Get the shovel: morphological and evolutionary complexities of belowground organs in geophytes

Herbaceous plants collectively known as geophytes, which regrow from belowground buds, are distributed around the globe and throughout the land plant tree of life. The geophytic habit is an evolutionarily and ecologically important growth form in plants, permitting novel life history strategies, enabling the occupation of more seasonal climates, mediating interactions between plants and their water and nutrient resources, and influencing macroevolutionary patterns by enabling differential diversification and adaptation. These taxa are excellent study systems for understanding how convergence on a similar growth habit (i.e., geophytism) can occur via different morphological and developmental mechanisms. Despite the importance of belowground organs for characterizing whole-plant morphological diversity, the morphology and evolution of these organs have been vastly understudied with most research focusing on only a few crop systems. Here, we clarify the terminology commonly used (and sometimes misused) to describe geophytes and their underground organs and highlight key evolutionary patterns of the belowground morphology of geophytic plants. Additionally, we advocate for increasing resources for geophyte research and implementing standardized ontological definitions of geophytic organs to improve our understanding of the factors controlling, promoting, and maintaining geophyte diversity.     

Effects of grazing on geophytes in Mediterranean vegetation

Abstract. The response of geophyte species diversity and frequency of individual geophyte species to cattle grazing was measured at 68 site pairs along fences separating ungrazed from grazed grassland and woodland on different geological formations in northern Israel. Over all site pairs, geophyte species density per 4 m² was significantly greater in grazed (2.37) than in protected (1.96) sites of the same site pair. There was considerable variation between site pairs in the magnitude and in the direction of the grazing effect. Part of this variation could be explained by differences in site altitude and in geological formation. The positive effect of grazing on geophyte diversity was lower in sites with low productivity. Of 22 geophyte taxa for which sufficient data were available, nine indicated greater frequency in grazed sites compared to only two in ungrazed sites. In 11 other taxa the response was not consistent. A positive response to grazing was most common in geophytes with narrow leaves of the Iridaceae, Liliaceae and allied families. Conservation of the entire geophyte flora in Mediterranean vegetation requires livestock grazing at moderate to high intensities in parts of the area of each community, and light or no grazing in other parts.     

Plant–Plant and Plant–Topography Interactions on a Rock Outcrop at High Altitude in Southeastern Brazil1

Plant establishment and growth on rocky outcrops in the Itatiaia massif (2400 m a.s.l.), southeastern Brazil, are limited by lack of soil and by freezing temperatures in winter nights. Mat‐forming species of different sizes and shapes on bare rock provide substrate for other plants to establish. The habitat preference of two geophytes, Stevia camporum (Asteraceae) and Alstroemeria foliosa (Alstroemeriaceae), was compared with regard to their association with the type of mat species and distinct rock topographies. The habitat preference of the mat species in regard to topography was also assessed. We found 1706 ramets of S. camporum and 1317 of A. foliosa in 253 vegetation islands ranging in size from 0.005 to 18.097 m². Mat species in these islands were Fernseea itatiaiae, Vriesea itatiaiae (both Bromeliaceae), Pleurostima gounelleana (Velloziaceae) and/or Campylopus pilifer (Dicranaceae) and other mosses. Mat species were segregated by topography. Geophytes were similarly distributed across types of topography but showed negative interspecific association. S. camporum occurred mostly on the moss‐dominated islands, whereas A. foliosa was more common in P. gounelleana islands. We found geophyte co‐occurrence in larger vegetation islands containing both P. gounelleana and mosses as mat species. Thus, the effect of topography on geophyte segregation was indirect, since topography affects mat species distribution, and the geophyte preference for distinct mat species as substrate resulted in their segregation.     

Floristic diversity of an eastern Mediterranean dwarf shrubland: the importance of soil pH

Questions: Does plant species richness and composition of eastern Mediterranean dwarf shrubland (phrygana) correlate with soil pH? How important is the effect of pH on species diversity in relation to other environmental factors in this ecosystem? What is the evolutionary background of the diversity–pH relationship? Location: Western Crete, Greece. Methods: Species composition of vascular plants, soil and other environmental variables were sampled in 100‐m² plots on acidic and basic bedrock in phrygana vegetation. The relationships between species composition and environmental variables (including climate) were tested using canonical correspondence analysis, and relationships between species richness and environment using correlation and regression analyses. Data were analysed separately for different plant functional types based on life form and life span. Results: Although soil pH varied across a narrow range (5.9‐8.1), species composition changed significantly along the pH gradient within all plant functional types. For most functional types, the effect of soil pH on species composition was stronger than that of other environmental variables. Species richness of annuals, geophytes and suffruticose chamaephytes increased with soil pH, while richness of hemicryptophytes and shrubs was not correlated with pH. Conclusions: The results are consistent with the evolutionary species pool hypothesis. High numbers of calcicole annuals, geophytes and suffruticose chamaephytes may be a result of the evolution of these groups on base‐rich dry soils in the Mediterranean climate. In contrast, hemicryptophytes, a life form typical of the temperate zone, evolved on both acidic and basic soils and therefore their species numbers do not respond to soil pH across the narrow range studied. The lack of a relationship between shrub species richness and pH is difficult to explain: it may reflect the more diverse or older origin of Mediterranean woody species and their conservative niches.     

Geophytes–herbivore interactions: reproduction and population dynamics of <Emphasis Type="Italic">Anemone coronaria</Emphasis> L.

Anemone coronaria, an attractive Mediterranean geophyte, seems to disappear from grazing-protected areas in Israel. We experimentally examined the ecological mechanism driving the decline of this geophyte. Ten plot-pairs were established, half we fenced as grazing exclosures and half were grazed by beef cattle. Grazing clearly reduced herbaceous biomass, increased relative solar photosynthetic active radiation (PAR) at ground level, but had almost no effect on soil properties. Grazing did not affect the number of flowers and young fruits produced by A. coronaria, nor the percentage fruit-set at the plot scale, indicating no effect on flowering, pollination, or on resource allocation to reproduction. Five years after grazing exclusion, Anemone seedling and adult plant densities were higher in grazed than in ungrazed plots. We propose a model explaining our results that can be applied also to other similar ecosystems: excluding grazing increased biomass and height of the herbaceous community and reduced relative PAR at ground level. Consequently, seedling, adult plant and flowering Anemone plant densities were lower in ungrazed plots. We recommend adding seasonal grazing as a management tool when vegetation outcompete light demanding geophytes that we wish to conserve.     

A study on nitrate reductase activity (NRA) of geophytes from Mediterranean environment

Nitrate reductase activity (NRA) in different compartments of 14 Mediterranean geophytes (bulbous, tuberous and rhizomatous) and actual mineral nitrogen (NO₃⁻ and NH₄⁺) in their soils were investigated. The nitrate reduction capacities of each species were determined as NRA per total plant material. Differences among compartments for NRA were significant in all species. The highest NRA was found in leaves of tuberous species (Anemone coronaria, Cyclamen coum) and of most bulbous species (Allium flavum, Allium guttatum, Bellevelia sarmatica, Galanthus plicatus, Leucojum aestivum, Ornithogalum nutans, Tulipa sylvestris). Therefore, in this group of species the contribution of the leaves to total plant NRA was the highest. The other bulbous species (Allium scorodoprasum, Crocus chrysanthus, Fritillaria bithynica, Muscari neglectum) and one rhizomatous taxon (Iris suaveolens) have a different NRA distribution within the plants. In these species the highest values of NRA were found in different organs. For example, in Allium scorodoprasum the highest NRA was in tunics, and in flowers in M. neglectum. Although leaves are the main compartments reducing nitrate in most of the studied geophytes, other compartments also contribute to total plant nitrate reduction.Our results show that the nitrate reduction capacity is different among geophyte species. Even if it roughly reflects the nitrogen supply in a habitat, differences in nitrate reduction capacities of different species collected from same sites indicate that the nitrate reducing capacity is species-specific.     

Foraging in the subterranean social highveld mole-rat (Cryptomys hottentotus pretoriae): an investigation into mass-dependent geophyte use and foraging patterns

The foraging behaviour of captive colonies of the highveld mole-rat Cryptomys hottentotus pretoriae was investigated in an artificial soil-filled burrow system provided with four trays (patches) that varied in geophyte density and mass. An initial trial involving empty trays (only soil) revealed that there was no preference for any specific tray. There were no statistically significant preferences for excavating in any of the patches of different geophyte density. No preferences were evident for excavation in patches containing geophytes of different mass classes. Empty patches seemed to be preferred over patches containing geophytes when combinations of geophyte density/mass were investigated. The duration of handling and the rate of consumption of geophytes were recorded for 23 individuals of two mass classes. Handling time of geophytes was not related to mole-rat sex, but was strongly linked to mole-rat mass class. Handling time of geophytes was related to geophyte mass class. Small geophytes were less profitable to consume. These findings are considered in light of optimal foraging theory and the situation in the field. It was concluded that the mole-rats generally followed the qualitative predictions of optimal foraging theory, although falling short of being energy maximizers.     

Investment in survival and reproduction along a semelparity–iteroparity gradient in the Beta species complex

The Beta species complex shows a gradient of life histories from pronounced semelparity (big‐bang reproduction) to pronounced iteroparity (repeated reproduction). Models assume a trade‐off between investment in reproduction and survival. Reproductive effort is thought to increase with decreasing life span, and to be invariable in semelparous plants and susceptible to environmental conditions in iteroparous plants. These assumptions and hypotheses were verified by a greenhouse experiment testing six different life cycles at three contrasting nutrient levels. This study suggests that reproductive effort is negatively correlated with mean life span along the life‐cycle gradient. Unlike semelparous beets, reproductive effort in iteroparous beets is extremely sensitive to nutrient level. Phenotypic correlation between allocation to reproduction and allocation to survival generally appeared significantly negative in the longest‐lived iteroparous beets, nonsignificant in intermediate life histories and obviously positive in semelparous beets (no trade‐off control).     

Intraspecific competition and light effect on reproduction of Ligularia virgaurea,an invasive native alpine grassland clonal herb

The relationship between sexual reproduction and clonal growth in clonal plants often shows up at the ramet level. However, only a few studies focus on the relationship at the genet level, which could finally account for evolution. The sexual reproduction and clonal growth of Ligularia virgaurea, a perennial herb widely distributed in the alpine grasslands of the Qinghai‐Tibetan Plateau of China, were studied under different competition intensities and light conditions at the genet level through a potted experiment. The results showed that: (1) sexual reproduction did not depend on density or light, and increasing clonal growth with decreasing density and increasing light intensity indicated that intraspecific competition and light intensity may affect the clonal life history of L. virgaurea; (2) both sexual reproduction and clonal growth show a positive linear relationship with genet size under different densities and light conditions; (3) a threshold size is required for sexual reproduction and no evidence of a threshold size for clonal growth under different densities and light conditions; (4) light level affected the allocation of total biomass to clonal and sexual structures, with less allocation to clonal structures and more allocation to sexual structures in full sunlight than in shade; (5) light determined the onset of sexual reproduction, and the genets in the shade required a smaller threshold size for sexual reproduction to occur than the plants in full sunlight; and (6) no evidence was found of trade‐offs between clonal growth and sexual reproduction under different densities and light conditions at the genet level, and the positive correlation between two reproductive modes indicated that these are two integrated processes. Clonal growth in this species may be viewed as a growth strategy that tends to maximize genet fitness.     

Persistence,multiple demographic strategies and conservation in long‐lived Mediterranean plants

Abstract. Persistence by longevity has been rarely considered as an alternative to regeneration by seeding for plants showing multiple demographic strategies. We propose a conceptual model of multiple demographic strategies for long‐lived plants in stable habitats, shifting from regeneration by seeding to persistence by longevity and/or vegetative reproduction, along gradients of abiotic stress or interspecific competition. Regeneration by seeding would be promoted under low abiotic stress or under low competition, whereas persistence by longevity and/or vegetative reproduction would predominate at high levels of abiotic stress or competition. We test this model with two threatened species of the Mediterranean region, the shrub Juniperus communis, a widely distributed species which maintains relict populations in the Mediterranean mountains thanks to great adult longevity and Pinguicula vallisneriifolia, a palaeo‐endemic herb relying on a perennial habit and vegetative reproduction under drought imposed stress or high competition at late successional phases. As a main consequence, multiple demographic strategies enhance a plant's ability to exploit environmental heterogeneity at different spatial (patches, localities, regions within the species’ distribution area) and temporal (individual life span, glacial‐interglacial cycles) scales. The potential of multiple demographic dynamics based on persistence and regeneration must be considered as a major ecological trait determining the long‐term viability of peripheral populations of relict species as well as the inertia against extinction of many threatened endemisms, thereby contributing to the maintenance of the high plant diversity characterizing the Mediterranean region.     

Does invasion by an alien plant species affect the soil seed bank?

Questions: How does invasion affect old‐field seed bank species richness, composition and density? How consistent are these effects across sites? Does the soil seed bank match vegetation structure in old‐fields? Location: Menorca, Balearic Islands, Spain, western Mediterranean basin. Methods: We monitored seed germination in soils from old‐fields that were both uninvaded and invaded (legacy effect) by the annual geophyte Oxalis pes‐caprae. We also added O. pes‐caprae bulbs to uninvaded soils to test O. pes‐caprae interference with seedling emergence (competitive effect). We compared species composition in the seed bank with that of the vegetation. Results: Species richness in the seed bank and in the vegetation was not significantly different between invaded and uninvaded areas. Uninvaded areas did not have larger seed banks than invaded areas. More seedlings, especially of geophytes, emerged when O. pes‐caprae bulbs were added to the soil. Species similarity between invaded and uninvaded areas was higher in the seed bank (74%) than in the vegetation (49%). Differences in species composition were as important as differences among sites. The degree of species similarity between the seed bank and the vegetation was very low (17%). Conclusions: Despite invasion by O. pes‐caprae not affecting species richness, the variation in the seed bank species composition in invaded and uninvaded areas, and the differences between the seed bank and the mature vegetation, highlights that even if the invader could be eradicated the vegetation could not be restored back to the exact composition as found in uninvaded areas.     

Biogeography of prostrate-leaved geophytes in semi-arid South Africa: hypotheses on functionality

Nowhere is the species diversity of geophytes greater than in the five mediterranean-climate ecosystems of the world. Of these, the Cape mediterranean zone of South Africa is the most speciose. While the relative diversity and importance of geophytes of all of the other four mediterranean regions of the world drops off sharply as one moves into adjacent winter-rainfall desert regions, geophytes in the semi-arid to arid Succulent Karoo (including Namaqualand) remain a very important component of the flora, both in terms of abundance and diversity (comprising 13 to 29% of the regional floras in this region). Apart from species richness, there are also a number of interesting geophyte growth forms in this region. One unusual growth form is geophytes with flattened leaves that lie prostrate on the soil surface. At least eight families (Amaryllidaceae, Colchicaceae, Eriospermaceae, Geraniaceae, Hyacinthaceae, Iridaceae, Orchidaceae and Oxalidaceae) exhibit this growth form. While this growth form is relatively common in many geophyte lineages in the Succulent Karoo biome and the Cape mediterranean zone (Fynbos biome), and occurs infrequently through the summer-rainfall temperate regions of Africa, it is virtually absent in other regions worldwide. A null hypothesis is that the prostrate leaved trait is a neutral characteristic, however biogeographical data do not support this. A neutral trait would be unlikely to show such a clear pattern of distribution. Several alternative hypotheses on the adaptive significance of this growth form are discussed. These include: avoidance of herbivory, reduction in competition from neighbors, creation of a CO₂ enriched environment below the leaves, reduction of water loss around the roots, reduction of water loss through transpiration, precipitation of dew on the leaves and maintenance of optimal leaf temperatures for growth.     

Plasticity in reproduction and growth among 52 range‐wide populations of a Mediterranean conifer: adaptive responses to environmental stress

A plastic response towards enhanced reproduction is expected in stressful environments, but it is assumed to trade off against vegetative growth and efficiency in the use of available resources deployed in reproduction [reproductive efficiency (RE)]. Evidence supporting this expectation is scarce for plants, particularly for long‐lived species. Forest trees such as Mediterranean pines provide ideal models to study the adaptive value of allocation to reproduction vs. vegetative growth given their among‐population differentiation for adaptive traits and their remarkable capacity to cope with dry and low‐fertility environments. We studied 52 range‐wide Pinus halepensis populations planted into two environmentally contrasting sites during their initial reproductive stage. We investigated the effect of site, population and their interaction on vegetative growth, threshold size for female reproduction, reproductive–vegetative size relationships and RE. We quantified correlations among traits and environmental variables to identify allocation trade‐offs and ecotypic trends. Genetic variation for plasticity was high for vegetative growth, whereas it was nonsignificant for reproduction. Size‐corrected reproduction was enhanced in the more stressful site supporting the expectation for adverse conditions to elicit plastic responses in reproductive allometry. However, RE was unrelated with early reproductive investment. Our results followed theoretical predictions and support that phenotypic plasticity for reproduction is adaptive under stressful environments. Considering expectations of increased drought in the Mediterranean, we hypothesize that phenotypic plasticity together with natural selection on reproductive traits will play a relevant role in the future adaptation of forest tree species.     

Demographic response to shading and defoliation in two woodland orchids

Many woodland herbs are long-lived, clonal geophytes that have evolved life histories favoring survival over reproduction. We examined the life history responses of natural populations of two woodland orchid species,Cypripedium calceolus andCephalanthera longifolia to defoliation and heavy shading conducted early in the growing seasons of 2002 and 2003. We asked whether, in view of the importance of growth for the survival of geophytes, treated plants were more likely to exhibit reduced flowering than reduced vegetative growth in the seasons following treatment. We also asked whether plants would suffer reduced ramet performance. Both treatments led to significant declines in flower number per ramet, number of leaves per ramet, and mean ramet height relative to controls inCypripedium. However, inCephalanthera, only shaded plants exhibited significant declines in flower number per ramet, and only defoliated plants exhibited declines in mean ramet height. The number of ramets per plant did not decline relative to controls in either species. Thus, these orchids, especiallyCypripedium, appeared to allocate resources preferentially to vegetative growth functions over sexual reproduction. Per-plant variation in leaf and flower number per ramet, as well as in mean ramet height, consistently declined in response to treatment, significantly so in the case of mean ramet height, suggesting that ramets became more similar within genets. These results suggest both similarities and differences in the ways in whichCephalanthera andCypripedium mobilize resources in response to stress.     

Floristic composition and floral phenology of the Mediterranean batha of Ariel,Samaria

Field observations over a three-year period were used to document the floristic composition, flowering and fruiting sequences of the Sarcopoterium spinosa Mediterranean batha of Ariel. This plant community was found to be species-rich, with high percentages of annuals, bushes and geophytes, low percentages of herbaceous plants, climbers and trees, and ten species considered rare in Israel. Typical Mediterranean characteristics were observed in the S. spinosa batha: spring flowering reaching a peak in April, small between-year variation in flowering and fruiting, most species having Mediterranean distribution, and partial resistance to fire.     

Widespread resistance of Mediterranean island ecosystems to the establishment of three alien species

Although some invasive plants are cosmopolitan, not all ecosystems are invaded to the same degree. Yet there is little experimental work on how ecosystem resistance to invasion at the establishment phase differs among ecosystems. We conducted two field sowing experiments in two consecutive years to examine establishment of the deciduous tree Ailanthus altissima, the succulent subshrub Carpobrotus spp. and the annual geophyte Oxalis pes‐caprae in coastal dunes, shrublands and oldfields in more than 200 sites across six Mediterranean Basin islands differing in climatic conditions and local species richness. Establishment success (i.e. percentage of plots with at least one seedling) and rates (i.e. seedling to sown seed ratio) were low, especially for Ailanthus even when accounting for differences in seed viability. Oxalis was capable of producing a new cohort of seedlings the year following planting. By contrast, all Ailanthus seedlings and half the Carpobrotus seedlings died following the first summer. Differences in establishment success and rates among ecosystems were species‐, island‐ and year‐dependent. Differences in precipitation and mean temperature were associated with differences in establishment rates across sites. Establishment rates tended to be positively correlated with cumulative precipitation and negatively with mean T^a. Unexpectedly, native species richness was not a good predictor of seedling establishment, except for higher Oxalis establishment success in species rich habitats. By conducting field sowing tests at multiple sites across a region we found that except for Oxalis, Mediterranean island ecosystems are quite resistant to invader establishment. These results suggest that differences in the degree of invasion between ecosystems and islands might be more dependent upon the influence of invasion event factors (e.g. propagule pressure) or factors acting at a later life‐history stages rather than differences in the resistance imposed by ecosystems to invader recruitment. Moreover, our results support the notion that in Mediterranean ecosystems invasions are highly idiosyncratic events and strongly dependent on water availability conditions.     

The Demography of Miconia prasina (Melastomataceae) During Secondary Succession in Puerto Rico1

Miconia prasina (Melastomataceae) is an important colonizing species during early stages of secondary succession in Puerto Rico but its abundance declines with increasing stand age. We studied its demography for 2 yr (1995–1997) in three populations across a pasture to forest chronosequence (1‐, 12‐, and 25‐yr old) and in one population in a hurricane‐disturbed older forest (>60‐yr old). Vegetative growth was positive in the two young sites, but negative in the two older sites. The highest percentage of plants that flowered occurred in the two younger sites but highest seed production occurred in the 25‐yr‐old site. Although seed germination occurred in the two older sites, no seedlings established. Vegetative reproduction (root sprouts) was found in all sites, with most sprouts occurring in young sites. The 2 yr of census data were used to construct stage‐based transition matrices. In the two young sites, the average population growth rate was positive (λ > 1), while average population growth was negative in the two older sites (λ < 1). Elasticity, a measure of the effects of proportional changes in life‐history transitions to the long‐term population growth rate, varied across both stages and sites. Elasticity shifted from large plants in young sites to small plants in older sites. Across all sites, elasticities related to survival were more important than those of growth or vegetative reproduction. The demographic analysis supported the previous observations of changes in the density of M. prasina based on the chronosequence and also suggests that establishment via seeds must occur prior to abandonment in active pastures.     

Role of selection and gene flow in population differentiation at the edge vs. interior of the species range differing in climatic conditions

Evaluating the relative importance of neutral and adaptive processes as determinants of population differentiation across environments is a central theme of evolutionary biology. We applied the Q_ST–F_ST comparison flanked by a direct test for local adaptation to infer the role of climate‐driven selection and gene flow in population differentiation of an annual grass Avena sterilis in two distinct parts of the species range, edge and interior, which represent two globally different climates, desert and Mediterranean. In a multiyear reciprocal transplant experiment, the plants of desert and Mediterranean origin demonstrated home advantage, and population differentiation in several phenotypic traits related to reproduction exceeded neutral predictions, as determined by comparisons of Q_ST values with theoretical F_ST distributions. Thus, variation in these traits likely resulted from local adaptation to desert and Mediterranean environments. The two separate common garden experiments conducted with different experimental design revealed that two population comparisons, in contrast to multi‐population comparisons, are likely to detect population differences in virtually every trait, but many of these differences reflect effects of local rather than regional environment. We detected a general reduction in neutral (SSR) genetic variation but not in adaptive quantitative trait variation in peripheral desert as compared with Mediterranean core populations. On the other hand, the molecular data indicated intensive gene flow from the Mediterranean core towards desert periphery. Although species range position in our study (edge vs. interior) was confounded with climate (desert vs. Mediterranean), the results suggest that the gene flow from the species core does not have negative consequences for either performance of the peripheral plants or their adaptive potential.     

Prolonged clonal growth: escape route or route to extinction?

Many plant species have the capability to reproduce sexually as well as clonally. The balance between clonal reproduction and sexual reproduction varies between different species. It was estimated that 66.5% of all central European flora may form independent but genetically identical daughter plants. Also within species there is great variation in the ratio clonal/sexual reproduction. Clonal reproduction can be considered as an alternative life cycle loop that allows persistence of a species in the absence of the ability to complete the normal life cycle (i.e. seed production, germination and recruitment). Plant populations exhibiting prolonged clonal growth have been referred to as 'remnant populations'. A remnant population in general is defined as "a population capable of persistence during extended time periods despite a negative population growth rate (λ<1) due to longlived life stages and life cycles, including loops, that allow population persistence without completion of the whole life cycle". Here we argue that prolonged and nearly exclusive clonal growth through environmental suppression of sexual reproduction can ultimately lead to local sexual extinction and to monoclonal populations of a species, and that this may imply significant consequences for population viability. Especially obligate or mainly outcrossing clonal plant species may be vulnerable for sexual extinction. We argue that the consequences of reduced sexual recruitment in clonally propagating plants may be understudied and underestimated and that a re-evaluation of current ideas on clonality may be necessary.     

Fire as a control agent of demographic structure and plant performance of a rare Mediterranean endemic geophyte

We examine the effects of fire and/or surrounding vegetation cover on demographic stage densities and plant performance for a rare endemic geophyte, Acis nicaeensis (Alliaceae), in Mediterranean xerophytic grasslands of the 'Alpes-Maritimes' French 'département', through sampling plots in unburned and burned treatments. Fire increases density of flowering individuals and seedling emergence, as well as clump densities and number of individuals per clump, per limiting vegetation height and cover, and increasing bare soil cover. In contrast, fire has no effect on reproductive success. Nevertheless, two growing seasons after fire, all parameters of demographic stages and plant performance do not significantly differ between the two treatments. Small-scale fire is beneficial for the regeneration of this threatened geophyte at a short-time scale. In this context, a conservation planning with small and controlled fires could maintain the regeneration window for populations of rare Mediterranean geophytes.