Heat-shock and seed germination of a group of Mediterranean plant species growing in a burned area: An approach based on plant functional types

Many plants from Mediterranean-type ecosystems show enhanced germination when exposed to heat and other fire-related cues. Whether Mediterranean-type plants are fire recruiters and need fire to germinate is subject of discussion. In the present work, we tested the above hypothesis by subjecting seeds from 57 species representative of a recently burned plant community to various heat pulses. Differences in germination among species, taking into account their phylogeny, were studied by dividing them according to their life-form (chamaephytes, hemicryptophytes), regeneration strategy (non-sprouters, sprouters) and geographical distribution range (Iberian Peninsula endemics, Mediterranean, widely distributed species). Seeds were exposed to 80, 100 and 120 °C during 10 min, and then, incubated at 17.5 °C for 6 weeks in darkness. Maximum germination was reached at 80 °C and control treatments and minimum germination was at 120 °C. Most species were tolerant to heat-shocks, only a few of them were stimulated by heat, and around 20% suffered a reduction in germination with heat. The germination response to heat-shock differed according to regenerative strategy and distribution range, but not to life-form. These results were supported independently of phylogeny. As expected, sprouters and Iberian endemics were more sensitive to heat-shock than non-sprouters and widely distributed species, respectively. As a conclusion, the studied plant group appears to exhibit resistance to heat but not dependence on it for germination. Therefore, recruitment could be high after low intensity fires, but might be significantly reduced for most species after more intense fires.     

The effect of short fire cycles on the cover and density of understorey sprouting species in South African mountain fynbos

Abstract. Two South African mountain fynbos sites were studied to determine the effect of short fire cycles on the cover and density of understorey sprouting species and their subsequent effect on plant-species richness. Frequent fires (4–6 years between burns) increased the cover of sprouting species by 32% when compared to an adjacent site where the penultimate fire was 28 years previously. There was little or no effect of fire frequency on the densities of understorey sprouters; however, individuals were larger at sites with short fire cycles. The response of individual species of sprouters was variable with one species, Hypodiscus striatus , showing no response to fire frequency. The impact of sprouting species on the species richness of the plant community was great. The mean number of species recorded in quadrats with a high cover of sprouters was 60% lower in comparison to quadrats with low covers or under the burned skeletons of overstorey proteas. The effect of sprouters was consistent for all functional groups of species (i.e. sprouters, non-sprouters, short-lived and long-lived species), in each case reducing the number of species present.     

Range-size,local abundance and germination niche-breadth in Mediterranean plants of two life-forms

The relationships between rarity (i.e., range-size, local abundance) and niche-breadth can be important to assess the risks the species face under global change, namely those resulting from fire regime change. In fire areas, germination is critical for establishing after fire for many species. We examined the relationships between rarity and germination niche-breadth for 53 plant species of two life-forms (chamaephytes and hemicryptophytes) growing in Eastern Spain. Rarity was measured as geographic range-size and as local abundance. Local abundance was evaluated at two sites differing in their post-fire successional status (a recently burned area and a long-unburned one). Germination niche-breadth was measured as the mean germination evenness index from four germination experiments that subjected the seeds to various germinating conditions characteristics of fire environments. Correlations between rarity (range-size, local abundance) and niche-breadth were calculated in cross-species mode or by way of phylogenetically independent contrasts, and either for the 53 species (all-species set) or for each of two life-form groups (chamaephytes, hemicryptophytes). In general, no significant correlations were found between the rarity measures and germination niche-breadth for the all-species set. However, significant correlations emerged when the analyses were done separating species by life-forms. Germination niche-breadth was positively correlated with range-size for chamaephytes, and negatively for hemicryptophytes. In addition, germination niche-breadth was uncorrelated, or negatively so, with local abundance for chamaephytes and hemicryptophytes, respectively. While no correlation between range-size and local abundance was found for chamaephytes, a negative one was obtained for hemicryptophytes. We conclude that rarity/germination-niche relationships varied by life-form. This pattern of relationships was obscured when all species were joined in a single group. Based on the contrasting patterns of correlations obtained for each life-form we argue that the changes in the germinating environment caused by alterations in fire regime are likely to differentially affect these two groups of species.     

Effects of Incubation Temperature on Seed Germination of Mediterranean Plants with Different Geographical Distribution Ranges

Little is known about consequences of the increasing global temperatures on seed germination and even less about the seed germination response of species with different geographical distribution ranges. In the present work, the effect of incubation temperature on seed germination of 31 species from the Iberian Peninsula is analyzed in relation to their geographical distribution range. We tested whether plants with different geographical distribution ranges (Iberian Peninsula endemics, Mediterranean, widely distributed species), respond similarly to incubation temperature using cross-species analysis as well as taking into account phylogenetic relationships among species. Seeds were incubated at 10°C, 15°C, 20°C and 25°C for 6?weeks in darkness. Distribution range appeared as a determinant of germination response to the temperature treatment independent of phylogeny. Widely distributed species were less sensitive than endemic or Mediterranean species to incubation temperatures, which may represent an advantage to colonize new areas or to tolerate changes in the environment. Although their ability to avoid germination during the warm and dry season might be favorable for endemic species in some situations, our results also suggest that under a future of temperature increase imposed by climate change, endemic plant species may be more vulnerable than widely distributed species.     

Profiling a besieged flora: endemic and threatened plants of the Cape Peninsula, South Africa

The Cape Peninsula (area: 471 km²), situated at the south-western extremity of the Cape Floristic Region, has exceptionally high plant species richness (2285 species and infraspecific taxa) and numbers of endemic (90; 88 species and two infraspecific) and threatened (141; 138 species and three infraspecific) taxa (termed species from here on). This biodiversity is threatened by urban development and the spread of invasive alien plants. Peninsula endemics are concentrated in a few, predominantly species-rich families and these correspond well with endemic-rich families in other areas of the Cape Floristic Region. A high level of similarity exists between families with threatened and families with endemic species. A frequency analysis of the biological traits of both endemic and threatened species shows that low growing, ant-dispersed shrubs are over-represented in both groups. Endemics are most likely to be non-sprouters, but threatened plants do not have a specific post-fire regeneration strategy. Threatened species have higher frequencies of geophytes, sprouters and wind-dispersed species compared to endemic species. Numbers of endemic and threatened species are not randomly distributed with regard to occurrence in vegetation types and patterns are similar for both groups. The habitat and biological profiles of both endemic and threatened species suggest that they are highly vulnerable to extinction as a result of increasing rates of alien plant infestation, urbanization and inappropriate fire regimes.     

Soil seed bank,fire season,and temporal patterns of germination in a seeder-dominated Mediterranean shrubland

Soil seed banks play a major role in the post-fire regeneration of Mediterranean shrublands. They vary throughout the year in species composition, abundance, and readiness to germinate. After fire, germination occurs mainly during the following fall to spring. Time of germination can determine recruitment success. It is unclear what factors control post-fire germination and its timing. We tested the effects of season and fire on the readily germinable soil seed bank of a seeder-dominated shrubland. Plots were burned early and late in the summer season (ES, LS). Soil samples were collected before and after fire, and germinated in a chamber simulating successively autumn, winter, and spring conditions. Samples were kept moistened at all times. Fire intensity was similar between ES and LS. Several species of Cistus and herbs, mostly annuals, were dominant. Most germination occurred during the simulated-autumn period, with little subsequent germination during the following two periods. Germination speed (T ₅₀) during simulated-autumn was similar for shrubs and herbs, and independent of season or fire. Germination was lower for two shrubs (Rosmarinus officinalis, Cistus salvifolius) and higher for herbaceous dicots in LS than in ES soils. Fire reduced monocots and enhanced Cistus. Germination period significantly interacted with fire and season in some groups or species, altering the simulated-autumn germination peak. We demonstrate that the seed bank can germinate swiftly under simulated-autumn conditions. Hence, water availability is the main controlling factor of germination. Fire season differentially affected some species or groups, and could affect the post-fire regeneration.     

Local versus regional intraspecific variability in regeneration traits

Intraspecific trait variability has a fundamental contribution to the overall trait variability. However, little is known concerning the relative role of local (e.g. disturbances and species interaction) and regional (biogeographical) processes in generating this intraspecific trait variability. While biogeographical processes enhance plant trait variability between distant populations, in fire-prone ecosystems, recurrent fires may have a preponderant role in generating variability at a local scale. We hypothesize that plants respond to the local spatio-temporal heterogeneity generated by fire by having a relatively large local variability in regeneration traits in such a way that overrides the variability at a broader biogeographical scale. We test this hypothesis by assessing the intraspecific variability in fire-related regeneration traits of two species (Cistus salviifolius and Lavandula stoechas) growing in fire-prone ecosystems of the Mediterranean Basin. For each species, we selected six populations in two distant regions, three in the east (Anatolian Peninsula) and three in the west (Iberian Peninsula). For each species and population, we analysed the following regeneration traits: seed size, seed dormancy and stimulated germination by fire-related cues (heat and smoke). To evaluate the distribution of the variability in these traits, we decomposed the variability of trait values at each level, between regions (regional) and between population within region (local), using linear mixed-effect models. Despite the biogeographical and climatic differences between regions, for the two species, intraspecific variability in regeneration traits was higher at a local (within regions) than at a regional scale (between regions). Our results suggest that, in Mediterranean ecosystems, fire is an important source of intraspecific variability in regeneration traits. This supports the prominent role of fire as an ecological and evolutionary process, producing trait variability and shaping biodiversity in fire-prone ecosystems.     

Ecophysiological effects of light quality and nitrate on seed germination in species from Western Australia

Germination occurs usually in response to multiple environmental cues. Seeds with the ecophysiological ability to simultaneously sense the previous presence of fire and appropriate levels of temperature, light and soil nitrate could restrict germination to postfire, winter and competition-free microhabitats, where the potential for seedling survival is enhanced. Germination responses of 16 species with a range of life forms, fire responses and seed weights were determined under controlled conditions of 15°C temperature, a 12 h light cycle, exposure to 1 g L^?1 nitrate solution, and six conditions of light quality (white, blue, yellow, red, far-red light and darkness). Germination in Oenothera stricta, a weedy naturalized ephemeral, and two small-seeded indigenous Asteraceae species of mulga woodlands, Leucochrysum fitzgibbonii and Craspedia sp., were enhanced by white, yellow or red light compared with germination achieved in the dark, or under far-red or blue light. In red light, KNO₃ further enhanced germination of these positively photoblastic species. The germination response of Trachyandra divaricata, a naturalized herb of sandy, seaside locations, and several native jarrah forest legumes (four Acacia species, Bossiaea aquifolium, Gompholobium marginatum and Sphaerolobium vimineum) proved to be negatively photoblastic. Of these seven negatively photoblastic herb and shrub species, exposure to KNO₃ overcame the inhibition of light only in the resprouter species, Acacia lateriticola. In the serotinous, negatively photoblastic tree species, Corymbia calophylla and Eucalyptus marginata, KNO₃ seemed to be required before the negative response to light exposure was recorded. A dose–curve experiment on two positively photoblastic and three negatively photoblastic species indicated that although KNO₃ exposure affected germination in all species, different concentrations of KNO₃ (0, 0.5, 1, 2, and 5 g L^?1) produced different levels of response. Detailed studies with additions of KNO₃ (1 g L^?1) and the growth hormone, gibberellic acid (GA₃; 50 mg L^?1), showed that increased germination percentages of the positively photoblastic species, Oenothera stricta, occurred in the light, but blocking endogenous gibberellic synthesis with paclobutrazol, or adding exogenous GA₃ or KNO₃ had no effect on the light-induced germination levels. In the negatively photoblastic species Trachyandra divaricata, additions of KNO₃ and GA₃ had no influence on the germination inhibition induced by exposure to light nor did blocking endogenous GA synthesis. The 16 species growing naturally in Western Australia, Australia show a range of germination responses to environmental conditions, but depending on their natural habitat, the ecophysiology of each species appears to be optimized for subsequent seedling survival.     

Competitive abilities of three narrowly endemic plant species in experimental neighborhoods along a fire gradient

We conducted field experiments manipulating lichens, shrubs, and herbs along a time-since-fire gradient and assessing effects on three endemic herbaceous species of Florida scrub: Eryngium cuneifolium, Hypericum cumulicola, and Polygonella basiramia. Responses included seed germination, survival, biomass, and fecundity. Transplants into recently burned patches generally had higher survival, larger biomass, and greater reproductive output than transplants into long-unburned patches. Open areas and sites near oaks frequently were more favorable than sites near Florida rosemary. Ground lichens did not affect germination but increased mortality rate of seedlings. Neighboring small shrubby and herbaceous species did not affect the performance of these species. Of the three species, naturally occurring E. cuneifolium were farthest from large shrubs, and their microhabitats had the least ground lichens and shrubs. Eryngium cuneifolium and H. cumulicola are capable of forming persistent seed banks and their recruitment after fire depends mostly on these dormant seeds. Polygonella basiramia relies on seed dispersal and immediate seed germination to colonize recently burned patches. Management for these species should involve variable fire regimes to allow all three species to persist along with many other scrub endemics.     

Shoot regeneration after fire or freezing temperatures and its relation to plant life-form for some heathland species

Shoot regeneration after prescribed burning or following the freezing temperatures of winter was monitored for nineteen heathland species present in an Arctostaphyleto-Callunetum community in northeast Scotland. Species whose renewal buds were near the surface of the ground started to grow earlier in the spring than species with renewal buds above the surface, but grouping species according to the position of their renewal bud (i.e. their life-form) did not account for all of the interspecific variation apparent. In the case of shoot regeneration after fire, species whose renewal buds were destroyed by fire because they were above-ground started to regenerate about the same time as species with belowground buds, protected from fire, but reached their maximum frequency of occurrence later. Grouping species by life-form was of limited value as a means of interpreting this interspecific variation in the timing of shoot regeneration after fire. It would be unwise to use plant life-form as the sole basis for interpreting or predicting a species' response to temperature stress when extreme temperatures occur regularly, as they do in heathland. The possible use of other plant traits to interpret and predict interspecific variation in the regeneration rate of heathland plants is discussed.Nomenclature follows Tutin et al. (1964–1980) for vascular plants. Acknowledgements. The Nature Conservancy Council and Mr J. J. Humphries kindly allowed Dinnet Moor to be used for the work presented here. One of us (RJR) received financial support for field work from the Natural Sciences and Engineering Research Council of Canada.     

Competitive interactions between overstorey proteas and sprouting understorey species in South African mountain fynbos

Abstract. Previous studies in the mountain fynbos of South Africa have demonstrated that short fire cycles favour the establishment of dense covers of understorey sprouters while longer fire intervals enable the establishment from seed of overstorey proteas and the formation of a overstorey. One consequence of these differences between fire cycle lengths is the effect that understorey sprouters and an overstorey protea canopy have on species richness. In the case of short fire intervals, species richness is decreased while longer intervals between fires allow species richness to decrease or increase depending on the patchiness of the overstorey canopy. Such results are suggestive of competitive effects between understorey sprouters and overstorey canopy proteas. In this study, data were collected from several pyric successional stages in mountain fynbos to study the effect of overstorey proteas on the growth and flowering of understorey sprouters since the last fire. Data were also collected to determine the effect that understorey sprouters had on the establishment and fecundity of overstorey protea species. Competitive interactions between overstorey proteas and sprouting understorey species were evident at all the sites studied. The vegetative growth and seed production of understorey sprouters, which grew under a canopy of overstorey proteas during the current interfire period, were significantly lower than that for plants growing in the open. In addition, the postfire growth and seed production of understorey sprouters were significantly lower for individuals, which grew under an overstorey protea canopy during the previous fire cycle, than for those individuals which grew in the open. The fecundity of overstorey proteas, which grew near understorey sprouters, was lower than that of plants which grew in the open. This effect was evident for up to the first 15 years after a fire. However, not all understorey sprouters affected the overstorey proteas equally. Also, seedlings of overstorey proteas established significantly less successfully in close proximity to understorey sprouters after a fire than in the open or under proteas. Finally, the results demonstrate that complex species‐specific, understorey–overstorey interactions are important in mountain fynbos. For example, some overstorey species depend on trophically similar species to reduce potential competition from understorey sprouters for their successful establishment at a site.     

Ecophysiology of fire-stimulated seed germination in Cistus incanus ssp. creticus (L.) Hey wood and C. salvifolius L.

Abstract. Seed germination characteristics were investigated in the most common Cistus species in Greece, namely C. incanus ssp. creticus and C. salvifolius. In addition to the soft seed subpopulation, both species produce a large fraction of hardcoated, water-impermeable seeds which can be softened and, thus, promoted to germinate by mechanical scarification and thermal pretreatment. Temperature and light control of seed germination are unimportant. In the ecological context of the Mediterranean ecosystems, the eventually advantageous, opportunistic strategy of germination is based on: (a) seed heterospermy (which allows the smaller, softcoated fraction to germinate promptly each year while the majority of the seeds, the hard ones, accumulate in the soil); (b) the seed population heterogeneity in relation to coat hardness (so that any heat conditions produced by fire induce the softening and germination of a certain seed fraction); (c) the notably low germination rate (which suppresses premature germination); (d) the wide, Mediterranean-type (relatively cool), temperature range of germination (while higher temperatures simply inhibit but do not induce any dormancy); and (c) the apparent lack of photo-sensitivity (enabling germination under every light regime). In non-fire years, the temporal distribution of field germination and seedling appearance might be partly determined by the seed dispersal strategy of the individual Cistus species. Nevertheless, the post-fire regeneration response is manifested in the form of a huge wave of germination (of practically all seeds softened by the fire heat), shortly after the onset of the rainy season.     

Seed germination of high mountain Mediterranean species: altitudinal,interpopulation and interannual variability

The germination response of 20 species from high altitude Mediterranean climates, most of them rare endemics, was studied. Our main goal was to model the germination response of a complete set of Iberian high mountain species. The effect of temperature and other parameters, such as spatial and temporal short gradients, on germination were also evaluated. Some seed features (mass and size) were also related to the germination response. Finally, we tested the effect of cold-wet stratification pretreatment when germination was low under natural conditions. Seeds were collected at four locations from 1,900 to 2,400 m a.s.l. in the Sierra de Guadarrama (Spanish Central Range) over two consecutive growing seasons (2001–2002) and submitted to different temperatures and a constant photoperiod of 16 h light/8 h darkness. Most plants readily germinate without treatment, reaching an optimum at relatively high temperatures in contrast to lowland Mediterranean species. Seeds seem to be physiologically prepared for rapid germination even though these plants usually face very intense summer droughts after ripening and dispersal. Germination was also highly variable among altitudes, populations and years, but results were inconsistent among species. Such flexibility could be interpreted as an efficient survival strategy for species growing under unpredictable environments, such as the Mediterranean climate. Finally cold-wet stratification increased germination capacity in five of nine dormant species, as widely reported for many arctic, boreal and alpine species. In conclusion, high mountain Mediterranean species do not differ from alpine species except that a relatively high number of species are ready to germinate without any treatment.     

Resprouting Ability of Six Atlantic Shrub Species

Resprouting from subterranean structures is a principal method of vegetative regeneration that many shrub species show after a disturbance. This study, therefore, aims to determine the resprouting capacity and intensity of six dominant species in an Atlantic shrubland area located in the NW of the Iberian Peninsula and compare their resprouting and germinating strategies. Resprouting intensity is measured using three variables: individual probability of resprouting, number of resprouts and length of these sprouts in three plant-age classes. The intensity for each species was calculated using a simple index (IRI) that included the three measured variables. All studied species, excepting Erica umbellata, could resprout. According to the IRI values, there are three groups of species: strong resprouters (Ulex europaeus, Ulex minor and Pterospartum tridentatum), weak resprouters (Ulex micranthus and Genista triacanthos) and non-resprouters (E. umbellata). The germination of strong resprouters is highly stimulated by fire. Frequent disturbances remove the non- and weak resprouter populations and promote the strong resprouter ones.     

Regeneration models and plant regenerative types related to the intensity of fire in Atlantic shrubland and woodland species

Question: Is it possible to model the germinative and resprouting behaviour of plant species in Atlantic shrublands and woodlands in relation to fire intensity? Is it possible to recognise different functional regenerative types in these plant species? Location: Galicia, NW Iberian Peninsula. Methods: We explored the patterns of germination and resprouting plant responses in relation to different intensities of fire using data from 37 trees, shrubs and herbaceous species growing in Atlantic shrublands and woodlands. Results: Synthesizing their germinative and resprouting behaviour, we created two graphical models: the Functional Germinative Model (FGM) and the Functional Sprouting Model (FSM). Integrating the germinative and resprouting responses, and taking into account fire intensity, we created the Functional Regenerative Model (FRM), which predicts the post‐fire recuperation of the populations of each species. The FRM has been validated with data from four Atlantic communities. We identified four plant functional regenerative types (PFRT) for Atlantic forest vegetation and we propose three intensities of response. Conclusions: The extracted models (FGM, FSM and FRM) and the grouping of species in four PFRTs could be applicable to more Atlantic species, to disturbance ecology in general and to population, community and landscape management.     

Tree diversity,composition, forest structure and aboveground biomass dynamics after single and repeated fire in a Bornean rain forest

Forest fires remain a devastating phenomenon in the tropics that not only affect forest structure and biodiversity, but also contribute significantly to atmospheric CO2. Fire used to be extremely rare in tropical forests, leaving ample time for forests to regenerate to pre-fire conditions. In recent decades, however, tropical forest fires occur more frequently and at larger spatial scales than they used to. We studied forest structure, tree species diversity, tree species composition, and aboveground biomass during the first 7 years since fire in unburned, once burned and twice burned forest of eastern Borneo to determine the rate of recovery of these forests. We paid special attention to changes in the tree species composition during burned forest regeneration because we expect the long-term recovery of aboveground biomass and ecosystem functions in burned forests to largely depend on the successful regeneration of the pre-fire, heavy-wood, species composition. We found that forest structure (canopy openness, leaf area index, herb cover, and stem density) is strongly affected by fire but shows quick recovery. However, species composition shows no or limited recovery and aboveground biomass, which is greatly reduced by fire, continues to be low or decline up to 7 years after fire. Consequently, large amounts of the C released to the atmosphere by fire will not be recaptured by the burned forest ecosystem in the near future. We also observed that repeated fire, with an inter-fire interval of 15 years, does not necessarily lead to a huge deterioration in the regeneration potential of tropical forest. We conclude that burned forests are valuable and should be conserved and that long-term monitoring programs in secondary forests are necessary to determine their recovery rates, especially in relation to aboveground biomass accumulation.     

Colonization of dynamic Mediterranean landscapes: where do birds come from after fire?

Aim Two main mechanisms may explain post‐disturbance species colonization patterns of early successional habitats such as those originated by wildfires. First, post‐disturbance colonization is not limited by the dispersal ability of the species to reach the newly created open areas and, secondly, colonization is limited by dispersal. Under the first hypothesis, we expect, at a regional scale, to find similar post‐disturbance communities to develop on recently burned sites. However, colonization limited by dispersal will lead to strong between‐site variations in species composition. Location To test these hypotheses, we studied the post‐fire colonization patterns of nine open‐habitat bird species in eight distantly located wildfires in the north‐eastern Iberian Peninsula. Methods We censused post‐fire bird composition by means of field transects and identified potential colonization sources from species–habitat suitability maps derived from atlas data. Results Our results showed strong significant differences in post‐fire species composition between burnt areas. Burnt areas located in areas with low probability of species presence before the fire event showed lower species occurrence and richness after the fire. Main conclusions These results do not support the idea that early successional stages and open habitats have a homogeneous community structure at regional scales and suggest that dispersal is a key constraint determining bird colonization of post‐fire habitats. Further attention should be paid to landscape heterogeneity as a key factor in determining population dynamics of open‐habitat species in the light of current and future land‐use changes in Mediterranean regions.     

Endemic species may have complex histories: within‐refugium phylogeography of an endangered Iberian vole

Glacial refugia protected and promoted biodiversity during the Pleistocene, not only at a broader scale, but also for many endemics that contracted and expanded their ranges within refugial areas. Understanding the evolutionary history of refugial endemics is especially important in the case of endangered species to recognize the origins of their genetic structure and thus produce better informed conservation practices. The Iberian Peninsula is an important European glacial refugium, rich in endemics of conservation concern, including small mammals, such as the Cabrera vole (Microtus cabrerae). This near‐threatened rodent is characterized by an unusual suite of genetic, life history and ecological traits, being restricted to isolated geographic nuclei in fast‐disappearing Mediterranean subhumid herbaceous habitats. To reconstruct the evolutionary history of the Cabrera vole, we studied sequence variation at mitochondrial, autosomal and sex‐linked loci, using invasive and noninvasive samples. Despite low overall mitochondrial and nuclear nucleotide diversities, we observed two main well‐supported mitochondrial lineages, west and east. Phylogeographic modelling in the context of the Cabrera vole's detailed fossil record supports a demographic scenario of isolation of two populations during the Last Glacial Maximum from a single focus in the southern part of the Iberian Peninsula. In addition, our data suggest subsequent divergence within the east, and secondary contact and introgression of the expanding western population, during the late Holocene. This work emphasizes that refugial endemics may have a phylogeographic history as rich as that of more widespread species, and conservation of such endemics includes the preservation of that genetic legacy.     

Determinant variables of Iberian Peninsula Aphodiinae diversity (Coleoptera, Scarabaeoidea, Aphodiidae)

Aim The aims of this paper are to examine diversity–variability patterns for species of Aphodiinae (Coleoptera, Scarabaeoidea, Aphodiidae) on the Iberian Peninsula, and to determine the factors that influence their geographic distribution. Location Iberian Peninsula (Spain and Portugal). Methods Data from 30 studies and their bibliographies on species of Iberian Peninsula Aphodiinae were compiled. The reliability of the inventories was evaluated using parametric species richness estimators. In addition, a further 11 variables related to rarity, geographic distribution, or phylogenetic diversity were considered. Diversity variables were analysed using principal components analysis to reduce the number of dependent variables. Subsequently, the effect of differences in locality size among the 30 studies was eliminated by calculating and retaining the residuals of the curvilinear relationship of each diversity variable with the area. Generalized linear models were used to examine the relationships between diversity and 17 environmental variables. The diversity variables and their residuals were also subject to trend surface analysis in order to identify the relevance of spatially structured variables that had not been considered. The contribution of explanatory variables was determined through hierarchical variance analysis. Results Principal components analysis of biodiversity variables revealed that most of the variability could be explained using three biodiversity indexes: BI1, correlated positively with species richness, widely distributed species, frequent species, abundant species, species occurring in North Africa, Europe and the Iberian Peninsula, and phylogenetic diversity; BI2, correlated positively with numbers of infrequent and African–Iberian species; and, BI3, correlated positively with numbers of endemic, non‐abundant, European, and Iberian‐restricted species. A latitudinal disjunction emerged in BI1, with maximum scores at the north‐western and southern corners, while maximum BI2 scores were found throughout the south, and maximum BI3 scores in the north‐west. For BI1, it was climate that had the greatest influence, followed by lithology, and livestock presence. Geographic variables were the most significant for BI2, followed by climate and livestock presence. Finally, for BI3, climate variables were the most important, while geography, lithology and livestock presence had some relevance. Main conclusions The relevance of geographic variables indicates that other unaccounted‐for factors that are spatially structured could possibly explain additional variation in Aphodiinae diversity. These factors may be historic in nature, relating to the species groups, namely the Ibero‐European and the Mediterranean or Afro‐Iberian. The northern pattern could reflect the fact that the Iberian Peninsula acted as a colonization route and as a refuge during the glacial/interglacial cycles, while the southern pattern could be a consequence of the connection between the Iberian Peninsula and North Africa during the Messinian crisis, and/or a historic relationship in common, related to human activity.