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Although most plants depend on different animals for pollination and seed dispersal, sometimes the same animal species provides both functions, being thus involved in what has been termed a ‘double mutualism’. Very little is known on the effectiveness of such species as both pollinators and seed dispersers, and even less on the intraspecific level differences at the contribution of an animal mutualist to plant fitness. In this study, we focused on the interaction between a Mediterranean shrub Cneorum tricoccon (Rutaceae) and the Balearic lizard Podarcis lilfordi (Lacertidae) in order to assess: 1) the role of this lizard as a legitimate pollinator of the plant (previously thought to be exclusively insect‐pollinated), and 2) the intraspecific variation in the use of flowers and fruits by lizard individuals, comparing males, females and juveniles. We further evaluated whether lizards show different fruit size selection depending on sex and age, with potential consequences for seed germination. Lizards visited more flowers and selected more hermaphrodite flowers than insects did, leading to relative increases in both fruit and seed set. Interestingly, female and juvenile lizards were the main flower visitors, whereas males were the main fruit consumers. Males selected the largest fruits (bearing the largest seeds) though this did not increase seed germination, which was only ca 15%. We concluded that P. lilfordi acts as a legitimate pollinator of C. tricoccon and, thus, confirm that this system constitutes a new case of double mutualism, especially common in island compared to mainland systems. Moreover, our findings show a large variation among conspecific individuals in their role as either pollinators or seed dispersers, with potential implications for plant reproductive success. Our study, therefore, highlights the importance of evaluating plant–animal interactions at the intraspecific level, and calls for more in‐depth studies on the consequences of such intraspecific variation., This work is framed within projects CGL2013‐44386‐P and CGL2017‐88122‐P, and FF is funded by a PhD fellowship (BES‐2014‐068207), all financed by the Spanish Government.

Almond trees are one of the most important crops in the Balearic Islands. The pollination of almonds is limited to the activity of insects, and cross-pollination is necessary for fruit development. Currently, honey bees and wild bee populations are declining considerably due to multiple causes, such as the use of pesticides, diseases and habitat loss. An alternative to increase the almond production is the use of commercial pollinators. In this long-term (3 years) study, the effect of the introduction of Bombus terrestris colonies on almond production was evaluated in two orchards. Two experimental designs were carried out to study the best management of this pollinator. For 2 years, all bumble bee colonies were placed in the middle of the plot and during the last year, the bumble bee colonies were distributed homogenously in the plot. Fruit set and the foraging behaviour of bumble bees during the blossoming period was determined, and the effect of different environmental variables on the visitation rate of bumble bees was assessed by means of a generalized linear mixed model (GLMM). Moreover, for the first time, the spatial distribution of fruit set was evaluated. Our results show that fruit set was significantly higher in the fields where B. terrestris had been introduced than in the control plots. This increased production resulted in a positive economic balance for the farmer. Moreover, bumble bees showed to prefer trees in a southwest orientation that were close to their colony. The activity of bumble bees showed to be significantly influenced by wind speed (the higher the speed the more flowers are visited by B. terrestris) and time after flowering (visitation rate decreased with days after flowering). In order to improve its management and obtain the highest possible almond production, it is important to understand the activity and behaviour of this pollinator., The authors would like to thank farmers of Son Marrano and Son Pons to have allowed us to realize the field work in their farms, and Fons de Garantia Agrària i Pesquera de les Illes Balears (FOGAIBA) BIA 7/17 to had funded this experiment. Elsevier Language Editing Services for editing this manuscript. Also, to the reviewers and the editor for their comments with which the paper has improved. Special thanks to students of the Laboratory of Zoology for their collaboration in the field.

Islands harbor a considerable portion of global biodiversity and endemic biota, and also are the recipients of the largest proportional numbers of alien invaders. Such invaders may jeopardize the performance of native species, through either their direct or indirect effects. In this study, we investigated the reproductive ecology of the endemic scrambling perennial herb Canarina canariensis in remnants of the former thermosclerophyllous woodland of Tenerife (Canary Islands), assessing how two widespread alien invasive species, the honeybee (Apis mellifera) and the black rat (Rattus rattus), affect its reproductive success. Apis mellifera visits its flowers whereas the black rat consumes both its flowers and fruits. Here, we compared the pollination effectiveness of different animal guilds (vertebrates vs insects) by means of selective exclosures and determined the level of floral herbivory. Three bird species (Phylloscopus canariensis, Cyanistes teneriffae and Sylvia melanocephala), a lizard (Gallotia galloti) and two insects (A. mellifera and the butterfly Gonepteryx cleobule) were the main flower visitors. Phylloscopus canariensis was the most frequent visitor in the early flowering season whereas A. mellifera predominated in the flowers during mid and late flowering periods. Birds increased fruit set, whilst lizards and insects had a negligible effect. Rats consumed about 10% of the flowers and reduced fruit set to one third. Besides contributing little to plant reproduction, A. mellifera might interfere with bird pollination by depleting flowers of nectar. We conclude that both alien species can threaten C. canariensis reproduction and hence population sustainability in the thermosclerophyllous vegetation. Apis mellifera, in particular, may become especially detrimental if apiculture keeps expanding, or if this bee becomes active earlier in the season due to global warming., Julia Jaca was funded by a predoctoral fellowship from the Ministerio de Educación, Cultura y Deporte [FPU13/05880] and by the unemployment benefit from the Ministerio de Trabajo, Migraciones y Seguridad Social; and the study was framed within a project financed by the Ministerio de Economía, Industria y Competitividad [CGL2013-44386-P], granted to Anna Traveset.

[Premise] Opportunistic nectar‐feeders may act as effective pollinators; nonetheless, we still lack information on whether these opportunistic species differ in their pollination effectiveness from specialized nectarivorous vertebrates and insects. Many nectar specialists have coevolved with the plants on which they feed; therefore, we would expect higher pollination effectiveness in specialists than in opportunistic feeders. Here, we assessed quantity and quality components of pollination effectiveness in specialist and opportunistic vertebrate nectarivores and insects, focusing on three plants from the Seychelles: Thespesia populnea, Polyscias crassa, and Syzygium wrightii., [Methods] We determined the quantity component (QNC) of pollination effectiveness with pollinator observations, and the quality component (QLC) by measuring fruit and seed set resulting from single visits by each pollinator. To detect potential negative effects of invasive ants on native plant‐pollinator interactions, we classified pollinator visits (quantity component) as disturbed (>6 ants/30 min) vs. undisturbed., [Results] All focal plants were visited by insects, and vertebrate specialist and opportunist nectarivores, yet their pollination effectiveness differed. Flying insects were the most effective pollinators of T. populnea. The other two plants were most effectively pollinated by vertebrates; i.e., sunbirds (nectar specialists) in S. wrightii and Phelsuma geckos (nectar opportunists) in P. crassa, despite marked variation in QNC and QLC. Ant presence was associated with lower pollinator visitation rate in P. crassa and S. wrightii., [Conclusions] Our study highlights the importance of all pollinator guilds, including opportunist nectarivorous vertebrates as pollinators of island plants, and the vulnerability of such interactions to disruption by nonnative species., This work was part of projects CGL2013‐44386‐P and CGL2017‐88122‐P awarded to A. T. and financed by the Spanish Government. F. F. was funded by the Spanish Government with a Ph.D. fellowship (BES‐2014‐ 068207). C. N. K.‐B. was supported by funding from the German Research Foundation (DFG; KA 3349/2‐1 and KA 3349/3‐1).

Premise of the study Relying on floral traits to assess pollination systems has been shown to underestimate the ecological service that novel mutualisms can provide. Although vertebrates with opportunistic food habits are common on islands, usually feeding upon flowers of entomophilous species (ES), little is known about how effective they are as pollinators. In a previous study, we had reported that native insectivorous and frugivorous Galapagos birds commonly visit ES flowers, without assessing whether they act as pollinators. Here we investigate this by focusing on three typically ES (Cryptocarpus pyriformis, Waltheria ovata, Cordia lutea) and one mostly ornithophilous species (OS), Opuntia echios. Methods The quantitative component (QNC: the product of floral visit frequency and number of flowers contacted) and qualitative components (QLC: fruit and seed set, fruit length, and mass and proportion of seedling emergence) of pollination effectiveness of birds was compared with that of insects. Key results Birds were not quantitatively important pollinators compared to insects. However, selective exclusion experiments in the four plant species revealed that all qualitative components of fitness improved when both birds and insects visited the flowers. Our study is the first to confirm pollination effectiveness of ES by native opportunistic birds. Conclusions The Galapagos pollination systems are probably more generalized than previous data suggested and, given that ES dominate the flora of this archipelago, we argue that, contrary to expectation, birds might have an important role in maintaining the reproductive success and diversity of plant communities.

Autochthonous plant species are heavily threatened by the increasing spread of invasive insects. The spatial distribution of invasive species' hosts is likely to play a pivotal role in the establishment and further expansion of the invading species; more specifically, distance and density dependent (DDD) processes linked to plant spatial arrangement are crucial in determining susceptibility to attack, but they have usually been overlooked in invasive research. We take a spatially explicit approach to evaluate potential DDD processes in the interaction between Paysandisia archon, a tropical lepidopteran recently introduced in Spain, and the endemic Mediterranean dwarf palm (Chamaerops humilis) in Mallorca (Balearic Islands). Specifically, we used spatial marked point pattern analyses, which allowed testing whether DDD processes affect attack probability and intensity on three dwarf palm populations corresponding to three different invasion phases (i.e., infestation core, early expansion, and expansion front). Our approach also allowed evaluation of whether and how intrinsic palm traits (size, sex) alter the proneness to P. archon attack over a range of spatial scales. The occurrence and nature of DDD effects on C. humilis performance varied largely among localities. At the infestation core, our analyses revealed positive density dependence, i.e., reduced damage at high densities or proximity to conspecific neighbors. By contrast, the early expansion locality showed negative density dependence of C. humilis performance, whereas at the expansion front, there was no evidence of DDD effects. Larger palms were consistently more prone to P. archon attack than small ones up to scales of 50 m. We found no evidence that palm sex altered the probability of attack by P. archon. Our results highlight the importance of spatially explicit analyses for assessing invasive events and point to the need of early interventions and prioritizing management efforts on larger palms in order to guarantee the conservation of autochthonous dwarf palm populations., This work was financed by the Conselleria de Medi Ambient of the Balearic Government and coordinated by AT. AL was supported by a postdoctoral contract co-funded by the Regional Government of the Balearic Islands and the European Social Fund 2014–2020. JMF was funded by a Portuguese FCT Grant (IF/00728/2013).

The annual export of the Mediterranean seagrass ( Posidonia oceanica ) litter to adjacent beaches and coastal dunes was quantified by examining the fortnight evolution of seagrass beach-cast volume on two beaches in the NW Mediterranean (Son Real and Es Trenc, Mallorca Island, Spain) for two years and analyzing the wind speed and direction obtained from the closest Meteorological Spanish Agency surface weather stations. The decomposition stage of the deposits was examined by analyzing the total hydrolysable amino acids, its percentage distribution and derived degradation indexes. Prevalent winds exceeding 6 m s −1 , the coastline morphology and type of terrestrial vegetation determine the annual dynamics of the seagrass beach-cast. In the most protected beach (Son Real) the seagrass beach-cast remained nearly stationary during the two studied years while it exhibited wide annual fluctuations in the less protected one (Es Trenc). The amounts of P. oceanica wrack washed on Son Real and Es Trenc beaches, respectively, were estimated at 309 kg DW m coastline −1 yr −1 and 1359 kg DW m coastline −1 yr −1 . They supplied between 20 kg CaCO 3 m coastline −1 yr −1 and 47 kg CaCO 3 m coastline −1 yr −1 . Between 54% (Son Real) and 70% (Es Trenc) of seagrass beach-cast, respectively accounting for 1.5 kg N m coastline −1 yr −1 and 8.6 kg N m coastline −1 yr −1 , were annually exported from the beaches to adjacent dune systems. Our results reveal that Mediterranean seagrass meadows might be an important source of materials, including sand and nutrients, for adjacent terrestrial systems, able to support their functioning.

© 2017 The Authors., The effect of population size on population genetic diversity and structure has rarely been studied jointly with other factors such as the position of a population within the species’ distribution range or the presence of mutualistic partners influencing dispersal. Understanding these determining factors for genetic variation is critical for conservation of relict plants that are generally suffering from genetic deterioration. Working with 16 populations of the vulnerable relict shrub Cneorum tricoccon throughout the majority of its western Mediterranean distribution range, and using nine polymorphic microsatellite markers, we examined the effects of periphery (peripheral vs. central), population size (large vs. small), and seed disperser (introduced carnivores vs. endemic lizards) on the genetic diversity and population structure of the species. Contrasting genetic variation (H: 0.04–0.476) was found across populations. Peripheral populations showed lower genetic diversity, but this was dependent on population size. Large peripheral populations showed high levels of genetic diversity, whereas small central populations were less diverse. Significant isolation by distance was detected, indicating that the effect of long-distance gene flow is limited relative to that of genetic drift, probably due to high selfing rates (F = 0.155–0.887), restricted pollen flow, and ineffective seed dispersal. Bayesian clustering also supported the strong population differentiation and highly fragmented structure. Contrary to expectations, the type of disperser showed no significant effect on either population genetic diversity or structure. Our results challenge the idea of an effect of periphery per se that can be mainly explained by population size, drawing attention to the need of integrative approaches considering different determinants of genetic variation. Furthermore, the very low genetic diversity observed in several small populations and the strong among-population differentiation highlight the conservation value of large populations throughout the species’ range, particularly in light of climate change and direct human threats., Funding was provided by the Spanish Ministry for Innovation and Science with the grants Consolider Montes (CSD2008_00040) and VULGLO (CGL2010‐22180‐C03‐03) provided to FV, and the Minsitry of Economy and Competitiveness, grant ECOMETAS (CGL2014‐53840‐REDT) and Junta de Andalucía Excellence project to Juan Arroyo and AT.

Special Issue: Animal‐plant interactions., Lizards have been reported as important pollinators on several oceanic islands. Here we evaluate the potential role of Galápagos lava lizards (Microlophus spp.) as pollinators across their radiation. Over 3 years, we sampled pollen transport by 9 lava lizard species on the 10 islands where they are present, including 7 single‐island endemics. Overall, only 25 of 296 individuals sampled (8.4%) transported pollen of 10 plant species, the most common being Prosopis juliflora, Exodeconus miersii, Sesuvium sp. and Cordia leucophlyctis. At least 8 of these plant species were native, and none were confirmed as introduced to the archipelago. Despite the low overall proportion of individuals carrying pollen, this was observed in 7 of the nine lizard species, and on 8 of the ten main islands (Española, Fernandina, Floreana, Isabela, Marchena, Pinta, Santa Cruz and Santiago), suggesting that this is a widespread interaction. The results reported here support the potential role of lava lizards as pollinators across their radiation, although they may represent a relatively modest contribution when compared with birds and insects. However, we cannot discard that lizards may be ecologically significant for particular plant species and ecosystems given the specific climatic condition and functional diversity of each island., This study is framed within the projects CGL2013-44386-P and CGL2015-67865-P funded by the Ministerio de Economía y Competitividad (Spain). We thank the Charles Darwin Foundation, especially Diana Flores and Patricia Jaramillo and the Galápagos National Park, especially Galo Quezada and Carlos Vera (research permit numbers: PC-46-14, PC-20-15, PC-22-16), for offering us information and logistic support.

Despite it is widely accepted that intrapopulation variation is fundamental to ecological and evolutionary processes, this level of information has only recently been included into network analysis of species/population interactions. When done, it has revealed non-random patterns in the distribution of trophic resources. Nestedness in resource use among individuals is the most recurrent observed pattern, often accompanied by an absence of modularity, but no previous studies examine bipartite modularity. We use network analysis to describe the diet composition of the Balearic endemic lizard Podarcis lilfordi in 2 islets at population and individual levels, based on the occurrence of food items in fecal samples. Our objectives are to 1) compare niche structure at both levels, 2) characterize niche partition using nestedness and modularity, and 3) assess how size, sex, season, and spatial location influence niche structure. At population-level niche width was wide, but narrow at the level of the individual. Both islet networks were nested, indicating similar ranking of the food preferences among individuals, but also modular, which was partially explained by seasonality. Sex and body size did not notably affect diet composition. Large niche overlap and therefore possibly relaxed competition were observed among females in one of the islets and during spring on both islets. Likewise, higher modularity in autumn suggests that higher competition could lead to specialization in both populations, because resources are usually scarce in this season. The absence of spatial location influence on niche might respond to fine-grained spatio-temporally distribution of food resources. Behavioral traits, not included in this study, could also influence resource partitioning., This work is framed within projects CGL2017-88122-P and BFU2009-09359 financed by the Spanish Government.

Nocturnal pollination plays an important role in sexual plant reproduction but has been overlooked, partially because of intrinsic difficulties in field experimentation. Even less attention has received the effect of within-inflorescence spatial position (distal or proximal) on nocturnal pollinators of columnar plants, despite numerous studies examining the relationship between such position and reproductive success. Woody endemic Echium simplex possesses large erect inflorescences bearing thousands of flowers which are visited by a wide array of diurnal and nocturnal animals. In this study, we identified nocturnal visitors and compared their pollination effectiveness with that of diurnal pollinators in different inflorescence sections by means of selective exclosures in NE Tenerife (Canary Islands). Nocturnal visitors included at least ten morphospecies of moths (such as Paradrina rebeli and Eupithecia sp.), two coleopteran species (mainly Alloxantha sp.), neuropterans (Chrysoperla carnea), dictyopterans (Phyllodromica brullei), dermapterans (Guanchia sp.) and julidans (Ommatoiulus moreletii). In general, plants excluded from pollinators set less fruits than open-pollination (control) plants which set fruits homogeneously across sections. Diurnally pollinated plants set more fruit in their upper parts whereas nocturnally pollinated plants set fruit in both upper and bottom sections. We conclude that although the frequency and diversity of diurnal pollinators is far higher than that of nocturnal pollinators, both exhibit different foraging behaviour that generates complementary effects on the reproductive success of E. simplex., Julia Jaca was funded by a predoctoral fellowship from the Ministerio de Educación, Cultura y Deporte (FPU13/05880) and by the unemployment benefit from the Ministerio de Trabajo, Migraciones y Seguridad Social. The study was framed within a project financed by the Ministerio de Economía, Industria y Competitividad (CGL2017-88122-P) to Anna Traveset.

Habitat loss and fragmentation are recognized as affecting the nature of biotic interactions, although we still know little about such changes for reptilian herbivores and their hindgut nematodes, in which endosymbiont interactions could range from mutualistic to commensal and parasitic. We investigated the potential cost and benefit of endosymbiont interactions between the spur-thighed tortoise (Testudo graeca L.) and adult oxyurid nematodes (Pharyngodonidae order Oxyurida) in scrublands of southern Spain. For this, we assessed the association between richness and abundance of oxyurid species with tortoises’ growth rates and body traits (weight and carapace length) across levels of habitat loss (low, intermediate and high). Furthermore, by using an intrapopulation ecological network approach, we evaluated the structure and diversity of tortoise–oxyurid interactions by focusing on oxyurid species infesting individual tortoises with different body traits and growth rates across habitats. Overall, tortoise body traits were not related to oxyurid infestation across habitats. Oxyurid richness and abundance however, showed contrasting relationships with growth rates across levels of habitat loss. At low habitat loss, oxyurid infestation was positively associated with growth rates (suggesting a mutualistic oxyurid–tortoise relationship), but the association became negative at high habitat loss (suggesting a parasitic relationship). Furthermore, no relationship was observed when habitat loss was intermediate (suggesting a commensal relationship). The network analysis showed that the oxyurid community was not randomly assembled but significantly nested, revealing a structured pattern for all levels of habitat loss. The diversity of interactions was lowest at low habitat loss. The intermediate level, however, showed the greatest specialization, which indicates that individuals were infested by fewer oxyurids in this landscape, whereas at high habitat loss individuals were the most generalized hosts. Related to the latter, connectance was greatest at high habitat loss, reflecting a more uniform spread of interactions among oxyurid species. At an individual level, heavier and larger tortoises tended to show a greater number of oxyurid species interactions. We conclude that there is an association between habitat loss and the tortoise–oxyurid interaction. Although we cannot infer causality in their association, we hypothesize that such oxyurids could have negative, neutral and positive consequences for tortoise growth rates. Ecological network analysis can help in the understanding of the nature of such changes in tortoise–oxyurid interactions by showing how generalized or specialized such interactions are under different environmental conditions and how vulnerable endosymbiont interactions might be to further habitat loss., The Dirección General de Gestión del Medio Natural de la Junta de Andalucía (SGB/FOA/AFR) and the Delegación General de Medio Natural de la Comunidad Autónoma de la Región de Murcia (AUT/ET/UND/48/2010) granted permission to sample the tortoises and nematodes.

[Premise] The endangered shrub Daphne rodriguezii (Thymelaeaceae) is endemic to the Balearic island of Menorca, where fragmentation and severe population decline are ongoing threats to this taxon. We developed a set of microsatellite markers to analyze the fine-scale genetics of its few extant populations., [Methods and Results] Fifteen microsatellite markers were obtained through Illumina high-throughput sequencing and tested in two populations. Twelve of these loci showed no evidence of null alleles and were highly polymorphic, with a mean number of 8.3 alleles per locus. Levels of observed and expected heterozygosity ranged from 0.100 to 0.952 and from 0.095 to 0.854, respectively. Seven to nine of these loci were successfully amplified in five other Daphne species., [Conclusions] This set of markers provides a useful tool for investigating the factors driving fine-scale population structure in this threatened species, and it represents a novel genetic resource for other European Daphne species., J.C.I. was funded by a GRUPIN research grant from the Regional Government of Asturias (Ref.: IDI/2018/000151), and C.G.‐V. was supported by a Vicenç Mut postdoctoral fellowship awarded by the Govern de les Illes Balears–Conselleria d'Innovació, Recerca i Turisme. The study was funded by the Ministerio de Economía, Industria y Competitividad (project Islet‐Foodwebs CGL2017‐88122‐P).

Original Research Megafauna extinctions often lead to the disruption of plant-animal interactions, such as the seed-disperser mutualisms, which might entail severe consequences for plant populations and entire communities. Interestingly, the contemporary persistence of anachronistic plant species might be possible thanks to surrogate dispersers or seed dispersal “rescuers”. We know very little on how these relevant functional replacements are contributing to the performance of present-day plant-frugivore networks. The dwarf palm Chamaerops humilis L. is a Mediterranean endemism with fleshy fruits and typically dispersed by mammals. Despite its ecological importance and wide distribution in some of the Mediterranean islands, no information exists about its seed dispersal on these depauperated-fauna systems. In this study, we aim at identifying and quantifying the relative importance of introduced frugivores on the island of Mallorca (Balearic Islands), where no native terrestrial mammals exist. Specifically, we assess for the first time the seed dispersal effectiveness (SDE) for C. humilis on islands; we evaluate the quantitative component by fecal and regurgitation sampling surveys, and the qualitative component by means of seed germination experiments and seedling growth measures. Introduced goats (Capra hircus L.) and pine martens (Martes martes L.) were the local mammal fruit consumers of C. humilis identified in our study sites. Results suggest that goats are much more important quantitatively than pine martens, due to the high number of fruits handled in each foraging bout and their extremely high abundance on the island. However, pine marten-ingested seeds showed the highest final seedling emergence success and seedling growth, thus its qualitative contribution on C. humilis seed dispersal is higher than that of goats. Overall, SDE was almost 9-fold higher for goats than for pine martens. We conclude that these two non-native mammal species are effective seed dispersers of C. humilis in this and probably other Mediterranean islands, where humans led to the extinction of its native seed dispersers, as it was probably the case of the goat-like Myotragus balearicus in the Balearic Islands info:eu-repo/semantics/publishedVersion

Many Mediterranean beaches accumulate wrack of Posidonia oceanica L. leaves and rhizomes, creating an ecotone between marine and terrestrial systems that provides essential ecosystem services (e.g. shoreline protection, nutrient supply to adjacent dune systems). However, little is still known about the diversity of macrofauna that lives in this wrack and how its community structure varies in space and time. In this study, we assessed the variation in macrofauna composition on three Mallorcan beaches (Balearic Islands, Western Mediterranean Sea) characterized by different geomorphological features. For an entire year, we seasonally sampled P. oceanica wrack on each beach and identified the animal species living in them. We found up to 20 different species belonging to four functional groups: (A) detritivores/fungivores/scavengers, (B) phytophagous, (C) secondary consumers, and (D) tertiary consumers. Species composition and diversity indices showed strong differences among beaches associated with wrack dynamism. Both variables revealed also a significant temporal variation, with highest individual abundance in summer and lowest in winter. Detritivores were the most abundant group, and their contribution to community structure represented over 75% through the study period. Springtails, in particular, contributed the most (45%) to spatial and temporal changes in community structure, followed by the oligochaete Enchytraeus sp. (12.6%) and the sand hopper Talitrus saltator (4.6%). Our study builds towards a more robust understanding of these marine-terrestrial ecotones, demonstrating the existence of complex and dynamic macrofauna communities which can provide more ecological information about the food webs of these ecosystems and the services they provide., This work was funded by the Spanish Ministry of Economy and Competitiveness (MedShift, CGL2015-71809-P) and the Government of the Balearic Islands (Competitive Group of Excellence GAEM). MAJ acknowledges financial support from the JAE-Doc program (CSIC, co-funded by FSE). We are very grateful to “Finca Pública de Son Real” staff for letting us work in their area and providing logistic support. We also thank Clara Vignolo for help both in the field and lab. Different taxonomists kindly aided us with the identification of the taxa: Lluc García (Isopoda), Xavier Canyelles (Coleoptera), and Dr. Antonio García- Valdecasas and Dr. Ilse Bartsch (Acari). The study is framed within project CGL-2017-88122-P financed by the Spanish Ministry of Science and Innovation, Spain.

Oceanic island ecosystems favour the appearance of novel interactions as a consequence of their depauperate and disharmonic flora and fauna. We investigated Echium simplex, endemic to the Anaga Biosphere Reserve in NE Tenerife, Canary Islands, belongs to the Canarian bird–flower element. Along two flowering seasons, we studied the breeding system of E. simplex, identified the floral visitors and compared the pollination effectiveness of different animal guilds (insects versus vertebrates) by means of selective exclosures. E. simplex is self-compatible but selfing significantly reduced fruit set. The flowers were visited by five bird species (mostly Phylloscopus canariensis and Serinus canarius, but also Cyanistes teneriffae, Sylvia atricapilla and Sylvia melanocephala), a lizard species (Gallotia galloti) and over a hundred insect species (mainly hymenopterans and coleopterans). Flying insects increased fruit set whereas small flower dwellers (mostly beetles) decreased both fruit and seed set. Vertebrates had a negligible effect on reproductive success. We conclude that although the floral resources provided by E. simplex may be important to some vertebrate species, these do not appear to contribute to increase fitness of the plant, which was more dependent upon flying insects for fruit and seed set. We additionally found that plant reproductive structures are heavily damaged by feral goats, which threaten the maintenance of this Canarian endemic species., The study was funded by a predoctoral fellowship to Julia Jaca from the Ministerio de Educación, Cultura y Deporte [FPU13/05880] and framed within a project financed by Ministerio de Economía, Industria y Competitividad [CGL2013‐44386‐P] to Anna Traveset.

An increasing number of double mutualisms (i.e. two interacting species benefiting each other in two different functions, e.g. pollination and seed dispersal) have been reported, mainly from island ecosystems, although we still lack much information on how effective such species are in both processes. Here, we assessed the pollination effectiveness of a double mutualism between an ancient Mediterranean gymnosperm, Ephedra fragilis, and a lizard, Podarcis lilfordi. On the one hand, we assessed the lizard contribution to different fitness measures (seed set and germination success), relative to that of insects and the wind effect; on the other, we determined the lizards’ seed removal rate (i.e. the quantity component of seed dispersal effectiveness). In both processes, we further tested for differences in their contributions among male, female and juvenile lizards. Ephedra fragilis showed to be mostly anemophilous, lizards and insects playing only a minor role on seed set. However, lizards qualitatively contributed to pollination success, as seeds coming from lizard-pollinated cones germinated at higher rates than those pollinated by wind or insects, although this was detected only for small seeds (, Our study provides evidence that the relationship between the Balearic endemic lizard Podarcis lilfordi and the plant Ephedra fragilis constitutes one of the five pollination/seed dispersal double mutualisms found around the World between an animal and a gymnosperm, besides representing the first double mutualism reported in the Mediterranean region. Although P. lilfordi does not contribute substantially to the pollination of E. fragilis, it does play a crucial role in the dispersal process. In turn, by means of the pollination drops and the fleshy cone scales, the plants feed its only seed disperser in this islet.

The disappearance of native seed dispersers due to anthropogenic activities is often accompanied by the introduction of alien species, which may to some extent replace the ecological service provided by the extinct ones. Yet, little empirical evidence exists demonstrating the evolutionary consequences of such alien “replacement.” Here, we document the conflicting selection exerted on seed size by two native lizards (Podarcis lilfordi and P. pityusensis) and an alien mammal species (Martes martes), all acting as legitimate seed dispersers of the Mediterranean relict Cneorum tricoccon. While lizards mostly exerted a negative directional selection on seed diameter, especially P. pityusensis, the much larger pine marten exerted positive selection on seed size. Our findings suggest that this among-disperser variation in the selection regimes, together with the occurrence of spatial variation in the presence of each seed disperser, help to create the geographical variation observed for seed size of C. tricoccon. To our knowledge, this is the first empirical evidence showing opposing selective pressures between native and alien species in the seed dispersal process in an invaded ecosystem., This study is framed within several projects funded by the Spanish Government (CGL2017‐88122‐P to AT, and CGL2L013‐47429‐P to AV). GEA was supported by Juan de la Cierva Programme.

Fern sporangia may provide an important source of energy for bird species, which in turn can act as potential dispersers of viable spores. This study reports the first case of fern spore dispersal by land birds. We document the consumption of fern sporangia and evaluate the potential spore dispersal by Galápagos finches on Santa Cruz Island. Overall, 18% of the 34 sampled individuals of three finch species, the Vegetarian Finch (Platyspiza crassirostris), the Small Ground Finch (Geospiza fuliginosa) and the Medium Ground Finch (Geospiza fortis), were found to disperse viable spores of two native ferns, Asplenium auritum and Asplenium feei., This research was undertaken within project CGL2017-88122-P funded by the Ministerio de Economía y Competitividad, Spain.

Many invasive plant species have fleshy fruits that are eaten by native frugivorous birds which disperse their seeds and may facilitate their germination, playing an important role in plant invasion success. The fleshy-fruited shrub Cotoneaster franchetii (Rosaceae) is an important invasive alien in the mountainous regions of central Argentina. To determine the role of avian frugivorous in fruit removal of this species, we conducted a frugivore exclusion experiment including bagged and unbagged branches in 75 plants of C. franchetii. At the end of the dispersal period, we compared the percentage of missing fruits (removed by birds + naturally dropped) in unbagged branches with the percentage of naturally dropped fruits in bagged branches. To assess whether any mechanism acting on seeds during their passage through bird guts (de-inhibition by pulp removal and/or seed scarification) affects seed germination of this species, we compared percentage and speed of germination among seeds obtained from faeces of the native frugivorous Turdus chiguanco, from manually de-pulped fruits, and from intact fruits. The percentage of missing fruits per shrub in unbagged branches was significantly higher than the percentage of naturally dropped fruits in bagged branches, suggesting that frugivorous birds play an important role in fruit removal of C. franchetii in the study area. Seeds from bird faeces and from manually de-pulped fruits germinated in higher percentage and faster than seeds from intact fruits. Germination percentage and speed of seeds from manually de-pulped fruits were significantly higher than those of gut-passed seeds. These results indicate that T. chiguanco increases and accelerates seed germination of C. franchetii through pulp removal, but not through seed scarification. Overall, our findings indicate that native frugivorous birds facilitate the dispersal and germination success of C. franchetii, likely playing an important role in its invasion throughout the mountainous region of central Argentina. Fil: Diaz Velez, Maria Celeste. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto Multidisciplinario de Biología Vegetal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto Multidisciplinario de Biología Vegetal; Argentina Fil: Sersic, Alicia Noemi. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto Multidisciplinario de Biología Vegetal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto Multidisciplinario de Biología Vegetal; Argentina Fil: Traveset, Anna. Consejo Superior de Investigaciones Cientificas. Instituto Mediterraneo de Estudios Avanzados; España Fil: Paiaro, Valeria. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto Multidisciplinario de Biología Vegetal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto Multidisciplinario de Biología Vegetal; Argentina

Alien invasive plant species can affect pollination, reproductive success and population dynamics of co-flowering native species via shared pollinators. Consequences may range from reproductive competition to facilitation, but the ecological drivers determining the type and magnitude of such indirect interactions remain poorly understood. Here, we examine the role of the spatial scale of invader presence and spatially contingent behavioural responses of different pollinator groups as potential key drivers, using the invasive Oxalis pes-caprae and the self-incompatible native annual Diplotaxis erucoides as a model system. Three treatments were assigned to native focal plants: (1) invader present at the landscape scale (hectares) but experimentally removed at the floral neighbourhood scale (pa); (2) invader present at both scales (pp); (3) invader absent at both scales (aa). Interestingly, we found pronounced spatially contingent differences in the responses of pollinators: honeybees and bumblebees were strongly attracted into invaded sites at the landscape scale, translating into native plant visitation facilitation through honeybees, while bumblebees almost exclusively visited Oxalis. Non-corbiculate wild bees, in contrast, showed less pronounced responses in foraging behavior, primarily at the floral neighborhood scale. Average heterospecific (Oxalis) pollen deposition onto stigmas of Diplotaxis was low (, This work was supported by a fellowship for prospective researchers by the Swiss National Science Foundation (Grant No. PBZHP3-131020).

Seed dispersal and seedling recruitment are crucial phases in the life cycle of all spermatophyte plants. The net contribution of seed dispersers to plant establishment is known as seed dispersal effectiveness (SDE) and is defined as the product of a quantitative (number of seeds dispersed) and a qualitative (probability of recruitment) component. In Galápagos, we studied the direct contribution to SDE (number of seeds dispersed and effect on seedling emergence) provided by the five island groups of frugivores (giant tortoises, lizards, medium-sized passerine birds, small non-finch passerine birds, and finches) in the two main habitats in this archipelago: the lowland and the highland zones, and found 16 vertebrate species dispersing 58 plant species. Data on frequency of occurrence of seeds in droppings and number of seeds dispersed per unit area produced contrasting patterns of seed dispersal. Based on the former, giant tortoises and medium-sized passerines were the most important seed dispersers. However, based on the latter, small non-finch passerines were the most important dispersers, followed by finches and medium-sized passerines. The effect of disperser gut passage on seedling emergence varied greatly depending on both the disperser and the plant species. Although the contribution to SDE provided by different disperser guilds changed across plant species, medium-sized passerines (e.g., mockingbirds) provided a higher contribution to SDE than lava lizards in 10 out of 16 plant species analysed, whereas lava lizards provided a higher contribution to SDE than birds in five plant species. While both the quantitative and qualitative components addressed are important, our data suggests that the former is a better predictor of SDE in the Galápagos archipelago.

The introduction of carnivorous mammals has led many native island species to extinction. In the Balearic Islands, the introduction of carnivorous mammals in the main islands has contributed to the extinction of the endemic lizard Podarcis lilfordi, which is an important seed disperser of many plant species. One of the introduced mammals, Martes martes (pine marten), is also known to disperse seeds and may replace the native lizard in this role in islands. However, little is known on the patterns of seed dispersal by these two different species and their possible implications for plant regeneration ability and population structure. We have compared the quality of seed deposition provided by both to the vulnerable Mediterranean shrub Cneorum tricoccon, by studying the pattern of seed distribution among sites generated by both dispersers as well as the suitability of those sites for seedling emergence and establishment. The study was carried out in two types of habitats (coastal shrublands and pine forests) and two islands (Mallorca and Dragonera). Lizards and mammals showed contrasting patterns of seed deposition and, where lizards were absent, mammals played their role as seed dispersers in pine forests but not in coastal shrublands. The lack of seed dispersal in the coastal shrubland seriously limits recruitment, by concentrating seeds under conspecifics and hindering colonisation opportunities, marking a long-term trend towards decline. The introduced predator replaces the native seed disperser in its service where it has gone extinct, although with important differences in the dispersal service provided. © 2013 Springer Science+Business Media Dordrecht., MCC was supported by an Isidro Parga Pondal Contract (Xunta de Galicia). AT was funded by Project CGL2010-18759BOS

© 2016 John Wiley & Sons Ltd, [Aim] The endangered Galápagos shrub snapdragon (Galvezia leucantha, Antirrhineae, Plantaginaceae) is restricted to small populations on four islands. In this study, we appraised results from taxonomy, genetics, phylogenetics, phylogeography and pollination ecology to reconstruct the evolutionary history of the genus Galvezia., [Location] Peru, continental Ecuador and Galápagos., [Methods] We sequenced the nuclear ribosomal ITS and two plastid regions, ndhF and ndhF-rpL32, to infer the origin of Galvezia and patterns of colonization to and across the Galápagos archipelago, based on Bayesian inference and statistical parsimony analyses. To investigate genetic diversity and differentiation within G. leucantha, we screened the genome of six populations and obtained 194 AFLP fingerprints. Autogamy tests and pollination network analyses were performed to evaluate the colonization potential and to investigate the structure of the pollinators’ assemblage of Galvezia., [Results] Relationships of seven nucleotide-substitution haplotypes and 11 nucleotide-substitution ribotypes of Galvezia revealed monophyly for the Galapagos species. Dating estimates indicated divergence of the insular Galvezia lineage in the Middle-Upper Pleistocene (0.66–0.09 Ma). In addition, distribution of genotypes (seven haplotypes, eight ribotypes) across the three continental species showed geographical differentiation, while low differentiation and distribution of G. leucantha. AFLP genetic diversity is relatively high (H = 0.109), but a low proportion of the total allelic variance is attributed to variation among subspecies/islands (H = 0.035, hierarchical AMOVA: 3.77% of total variance). The endemic bee (Xylocopa darwinii) accounted for 87.30% of the floral visits to G. leucantha., [Main conclusions] We inferred a single origin for an insular lineage that colonized the Galápagos Islands from northern Peru in the Pleistocene. Recent colonization of the archipelago, Pleistocene land bridges between islands and active gene flow promoted by X. darwinii may account for the low‐moderate genetic differentiation of G. leucantha subspecies. An unusual pollination shift from ornithophily (hummingbirds on the continent) to entomophily (Xylocopa in the Galápagos Islands) is supported., This research was supported by the Spanish Ministry of Science and Innovation through funding conceded to PV(CGL2009-10031) and a Juan de la Cierva fellowship to BG.

Questions How does salinity affect germination, seedling performance and survival of the critically endangered Limonium barceloi, compared to ruderal species invading its salt marsh habitat? Is salt application an effective management tool for controlling invaders and favouring endangered halophytes? Location Ses Fontanelles, Mallorca, Balearic Islands, Spain. Methods The germination of seeds of L. barceloi and nine ruderal species was tested in five salinity levels (0–37 g·L−1), and also in freshwater after a pre-treatment with seawater. The ruderal study species were: Asphodelus fistulosus, Avena barbata, Geranium molle, Hypochoeris achyrophorus, Oryzopsis miliacea, Plantago coronopus, Reichardia picroides, Sonchus asper and Sonchus tenerrimus. Two salt concentration levels (18 and 37 g·L−1) were applied twice monthly, weekly or twice weekly to examine the effect of salt irrigation schemes on the performance and seedling survival of L. barceloi compared to two of the principal species invading these salt marshes, S. tenerrimus and A. fistulosus. Results Germination of L. barceloi and most ruderal species was highest at low salinity (0–8 g·L−1). Seeds of L. barceloi, A. fistulosus and S. tenerrimus were also able to germinate at the highest salinity level, in contrast to the other species. Germination in freshwater after seawater pre-treatment was highest in L. barceloi (72 ± 6%). Seedling performance and survival of A. fistulosus and S. tenerrimus decreased with frequency and concentration of the salt treatment. An application of 37 g·L−1 salt twice a week reduced seedling survival of A. fistulosus and S. tenerrimus by 83.3 and 91.6%, respectively. In contrast, 100% of the L. barceloi seedlings survived and showed similar performance among treatments. Conclusions Limonium barceloi had higher seedling growth and survival than ruderal invaders when salinity levels were high (37 g·L−1). Nevertheless, at lower salinity, ruderal species may germinate and grow better and thus salinity level changes may represent a threat to the survival of this critically endangered species. Although further field testing is required, our experiments suggest that salt application could be an effective measure to protect L. barceloi and other endangered halophytic plant species from less salt-tolerant invaders.

Predispersalseedpredation (PSP) by insects was studied in a plant community of the VenezuelanCentral Plain (VCP). The main goal was to examine to what extent vegetation structure and fruit–seed attributes determined the incidence of PSP by insects at the community level. We studied a total of 187 species from 59 families, in five habitat types. The proportion of seed-predated plant species (N=89; 47.6% of the total) was explained by different factors such as the abundance of legume families, plant species richness, fruit dehiscence, seed biomass, and starch content. Coleoptera was the most diverse taxonomic order of insectseed predators, with Bruchidae and Curculionidae showing the largest number of genera and species, followed by Lepidoptera (Pyralidae). Bruchidae and Pyralidae, but not Curculionidae, tended to be separated according to vegetation attributes, such as vegetation structure and flowering and fruiting phenology. In addition, Bruchidae was associated with dehiscent fruits, legumes, epizoochory, and granivory, whereas Curculionidae was more related with indehiscent fruits and endozoochory, and Pyralidae with abiotic seed dispersal. Bruchid larvae tended to prey upon single seeds, whereas larvae of Lycaenidae usually preyed upon more than one seed per fruit; Curculionidae did not show any clear pattern on this. One larva developing in a single seed (Type I) was the most common type of PSP, closely followed by one or more larvae developing outside seeds within the fruit (Type II). Type III (seedpredation by adult insects inside the fruit) was the least common. Type I appears to occur most frequently in climbers, in the forest–savanna transition habitat, and in those cases in which insect adults emerged during the rainy season. By contrast, Types II and III tended to be associated with annual herbs, the lowest stratum (0.05–0.6 m), and disturbed areas. Moreover, Type I was usually found in samaras, drupes, and indehiscent fruits, whereas Type II and Type III were more commonly observed in capsules. Type I and Type II seeds differed in their nutritional composition, which in turn appears to affect seedpredation specialization. The number of insectseed predator species and plant richness per habitat were positively correlated. The number of seed predator species was associated with the abundance of trees and climbers because more complex plant life forms offer a large variety of resources, woody species frequently had fruits and seeds larger than herbaceous species, and trees and climbers showed the lowest values of host specificity. To our knowledge, this study is the first one that characterizes the community of predispersalseed predators and simultaneously evaluates ecological, morphological, and nutritional factors determining the groups of seed predators and how PSP occurred.

The main goal of the study was to shed light on the enigmatic world distribution of the Cneoraceae family, frequently, and until very recently, used as an example of the connection between the Mediterranean and the Caribbean floras. Specifically, we wanted to document the presence/absence of an endemic taxon, first collected in 1861 by Charles Wright from Cuba ( Cneorum trimerum ). During the last five years, we have visited all the localities where this taxon was originally collected, at Sierra Maestra in eastern Cuba, and revised all available herbarium material. Moreover, we checked all botanical and historical references, and also interviewed local people, to whom we showed pictures of the Mediterranean Cneorum . The conclusions of our survey are the following: i) nowadays, there are no Cneorum plants growing in their type localities in Cuba; ii) all the collected material from Cuba and deposited in herbaria by Wright, and also others studied by Carlquist, correspond unequivocally to C. tricoccon . The material collected by Ekman in 1922, and identified as C. trimerum , is sterile, and cannot be identified with certainty, however we think it is unlikely to be C. tricoccon; iii) this plant was probably introduced during the mid 19th century from Southern France, although it never become naturalized. The presence of French colonist settlements in coffee plantations in those areas where Cneorum was collected, together with the fact that the plant has medicinal properties, lead us to suggest that the colonist themselves were those introducing the plant in Cuba. Lastly, iv) Schoepfia stenophylla (Olacaceae) has also been recently misidentified as Cneorum tricoccon due to the resemblance of their leaves. Schoepfia stenophylla is an endemic species currently threatened and for which there is an in situ conservation plan. Such confusion between these species has contributed to the maintenance and spreading of the mistakes through the long taxonomic history of the presence of Cneorum in Cuba. Hence, we conclude that Cneorum can no longer be used as an example of biogeographical disjunction between the floras of the Mediterranean and the Caribbean.

In this study we show that spatial scale of investigation affects the estimated strength of competition for pollinators between an invasive and a native plant species. The effect of the invasive herb Oxalis pes-caprae on pollinator visits to the native herb Diplotaxis erucoides was studied when the invader was (1) totally present, (2) present on a large scale (of hectares) but absent on a small scale (of square metres), and (3) totally absent. No difference in number of pollinator visits to D. erucoides was found between treatment 1 and 3, i.e., between total presence and total absence of the invader. However, when the invader was removed in the small scale while still remaining in the large scale, a higher number of visits to the native was recorded. Our study thus shows the importance of incorporating multiple spatial scales to allow for investigation of hierarchical effects on competition for pollinators, and it suggests that small-scale studies of effects of invasive plants on pollinator visitation might risk overstating negative effects of the invader.

Since the success of an invasive species depends not only upon its intrinsic traits but also on particular characteristics of the recipient habitat, assessing the performance of an invader across habitats provides a more realistic analysis of risk. Such an analysis will not only provide insights into the traits related to invasiveness, but also the habitat character- istics that underpin vulnerability to invasion that, taken together, will facilitate the selection of management strategies to mitigate the invader’s effect. In the present study, we considered the Mediterranean basin islands as an excellent study region to test how the same invasive species perform in different habitats within a single island, and to scale up differences among islands with similar climate. We tested how the performance of three widespread plant invaders with clonal growth but contrasting life-history traits, a deciduous tree Ailanthus altissima, a succulent subshrub Carpobrotus spp., and an annual geophyte Oxalis pes-caprae, varied depending upon the species identity, habitat, and invaded island. The environmental parameters consid- ered were habitat type, elevation, species diversity in the invaded plot, and several soil traits (% C, % N, C/N, pH, and relative humidity). The study documents that the performance of these three important and wide- spread plant invaders is dependent mainly on species identity, and less upon the invaded island’s general features. Likewise, differences in performance among habitats were only significant in the case of Ailanthus, whereas Carpobrotus and Oxalis appear to perform equally well in different environments. Ailanthus thus appears to have a broader spectrum of invasiveness, being able to invade a larger number of habitat types. On the contrary, Carpobrotus spp. have not yet invaded habitats different from those where the species have been originally introduced and where they are still commonly spread by humans. Oxalis distribution is mainly related to agricultural activities and disturbed sites, and the total area infested by this geophyte may be more reflection of the extent of suitable habitats than of invasiveness or ecological impact. Our results confirm the potential for these species to significantly alter the functioning of ecosystems in the Mediterranean islands and highlight the risk to other islands not yet invaded

Question: Do specific environmental conditions affect the performance and growth dynamics of one of the most invasive taxa (Carpobrotus aff. acinaciformis) on Mediterranean islands? Location: Four populations located on Mallorca, Spain. Methods: We monitored growth rates of main and lateral shoots of this stoloniferous plant for over two years (2002–2003), comparing two habitats (rocky coast vs. coastal dune) and two different light conditions (sun vs. shade). In one population of each habitat type, we estimated electron transport rate and the level of plant stress (maximal photochemical efficiency Fv/Fm) by means of chlorophyll fluorescence. Results: Main shoots of Carpobrotus grew at similar rates at all sites, regardless habitat type. However, growth rate of lateral shoots was greater in shaded plants than in those exposed to sunlight. Its high phenotypic plasticity, expressed in different allocation patterns in sun and shade individuals, and its clonal growth which promotes the continuous sea...

The influence of invasive plant species on native community composition is well-documented, but less is known about underlying mechanisms. Especially scarce is knowledge about effects on biotic interactions such as relationships between native plants and their pollinators. In this study we investigate if pollen transfer from the invasive and highly pollen productive Carpobrotus spp. affects seed production and/or seed quality in three native species. We monitored pollinator movements and pollen loads on pollinators and native stigmas, and in a field pollination experiment we investigated the effect of invasive pollen on reproduction. Invasive pollen adhered to pollinators, pollinators switched from Carpobrotus spp. to natives, invasive pollen was transferred to native stigmas, and it affected seed production in one species. Although all possible steps for interference with seed production were found to be qualitatively taken, invasive pollen has probably little impact on the native community because the frequency of invasive pollen transfer to natives was low. However, pollination interactions may change with plant abundance and our study provides evidence that pollen transfer from Carpobrotus spp. to natives does occur and have the potential to affect seed production. We found the species identity of shared pollinators to be of importance, higher flower constancy and lower capacity of pollen adherence are likely to result in less invasive pollen transfer. © 2007 Elsevier Ltd. All rights reserved., This research was supported by a post-doc grant to A. Jakobsson from the Swedish Council for Forestry and Agricultural Research

A fundamental way in which animal-dispersed plants can influence the viability and distribution of dispersed seeds is through control of retention time in the guts of dispersers. Using two species of wild chilies and their dispersers, we examined how chemical and physical properties of fruits and seeds mediate this interaction. Capsicum chacoense is polymorphic for pungency, occurs in Bolivia, and is dispersed mostly by elaenias. Capsicum annuum is not polymorphic, occurs in Arizona (USA), and is dispersed mostly by thrashers. We first tested whether capsaicin, the substance responsible for the pungency of chilies, affects gut retention time of seeds in primary dispersers. Capsaicin slowed gut passage of seeds but did so in a manner that differed greatly between bird species because the constipative effects of capsaicin occurred only after an 80-minute time lag. Elaenias in Bolivia held only 6% of C. chacoense seeds for >80 minutes, whereas thrashers in Arizona held 78% of C. annuum seeds for >80 minutes. Next we examined the effects of retention time on seed viability and germination. Increased retention resulted in a greater proportion of seeds germinating in C. annuum, had no effects on non-pungent C. chacoense, and had negative effects on pungent C. chacoense. These divergent effects are explained by differences in seed coat morphology: seed coats of pungent C. chacoense are 10-12% thinner than those of the other two types of seeds. Thus, longer retention times damaged seeds with the thinnest seed coats. In C. annuum, seed viability remained high regardless of retention time, but germination increased with retention, suggesting a role for scarification. Thus, in C. annuum, fruit chemistry appears well matched with seed morphology and disperser physiology: capsaicin extends gut retention for most seeds, resulting in greater seed scarification and higher germination rates. Increased retention of pungent C. chacoense seeds is detrimental, but because the primary consumers have short retention times, capsaicin slows only a small proportion of seeds, minimizing negative effects. These results illustrate the importance of context in studies of fruit secondary metabolites. The same chemical can have different impacts on plant fitness depending on its morphological, physiological, and ecological context. © 2008 by the Ecological Society of America., This research was supported by grants from the National Science Foundation (DEB 0129168) and the National Geographic Society (CRE 7190-02)

We evaluated the structure of intrapopulation howler monkey-plant interactions by focusing on the plant species consumed by different sex and age classes in continuous and fragmented forests in southern Mexico. For this we used network analysis to evaluate the impact of fragmentation on howler population traits and on resource availability and food choice. A total of 37 tree and liana species and seven plant items (bark, immature fruits, flowers, mature fruits, immature leaves, mature leaves and petioles) were consumed, but their relative consumption varied according to sex and age classes and habitat type. Overall, adult females consumed the greatest number of plant species and items while infants and juveniles the lowest. For both continuous and fragmented forests, we found a nested diet for howler monkey-plant networks: diets of more selective monkeys represent subsets of the diets of other individuals. Nestedness was likely due to the high selectivity of early life stages in specific food plants and items, which contrasts with the generalized foraging behaviour of adults. Information on the extent to which different plant species and primate populations depend on such interactions in different habitats will help to make accurate predictions about the potential impact of disturbances on plant-animal interaction networks., This research was supported by grants from Consejo Nacional de Ciencia y Tecnología (CONACyT, CB2005-C01-51043, CB2007-79121 to JBM) and Universidad Nacional Autónoma de México (UNAM, IN206111 to JBM). The Posgrado en Ciencias Biológicas (UNAM) and CONACyT are greatly acknowledged for providing a doctorate scholarship to AMG and UNAM for a post-doctoral scholarship to APM. This manuscript was partially written while JBM was on sabbatical at IMEDEA and supported by DGAPA (UNAM).

Biodiversity loss, including the decrease of diversity of ecological interactions, is known to reduce the capacity of ecosystems to cope with the effects of global change. Here we assessed whether the distribution and survival of two declining relict plant species, Daphne rodriguezii and Cneorum tricoccon, were affected by the mutualism disruption with their only seed-disperser and by climatic and habitat conditions. Due to the lack of data on demographic rates, we used an indirect approach to test these hypotheses. We used presence-absence data as response variables and took two distinct sets of predictors (i.e. habitat and topoclimatic conditions), which were hypothesized to be the main determinants of the demographic rates under question. With these two datasets we fitted species distribution models by means of MaxEnt. Such models were later used to build Combined Species Distributions Models (CM). For each plant species, these CM allowed evaluating the role of both climatic and non-climatic factors, such as the mutualism disruption. Results showed that both climate and habitat conditions determined the current distribution of the two species at a landscape scale. Additionally, the mutualism disruption between C. tricoccon and its native seed-disperser affected plant distribution, moving it to areas where a new alien disperser was present. This alien disperser modified the pattern of habitat selection by plants in terms of habitat quality, reducing their presence in suboptimal areas, which might be the determinant for their survival. Our findings highlight the need of a better understanding of the role of mutualisms within natural communities in order to undertake appropriate conservation actions on threatened plant populations. Furthermore, acting on key factors affecting plant-disperser disruptions (e.g. controlling invasive species) might help to mitigate the effects of global change on declining relict plants., The study is framed within projects GAEM financed by the Balearic Government and CGL2013-44386-P financed by the Ministerio de Economía y Competitividad

The role of the most common land iguana (Conolophus subcristatus) in the Galápagos Islands as an effective seed disperser is explored in this study. A total of 5705 seeds of 32 plant species were identified from 160 scats, 4545 of which (80%) appeared visually undamaged. Germination trials of 849 seeds from 29 species revealed that at least 10 species remained viable after passing through the iguana's gut, although only a small proportion of those seeds (4%) germinated. In any case, we argue that C. subcristatus exerts an important role on the 7 Galapagos islands where it occurs because of its abundance and capacity to ingest and disperse seeds at long distances. Our results strongly suggest that the Galápagos C. subcristatus plays an important role as a seed disperser of not only of native species but also some introduced plants in the Galápagos Islands.

[Questions] Many studies report a decrease of native species richness in communities after plant invasion by exotic taxa, but the implications of species losses on community functional structure and ecosystem processes have been less explored. The questions addressed are: (1) what are the impacts of invasive plant taxa on the functional structure of the recipient community; and (2) are there links between such functional structure and ecosystem properties representing key ecosystem processes?, [Location] Costal habitats of Mallorca and Menorca, Balearic Islands, Spain., [Methods] In paired invaded and non-invaded plots, species frequencies and ecosystem properties related to biogeochemical cycles (soil C and N content, soil moisture and decomposition rate) were assessed. For every native species we obtained values of several functional traits (plant height, woodiness, N fixation and evergreenness). Then we calculated indices of community functional structure (community-weighted means for each trait; functional diversity: richness, evenness, divergence and dispersion; and functional redundancy) and compared them between paired invaded and non-invaded plots. The relationship with ecosystem properties was assessed using GLM., [Results] The spectra of trait distribution changed in the invaded assemblages towards a higher frequency of woodiness and evergreenness. Invaded communities showed decreased species richness, and lower functional richness, divergence, dispersion and redundancy. In contrast, ecosystem properties were scarcely sensitive to invasion. Functional redundancy was negatively related to soil N of invaded plots and soil moisture in the dry season, which suggests higher depletion of soil resources when several species share the same combination of trait values., [Conclusions] Carpobrotus invasion led to a functional homogenization within communities, which probably reduces their resilience. The functional structure of the community was more responsive to invasion than the ecosystem properties. The weak relationship between functional indices and ecosystem properties suggests that they may respond to invasion at different rates., This study was supported by projects CGL2015-65346-R and CGL2011˗16388/BOS of the Ministerio de Economía y Competitividad of Spain and POII10˗0179˗4700 of the Junta de Comunidades de Castilla˗La Mancha. Field sampling was funded by the 5th EU Framework project EPIDEMIE (Contract no. EVK2-CT-2000-00074). PCD acknowledges CONICYT (Ministry of Education, Chile Government) for granting her stay at Concepción University (Concurso de atracción de Capital Humano Extranjero-MEC programme) and support of the REMEDINAL3-CM MAE-2719 network (Comunidad de Madrid). AP was partially funded by the Institute of Ecology and Biodiversity, with the grants from the Ministry of the Economy ICM P05-002 and CONICYT PFB-23. AT acknowledges support from the LINCGLOBAL project between CSIC (Spain) and U. Católica de Chile. MV acknowledges support through the Severo Ochoa Program for Centers of Excellence in R+D+I (SEV-2012-0262)

Summary 1 Secondary seed dispersal occurs whenever a seed is dispersed in two or more different dispersal events, so that different dispersal agents (e.g. animal frugivores or invertebrates) contribute to different events. Three secondary seed dispersal systems, in which lizards and predatory birds participate, are studied in Lanzarote (Canary Islands). 2 Seeds from all three plant species studied (Lycium intricatum, Rubia fruticosa and Asparagus nesiotes) were found with the remains of lizards (Gallotia atlantica) when they appeared inside shrike (Lanius meridionalis) and kestrel (Falco tinnunculus) pellets. This suggests that these two avian predators might secondarily disperse these plants by incidentally ingesting seeds when they prey upon the frugivorous lizards. 3 Lycium and Rubia seeds dispersed by Falco had significantly thinner seed coats compared to the other treatments (Gallotia and Lanius) and to uningested seeds. Correlated with this result, seed hardness increased from Lycium (the softest coat), to Rubia (intermediate values), to Asparagus. 4 Seed viability was high in all three species, although both viability and germination were significantly reduced in seeds of Lycium and Rubia dispersed by Falco. 5 The seed distribution was markedly different, Gallotia being an important disperser in the open ground microhabitats, Lanius in hillocks and Falco in hills. Because all three plant species were present in all these microhabitats, our observations suggest that the dispersal of these plants might be associated with the differential use of the habitat by the different dispersers. 6 Synthesis: Contrary to some long-distance seed dispersal (LDD) paradigms, the complex seed dispersal systems we describe are common and affect an important number of seeds that are moved each year to particular microhabitats. Finally, the simultaneous ecological factors analysed in this study support the hypotheses that diplochory by double endozoochory could have played a more important role in LDD events than is currently recognized, both in recent volcanic areas (lowlands) and also probably in the colonization of other subtropical islands.

9 pages.-- Final full-text version of the paper available at: http://dx.doi.org/10.1016/j.ecolmodel.2007.03.027., Carpobrotus affine acinaciformis is one of the most harmful invasive plants in the Mediterranean basin. In this study, we built a numerical growth model containing simple, and essentially linear, growth rules that provided us a tool capable to analyse the complex non-linear behaviour observed in the dynamic growth of Carpobrotus spp. This model includes a set of ecologically relevant parameters such as: (1) the stolon elongation rate, setting the upper limit of the horizontal spreading of the clone, (2) the branching rate, closely related to the capacity of the plant to occupy the space forming a dense and complex network, (3) the branching angle, which is determinant in the efficiency of the space occupation during the growth process, (4) the node mortality, and (5) the node branching age. We found that young patches of Carpobrotus (50 years old; speed keeps oscillating around 0.27 m/year further on) with a dimension equal to the euclidean one (Df = d = 2). Regarding the implications for the management of Carpobrotus, it is remarkable that with few relevant parameters we have been able to reproduce the patch dynamics of this plant, which may indeed be quite useful to improve the yield prediction of Carpobrotus growth in coastal Mediterranean zones. Our modelling approach can further be extended to interdisciplinary problems in which dendritic patterns and branched structures are developed., This paper is framed within projects EPIDEMIE (Exotic Plant Invasions: Deleterious Effects on Mediterranean Island Ecosystems) a research project supported by the European Commission under the V Framework Program, and within projects FIS2004-00953, FIS2006-09966 and CGL2004-04884-C02-01/BOS supported by the Spanish Ministry of Education and Science.

Plants that flower for long periods are ideal organisms in which to test whether the timing and duration of flowering adjust to the seasonal timing of biotic and abiotic resources, and whether that influences their reproductive success. Hypericum balearicum is an evergreen endemic shrub that flowers all year round but mainly during spring and summer. We found that those plants inhabiting a locality with lower rainfall had the capacity to flower longer. During spring and summer, the reproduction of plants that flowered earlier depended on both pollinators and rainfall, whereas the reproduction of plants that flowered later depended more upon rainfall scarcity., Flowering phenology and synchrony with biotic and abiotic resources are crucial traits determining the reproductive success in insect-pollinated plants. In seasonal climates, plants flowering for long periods should assure reproductive success when resources are more predictable. In this work, we evaluated the relationship between flowering phenology and synchrony and reproductive success in Hypericum balearicum, a shrub flowering all year round but mainly during spring and summer. We studied two contrasting localities (differing mostly in rainfall) during 3 years, and at different biological scales spanning from localities to individual flowers and fruits. We first monitored (monthly) flowering phenology and reproductive success (fruit and seed set) of plants, and assessed whether in the locality with higher rainfall plants had longer flowering phenology and synchrony and relatively higher reproductive success within or outside the flowering peak. Secondly, we censused pollinators on H. balearicum individuals and measured reproductive success along the flowering peak of each locality to test for an association between (i) richness and abundance of pollinators and (ii) fruit and seed set, and seed weight. We found that most flowers (∼90 %) and the highest fruit set (∼70 %) were produced during the flowering peak of each locality. Contrary to expectations, plants in the locality with lower rainfall showed more relaxed flowering phenology and synchrony and set more fruits outside the flowering peak. During the flowering peak of each locality, the reproductive success of early-flowering individuals depended on a combination of both pollinator richness and abundance and rainfall; by contrast, reproductive success of late-flowering individuals was most dependent on rainfall. Plant species flowering for long periods in seasonal climates, thus, appear to be ideal organisms to understand how flowering phenology and synchrony match with biotic and abiotic resources, and how this ultimately influences plant reproductive success.

Aims Although biological invasions occur throughout the world, and some invaders are widespread in many habitats, few studies on the ecological impact of invaders have examined multiple sites. We tested how the impact of three widespread plant invaders changed depending on the identity of the species and the invaded island. We also tested whether relative species loss was lower in species-rich communities than in species-poor ones. Location We conducted floristic surveys and soil analyses in eight Mediterranean Basin islands: Crete and Lesbos (Greece), Sardinia (Italy), Corsica, Bagaud and Porquerolles (France), and Mallorca and Menorca (Spain). Methods We compared native species richness and diversity, proportion of life forms, soil percentage nitrogen, percentage organic carbon, C/N, and soil pH in nearby paired plots of 2 × 2 m: one control and one invaded by either the deciduous tree Ailanthus altissima, the succulent subshrubs Carpobrotus spp. or the annual geophyte Oxalis pes-caprae, across eight Mediterranean Basin islands. Results On average, the presence of invaders reduced species diversity, Carpobrotus spp. exhibiting the largest impact and Oxalis the least. However, the relative impact was island-dependent, and was positively but weakly associated with the species richness of the recipient community. Therophytes were the life form that experienced the largest decrease across islands. The effects of invasion on soil properties were very variable. Total N changed (increased) only in plots invaded by Ailanthus, significantly decreasing the C/N ratio. The presence of this tree increased soil pH, whereas the opposite was found in plots invaded by the other two species. Organic C increased in plots invaded by Ailanthus and Carpobrotus species. Main conclusions By conducting an analysis at multiple sites, we found that the three plant invaders had an impact on plant community structure not entirely concordant with changes in soil properties. The impacts depended on the identity of the species and of the invaded island, suggesting that impact of invaders is context-specific. The impact in terms of species loss was not lower in species-rich than in species-poor communities.

Thymelaea velutina (Thymelaeaceae) is a unique dioecious and heterocarpic shrub, each female producing both dry and fleshy fruits. It is endemic to the Balearic Islands (western Mediterranean) and has suffered an important regression in recent decades, especially due to habitat loss. It lives in coastal areas, mainly in dunes, but a few populations occur up to 1300 m in the mountains of Mallorca. In the present study, we examined its breeding system and pollen vectors, determined whether seed production is pollen-limited, and quantified the relative importance of different insect visitors at coastal and mountain sites. Selective exclusions showed no differences between populations in the relative importance of insects and wind as pollen vectors. Pollen was limiting at both populations, though not consistently in time. Flowers of T. velutina were visited mostly by generalist insects, and species composition and abundance of flower visitors varied both in space and time. The flowering period is about a month later in the mountain than in the dune, and this results in a higher frequency of insect visits to plants in the mountain due to the higher insect abundance; insects were also more diverse in the mountain, although they visited a proportionally lower number of flowers than in the dune. Despite this spatio-temporal variation in the frequency of insect visits and in the importance of a particular insect group, the ambophilous system in this species (previously described as anemophilous) ensures a relatively high fruit set at the two different sites. © Springer-Verlag 2005., This work was partially supported by projects PB97-1174 and BOS2001-0610 financed to A.T

Mixed-pollination systems may allow plants to achieve stable seed production when unpredictable conditions cause variation in the relative success of different pollination modes. We studied variation in time (two years) and space (in five populations, three from an island and two from mainland) in the pollination mode of Buxus balearica, an ambophilous (i.e. pollinated by wind and insects) and selfing species distributed in the Mediterranean Basin, by means of direct observations and experimental manipulations (bagging with different material). The relative importance of each pollination mode differed among populations; however, levels of selfing and wind pollination were similar between island and mainland. Flowers of B. balearica were visited only by generalist insects, and species composition and abundance of flower visitors varied both in space and time. Frequency of insect visits to plants were not higher in mainland than island populations, although insects on the mainland were more diverse, visited a proportionally greater number of flowers, and remained longer on the plants than insects on the island. Frequency of insect visits was negatively correlated with flowering synchrony (all populations pooled) and was found to increase seed set in one of the mainland populations (that with highest frequency of insect visits and highest flower visitation rate). Fruit and seed mass were found to be not affected by pollination mode. Scarcity of pollinators in the island seems to have an effect on the pollination mode, although the greatest variation in breeding system was found at a more local scale. Copyright © Ecography 2005., This work was supported by a FPU fellowship (Spanish Ministry of Education, MECD) granted to A. L. and by project BOS2001-0610 (Spanish Ministry of Science and Technology, MCyT) financed to A. T

Summary 1. This study shows the first results obtained concerning the relative importance and effect on the seeds of a particular plant species, Rubia fruticosa Ait. (Rubiaceae), of passage through the digestive tube of the three main groups of vertebrate seed dispersers: native reptiles and birds, and alien mammals. 2. Native birds and reptiles dispersed a high proportion of undamaged seeds of R. fruticosa . 3. In contrast, seeds defecated by the two alien mammals showed significantly greater proportions of damaged seeds. Seed viability was much higher for reptiles and birds than for mammals, which was consistent with germination results. 4. Ingested seeds had a significantly thinner coat than uningested control seeds. Seeds that had passed through native dispersers had a significantly thicker coat than those passed through one of the alien mammals (the squirrel). 5. Considering both quantitative and qualitative components of seed dispersal, we conclude that native reptiles and birds are the most important seed dispersers in this multidisperser system.

Plants can respond to climate change by either migrating, adapting to the new conditions or going extinct. Relict plant species of limited distribution can be especially vulnerable as they are usually composed of small and isolated populations, which may reduce their ability to cope with rapidly changing environmental conditions. The aim of this study was to assess the vulnerability of Cneorum tricoccon L. (Cneoraceae), a Mediterranean relict shrub of limited distribution, to a future drier climate. We evaluated population differentiation in functional traits related to drought tolerance across seven representative populations of the species’ range. We measured morphological and physiological traits in both the field and the greenhouse under three water availability levels. Large phenotypic differences among populations were found under field conditions. All populations responded plastically to simulated drought, but they differed in mean trait values as well as in the slope of the phenotypic response. Particularly, dry-edge populations exhibited multiple functional traits that favored drought tolerance, such as more sclerophyllous leaves, strong stomatal control but high photosynthetic rates, which increases water use efficiency (iWUE), and an enhanced ability to accumulate sugars as osmolytes. Although drought decreased RGR in all populations, this reduction was smaller for populations from the dry edge. Our results suggest that dry-edge populations of this relict species are well adapted to drought, which could potentially mitigate the species’ extinction risk under drier scenarios. Dry-edge populations not only have a great conservation value but can also change expectations from current species’ distribution models., This study was supported by a JAE-predoc fellowship from the LINCGlobal. Funding was provided by the Spanish Ministry for Innovation and Science with the grants Consolider Montes (CSD2008_00040), VULGLO (CGL2010-22180-C03-03), PATSIMON (CGL2010-18759) and REMEDINAL III (CM-S2013/MAE-2719)

© 2015 Vargas et al. Colonization across the Galápagos Islands by the carpenter bee (Xylocopa darwini) was reconstructed based on distribution of mitochondrial haplotypes (cytochrome oxidase II (COII) sequences) and haplotype lineages. A total of 12 haplotypes were found in 118 individuals of X. darwini. Distributional, phylogenetic and phylogeographic analyses suggest early colonization of most islands followed by historical isolation in two main groups: eastern and central-western islands. Evidence of recurrent inter-island colonization of haplotypes is largely lacking, despite strong flight capability and ecological amplitude of the species. Recent palaeogeographic data suggest that several of the current islands were connected in the past and thus the isolation pattern may have been even more pronounced. A contrast analysis was also carried out on 10 animal groups of the Galápagos Islands, and on haplotype colonization of seven animal and plant species from several oceanic archipelagos (the Galápagos, Azores, Canary Islands). New colonization metrics on the number of potential vs. inferred colonization events revealed that the Galápagos carpenter bee shows one of the most significant examples of geographic isolation.

© 2014 The Authors. Most flowering plants depend on animal pollination. Several animal groups, including many birds, have specialized in exploiting floral nectar, while simultaneously pollinating the flowers they visit. These specialized pollinators are present in all continents except Europe and Antarctica, and thus, insects are often considered the only ecologically relevant pollinators in Europe. Nevertheless, generalist birds are also known to visit flowers, and several reports of flower visitation by birds in this continent prompted us to review available information in order to estimate its prevalence. We retrieved reports of flower-bird interactions from 62 publications. Forty-six bird species visited the flowers of 95 plant species, 26 of these being exotic to Europe, yielding a total of 243 specific interactions. The ecological importance of bird-flower visitation in Europe is still unknown, particularly in terms of plant reproductive output, but effective pollination has been confirmed for several native and exotic plant species. We suggest nectar and pollen to be important food resources for several bird species, especially tits Cyanistes and Sylvia and Phylloscopus warblers during winter and spring. The prevalence of bird flower-visitation, and thus potential bird pollination, is slightly more common in the Mediterranean basin, which is a stopover to many migrant bird species, which might actually increase their effectiveness as pollinators by promoting long-distance pollen flow. We argue that research on bird pollination in Europe deserves further attention to explore its ecological and evolutionary relevance., LPS was supported by the grant SFRH/BD/77746/2011 of the Fundaçao para Ciência e Tecnologia (FCT). RH was supported by the grant SFRH/BPD/76037/2011 of the FCT and the Marie Curie Action FP7-PEOPLE-2012-CIG-321794

Summary 1 Seed germination characteristics are often modified after seeds are ingested by frugivores. Factors that are intrinsic either to the plant or to the frugivore’s digestive tract are responsible for the great variation observed in germination response. 2 Our objectives were to determine whether and how the seed germination patterns of five common western Mediterranean plant species are affected by seed passage through the guts of their major dispersers, and to elucidate the mechanism by which such patterns are changed. 3 We used captive birds (Turdus merula and Sylvia melanocephala) to obtain ingested seeds and compared their germination rate (speed) and germinability (final percent germination) with those of controls (uningested, pulp-removed seeds), controlling for seed age, size and source. Germination was monitored for 2 years in an experimental garden. We evaluated the possible changes in seed traits after ingestion by measuring weight and coat thickness, and by observing seed coat sculpture. 4 Rate of seed germination, but not germinability, changed in all species after gut treatment. The greatest effect was in Osyris, in which germination was much enhanced. A great acceleration of germination, which is likely to translate into a seedling size advantage, was also found in Asparagus. In the other three species tested, germination was slower for ingested than for control seeds. 5 For Rubus and Rubia seeds, we found a different germination response depending upon the frugivore species tested. A different degree of seed coat scarification caused by differences in gut retention time, chemical and/or mechanical abrasion probably account for such responses. 6 In three of the species (Osyris, Rubia and Phillyrea), seed weight decreased after gut treatment. Such weight loss was not caused by any change in coat thickness, but may have been because of the scarification and consequent alteration of the seed coat structure. 7 The five Mediterranean species studied germinate when rains are most likely to fall (mostly autumn and spring). The different speed of germination promoted by gut treatment within frugivores may increase the probability that seeds can recruit successfully at a given time and in a given place. 8 This study suggests that frugivores contribute to the heterogeneity in germination characteristics not only within plant populations but also within plant communities, each frugivore species having a particular effect on the seeds of each plant consumed.

Summary 1 The fruit-colour polymorphism of Myrtus communis, a common Mediterranean shrub, is examined. We investigate whether frugivores affect the maintenance of the polymorphism, whether morphs differ in germination or seedling growth, and whether passage through frugivores’ gut affects seed behaviour and seedling growth. 2 Blue berries are very similar in morphological traits and nutrient composition to the rarer white morph. Rates of fruit removal by birds, the main dispersers of this species, did not differ between morphs, suggesting that a colour preference is not involved in maintenance of the polymorphism. 3 Seeds of the two morphs showed the same germinability (final percentage germination) as well as similar rates of germination under controlled conditions (in growth chamber and greenhouse). Outdoors, seeds from blue berries tended to germinate slightly faster (which might give them an early advantage) but differences between morphs disappeared after several weeks of growth. 4 The germination of myrtle seeds is differently affected by passage through the digestive tract of birds, which appeared to significantly increase germinability, and of the carnivorous pine marten, Martes martes, which did not. The different seed retention time in the gut or the chemical composition of the food ingested along with the seeds may be responsible for such differences. 5 The detection of a positive effect of bird ingestion in outdoor conditions, but not in the growth chamber or in the greenhouse, demonstrates the importance of performing germination tests in the natural habitat. 6 Seed size, irrespective of morph, affects the response to passage through the bird gut. Large seeds, which are more rapidly ejected, and are thus less abraded, germinated at a similar speed to non-ingested controls. Variation in seed size within a single species should thus be considered in future studies.

Theidea that fecal material accompanying vertebrate-dispersed seeds at depositionsites plays an important role in enhancing seed germination and seedlingsurvival has, surprisingly, little empirical support. The present studyattemptsto experimentally test this hypothesis. We examined the effect that manurecomposition from brown bears (Ursus arctos), importantseeddispersers of Rubus spectabilis and Vacciniumovalifolium/alaskaense in the temperate rainforests of SoutheastAlaska, has on seedling emergence and growth of these two fleshy-fruitedspeciesin their natural habitat. The seeds of Rubus spectabilisshowed a significantly higher germination rate in manure composed of animalmaterial (mainly deer hair and bones) than in manure consisting of either fruitpulp or vegetation fiber and than in controls (potting soil with no manureadded). The final number of germinated Rubus seeds wassimilar between the animal material and the fruit pulp treatments, perhaps dueto similar water retention capacities, but was significantly higher than in thevegetation fiber treatment and the control. The germination patterns ofVaccinium seeds, in contrast, appeared to be unaffected bythe composition of manure in which they were embedded. Seedlings of bothspeciesgrew faster, and in the case of Rubus produced moreleaves,when manure consisted of animal material. A principal constituent of bone iscalcium phosphate, which may provide important minerals to growing plants.Manure containing vegetation fiber also enhanced seedling growth compared tomanure with fruit pulp or the control. For Rubus, manurewith fruit pulp did not affect seedling length significantly, although thenumber of leaves per seedling was greater in this treatment than in thecontrol. We conclude that the influence of frugivores on the final fate of seedsof fleshy-fruited plants appears to depend not only upon commonly consideredfactors such as distance of dispersal, treatment in the digestive tract, andlocation of deposition, but also on what material the dispersed seeds areembedded in, i.e., on what other food frugivores have consumed along with thefruits.

We observed wild banana slugs (Ariolimax columbianus) eating fruits of several Pacific Northwest plant species. Slime trails and direct observations indicated that slugs are capable of reaching the fruits of many wild plants. This motivated us to test the hypothesis that slugs may act as seed dispersers, provided that they defecate viable seeds. We fed captive slugs the fruits of Rubus spectabilis, R. discolor, Vaccinium ovatum, V. parvifolium, Gaultheria shallon and Disporum smithii to determine the effects of slug ingestion on seed germination. At least some seeds of each species germinated after the fruits were consumed by the slugs, but the effects on germination were species-specific. Seeds of Rubus spectabilis were less likely to germinate after passage through the guts of slugs, and we found significant evidence that the two fruit color morphs reacted differently over time. Disporum smithii seeds did not statistically differ in germination behavior between treatments, although the trend su...