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In a unique study, Luna (Luna, Sci Rep 10:1–10, 2020) examined the viability and germination of 12 hard-seeded Cistaceae in the Mediterranean Basin by alternating a prolonged summer-type temperature (50/20 °C at 12 h cycles) treatment with a fire-type heat pulse. A re-analysis of their data shows that the summer treatment applied before the heat pulse was superfluous as similar high levels of germination under ambient conditions were attained with the heat pulse only. Additional tests using the hard seeds of Acacia showed that the water gap opened at once in the presence of dry heat such that contact with moisture is not required to complete the process of softening. The abundance of hard seeds remaining when the summer treatment was applied after the heat pulse is better explained by ungerminated seeds having become hard again under such dry conditions rather than remaining dormant, i.e., acquiring secondary physical dormancy, and thus becoming ‘desensitized’ to their environment. While this response may be adaptive, such a retarding effect will be limited in practice as most fires are expected in autumn, at least historically, and are thus close to the start of optimal winter conditions for germination. Future studies should concentrate on the fate of the water gap plug during such alternating treatments and also ensure that realistic summer temperature regimes are used.

The occurrence of Helianthemum jonium Lacaita & Grosser ex Bég. in Fiori & Bég. (Cistacaeae) is reported and documented here for the first time in Croatia and Montenegro, and it is confirmed in the Albanian flora after 80 years. The species grows on sandy soils within maquis and garrigue communities in four Adriatic islands: Vis, Biševo, Hvar and Lopud, and within psammophilous vegetation near Ulcinj in Montenegro and in Hamallaj, north of Durrës in Albania. The morphological relationship with the allied H. leptophyllum Dunal and H. apenninum (L.) Mill. is here briefly discussed. Remarks about its taxonomy, lectotypification, habitat and ecology in the Adriatic region are also provided. The species H. jonium was previously known only from the Apennine Peninsula; hence these new eastern Adriatic records extend its distribution area to the east, and it can therefore be considered an amphi-Adriatic endemic element., Prisutnost vrste Helianthemum jonium Lacaita & Grosser ex Bég. in Fiori & Bég. (Cistacaeae) je ovdje po prvi puta zabilježena i dokumentirana u flori Hrvatske i Crne Gore te potvrđena u flori Albanije nakon 80 godina. Vrsta raste na pješčanim tlima unutar vegetacije makije i gariga na četiri jadranska otoka: Vis, Biševo, Hvar i Lopud te unutar psamofilne vegetacije kod Ulcinja u Crnoj Gori i Hamallaju, sjeverno od Drača u Albaniji. Morfološka sličnost sa srodnim vrstama, H. leptophyllum Dunal i H. apenninum (L.) Mill. je ovdje ukratko diskutirana. Također se daju napomene o njezinoj morfologiji, lektotipifikaciji, staništu i ekologiji u Jadranskom području. Vrsta H. jonium prethodno je bila poznata jedino s Apeninskog poluotoka, a nova nalazišta u istočnom Jadranu pridonose istočnom proširenju granice areala vrste te se ista mora smatrati amfi-jadranskim endemičnim elementom.

Impoverished pinweed (Lechea intermedia var. depauperata Hodgdon, Cistaceae) is a narrow endemic restricted to the dry, fire-prone Athabasca Plain in northeastern Alberta and northwestern Saskatchewan, Canada. Little is known about the ecology or life history of the taxon that could be used to inform conservation or management decisions. We investigated the reproductive biology of impoverished pinweed to determine whether it exhibits the suite of traits characterizing the temporal disperser syndrome, which is prevalent among Cistaceae species inhabiting fire-prone environments. Specifically, we examined the soil seedbank, characterized seed production, and conducted three sets of germination trials examining the interaction of heat and scarification, burial, and alternate wetting and drying. Our results show that impoverished pinweed is similar to other Cistaceae with respect to seed production, dispersal, and germination, supporting the classification of impoverished pinweed as a temporal disperser. Exposing seeds to temperatures between 125 and 175 °C significantly increased germination as compared with the untreated controls. We discovered an additional cause of seed loss incidentally; here we report the first western Canadian records of Anthonomus pusillus Le Conte, a seed-predating weevil. Our results suggest that the temporal and spatial dynamics of impoverished pinweed populations are strongly shaped by fire and the distribution of soil seedbanks.

11 páginas, 5 tablas y 4 figuras, Seed shape in the Malvaceae and other families of the order Malvales was investigated. Seed shape was quantified by comparison with the cardioid. The J index is the percent similarity between both images, the seed and the cardioid, and similarity is considered in cases where the J index is over 90. Seed shape was analysed in 73 genera, and seeds resembling the cardioid were found in 10 genera, eight in the Malvaceae and two in the Bixaceae and Cistaceae. Seed shape was quantified by comparison with the cardioid in 105 species. A correlation was found between the values of the J index and plant form, with higher values of the J index in the seeds of herbs, intermediate – in bushes, and lower values in trees. The results suggest a relationship between seed shape and plant form, where seeds resembling the cardioid are associated with plants having small size.

Mediterranean shrublands are water-limited and fire-prone ecosystems. Post-fire seed regeneration occurs under a rainfall regime that is highly variable, with frequent drought periods. It has been shown that smoke can alter germination performance in numerous species, but little is known about its role during germination under water stress. Here, we studied the germination response to water stress and smoke of nine abundant Mediterranean shrubs. Seeds of six species with hard coats (three Cistaceae and three Fabaceae) and three with soft coats (Lamiaceae), exposed or not to smoke, were set to germinate under four levels of water availability (0; − 0.15; − 0.30; − 0.45 MPa). Seeds of hard-coat species were previously mechanically scarified. Onset of germination, final germination and viability were recorded. Water stress reduced final germination in five of the six hard-coat species and in one soft-coat species, delayed the onset of germination in all species and decreased viability in several hard-coat species. Smoke increased final germination (10%) in one soft-coat species, not affecting the rest. Additionally, smoke reduced the onset of germination in one hard- and one soft-coat species. However, there was no interaction between smoke and water stress. Hydrotime modeling of the germination responses confirmed the above findings. In conclusion, the species studied differed in germination sensitivity to water stress, with hard-coat species being more sensitive than soft-coat species. Exposing the seeds to smoke generally did not change this sensitivity.

Background: Species of the Cistaceae family have hard-coated seeds and consequently require scarification for germination. It is widely assumed that the high temperature caused by fire is a scarify...

The meadow spittlebug, Philaenus spumarius (L.) (Hemiptera: Aphrophoridae), is the main vector in Europe of the recently detected plant pathogen bacterium Xylella fastidiosa Wells et al. (Xanthomonadales: Xanthomonadaceae). While the ecology of continental populations is well documented, nothing is known about the insular populations of P. spumarius, such as in Corsica, where the bacterium was detected in 2015. Hence, in an epidemiological context, the ecology of P. spumarius has been studied in a maquis landscape in the Ajaccio region between 2017 and 2019. Adults and nymphs were almost exclusively collected on Cistus monspeliensis L. (Cistaceae). However, very few specimens were collected in summer, suggesting a movement of the adults to sheltered habitats. Unfortunately, despite several trapping methods used, the location of adult summer habitat remains unknown for the studied population. It might be tempting to destroy the central plant host of P. spumarius populations. However, as spittlebug nymphs are highly polyphagous on low-growing plant species and as the females can lay eggs in any dead plant tissues, such practice could have limited the impact. Instead, the strong relationship between P. spumarius and C. monspeliensis could be used to monitor spittlebug populations, to limit/concentrate the means of insect control, or in an agronomic context to lure insects away from crops. Maintaining natural arboreal vegetation around agronomic systems could help decrease insect abundance – and potentially, pathogen load – on cultivated species. Such hypotheses need to be further studied by landscape experiments.

Natural rangelands occupy about 5.5 million hectares of Tunisia’s landmass, and 38% of this area is in Tataouine governorate. Although efforts towards natural restoration are increasing rapidly as a result of restoration projects, the area of degraded rangelands has continued to expand and the severity of desertification has continued to intensify. Any damage caused by disturbances, such as grazing and recurrent drought, may be masked by a return of favorable rainfall conditions. In this work, conducted during March 2018, we surveyed the botanical composition and species diversity of natural rangelands in Tataouine in southern Tunisia. The flora comprised about 279 species belonging to 58 families, with 54% annuals and 46% perennials. The Asteraceae family had the greatest richness of species, followed by Poaceae, Fabaceae, Amaranthaceae, Brassicaceae, Boraginaceae, Caryophyllaceae, Lamiaceae, Apiaceae, and Cistaceae. Therophytes made the highest contribution, followed by chamaephytes and hemicryptophytes. Of all these species, 40% were palatable to highly palatable and more than 13% are used in both traditional and modern medicine.

Accurate inference in phylogeography requires appropriate sampling strategies. Complex questions demand a large sample size at both the population and genetic levels to obtain precise reconstructions. This is the case of the phylogeographic history of Cistus monspeliensis, a plant that displays low plastid (cpDNA) diversity in the Mediterranean Basin but high diversity in the Canary Islands. Here, we aimed to identify Mediterranean refugial areas and to accurately quantify inter-island colonization events in the Canaries. Using a previous study as starting point, we increased sample size in two ways: (i) additional sampling of plastid genetic markers (from 1041 to 1899 bp); and (ii) additional sampling of populations (from 47 to 69) in long-term persistence areas suggested by species distribution modeling (SDM). The synergy between SDM and extended population sampling helped find higher genetic diversity. Our deeper phylogeographic sampling of C. monspeliensis revealed the following: (i) potential refugia in long-term persistence areas with high cpDNA diversity in western Europe and the Canary Islands; and (ii) a significant increase (from 7 to 12) in the number of inferred inter-island colonization events across the archipelago. Our results stress the usefulness of SDM to identify the genetic signature associated with potential refugial areas. We herein propose a field sampling approach based on SDM that, in combination with a larger cpDNA sampling, can help answer a wide array of phylogeographic questions, such as the location of Quaternary refugia and number of colonizations across archipelagos.

Some fire ecology studies that have focused on garrigue-like vegetation suggest a weak selective pressure of fire in the Mediterranean Basin compared to other Mediterranean-type regions. However, fire-prone Mediterranean heathland from the western end of the Mediterranean Basin has been frequently ignored in the fire ecology literature despite its high proportion of pyrogenic species. Here, we explore the evolutionary ecology of seed traits in the generalist rockrose Cistus salviifolius L. (Cistaceae) aiming to ascertain the role of the Mediterranean heathland for fire adaptations in the Mediterranean Region. We performed a germination experiment to compare the relationship of seed size to (i) heat-stimulated germination, (ii) dormancy strength, and (iii) heat survival in plants from &lsquo, high-fire&rsquo, heathland vs. &lsquo, low-fire&rsquo, coastal shrubland. Germination after heat-shock treatment was higher in large seeds of both &lsquo, and &lsquo, habitats. However, dormancy was weaker in small seeds from &lsquo, habitats. Finally, seed survival to heat shock was positively related to seed size. Our results support that seed size is an adaptive trait to fire in C. salviifolius, since larger seeds had stronger dormancy, higher heat-stimulated germination and were more resistant to heat shock. This seed size&ndash, fire relationship was tighter in &lsquo, Mediterranean heathland than &lsquo, coastal shrubland, indicating the existence of differential fire pressures and evolutionary trends at the landscape scale. These findings highlight the Mediterranean heathland as a relevant habitat for fire-driven evolution, thus contributing to better understand the role of fire in plant evolution within the Mediterranean region.

Hardseededness is a common trait in Mediterranean plant communities, although the origin of its selection is controversial. It may be a mechanism of persistence to fire temperatures, but could also form part of a gap-detecting mechanism to provide germination cues under arid conditions. To disentangle this, we studied the phylogenetic structure of plant communities against fire frequency and aridity gradients. The phylogenetic structure in Mediterranean Basin ecosystems was analysed for the hardseededness trait as a whole and was separated by the families composing this trait (Fabaceae and Cistaceae). This study focused on woody perennial species. The phylogenetic structure was also contrasted against soil classes. Hardseededness on the whole, and for the Fabaceae family alone, showed phylogenetic clustering as aridity increased. Cistaceae displayed the opposite pattern with phylogenetic clustering in most humid areas, together with a significant soil effect. Surprisingly, fire frequency had no influence in any case. This climate-driven phylogenetic clustering indicates that the hardseededness trait could confer some fitness advantage under dry conditions. For this reason, coexisting species were more closely related in the community with increasing aridity. This effect was especially evident for the Fabaceae family. These results shed some light on the evolutionary selection of this adaptive trait under Mediterranean conditions. Our results question the role of fire in the selection of the hardseededness trait in Mediterranean Basin ecosystems and indicates that climate is the most important factor. Therefore, we should be cautious in assigning to fire a preponderant role in the selection of some plant traits. This research was financed by the IMAGINA (PROMETEO/2019/110) project. VMS was supported by a “Beatriu de Pinós” fellowship (2014BP-B-00056) from the Generalitat de Catalunya. JGA was supported by a Ramón y Cajal fellowship (RYC-2016-20528). IA was supported by a Horizon 2020 research and innovation programme with Marie Sklodowska-Curie Cofund grant agreement No. 801596.

Natural and anthropogenic fires have historically been part of the Mediterranean Basin (MB). As a result of this long exposure to fires, MB flora has developed various strategies to persist under recurrent fires. One of these strategies is to survive and regenerate by resprouting and another is through post-fire seedling recruitment. For species that recruit seedlings after a fire there is evidence that fire-related germination cues are implicated in the process of triggering seedling emergence from soil-stored seeds. In some species, particularly in Cistaceae and Fabaceae, germination is stimulated by heat. Fire-related chemical germination cues, such as smoke, charcoal, and nitrogenous compounds (collectively termed ‘smoke’), have also been recognised as promoting seed germination in MB flora. However, the role of smoke in the germination of MB plant species has received less attention and recognition than the role of heat-shock and is generally seen as having a relatively limited role as a post-fire germination cue in MB flora. Our hypothesis is that this is due to the fact that research on smoke-stimulated germination in the MB region has been limited and poorly addressed, and this is biasing our view on the role of smoke in MB flora. Using available literature, we first analyse the role of smoke in the germination of MB flora and discuss some of the issues that limit our understanding of its role on the flora of the region. We point to gaps in the literature and provide suggestions for further research. We conclude that differences in evidence of smoke-stimulated germination in MB flora when compared with other Mediterranean regions may not reflect real ecological or evolutionary differences among these regions but are likely the consequence of the limited number of well-designed studies in the MB., BM was supported by the Fundação para a Ciência e a Tecnologia [grant number SFRH/BPD/90277/2012]. This work was performed under the framework of the TREVOL and FILAS projects (CGL2012-39938-C02-01, CGL2015-64086-P) from the Spanish Government and the PROMETEO/2016/021 project from the Valencia government (Generalitat Valenciana). CIDE is supported by CSIC, Generalitat Valenciana and the University of Valencia.

The biogeographic history and the degree of environmental niche conservatism provide essential clues to decipher the underlying macroevolutionary processes of species diversification and to understand contemporary patterns of biodiversity. The genus Helianthemum constitutes an excellent case study to investigate the impact of the geo-climatic changes and the environmental niche shifts on the origins of plant species diversity in the Mediterranean hotspot. It is a palearctic species-rich lineage with c. 140 species and subspecies mostly belonging to three distinct evolutionary radiations, almost confined to the Mediterranean region and occurring across varied environmental conditions. In this work, we studied the ample and rapid diversification of the genus Helianthemum across its whole distribution range by performing phylogenetic reconstructions of ancestral ranges and environmental niche evolution. We observed a striking synchrony of biogeographic movements with niche shifts between the three major clades of the genus Helianthemum, likely related to the geo-climatic events occurred in the Mediterranean Basin since the Upper Miocene. In particular, Late Miocene and Early Pliocene were dominated by episodes of range expansions, the Late Pliocene by range contraction and vicariance events, and Pleistocene by most intense environmental niche shifts and in-situ diversification. Our study also provides evidence for four main environmental niches in Helianthemum (i.e., Mediterranean, subdesert, humid-montane and subtropical-insular) and a tendency toward environmental niche conservatism within different subclades, with few niche shifts mostly occurring from Mediterranean ancestors. The relative longer time spent in Mediterranean areas by the ancestors of Helianthemum suggests that the larger species diversity observed in the Mediterranean (i.e. Northern Africa and Southern Europe) may have been generated by a time-for-speciation effect reinforced by environmental niche conservatism. Overall, our work highlights the role of the Mediterranean Basin as a ‘cradle of diversity’ and an ‘evolutionary hub’, facilitating the environmental transitions and determining the building up of a global plant biodiversity hotspot. Ministerio de Economía y Competitividad CGL2014-52459-P, CGL2017-82465-P

Isolated populations are usually subject to low fitness and reduced genetic diversity, both of which may negatively affect their survival and adaptive potential. Hence, these issues cannot be neglected when planning conservation actions for isolated populations. The Italian population of Cistus laurifolius subsp. laurifolius is extremely isolated. Furthermore, it is affected by fragmentation, being constituted by a single larger subpopulation, surrounded by three much smaller subpopulations, a few hundred metres to a few kilometres apart. In order to fill gaps in demographic and genetic knowledge concerning the Italian population, its area of occupancy, size, age-stage structure and phenology were investigated and its reproductive fitness, pollination strategies and genetic variability were assessed. The population was inferred as fully xenogamous and showed good reproductive performance. Despite this, its genetic variability was low and it showed relatively high levels of inbreeding depression (Fis), seemingly not affected by sub-population size. These results suggest that the Italian population recently suffered fragmentation and reduction in size. The low genetic diversity observed could be explained by the high percentage of mature individuals found in the population, possibly established before fragmentation. For these reasons, the Italian population of C. laurifolius subsp. laurifolius should be monitored and concrete actions aimed at its conservation planned.

Plant’s reproductive success is determined by intrinsic characteristics of the reproductive system of the species, as well as by extrinsic abiotic and biotic factors. Reproductive traits can vary along different gradients (e.g. ecological, geographical, etc.) and several studies revealed that the most favourable conditions (i.e. greater densities, fitness and genetic diversity) will be found at the core of a plant distribution range than in marginal or isolated populations. In this study the phenological and reproductive traits of several fragmented populations of Helianthemum caput-felis Boiss. (Cistaceae), a perennial Mediterranean plant were investigated. Six populations, covering the entire distribution range of the species were studied over two years (2013–2014). The phenology and reproductive outputs were analysed considering ecological and climatic factors, and human trampling intensity by fitting Generalised Linear Mixed Models (GLMMs). The results indicated that geomorphology, substrate, slope and human trampling intensity affected the reproductive traits of H. caput-felis. An ecological gradient, mainly linked by the interaction of temperature and precipitation regimes could explain this finding. Plants growing at the eastern edge of the distribution range (Sardinia; geographically marginal) produced more fruits than those growing in the other populations. Conversely, plants growing in arid substrate showed a negative correlation with fruit production and viable fruits. The reproductive outputs of the populations located in the core of the distribution range (Cabo Roig and Moraira; ecologically marginal) were lower than those of the other populations. On the other hand an ecological gradient dominated by the precipitation and temperature regimes was detected. In addition, a positive effect of the high and moderate level of human trampling on the reproductive output was found. Further studies, based in particular on the habitat fragmentation, are needed to understand the complex interactions that govern the reproductive output of H. caput-felis in the Mediterranean region.

Premise of the study Species differ in their temperature germination niche. Populations of a species may similarly differ across the distribution range of the species. Anticipating the impacts of climate variability and change requires understanding the differential sensitivity to germination temperature among and within species. Here we studied the germination responses of four hard-seeded Cistaceae seeders to a range of current and future temperatures. Methods Seeds were collected at sites across the Iberian Peninsula and exposed or not exposed to a heat shock to break dormancy, then set to germinate under four temperature regimes. Temperatures were varied daily and seasonally, simulating the temperature range across the gradient, plus an increased temperature simulating future climate. Time to germination onset and cumulative germination at the end of each season were analyzed for the effects of temperature treatments, seasons, and local climate (temperature of the germination period, Tgp) at each site. Key results Tgp was a significant covariate of germination in all species but Cistus populifolius. Temperature treatments significantly affected Cistus ladanifer, C. salviifolius, and Halimium ocymoides. Germination occurred in simulated autumn conditions, with little germination occurring at later seasons, except in unheated seeds of H. ocymoides. Exposure to a heat shock changed the sensitivity to temperature treatments and the relationships with Tgp. Conclusions Germination responses to temperature differ not only among species but also within species across their latitudinal range. The responses were idiosyncratic and related to the local climate of the population. This germination variability complicates generalizing the impacts of climate variability and climate change.

Terfezia claveryi Chatin is a mycorrhizal fungus that forms ectendomycorrhizal associations with plants of Helianthemum genus. Its appreciated edibility and drought resistance make this fungus a potential alternative crop in arid and semiarid areas of the Mediterranean region. In order to increase the knowledge about the biology of this fungus in terms of mycorrhiza formation and response to drought stress, a catalase from T. claveryi (TcCAT-1) has been purified to apparent homogeneity and biochemically characterized; in addition, the expression pattern of this enzyme during different stages of T. claveryi biological cycle and under drought stress conditions are reported. The results obtained, together with the phylogenetic analysis and homology modeling, indicate that TcCAT-1 is a homotetramer large subunit size monofunctional-heme catalase belonging to Clade 2. The highest expression of this enzyme occurs in mature mycorrhiza, revealing a possible role in mycorrhiza colonization, but it is not upregulated under drought stress. However, the H2O2 content of mycorrhizal plants submitted to drought stress is lower than in well watered treatments, suggesting that mycorrhization improves the plant's oxidative stress response, although not via TcCAT-1 upregulation.

[EN] In animal-pollinated hermaphroditic species, larger and xenogamous flowers increase male-biased resource allocation, whereas smaller and selfing flowers invest disproportionally more resources to female function. In Cistaceae, an entomophilous and hermaphroditic Mediterranean family, this pattern generally follows a phylogenetic signal. However, resource allocation to carpels is independent of phylogeny, which suggests trait divergences among closely related species during the diversification into different environmental conditions. We tested this hypothesis across 37 species of Cistaceae along a temperature and precipitation gradient, including semiarid, dry, subhumid and humid sites. We quantified the proportions of dry mass and nutrient investment to carpels and tested the influence of the climatic gradient and site-specific precipitation on the interspecific variation in carpel resource allocation. Lowest and highest percentages of resource allocation to carpels ranged from 1.5–4.2% to 24.2–36.6%, respectively. The proportion of resources comprised in carpels significantly decreased with increasing precipitation/decreasing temperature. Thus, carpels comprised proportionally more resources under drier and hotter conditions, especially in semiarid sites. Our results demonstrate how the extent of climatic constraints is more important than phylogenetic relationships in determining stress-induced differences in carpel resource allocation across species of Cistaceae in a Mediterranean environment. We suggest that allocation of proportionally more resources to carpels in drier and hotter sites lies within a strategy to deal with the most stressful conditions by means of a high reproductive effort., We thank V.G. Staggemeier and J.M.F. Penha for statistical advice, and two anonymous reviewers for their valuable comments. We also thank J. Arroyo, E. Carrio, J.P. Gonzalez-Varo, J. Güemes, J. Herrera, A.L. Luzuriaga, M. Méndez, M.A. Rodríguez-Gironés, A. Sánchez, S. Santamaría, E. Triano and R. Torices for collecting flower buds from several species. We are also grateful to E. Galisteo and Y. Valinani for lab assistance. This study was supported by the Comunidad de Madrid grant REMEDINAL (TE_CM S2018/EMT-4338) and COMEDIAS FEDER/State Research Agency, Spanish Ministry of Science, Innovation and Universities (Project CGL2017-83170-R)

Demographic rates in plants are usually assumed to be more stage or size dependent than age dependent, and aging is therefore not considered in demographic models. However, little is known about the effect of age on demographic rates, as there still are few studies based on long-term individual-based plant population data that consider both individual age and size. In addition, little is known about how aging of individuals may affect population dynamics. We present analyses of demographic data for three populations of Fumana procumbens collected 1985–2013, on individuals with known year of germination. We modeled age- and size-dependence of the vital rates of survival, growth, fruiting probability, and fruit number using thin plate spline regressions, and constructed an age × size integral projection model (IPM) to project population-level effects of aging. We found strong correlations between age and vital rates in solely age-based vital rate models, where vital rates initially increased with age, after which they stabilized and, in some cases, eventually declined. In survival models with both age and size, the effects of age were statistically significant, whereas size effects were insignificant at two of the sites. For other vital rates, most of the effect of age could be explained by size alone. In addition, including the age effects on survival in the IPM led to lower population growth rates compared to predictions of a size-only IPM. These results illustrate that demographic senescence does occur in perennial plants, which has only been demonstrated clearly in a few recent detailed studies. Moreover, we show that population projections may be overly optimistic if they do not consider plant age. We conclude that the possibility of demographic senescence should be considered in demographic population models, such as those used in viability analyses of threatened species.

Wild edible mycorrhizal mushrooms are among the most appreciated and prized mushrooms in the world. Despite the cultivation of ectomycorrhizal (ECM) mushrooms has been a growing subject of study worldwide, it has been hampered by the mutualistic lifestyle of the fungi. Although not being obligate symbionts, most of the species of ECM mushrooms only produce fruit bodies in association with trees or shrubs. In the present study, we aimed at understanding certain aspects of the ecology of four different edible ECM fungi: Lactarius deliciosus, Tricholoma equestre, T. portentosum and Boletus fragrans. Despite having a broad distribution worldwide, these fungi inhabit also Mediterranean habitats with understories typically dominated by rockroses (Cistaceae). Studying the ecology of these mutualistic fungi as well as the interaction with these species of shrubs is not only scientifically relevant but also pivotal for the discovery of profitable cultivation protocols. We evaluated the compatibility of these ECM species with five species within Cistaceae family - Cistus ladanifer, C. psilosepalus, C. salviifolius, Halimium halimifolium and Tuberaria lignosa. Each species of fungi proved to be able to establish mycorrhizas with at least 2 different plants species but varied in their host range of the tested Cistaceae. The dissimilarity in terms of host specificity between some fungal species seemed to be connected with the phylogenetic distances of the fungi. A correlation between the colonization percentage of the root systems and the mycelial growth rates in pure culture was found. The connection of these traits might be an important key to understanding the ecological competitor-colonizer tradeoffs of these ECM fungal species. Altogether, our study reports unknown plant-fungi combinations with economical relevance and also adds new insights about the ecology of these species of ECM fungi.

Chrysoperla carnea s.l. (Stephens) larvae are important natural enemies in agroecosystems. However, adults feed on honeydew excreted by hemipterans, and pollen and nectar from flowering plants. Pollen is essential for egg production, but to our knowledge, the pollen diet of C. carnea in perennial crops has never been addressed. In this work, the objective was to study the diversity and potential selection of pollen types consumed by C. carnea in the olive grove agroecosystem. For this study, C. carnea adults were captured from April to December of 2012 and 2013, and simultaneously, inventories of the plant diversity were obtained in olive orchards and adjacent scrubland and herbaceous patches. The pollen types contained in C. carnea guts were identified by microscopy and compared with the pollen types in the environment using Jacobs`s second selection index. The results indicated that (i) C. carnea females and males captured in the olive tree canopy visited scrub and herbaceous vegetation patches; (ii) they fed on different anemophilous and entomophilous pollen types from tree and scrub (Olea europaea, Fabaceae, Pinaceae, Cistaceae or Ericaceae) and herbaceous (Asteraceae, Apicaceae, Brassicaceae, Poaceae, Rumex type or Plantago type) strata; and (iii) adults fed not only on flowers but also on pollen settled on vegetation surfaces. Here, we demonstrated that C. carnea could benefit not only from the diversity of entomophilous pollen during the plant flowering periods but also from other pollens that could remain in the environment in different periods. This has important implications for the management of the floral diversity adjacent to the crop. The authors are grateful to the Portuguese Foundation of Science and Technology for financial support through the project EXCL/AGRPRO/ 0591/2012 “Olive crop protection in sustainable production under global climatic changes: linking ecological infrastructures to ecosystem functions.” M.V. thanks FCT, PostDoc grant (SFRH/BPD/ 119487/2016). info:eu-repo/semantics/publishedVersion

Maritime pine forests are a major ecosystem throughout the Portuguese coast and are severely affected by the invasion of Acacia longifolia. The presented study investigated the diversity of ectomycorrhizal fungi (ECM) of major plant species in these ecosystems to find possible links between Pinus pinaster, three native Cistaceae shrubs and the Acacia invasive species. We successfully identified 13 ECM fungal taxa. Within those, two species from the order Helotiales were found in all plant species, and over half of the fungal ECM species found in Pinus pinaster were also common to the Cistaceae shrubs. Network analysis points to the Cistaceae shrubs having a central role in these below-ground communities, therefore enforcing the idea that they are key to these communities and should not be underestimated. Our results also point to the evolving role of invasive plant species in the ecosystem dynamics in the rhizosphere, which host fungal species that are common to native plants, although it is not yet clear whether these fungal taxa are native or a consequence of the presence of Acacia longifolia.

Plant-derived smoke promotes germination in Mediterranean-like environments, but its effect is unclear in the Mediterranean Cistaceae. This article investigates the role of smoke in the comparative germination ecology of five Helianthemum taxa. Laboratory germination experiments were conducted using seeds collected in the field and stored in a seed bank. All seeds were mechanically scarified prior to testing. Various pre-treatments with smoke solutions and gibberellins were applied, and seeds were incubated in three germination temperature regimes (30/20 °C, 22/12 °C, 14/4 °C). In H. cantabricum , H . nummularium , H. oelandicum and H. urrielense germination was very high at the three temperatures, and no effect of smoke was detected. In H. tinetense germination was generally poor and a significant effect of smoke was found, but the effect size was small. The thermal niche of Helianthemum appears to follow an opportunistic strategy, being limited only by physical dormancy at the time of dispersal. H. tinetense follows a more conservative strategy, with physiological dormancy retarding germination. In the Mediterranean basin and surrounding areas, smoke appears to be one of a multitude of environmental signals controlling germination. The smoke effect might be more relevant in certain species such as H. tinetense .

Helianthemum nummularium s.l. is a young, morphologically diverse species distributed from western Europe to the Caucasus and the Southern Urals in the east. We analysed the rps16-trnK plastid intergenic spacer sequences from 85 localities covering most of the range of H. nummularium. Thirteen haplotypes were very unevenly distributed throughout the range of the species, and exhibited a strong phylogeographic signal. The results confirm range expansions of H. nummularium from Mediterranean refugia northwards, but also show the major role of eastern European (the Caucasus and the Southern Urals) refugia in rapid postglacial colonization of east, north and central Europe. The plastid haplotypes form distinct clades, one representing an eastern European lineage with few haplotypes and the other representing a western European lineage with many haplotypes. Parallel to this split in haplotype diversity is the pronounced differentiation in morphological variation displayed by the taxa found in west and east Europe. We discuss the role of topography in generating differences in morphological and genetic diversity between these two groups. We also discuss the taxonomical status of Helianthemum arcticum, which is regarded as an endangered local endemic of the Kola Peninsula. Helianthemum arcticum appears to represent an outlying peripheral population of H. nummularium preserved since the last postglacial major range expansion of this species, and bears the same plastid haplotype as the bulk of east and north European populations.

Mancilla-Leyton, J.M., Fernandez-Ales, R., Martin Vicente, A. 2015. Seed dispersal effectiveness of shrubs by goat. Ecosistemas 24(3): 22-27. Doi.: 10.7818/ECOS.2015.24-3.04 We evaluated the role of domestic goats as seed dispersers of eight representative Mediterranean shrub species ( Cistus salviifolius, C. libanotis, Halimum halimifolium, H. calycinum, Myrtus communis, Phillyrea angustifolia, Pistacia lentiscus and Rhamnus lycioides ). Six female goats of similar size and age were fed with fruits of study species. Following fruits ingestion by goats, total faeces were collected at 24-h intervals. The seeds retrieved were tested for germination and viability, along with seeds not eaten by the goats. Less than 30% of the seeds eaten were retrieved from the dung, with significant differences between species. C. salviifolius showing the maximum recovery percentage (30%), and P. lentiscus the minimum (0%). Seed recovery was significantly related to seed hardness. In most species, the maximum amount of seeds recovered occurred 24-72 h after ingestion. The passage through the goat gut significantly increased seed germination in C. salviifolius, H. halimifolium, H. calycinum and P. angustifolia . The results of this study show that goats can potentially favour (Cistaceae) or inhibit (fleshy fruits) seed dispersal of browsed Mediterranean shrub species. Therefore, these results should be considered when developing conservation and restoration plans of natural vegetation in Mediterranean areas.

Plant population dynamics is fundamental to identify which parts of the life cycle are important when designing management actions. In this study, the short-term population dynamics of Helianthemum caput-felis (Cistaceae), a perennial plant distributed throughout the western Mediterranean Basin, was investigated by monitoring 98 permanent plots randomly placed along the overall distribution range for three years (2013–2015). Demographic surveys were analysed by Integral Projection Models (IPMs), and the analysis of population growth rates, elasticities, and life table response experiment (LTRE) analyses were performed. Under deterministic conditions, the H. caput-felis population showed a slight increase in the first censuses, but a general population decline followed. This trend was probably due to the shrinkage of plants in the last year, which correlated negatively with reproductive traits. Despite this decline, the population was at equilibrium under the simulated environmental stochasticity. The population dynamics of H. caput-felis followed the general pattern typical for long-lived Mediterranean plants: populations with growth rates closer to the equilibrium in the long term, determinant role of the survival-growth transitions for the population growth rate, slow growth and stasis, longevity of established individuals, shrinkage of medium-large plants, low recruitment rate, high generation times and mean long lifespans. The results of this study, which highlight how conservation measures can be focused on protecting and increasing the number of plants of medium-large size, provide important information for the planning management conservation actions for H. caput-felis and for several Mediterranean plants that show a similar life-history strategy.

Environmental changes can affect the distribution and conservation of plant populations, therefore the study of the ability to modulate reproductive and vegetative traits is of large interest. Cistus laurifolius is a shrub typically occurring in Mediterranean open vegetation, often under different degrees of habitat light intensity. In the present work, we investigated reproductive and leaf traits to assess the ability of this species to cope with unfavorable / suboptimal conditions. We quantified fruit set, seed set and seed mass under different conspecific plant density and light conditions, whereas vegetative traits were assessed through the measurement of plant height, Specific Leaf Area (SLA), and Leaf Dry Matter Content (LDMC). Results show that plants of Cistus laurifolius are able to adjust both SLA and LDMC under shaded conditions, but produce fewer seeds with reduced seed mass compared with plants under full light. We conclude that plants are able to modulate their vegetative traits maximizing the carbon gain under shaded conditions, but they are not able to offset the carbon balance deficit and therefore fail to convoy adequate resources to reproductive traits. Our results also highlight that the reproductive fitness is positively affected by conspecific plant density, but negatively affected by canopy coverage.

Cistus ladanifer has a well-defined taxonomic identity. 2,2,6-trimethylcyclohexanone may be an authenticity and taxonomic marker. Its traits and applications make it a possible economic resource fitted for Mediterranean areas. Cistus ladanifer is a dominant shrub species endemic to the western Mediterranean region. Due to its dominant nature and its potential ecological, aromatic or pharmacological applications, C. ladanifer has been the object of numerous studies. In this review current knowledge on different aspects of this species is summarized, from its taxonomy to its chemical characterisation or its competitive traits. There are no doubts about the taxonomic entity of C. ladanifer, although the recognition of infraspecific taxa deserves more attention. Given that the fragrant exudate of C. ladanifer holds a very specific composition, one species specific carotenoid, 2,2,6-trimethylcyclohexanone, derivative is proposed as an authenticity marker for uses of C. ladanifer in pharmacological or aromatic industries. Evidence is also gathered on the extreme adaptation of C. ladanifer to stressful conditions in the Mediterranean region, such as the ability to survive in low hydric and high solar exposition conditions, presistence in poor and contaminated soils, and growth inhibition of several other plants through the release of allelochemicals. Thus, the finding of potential applications for this plant may contribute to enhance the economic dimension of derelict lands, such as mine tailings or poor agricultural Mediterranean areas.

Helianthemum songaricum is an ecologically important endangered relic plant species growing in the arid and semi-arid regions of northwestern China. It is also a valuable germplasm resource. Conseq...

Border and isolated plant populations represent an interesting target for ecological and conservation issues. We analysed the ecological constraints and the conservation status of the eastern population of Helianthemum caput-felis Boiss. (Cistaceae), located in Sardinia. The distribution of H. caput-felis was verified via field surveys; ecological data, morphological and reproductive traits, were recorded in 40 permanent plots randomly established; the human trampling effects on plant density, plant size and plant performance were analysed. Plant density was higher in bedrock and lowland areas, in garrigue and maquis habitats; however, the differences among plants growing in different ecological conditions were not statistically significant; only human trampling intensity significantly affected plant density and lowest values were observed in areas with intense trampling pressure. All ecological variables analysed had a statistically significant effect on plant size and on the number of fruits per...

Most habitat fragmentation studies have focused on the effects of population size on reproductive success of single species, but studies assessing the effects of both fragment size and connectivity, and their interaction, on several coexisting species are rare. In this study, we selected 20 fragments along two continuous gradients of size and degree of isolation in a gypsum landscape in central Spain. In each fragment, we selected 15 individuals of each of three dominant gypsophiles (Centaurea hyssopifolia, Lepidium subulatum and Helianthemum squamatum, 300 plants per species, 900 plants in total) and measured several reproductive traits: inflorescence number, fruit set, seed set and seed mass. We hypothesised that plant fitness would be lower on small and isolated fragments due to an interaction between fragment size and connectivity, and that response patterns would be species-specific. Overall, fragment size had very little effect on reproductive traits compared to that of connectivity. We observed a positive effect of fragment connectivity on C. hyssopifolia fitness, mediated by the increased seed predation in plants from isolated fragments, resulting in fewer viable seeds per capitulum and lower seed set. Furthermore, seed mass was lower in plants from isolated fragments for both C. hyssopifolia and L. subulatum. In contrast, few reproductive traits of H. squamatum were affected by habitat fragmentation. We discuss the implications of species-specific responses to habitat fragmentation for the dynamics and conservation of gypsum plant communities. Our results highlight the complex interplay among plants and their mutualistic and antagonistic visitors, and reinforce the often-neglected role of habitat connectivity as a key component of the fragmentation process.

The importance of shrub formations in the Mediterranean area both in terms of area occupied and carbon sequestered by them is currently being recognized. However, due to the lack of suitable models to estimate biomass accumulation and growth rate in this region, the carbon sequestered by these formations is not included when computing the total carbon stocks in aboveground biomass in Mediterranean forest ecosystems, according to the IPCC guidelines. The aim of the present study is to develop equations to predict biomass accumulation and growth rate for the main shrub formations in the region of Andalusia (Southern Spain), using the fraction of canopy cover (FCC m ) and the average height of the shrub formations (H m ) as predictors. To build these models, more than 800 plots were inventoried using destructive sampling; the mean value found in the region for biomass accumulation and annual growth rate being 16.73 Mg ha−1 and 1.14 Mg ha−1 year−1, respectively. Heathers and big-size Cistaceae formations were the ones that presented the highest values of biomass accumulation (24.99 and 21.01 Mg ha−1, respectively), while the highest values for annual growth rate were achieved by Leguminous gorse shrubs and, again, big-size Cistaceae bushes (1.49 and 1.64 Mg ha−1 year−1, respectively). The carbon content for the main shrub species and formations in the area was also obtained. The developed models provide the opportunity to estimate shrub carbon stocks in Mediterranean forest management from easy-to-obtain variables, namely FCC m and H m . Moreover, the used shrub formations classifications and model structure allow their applicability to compute biomass accumulation and growth rate at regional and national level using as input data from the Spanish National Forest Inventory.

An up-to-date report on the distribution of Helianthemum lippii in Italy is provided also on the basis of the findings by Lojacono-Pojero in 1889 who reported the existence of a relic population in Sicily (southern Italy), which is now thought to be extinct because of human disturbance. Field excursions permitted to confirm the locus classicus reported by Lojacono-Pojero as the other Sicilian localities that have been reported more recently. Based on a careful analysis of the literature the presence of H. lippii is currently excluded in Calabria and Apulia. The main causes that threaten the habitat of H. lippii in Sicily are pointed out according to IUCN Threats Classification Scheme.

Camptopoeum (Camptopoeum) baldocki spec. nov., a new European panurgine bee species is described and diagnosed. It is currently known only from saltmarshes along the southern coast of the central and eastern Algarve, Portugal. Observations and analysis of scopal pollen loads suggest narrow oligolecty on the similarly halophilous Frankenia laevis (Frankeniaceae). In addition, the allotype male of the Portuguese endemic Flavipanurgus fuzetus Patiny is described and the pollen preferences of Flavipanurgus are reviewed with the addition of new data from Portugal. As a genus, Flavipanurgus species appear to be narrowly oligolectic on a range of flowers from the botanical families Caryophyllaceae, Cistaceae and Crassulaceae.

In animal‐pollinated hermaphroditic plants, optimal floral allocation determines relative investment into sexes, which is ultimately dependent on flower size. Larger flowers disproportionally increase maleness whereas smaller and less rewarding flowers favour female function. Although floral traits are considered strongly conserved, phylogenetic relationships in the interspecific patterns of resource allocation to floral sex remain overlooked. We investigated these patterns in Cistaceae, a hermaphroditic family. We reconstructed phylogenetic relationships among Cistaceae species and quantified phylogenetic signal for flower size, dry mass and nutrient allocation to floral structures in 23 Mediterranean species using Blomberg's K‐statistic. Lastly, phylogenetically‐controlled correlational and regression analyses were applied to examine flower size‐based allometry in resource allocation to floral structures. Sepals received the highest dry mass allocation, followed by petals, whereas sexual structures increased nutrient allocation. Flower size and resource allocation to floral structures, except for carpels, showed a strong phylogenetic signal. Larger flowered species allometrically allocated more resources to maleness, by increasing allocation to corollas and stamens. Our results suggest a major role of phylogeny in determining interspecific changes in flower size and subsequent floral sex allocation. This implies that flower size balances the male–female function over the evolutionary history of Cistaceae. While allometric resource investment in maleness is inherited across species diversification, allocation to the female function seems a labile trait that varies among closely related species that have diversified into different ecological niches., This study was supported by the Spanish‐funded projects REMEDINAL3eCM (S2013/MAE‐2719) and Ecometas excellence network (CGL2014‐53840‐REDT). ALT held a doctoral fellowship at Rey Juan Carlos University, Spain. BG received a Juan de la Cierva fellowship, and VGS a post‐doctoral fellowship (grant #2014/13899‐4) from São Paulo Research Foundation (FAPESP).

To date, only the three syntipic specimens of the Iberian endemism Strophosoma (Neliocarus) burdigalense (Escalera, 1928), were known, from Burgos (without more data). The species has been found again in the neighbouring province of Soria (Castilla and Leon), permitting the finding that its biological cycle is completed in Helianthemum hirtum (Cistaceae). The differential characters of the species are commented and biometrical data, images of the adult, aedeagus, its habitat and food plant are provided.

Authors briefly present some observations and reflections on the “seeding strategy” of some plant species (Cistus spp., Cistaceae, in particular) in the Mediterranean area in relation to fire: they conclude it does not seem a specific adaptation to wildfires (as suggested in a relevant part of the recent literature), but a generalistic adaptation to open, high energy and variable habitats with aleatory fluctuations. The difference is important both for theoretical and applied aspects, e.g. for the correct management of Cistus species and of other Mediterranean species and habitats. Fire is undoubtedly a major ecological factor in the Mediterranean area. Many botanists have postulated the relevant role of fire in the evolution of Mediterranean vegetation and flora, beginning with the milestone papers of Le Houerou (1973) and Naveh (1975) and up to the most recent works (e.g. Keeley et al., 2011; Pausas and Schwilk, 2012). Consequently, pyrophytism (or adaptation of plants to wildfires) in Mediterranean vegetation and flora has been the subject of many studies and papers, with many consequences also

Crocanthemum greenei (B.L.Rob.) Sorrie (Cistaceae), a perennial sub-shrub, was measured as part of a demographic monitoring effort on Santa Catalina Island, California, USA (hereafter, Catalina). Introduced ungulate browsers remain present on Catalina. Consequently, many palatable plant taxa on the island are subject to and putatively limited by top-down browsing forces. Historically, introduced ungulates have also been present on each island throughout the range of Crocanthemum greenei. Habitat conservation work, resulting in the construction of ungulate exclosures on Catalina, has now allowed us to measure individuals in their mature, non-browsed form. The published value for Crocanthemum greenei stem (height) is usually 15–30 cm. While the original description hints at a greater potential size, recent descriptions appear to be influenced by observations made during the decades when plants would have been impacted by introduced ungulate herbivores. Here we present stem measurements of 81 adult individuals, with a median of 49 cm and an interquartile range of 42–56 cm. These measurements suggest an expanded stem (height) range of 15–60 cm better describes the taxon and shed light on the historical impacts of invasive ungulates across the islands and those continuing on Catalina.

Extreme specialization is a common phenomenon in antagonistic biotic interactions but it is quite rare in mutualistic ones. Indeed, bee specialization on a single flower species (monolecty) is a questioned fact. Here, we provide multiple lines of evidence on true monolecty in a solitary bee (Flavipanurgus venustus, Andrenidae), which is consistent across space (18 sites in SW Iberian Peninsula) and time (three years) despite the presence of closely related congeneric plant species whose flowers are morphologically similar. The host flower (Cistus crispus, Cistaceae) is in turn a supergeneralist, visited by at least 85 insect species. We uncover ultraviolet light reflectance as a distinctive visual cue of the host flower, which can be a key mechanism because bee specialization has an innate basis to recognize specific signals. Moreover, we hypothesized that a total dependence on an ephemeral resource (i.e. one flower species) must lead to spatiotemporal matching with it. Accordingly, we prove that the bee's flight phenology is synchronized with the blooming period of the host flower, and that the densities of bee populations mirror the local densities of the host flower. This case supports the `predictable plethora' hypothesis, that is, that host-specialization in bees is fostered by plant species providing predictably abundant floral resources. Our findings, along with available phylogenetic information on the genus Cistus, suggest the importance of historical processes and cognitive constraints as drivers of specialization in bee-plant interactions.

Flower size is a key trait in the reproductive ecology of animal-pollinated plants. However, pollinator-mediated selection does not always modulate this trait and environmental conditions and/or antagonist interactions may favor smaller flowers. We evaluate the occurrence of a large-flowered family in a hot and dry Mediterranean environment, mediated by a cost-benefit balance and a male–female conflict. Large flowers have sizeable benefits in terms of pollination and reproductive success and pollinators mediate selection through male function, but female fitness is context-dependent. High floral production and maintenance costs and florivore incidence in large flowers limit female function, which counteracts pollinator-mediated selection. Large flowers are highly costly in the Mediterranean and flower size is mediated by a sexual conflict between the benefits of male function and the costs of the female one. However, a short floral longevity, occasional pollen limitation and selection through maleness keep the existence of large flowers in these environments.

Identifying the internal and external drivers of population dynamics is a key objective in ecology, currently accentuated by the need to forecast the effects of climate change on species distributions and abundances. The interplay between environmental and density effects is one particularly important aspect of such forecasts. We examined the simultaneous impact of climate and intraspecific density on vital rates of the dwarf shrub Fumana procumbens over 20 yr, using generalized additive mixed models. We then analyzed effects on population dynamics using integral projection models. The population projection models accurately captured observed fluctuations in population size. Our analyses suggested the population was intrinsically regulated but with annual fluctuations in response to variation in weather. Simulations showed that implicitly assuming variation in demographic rates to be driven solely by the environment can overestimate extinction risks if there is density dependence. We conclude that density regulation can dampen effects of climate change on Fumana population size, and discuss the need to quantify density dependence in predictions of population responses to environmental changes.

The Helianthemum origanifolium aggregate is widely diversified, with several subspecies distributed in the western Mediterranean basin. Populations from Morocco and Algeria have been usually referred either to H. origanigolium subsp. origanifolium or to H. origanifolium subsp. molle, a taxon usually considered an eastern Iberian endemic. Detailed studies of those Moroccan and Algerian populations revealed a unique combination of constant macro- and micromorphological traits, allowing the description of a new subspecies to accommodate them, H. origanifolium subsp. africanum. Some floral and fruit characters, together with SEM observations of the indumentum, were found to be reliable and diagnostic for differentiating the new taxon with regard to the other subspecies of the aggregate. Data on the ecology and distribution are also reported for this new subspecies, as well as affinities and divergences with other closely allied taxa are also discussed. This work was partly supported by the research grants ACIE13-01 and ACIE14-01 from Universidad de Alicante (Spain).

Plant–animal interactions imply costs and benefits with net balance depending on interacting species and ecological context. Ungulates, in particular, confer costs (e.g., plant leaf consumption, flower bud predation) and benefits (e.g., plant overcompensation, seed dispersal) to plants. Magnitude of costs and benefits may be altered by habitat management or ecological conditions favoring high density ungulate populations. Little is known however on whether plant costs or benefits predominate over the years, or the long-term outcomes of plant-animal interactions in habitat types sustaining high density ungulate populations. We investigated how high density ungulate populations alter plant costs and benefits by quantifying ungulate long-term effects on the shrub Cistus ladanifer (Cistaceae) individual size, seed weight and number, seed bank, and population density, through a 12-year ungulate exclusion experiment in a Mediterranean scrubland. We monitored plant size and flower buds in plants exposed or protected from ungulates and number of developed capsules and seeds consumed (potential seed dispersal) by ungulates during three reproductive seasons. We found that ungulates negatively affected shrub size and led to a dramatically decline of shrub reproductive structures and seed production, affecting the plant reproductive cycle. Number of buds was 27 times higher and number of developed seed 5 times higher in ungulate-excluded as compared to ungulate-exposed plots. After 9 years of ungulate exclusion, the C. ladanifer seed bank was 2.6 times higher in ungulate-excluded plots. The population density of C. ladanifer was 4 times higher in ungulate-excluded plots. Our long-term experiment showed that high density ungulate populations can alter plant-animal interactions by reducing plant benefits and increasing plant costs.

This paper addresses section Theobromina within the genus Hebeloma (Agaricales). We recognise seven European species within this section, three of which are de- scribedasnew:Hebelomaalboerumpens,H.griseopruinatum and H. parvicystidiatum. The first two of these species appear to be ectomycorrhizal with Cistaceae: Cistus and Helianthemum. Hebeloma parvicystidiatum is more likely to be in mycorrhizal association with Quercus spp. We also provide a key to the European species within sect. Theobromina and an updated key of known Hebeloma asso- ciates of Cistus. Molecular analyses based on multiple loci further illustrate the distinctness of the newly described taxa and provide molecular evidence, supporting the morphologi- cal evidence, for the relationship that exists among species of this section. The ITS is the only one from the sequenced loci that, alongside with morphology, distinguishes among all of the speciesofsect.Theobromina. Thesection gains mostofits molecular support from the MCM7 locus, followed by RPB2.

Summary Fumana laevis (Cav.) Pau. is reported for the Flora of the Maltese islands. An account on the taxonomy of Fumana laevis in relation with the other three Fumana spp. occurring in Malta is presented and summarised in an identification key. Notes on its local and general distribution, habitat and local status is given.

Helianthemum caput-felis is an endangered plant species growing in fragmented habitats in the western Mediterranean basin. Reproductive traits, breeding system and pollinator assemblage were studied in its largest known (mainland) European population to improve knowledge on the reproductive biology of the species. Hand-pollination experiments were carried out to determine the breeding system. Generalized linear mixed models (GLMM) were fitted to the data to evaluate the effect of treatment on fruit set and number of viable seeds per fruit. We also investigated the composition of the pollinator assemblage by direct observations, and studied their pollen load. Results were then compared to previous studies conducted in an island population. In the studied population, H. caput-felis is partially self-compatible, but mostly an outbreeder species, since outcrossed flowers produced many more fruits and seeds than self-pollinated ones. Conversely, pollination treatments did not affect reproductive output in the island population. We also found several differences between island and mainland composition of floral visitors, as it was expected. The study of pollen loads revealed that insects were mostly visiting H. caput-felis. Despite the low capacity to produce fruits with self-pollination, H. caput-felis presented no reproductive limitations in its main inland population. Reproductive characteristics along with differences among populations should be taken into account for adequate management and conservation practices. This work was partly carried out in the frame of project GVPRE/2008/197 (Generalitat Valenciana, Spain).

In a recent study (American Journal of Botany 97: 730-737), we described the first case of a tripartite association in natural conditions among a holoparasitic plant (Cytinus), its host Cistaceae species, and mycorrhizal fungi at an anatomical level. In a letter to the editor, Brundrett (American Journal of Botany 98: 595-596) commented on our manuscript and questioned our conclusions, arguing that they are not adequately supported by the data. We reject this point of view and believe that the controversy has arisen because of the parasitic way of life of Cytinus. We maintain and demonstrate that there is enough evidence in the data that we presented to confirm the existence of mycorrhizal associations in the Cytinus-Cistaceae complex, supporting the functionality of the tripartite association. Most holoparasitic plants have been considered as nonmycorrhizal. However, it is not advisable to be categorical in drawing conclusions about the mycorrhizal status of a group of plants that has not been fully studied.

Helianthemum motae Sanchez-Gomez, J.F. Jimenez & J.B. Vera (Cistaceae) is described as a new species from southeastern Spain. It belongs to the section Lavandulaceum and appears to be close to H. syriacum. The diagnostic characters, a phylogenetic study as well as a full description, a dichotomic key, and illustrations are provided. Both macromorphological and micromorphological characters are discussed and compared with the morphologically related species. The conservational status of H. motae is discussed.