, N inputs caused aboveground production to increase only during wet years, and this interaction between added N and precipitation was due in part to a highly flexible growth response of CSS shrubs to increases in N and water availability and to a shift from slower-growing native shrubs to fast-growing introduced annuals. Together, these results indicate that long-term N inputs will lead to complex, spatially and temporally variable growth responses for these, and similar, Mediterranean-type shrublands.

Field comparison of ecophysiological traits between an invader and a native species in a Mediterranean coastal dune.

Photochemical efficiency, gas exchange, leaf water potential, pigment content and free proline content of Oenothera drummondii subsp. drummondii Hook (an invasive non-native species) and Achillea maritima (L.) Ehrend. & Y.P. Guo, (an ecologically similar native species) were explored to understand the success of invasive non-native species in Mediterranean coastal dunes of southwest Spain. We have conducted a field study during a complete annual cycle, comparing both species. Fifteen pairs of neighbouring plants of the two study species of similar size were selected and measured seasonally. The results show that in spring and summer, assimilation rates of O. drummondii were significantly higher than those of the native, A. maritime, even though the native species had higher photochemical efficiency. Additionally, the non-native species presented better water content regulation than the native one, surely related to better water use efficiency and maybe linked to greater root development. The differences in leaf dry matter content values for both species might indicate a different strategy of resource use; with A. maritima displaying a more conservative strategy and O. drummondii presenting a rapid resources acquisition and use strategy as predictors of rapid growth and soil fertility. We conclude that O. drummondii utilizes light, water and probably nutrients more efficiently than the native A. maritima and suffers lower stress in Mediterranean coastal dunes where water availability is reduced (44 mm from May to October in the study area) and light radiation levels are high.

Environmental niche conservatism explains the accumulation of species richness in Mediterranean-hotspot plant genera.

The causes of exceptionally high plant diversity in Mediterranean-climate biodiversity hotspots are not fully understood. We asked whether a mechanism similar to the tropical niche conservatism hypothesis could explain the diversity of four large genera (Protea, Moraea, Banksia, and Hakea) with distributions within and adjacent to the Greater Cape Floristic Region (South Africa) or the Southwest Floristic Region (Australia). Using phylogenetic and spatial data we estimated the environmental niche of each species, and reconstructed the mode and dynamics of niche evolution, and the geographic history, of each genus. For three genera, there were strong positive relationships between the diversity of clades within a region and their inferred length of occupation of that region. Within genera, there was evidence for strong evolutionary constraint on niche axes associated with climatic seasonality and aridity, with different niche optima for hotspot and nonhotspot clades. Evolutionary transitions away from hotspots were associated with increases in niche breadth and elevated rates of niche evolution. Our results point to a process of "hotspot niche conservatism" whereby the accumulation of plant diversity in Mediterranean-type ecosystems results from longer time for speciation, with dispersal away from hotspots limited by narrow and phylogenetically conserved environmental niches.

Plant stored reserves do not drive resprouting of the lignotuberous shrub Erica australis.

• Lignotuberous plants store carbohydrates and mineral nutrients within the lignotuber. Resprouting vigour may depend on stored reserves, as well as on the availability of soil mineral nutrients and water. • Here the role played by plant reserves and soil resources on the resprouting response of Erica australis was analysed after clipping plants in 13 different stands, varying in soil resource availability and in plant reserves. • There were significant among-site differences for resprout biomass and maximum length, but not for resprout number, 1 yr after clipping. Plant reserves at the time of clipping were not significantly correlated with resprout number, length or biomass. However, resprouting variables were significantly correlated with soil nitrogen or extractable cations, or plant water potentials. Resprout biomass and maximum length were negatively correlated with lignotuber size. • These findings indicate that the assumption that resprouting vigor in lignotuberous plants is primarily dependent on the amount of reserves stored in the lignotuber must be revised, as well as the overall role of lignotubers in resprouting.