Adaptation to hummingbird pollination is associated with reduced diversification in Penstemon

This work is licensed under a Creative Commons Attribution 4.0 International License.A striking characteristic of the Western North American flora is the repeated evolution of hummingbird pollination from insect‐pollinated ancestors. This pattern has received extensive attention as an opportunity to study repeated trait evolution as well as potential constraints on evolutionary reversibility, with little attention focused on the impact of these transitions on species diversification rates. Yet traits conferring adaptation to divergent pollinators potentially impact speciation and extinction rates, because pollinators facilitate plant reproduction and specify mating patterns between flowering plants. Here, we examine macroevolutionary processes affecting floral pollination syndrome diversity in the largest North American genus of flowering plants, Penstemon. Within Penstemon, transitions from ancestral bee‐adapted flowers to hummingbird‐adapted flowers have frequently occurred, although hummingbird‐adapted species are rare overall within the genus. We inferred macroevolutionary transition and state‐dependent diversification rates and found that transitions from ancestral bee‐adapted flowers to hummingbird‐adapted flowers are associated with reduced net diversification rate, a finding based on an estimated 17 origins of hummingbird pollination in our sample. Although this finding is congruent with hypotheses that hummingbird adaptation in North American Flora is associated with reduced species diversification rates, it contrasts with studies of neotropical plant families where hummingbird pollination has been associated with increased species diversification. We further used the estimated macroevolutionary rates to predict the expected pattern of floral diversity within Penstemon over time, assuming stable diversification and transition rates. Under these assumptions, we find that hummingbird‐adapted species are expected to remain rare due to their reduced diversification rates. In fact, current floral diversity in the sampled Penstemon lineage, where less than one‐fifth of species are hummingbird adapted, is consistent with predicted levels of diversity under stable macroevolutionary rates

Assessment of pollen rewards by foraging bees

This is the author accepted manuscript. The final version is available from Wiley via the DOI in this record.The removal of pollen by flower-visiting insects is costly to plants, not only in terms of production, but also via lost reproductive potential. Modern angiosperms have evolved various reward strategies to limit these costs, yet many plant species still offer pollen as a sole or major reward for pollinating insects. 2. The benefits plants gain by offering pollen as a reward for pollinating are defined by the behaviour of their pollinators, some of which feed on the pollen at the flower, while others collect pollen to provision offspring. 3. We explore how pollen impacts on the behaviour and foraging decisions of pollen-collecting bees, drawing comparisons with what is known for nectar rewards. This question is of particular interest since foraging bees typically do not ingest pollen during collection, meaning the sensory pathways involved in evaluating this resource are not immediately obvious. 4. Previous research focussed on whether foraging bees can determine the quality of pollen sources offered by different plant species, and attempted to infer the mechanisms underpinning such evaluations, mainly through observations of collection preferences in the field 5. More recent experimental research has started to focus on if pollen itself can mediate the detection of, and learning about, pollen sources and associated floral cues. 6. We review advancements in the understanding of how bees forage for pollen and respond to variation in pollen quality, and discuss future directions for studying how this ancestral floral food reward shapes the behaviour of pollinating insects

Pollination effectiveness of specialist and opportunistic nectar feeders influenced by invasive alien ants in the Seychelles

This is the author accepted manuscript. The final version is available from the Botanical Society of America via the DOI in this recordPremise: 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.Spanish GovernmentGerman Research Foundatio

Aquatic community response to volcanic eruptions on the Ecuadorian Andean flank: evidence from the palaeoecological record

Aquatic ecosystems in the tropical Andes are under increasing pressure from human modification of the landscape (deforestation and dams) and climatic change (increase of extreme events and 1.5 °C on average temperatures are projected for AD 2100). However, the resilience of these ecosystems to perturbations is poorly understood. Here we use a multi-proxy palaeoecological approach to assess the response of aquatic ecosystems to a major mechanism for natural disturbance, volcanic ash deposition. Specifically, we present data from two Neotropical lakes located on the eastern Andean flank of Ecuador. Laguna Pindo (1°27.132′S–78°04.847′W) is a tectonically formed closed basin surrounded by a dense mid-elevation forest, whereas Laguna Baños (0°19.328′S–78°09.175′W) is a glacially formed lake with an inflow and outflow in high Andean Páramo grasslands. In each lake we examined the dynamics of chironomids and other aquatic and semi-aquatic organisms to explore the effect of thick (> 5 cm) volcanic deposits on the aquatic communities in these two systems with different catchment features. In both lakes past volcanic ash deposition was evident from four large tephras dated to c.850 cal year BP (Pindo), and 4600, 3600 and 1500 cal year BP (Baños). Examination of the chironomid and aquatic assemblages before and after the ash depositions revealed no shift in composition at Pindo, but a major change at Baños occurred after the last event around 1500 cal year BP. Chironomids at Baños changed from an assemblage dominated by Pseudochironomus and Polypedilum nubifer-type to Cricotopus/Paratrichocladius type-II, and such a dominance lasted for approximately 380 years. We suggest that, despite potential changes in the water chemistry, the major effect on the chironomid community resulted from the thickness of the tephra being deposited, which acted to shallow the water body beyond a depth threshold. Changes in the aquatic flora and fauna at the base of the trophic chain can promote cascade effects that may deteriorate the ecosystem, especially when already influenced by human activities, such as deforestation and dams, which is frequent in the high Andes

Sporormiella as a tool for detecting the presence of large herbivores in the Neotropics

The reliability of using the abundance of Sporormiella spores as a proxy for the presence and abundance of megaherbivores was tested in southern Brazil. Mud-water interface samples from nine lakes, in which cattle-use was categorized as high, medium, or low, were assayed for Sporormiella representation. The sampling design allowed an analysis of both the influence of the number of animals using the shoreline and the distance of the sampling site from the nearest shoreline. Sporormiella was found to be a reliable proxy for the presence of large livestock. The concentration and abundance of spores declined from the edge of the lake toward the center, with the strongest response being in sites with high livestock use. Consistent with prior studies in temperate regions, we find that Sporormiella spores are a useful proxy to study the extinction of Pleistocene megafauna or the arrival of European livestock in Neotropical landscapes

Revealing the last 13,500 years of environmental history from the multiproxy record of a mountain lake (Lago Enol, northern Iberian Peninsula)

This is the author's accepted manuscript. The final publication is available at Springer via http://dx.doi.org/10.1007/s10933-009-9387-7.We present the Holocene sequence from Lago Enol (43°16′N, 4°59′W, 1,070 m a.s.l.), Cantabrian Mountains, northern Spain. A multiproxy analysis provided comprehensive information about regional humidity and temperature changes. The analysis included sedimentological descriptions, physical properties, organic carbon and carbonate content, mineralogy and geochemical composition together with biological proxies including diatom and ostracod assemblages. A detailed pollen study enabled reconstruction of variations in vegetation cover, which were interpreted in the context of climate changes and human impact. Four distinct stages were recognized for the last 13,500 years: (1) a cold and dry episode that includes the Younger Dryas event (13,500–11,600 cal. year BP); (2) a humid and warmer period characterizing the onset of the Holocene (11,600–8,700 cal. year BP); (3) a tendency toward a drier climate during the middle Holocene (8,700–4,650 cal. year BP); and (4) a return to humid conditions following landscape modification by human activity (pastoral activities, deforestation) in the late Holocene (4,650–2,200 cal. year BP). Superimposed on relatively stable landscape conditions (e.g. maintenance of well established forests), the typical environmental variability of the southern European region is observed at this site.The Spanish Inter-Ministry Commission of Science and Technology (CICYT), the Spanish National Parks agency, the European Commission, the Spanish Ministry of Science, and the European Social Fund

Revealing the last 13,500 years of environmental history from the multiproxy record of a mountain lake (Lago Enol, northern Iberian Peninsula)

This is the author's accepted manuscript. The final publication is available at Springer via http://dx.doi.org/10.1007/s10933-009-9387-7.We present the Holocene sequence from Lago Enol (43°16′N, 4°59′W, 1,070 m a.s.l.), Cantabrian Mountains, northern Spain. A multiproxy analysis provided comprehensive information about regional humidity and temperature changes. The analysis included sedimentological descriptions, physical properties, organic carbon and carbonate content, mineralogy and geochemical composition together with biological proxies including diatom and ostracod assemblages. A detailed pollen study enabled reconstruction of variations in vegetation cover, which were interpreted in the context of climate changes and human impact. Four distinct stages were recognized for the last 13,500 years: (1) a cold and dry episode that includes the Younger Dryas event (13,500–11,600 cal. year BP); (2) a humid and warmer period characterizing the onset of the Holocene (11,600–8,700 cal. year BP); (3) a tendency toward a drier climate during the middle Holocene (8,700–4,650 cal. year BP); and (4) a return to humid conditions following landscape modification by human activity (pastoral activities, deforestation) in the late Holocene (4,650–2,200 cal. year BP). Superimposed on relatively stable landscape conditions (e.g. maintenance of well established forests), the typical environmental variability of the southern European region is observed at this site.The Spanish Inter-Ministry Commission of Science and Technology (CICYT), the Spanish National Parks agency, the European Commission, the Spanish Ministry of Science, and the European Social Fund

Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon

The nature and extent of pre-Columbian (pre-1492 AD) human impact in Amazonia is a contentious issue. The Bolivian Amazon has yielded some of the most impressive evidence for large and complex pre-Columbian societies in the Amazon basin, yet there remains relatively little data concerning the land use of these societies over time. Palaeoecology, when integrated with archaeological data, has the potential to fill these gaps in our knowledge. We present a 6,000-year record of anthropogenic burning, agriculture and vegetation change, from an oxbow lake located adjacent to a pre-Columbian ring-ditch in north-east Bolivia (13°15’44” S, 63°42’37” W). Human occupation around the lake site is inferred from pollen and phytoliths of maize (Zea mays L.) and macroscopic charcoal evidence of anthropogenic burning. First occupation around the lake was radiocarbon dated to ~2500 years BP. The persistence of maize in the record from ~1850 BP suggests that it was an important crop grown in the ringditch region in pre-Columbian times, and abundant macroscopic charcoal suggests that pre-Columbian land management entailed more extensive burning of the landscape than the slash-and-burn agriculture practised around the site today. The site was occupied continuously until near-modern times, although there is evidence for a decline in agricultural intensity or change in land use strategy, and possible population decline, from ~600-500 BP. The long and continuous occupation, which predates the establishment of rainforest in the region, suggests that pre-Columbian land use may have had a significant influence on ecosystem development at this site over the last ~2000 years

Palynological evidence for environmental and climatic change in the lower Guadiana valley, Portugal, during the last 13 000 years

Pollen analysis of a 48 m AMS radiocarbon-dated sediment sequence from the Guadiana estuary provides the first record of Lateglacial and Holocene vegetation history in the Algarve province of Portugal. This paper focuses on the record of terrestrial pollen taxa, which document a series of forest expansions and declines during the period 13 000 cal. BP to 1600 cal. BP and provide insights into climate evolution in southwestern Iberia. The main vegetation phases identified in the Guadiana valley are (1) Lateglacial interstadial (Allerød) forest with Quercus and Pinus under a temperate, moist, continental climate; (2) a Younger Dryas forest decline (Quercus) and expansion of pinewoods, xeric scrub and open ground habitats (with Juniperus, Artemisia, Ephedra distachya type, Centaurea scabiosa type) under arid and cold conditions; (3) an early Holocene forest/scrub/open-ground vegetation mosaic developing under a warm, dry and continental climate; (4) a maximum of Quercus forest and thermomediterranean evergreen taxa (Olea, Phillyrea, Pistacia) reflecting a warm, moist oceanic climate between c. 9000 cal. BP and c. 5000 cal. BP; and (5) the expansion of shrublands with Cistaceae and Ericaceae under a drier climatic regime and increasing anthropogenic activity since c. 5000 cal. BP. Holocene episodes of maximum climatic aridity are identified in the record of xerophytic taxa (Juniperus, Artemisia, Ephedra distachya type) centred around 10 200 cal. BP, 7800 cal. BP, 4800 cal. BP, 3100 cal. BP and 1700 cal. BP. Regional comparisons suggest a correlation of arid phases across southern Iberia and northwest Africa, which can be related to abrupt North Atlantic coolings (Bond events).Research Council studentship for WJF (NERC/S/A/2001/06109), with the support of Trinity College and the Department of Geography, University of Cambridg