Tree regeneration and future dynamics of the laurel forest on Tenerife,Canary Islands

Abstract. We studied two sites in the laurel forest of Tenerife to predict future changes in canopy composition. We used two projection methods: Horn's ‘Markovian Projection’, which utilizes information on juveniles in the vicinity of canopy trees, and a 'Stand Projection’, which ignores such information. We performed these projections both including and excluding a-sexual regeneration. Although all of our projections predict a change in species composition, inclusion of a-sexual reproduction decreased the magnitude of successional change. The persistence of Prunus lusitanica and Ilex canariensis appears to be highly dependent on a-sexual regeneration. Both the Markov- and stand projections predict a slight convergence in species composition between the two sites when only sexual regeneration is considered, and also a higher dominance of the shade-tolerant species Laurus azorica. When a-sexual regeneration is also considered, some divergence is shown, with less projected change in the stand projection than in the Markov projection. In spite of some differences between the models, general patterns such as an increase of shade-tolerant species (Laurus azorica and Prunus lusitanica) and a decrease of shade-intolerant species (Erica arborea, Erica scoparia and Myrica faya) are consistent.     

Altitudinal pattern of vegetation variation on Tenerife

Abstract. The variation in structure and floristic composition of the vegetation of Tenerife is related to different environmental factors. 200 sample plots, stratified according to altitude, exposure to wind, kind and age of parent rock, inclination and degree of disturbance were analysed throughout the island. The resulting data were analysed using univariate statistics (altitudinal distribution profiles of the more frequent species) and multivariate methods (Correspondence, Cluster and Pattern Analysis). Altitude, exposure to wind and, to a lesser extent, disturbance were found to be the major determinants of vegetational variation, whereas parent rock and inclination were less important. Ordination results show a continuum in structural and floristic variation of the vegetation with species distributions varying continually along the altitudinal gradient. Nevertheless, altitudinally bound plant community types were recognized with Cluster Analysis. Vegetational discontinuities, which were also detected by pattern analysis, are more evident on the windward slope of the island; they have been attributed to the existence of a major underlying environmental discontinuity the ‘cloud sea’-which is associated with a climatic inversion. On the leeward slope, beyond the influence of the cloud sea, discontinuities are less evident and vegetational variation seems to be closer to a coenocline, i.e. a vegetational continuum.     

Distribution of alien vs. native plant species in roadside communities along an altitudinal gradient in Tenerife and Gran Canaria (Canary Islands)

Roadside plant communities were studied along two roads following an altitudinal gradient in Gran Canaria and Tenerife (Canary Islands). Our aim was to investigate variation in plant species richness, particularly of the alien flora, along a gradient from coastal shrubland to summit vegetation (1950 m a.s.l. in Gran Canaria, 2300 m in Tenerife) in relation to variation in habitat factors (altitude, habitat structure, roadside disturbance, distance to urban nuclei). We compared different species groups that were classified in terms of their biogeographical status, origin and life form. Altitude was the most important factor determining species richness and composition along both roadside transects. Alien plants showed a unimodal distribution pattern along the altitudinal gradient, with less species and lower abundance at low and high altitudes, and highest abundance at intermediate altitude. Alien plant species were also relatively more frequent near urban centres. The number of native and alien species was significantly positively correlated along the altitudinal gradient. Both alien and native, non-endemic species showed differences in their distribution along the altitudinal gradient according to their biogeographical affinities and climatic tolerances. Despite considerable differences in species pools these patterns were consistent among the two islands. Environmental (abiotic) stress is proposed as a primary, altitude-related factor acting as a filter against most alien plants at coastal and high-mountain altitudes. A higher frequency or intensity of disturbance at intermediate altitudes may be a further causal factor promoting alien plants in this zone. Future management efforts to control alien plants along roads should, therefore, concentrate on intermediate altitudinal zones of the higher Canary Islands.     

Contrasting patterns of intraspecific trait variability in native and non-native plant species along an elevational gradient on Tenerife,Canary Islands

Background and AimsNon-native plant species are not restricted to lowlands, but increasingly are invading high elevations. While for both native and non-native species we expected variability of plant functional traits due to the changing environmental conditions along elevational gradients, we additionally assumed that non-native species are characterized by a more acquisitive growth strategy, as traits reflecting such a strategy have been found to correlate with invasion success. Furthermore, the typical lowland introduction of non-native species coming from multiple origins should lead to higher trait variability within populations of non-native species specifically at low elevations, and they might therefore occupy a larger total trait space.MethodsAlong an elevational gradient ranging from 55 to 1925 m a.s.l. on Tenerife, we collected leaves from eight replicate individuals in eight evenly distributed populations of five native and six non-native forb species. In each population, we measured ten eco-morphological and leaf biochemical traits and calculated trait variability within each population and the total trait space occupied by native and non-native species.Key ResultsWe found both positive (e.g. leaf dry matter content) and negative (e.g. leaf N) correlations with elevation for native species, but only few responses for non-native species. For non-native species, within-population variability of leaf dry matter content and specific leaf area decreased with elevation, but increased for native species. The total trait space occupied by all non-native species was smaller than and a subset of that of native species.ConclusionsWe found little evidence that intraspecific trait variability is associated with the success of non-native species to spread towards higher elevations. Instead, for non-native species, our results indicate that intermediate trait values that meet the requirements of various conditions are favourable across the changing environmental conditions along elevational gradients. As a consequence, this might prevent non-native species from overcoming abruptly changing environmental conditions, such as when crossing the treeline.     

Spatial patterns of trees and juveniles in a laurel forest of Tenerife, Canary Islands

Spatial patterns are important characteristics of the forest and theycan reveal such things as successional status and ecological characteristics ofthe species. We tested the hypothesis that spatial distribution will bedifferent, depending on whether the species is intolerant or tolerant to shade.We assessed the spatial distribution of trees (> 4 cm dbh) andjuveniles in eight laurel forest plots. A univariate spatial analysis(performed with Ripley's K₁) showed that all tree species havesignificantaggregation at short distances (2 m). Nevertheless, two groups ofspecies could be differentiated: Erica scoparia,Myrica faya and Ilex canariensisshowed a tendency for aggregation at large distances (larger than 6m)while L. azorica and Prunuslusitanicashowed aggregation only at shorter distances. Ripley's BivariateK_1,2 analyses showed no significant differences in the spatialdistribution ofanalyzed species pairs from a null model. Only Laurusazoricahad a sufficient sample size for analysis of juvenile distribution. Aunivariateanalysis revealed thatL. azorica seedlings (stems < 50 cm high)were clumped in some plots up to 5 m, but this was not consistent.Saplings (stems > 50 cm high and < 4 cm dbh)didnot show strong clumping even at short distances. L.azoricasaplings had no significant aggregation with, nor repulsionfrom, adults of the same or different species. Spatial patterns of the speciesshould be considered in the development of restoration plans of the laurelforest 90%of which has disappeared or been intensively disturbed on Tenerife Island.     

Variation in species composition and vegetation structure of succulent scrub on Tenerife in relation to environmental variation

Abstract. On Tenerife, the occurrence of environmental gradients over short distances provides a unique opportunity to investigate the relationship between vegetation and environmental factors. In the semi‐arid coastal region of Tenerife, floristic composition, species richness and vegetation structure of perennial plants have been studied in 67 locations covering the existing precipitation gradient. On the island as a whole, variation in species composition could be best explained by mean annual precipitation; at coastal sites, substrate age and soil characteristics also played a significant role. On the other hand, substrate chemistry and the type of eruptive material explained little of the floristic variation. Stand biomass was strongly correlated with mean annual precipitation and was, on the youngest lava flows studied, also affected by substrate age. The native stem succulent species made up the bulk of total biomass along the whole precipitation gradient. Disturbed and undisturbed sites differed significantly in stand biomass and cover. Species richness was correlated with precipitation and substrate age. Distribution of plant functional types was also related to the precipitation gradient. The relative abundance of hemicryptophytes and shrubs with non‐hairy leaves increased with increasing precipitation whereas the ratio of shrubs with hairy/non‐hairy leaves and succulent plants decreased. Some alien plants were quite frequent at disturbed sites but, on the whole, they contributed little to the species spectrum and to the stand biomass. Undisturbed sites remained almost free of introduced species not considering annuals.     

Regeneration strategies of tree species in the laurel forest of Tenerife (The Canary Islands)

The laurel-forest of the Canary Islands is a montane cloud-forest. In order to gain some knowledge on the processes that maintain tree species diversity, we conducted an analysis of three different laurel-forest plots of the Anaga massif (Tenerife), varying in canopy composition but growing under similar environmental conditions. For each plot we recorded basal area of the canopy trees (h<1.30 m), the density of suckers and seedlings (h>1.30 m), as well as seed-bank composition. The plots have similar regeneration composition, which appears to be independent of differences in canopy composition. Laurus azorica is the most common seedling species, whereas Prunus lusitanica is the most abundant species among suckers and basal shoots. Neither Erica arborea nor Myrica faya, the two main canopy trees in one of the plots, were found in any of the stands as seedlings or suckers, despite their existence as viable seeds in the seed-bank. The regeneration composition and the canopy composition in one of the plots is remarkable different, revealing differents dynamics processes in the three plots. The results suggest the existence of three well-defined ecological groups: pioneer (regeneration primarily by seedlings), non-pioneer (regeneration by seedlings and suckers) and remnant species (regeneration primarily by suckers).These three groups and the effect of small scale disturbances (natural and human-induced), could help to understand the maintenance of tree species richness.     

The long-term ecology of the lost forests of La Laguna, Tenerife (Canary Islands)

Aim We report the first analysis of the long‐term ecology of Tenerife, in order to establish a pre‐colonization base‐line and to assess the effect of human activity and the role of climatic variation on vegetation during the Late Holocene. Location A former lake bed in the city of La Laguna (Tenerife, Canary Islands, Spain). Methods A sedimentary sequence of over 2 m was obtained from the former lake bed. Fossil pollen and microfossil charcoal concentrations were analysed. Radiocarbon dating of the sequence indicates that it spans approximately the last 4700 years. The pollen diagram was zoned using optimal splitting within psimpoll 4.25. Results Three pollen zones were differentiated: (1) in Zone L1 (c. 4700–2900 cal. yr bp ) a mixed forest was dominated by Quercus, Carpinus, Myrica and Pinus; (2) in Zone L2 (c. 2900–2000 cal. yr bp ) the laurel forest taxa increased, while Pinus, Juniperus and Phoenix declined; and (3) Zone L3 (c. 2000–400 cal. yr bp ) was characterized by the decline of Carpinus and Quercus and the abundance of laurel forest taxa (e.g. Myrica). Neither Carpinus nor Quercus was hitherto considered to be native to the Canary Islands. Their decline started c. 2000 years ago, coinciding with microfossil charcoal evidence of increased burning and with archaeological evidence for the first human settlement on Tenerife. Main conclusions Between c. 4700 and 2000 cal. yr bp , the composition of the forest in the valley of La Laguna was very different from what it is at present. In particular, Quercus and Carpinus appear to have been significant components, alongside components of the present‐day laurel forest, and the native pine (Pinus canariensis) forest and thermophilous woodland were also more prevalent in the region (but probably not within the lake basin itself) until 3000 cal. yr bp . The subsequent decline of Quercus and Carpinus led to the establishment of the present laurel forest in the region and a shift to more open vegetation types. These changes indicate that the aboriginal inhabitants of the islands, the Guanches, had a far more profound impact on the vegetation of Tenerife than hitherto realized.     

A new species of Pseudoniphargus (Crustacea,Amphipoda) from subterranean waters in Tenerife (Canary Islands)

Pseudoniphargus candelariae n. sp. is described from a ground-water-fed well in Tenerife (Canary Islands) and is compared with all species known from Macaronesia. The new species has one unique character (spination of palmar angle of gnathopod 1 sexually dimorphic) and presents furthermore a new combination of morphological features not found in other species of the genus in the area studied.     

Patterns of plant species distribution in the Trans‐Himalayan region of Ladakh,India

Abstract. We studied the vegetation distribution in eight landscape types distinguished along an altitudinal gradient in the Trans‐Himalayan region of Ladakh, India. The point‐intercept method was used for vegetation sampling. Six plant communities were distinguished by cluster analysis. Of these 6 communities, three communities were dominated by shrub species. Table lands are the landscape type with the highest species diversity followed by undulating areas and river beds. Most plant species were restricted to one landscape type.     

Genetic structure, phylogeography and demography of two ground-beetle species endemic to the Tenerife laurel forest (Canary Islands)

The volcanic island of Tenerife (Canary archipelago) was formerly covered at 600-1200 m above sea level on most of its northern side by a cloud forest holding much of the endemic insect fauna. In the most significant surviving patches of this laurel forest at the eastern and western tips of the island occur two forest-specialist, closely related species of Eutrichopus (Coleoptera, Carabidae); here we present data on mitochondrial DNA variation among populations of these species. In total, 116 individuals from 16 localities were sampled and a 638 bp fragment of the cytochrome oxidase subunit II gene was sequenced, obtaining evidence for two distinct evolutionary lineages, in accordance with morphological and biogeographical data. Volcanic events at approximately 0.7 Ma might be responsible for vicariance and the fragmentation of the geographical range of an ancestral species, causing the establishment of two matrilineal lineages. Using nested clade and historical demography analyses we infer past cycles of demographic bottlenecks followed by population expansion, mostly in agreement with the geological time scale of volcanic events. Recent trends, however, refer to fragmentation of the cloud forest due to human intervention.     

A molecular study of hybridization and homoploid hybrid speciation in Argyranthemum (Asteraceae) on Tenerife,the Canary Islands

Examples of recurrent homoploid hybrid speciation are few. One often‐cited example is Argyranthemum sundingii. This example includes two described species, A. lemsii and A. sundingii, resulting from reciprocal hybridization between A. broussonetii and A. frutescens on Tenerife. The four species and artificial F₁ and F₂ hybrids have previously been investigated morphologically and cytologically. Here, we examine population differentiation based on amplified fragment length polymorphism to get a better understanding of the genetic relationships among the species and the extent of hybridization. We aim to investigate if there is molecular support for treating the hybrid species as one taxon. Seven parental and four hybrid species populations (149 individuals) were analysed and we scored 85 polymorphic markers. A few (2–5) were private to each species but variably present and mostly rare. Our principal coordinate, STRUCTURE and BAPS analyses and AMOVA resulted in a clear separation of the parental species. The hybrid species were genetically less divergent but not identical. Our data indicate that hybridization and introgression are common in all these species on Tenerife and support the hypothesis that homoploid hybrid speciation has occurred repeatedly. Intrinsic post‐zygotic barriers are notoriously weak in Argyranthemum and reproductive isolation and speciation result primarily from strong ecological selection. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 19–31.     

Changes in plant species composition and litter production in response to roads and trails in the laurel forest of Tenerife (Canary Islands)

Abstract

Road density has increased in the Canary Islands' forests during the last century, affecting an unknown amount of forested area. We studied road effects on vegetation in the relict laurel forest of Tenerife. We assessed edge effects on plant species richness, plant composition and litter production. Effects of anthropogenic corridors on vegetation differed between paved roads and unpaved trails. Opportunistic species (shade intolerant) dominated road edges, but composition differed among all sites. Multivariate analysis revealed convergence in species composition along the corridor-interior gradient. For trails, both species richness and litter production did not differ significantly between edge and interior. Road edge effects on vegetation were detectable only within the first 10 m towards the interior. This suggests that the main effects of roads and trails on species richness are limited to the immediate edge of the laurel forest. Litter fall along road edges was half that of the interior. However, no significant differences were detected due to the high variability of the data. A buffer of approximately 10 m would result in the reduction of the total area of the remaining undisturbed laurel forest. Based on these results, the building of new paved roads should not be considered. Low human population inflow into the Anaga Rural Park needs to be maintained on a sustainable basis. Forest managers should take these road/trail effects into account when planning new road openings in this ecosystem.     

Effects of climate change on the distribution of plant species and plant functional strategies on the Canary Islands

Aim

Oceanic islands possess unique floras with high proportions of endemic species. Island floras are expected to be severely affected by changing climatic conditions as species on islands have limited distribution ranges and small population sizes and face the constraints of insularity to track their climatic niches. We aimed to assess how ongoing climate change affects the range sizes of oceanic island plants, identifying species of particular conservation concern.

Location

Canary Islands, Spain.

Methods

We combined species occurrence data from single-island endemic, archipelago endemic and nonendemic native plant species of the Canary Islands with data on current and future climatic conditions. Bayesian Additive Regression Trees were used to assess the effect of climate change on species distributions; 71% (n = 502 species) of the native Canary Island species had models deemed good enough. To further assess how climate change affects plant functional strategies, we collected data on woodiness and succulence.

Results

Single-island endemic species were projected to lose a greater proportion of their climatically suitable area (x ̃ = −0.36) than archipelago endemics (x ̃ = −0.28) or nonendemic native species (x ̃ = −0.26), especially on Lanzarote and Fuerteventura, which are expected to experience less annual precipitation in the future. Moreover, herbaceous single-island endemics were projected to gain less and lose more climatically suitable area than insular woody single-island endemics. By contrast, we found that succulent single-island endemics and nonendemic natives gain more and lose less climatically suitable area.

Main Conclusions

While all native species are of conservation importance, we emphasise single-island endemic species not characterised by functional strategies associated with water use efficiency. Our results are particularly critical for other oceanic island floras that are not constituted by such a vast diversity of insular woody species as the Canary Islands.     

Global patterns of genetic variation in plant species along vertical and horizontal gradients on mountains

Aim To understand global patterns of genetic variation in plant species on mountains and to consider the significance of mountains for the genetic structure and evolution of plant species. Location Global. Methods We review published studies. Results Genetic diversity within populations can vary along altitudinal gradients in one of four patterns. Eleven of 42 cited studies (26% of the total) found that populations at intermediate altitudes have greater diversity than populations at lower and higher altitudes. This is because the geographically central populations are under optimal environmental conditions, whereas the peripheral populations are in suboptimal situations. The second pattern, indicating that higher populations have less diversity than lower populations, was found in eight studies (19%). The third pattern, indicating that lower populations have lower diversity than higher populations, was found in 10 studies (24%). In 12 studies (29%), the intrapopulation genetic variation was found to be unaffected by altitude. Evidence of altitudinal differentiation was found in more than half of these studies, based on measurements of a range of variables including genome size, number of chromosomes or a range of loci using molecular markers. Furthermore, great variation has been found in phenotypes among populations at different altitudes in situ and in common garden experiments, even in cases where there was no associated variation in molecular composition. Mountains can be genetic barriers for species that are distributed at low elevations, but they can also provide pathways for species that occupy high‐elevation habitats. [Correction added after publication 9 October 2007: ‘less diversity’ changed to ‘greater diversity’ in the second sentence of the Results section of the Abstract] Main conclusions Genetic diversity within populations can vary along altitudinal gradients as a result of several factors. The results highlight the importance of phenotypic examinations in detecting altitudinal differences. The influence of mountain ridges on genetic differentiation varies depending, inter alia, on the elevation at which the species occurs. Based on these findings, zoning by altitudes or ridges would be helpful for the conservation of tree populations with the onset of global warming.     

Evolution and island endemism of morphologically cryptic Baetis and Cloeon species (Ephemeroptera,Baetidae) on the Canary Islands and Madeira

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Germination patterns throughout an insular altitudinal gradient: The case of the Macaronesian endemic plant Rubia fruticosa Ait. (Rubiaceae) in El Hierro (Canary Islands)

Seed germination percentage and rate of Rubia fruticosa, an endemic Macaronesian shrub was assessed with seed lots collected at three levels of an altitudinal gradient (200–800 m asl) located in El Hierro Island (Canarian Archipelago). Both seeds collected from control plants and those found in lizard droppings were studied. A significant decrease in the germination percentage was observed with control seeds and those seeds which are previously eaten by lizards if collected at 800 m asl. This indicates that R. fruticosa is a plant whose optimal germination environment is located between 200 and 500 m asl, occurring in higher altitudes only under sub-optimal conditions. The effect of lizard gut on R. fruticosa seeds was variable in the two study years: while germination percentage of seeds having passed through the lizards was not significantly different to that of control seeds in the first year, an increased germination was found for the animal-treated seeds in all three altitudinal zones in the second year. In general, germination rate of control seeds was rather variable between years. However, improvement of R. fruticosa seed germination caused by lizards may be important for its fitness; and a co-evolution may have occurred, since lizards and seeds of this endemic plant have been intensively interacting for millions of years in the lower zones of the Canary Islands.     

The effect of spring burning on competitive ranking of prairie species

Abstract. A common explanation for the changes in species abundance following a fire is a shift in competitive ranking. However, experimental tests have been inconsistent and generally do not support this explanation. I examined the competitive ability of an abundant C₄ grass, Andropogon gerardii, and a C₃ forb, Ratibida pinnata, in a prairie remnant in northern Ohio, USA, for each of three years following a spring burn in 1996. While the abiotic environment directly influenced both species similarly, relative competitive abilities in terms of growth changed markedly: in 1996 Andropogon was less inhibited by neighbors; in 1997 both Andropogon and Ratibida had similar competitive abilities; and in 1998 Ratibida was less inhibited by neighbors. This shift in competitive response ranking paralleled the changes in relative abundance for the two species. In contrast, the effect of neighbors on survival changed markedly over time but did not differ among the two species. Thus, fire may influence species abundance through changing species competitive response ranking, at least in terms of growth.     

Climatic range and distribution of Chinese Fagus species

Abstract. The distribution of five Chinese Fagus species was mapped, and their geographical range clarified. Their climate range was characterized following Thornthwaite (1948) for moisture and temperature (Thornthwaite's moisture index ranging from 26 to 320), and Kira (1945) for temperature (mean annual temperature from 5 to 17 °C), and compared with the range of other beeches. Chinese beeches occur only in S. China, mainly between 700 m and 2500 m altitude. They grow abundantly only in perhumid climates. The Northern Hemisphere beeches occur mainly in temperate climates; however, they reach warmer extremes in North America and China than in Europe and Japan. Beech habitats in montane China are more humid than in lowland Europe and North America. Chinese beeches are limited by a moisture deficit in the north, and by high temperatures plus relatively low moisture levels in the south. However, they tolerate a certain water deficit in the growing season. The southern geographical and low altitudinal limits of Chinese Fagus species are probably largely determined by the lack of competition with evergreen dicotyledonous trees, together with human influences.     

Effects of temperature,light and gibberellic acid on the germination of seeds of 43 species native to Western Australia

Abstract. Species native to the southwest of Western Australia, representing a range of plant families, life-history strategies, fire-response syndromes, seed-store types and seed weights, were tested for viability using tetrazolium chloride and for germination under combinations of constant temperatures of 15 °C or 23 °C, constantly dark or 12 h diurnal whitelight conditions, and with, or without, addition of gibberellic acid (GA₃, 50 mg/l). Species previously known to require a heat-shock treatment to overcome dormancy due to an impervious testa were pre-treated prior to imposition of temperature, light and GA₃ conditions. The test environmental conditions related to differences between winter and autumn temperatures and surface and buried seed germination positions of post-fire habitats. The viability of the selection of native Western Australian species ranged from 0 to 100 %, averaging 71 %. For all taxa, no combination of temperature, light and gibberellic acid treatment induced all viable seeds to germinate. The greatest percentage germination achieved in any combination of treatments averaged 71 % of all viable seeds for all species. Larger seeds (> 10 mg seed weight) tended to have greater viability percentages, but no overall patterns of viability or germinability were attributable to seed storage syndrome, strategy of fire recovery response or life-form type. Germination of most of the dominant tree representatives (Eucalyptus calophylla, E. diversicolor, E. erythrocorys, E. gomphocephala, and E. patens) was indifferent to the trial conditions of temperature, light and GA₃. However, Eucalyptus marginata showed reduced germination in the light, which was overcome with GA₃. GA₃ also overcame the inhibition resulting from exposure to light in some understorey species (e.g. Allocasuarina campestris, Regelia ciliata, Xanthorrhoea gracilis and X. preissii). Germination of many hard-seeded, understorey shrub and herbaceous perennial species, especially those with small (< 10 mg) seeds (e.g. Bossiaea ornata, B. aquifolium and Acacia drummondii ssp. candolleana) was greater at the lower trial temperature and in the dark. Some large (> 10 mg) seeded, understorey species (e.g. Acacia extensa, Kennedia coccinea, K. prostrata, Hovea trisperma and Hardenbergia comptoniana) germinated in high percentages in both temperatures, but maximum germination percentages still tended to be at 15 °C. Large-seeded species were less sensitive to exposure to light compared to the smaller seeded species. The largest seeded species tested, Paraserianthes lophantha, germinated best in the warmer incubation temperature and in the light. The ecological significance of the tests would be that species which have seed dormancy mechanisms capable of delaying germination until the cool temperature, winter rainy period of this mediterranean-type climate would be more likely to survive than if germination followed summer rain showers or the first, intermittent rains of autumn. Burial of seeds becomes more important if germination occurs when rains first begin as this period has less available soil moisture and temperatures are high. Also survival of seedlings could be enhanced if germination of seed was restricted to the positions protected from high light, higher temperatures and lower soil moisture by the presence of a forest canopy. Therefore, seeds which have an ability to sense the presence of a previous fire in the habitat, conditions in light environment and appropriate temperature level have an adaptive advantage to time emergence to situations of time and space where survival is maximized. Variation in viability and germination percentages were apparent in some cases where more than one seed collection of available for testing, indicating that further aspects, such as seed age, maturity at collection, storage conditions and depth of seed dormancy, remain to be considered.