Unexpected costs of extended phenotypes: nest features determine the effect of fires on leaf cutter ant's demography.

A key principle of the extended phenotype concept is that the benefit of the structures that an animal builds exceeds its cost. However, some contexts may enhance the costs of structures that often represent a benefit, reversing their adaptive nature. In leaf-cutting ant nests, thatched mounds are extended phenotypes that offer a stable microclimate for the growth of the fungus culture. We hypothesized that fires will affect the species that build external, easily flammable thatch mounds (Acromyrmex lobicornis) more than colonies that build subterranean nests in the less-flammable bare ground (Amoimyrmex striatus). We use a stochastic matrix demographic model parameterized with 4 years of data in pre- and post-fire scenarios. Before fires, Ac. lobicornis showed higher stochastic population rate (λs) than Am. striatus. However, fire frequency every 2 years completely reversed this trend, showing population decline only in Ac. lobicornis. Small nests were the stage that most contributed to λs and the most sensitive in all the species and fire scenarios. This illustrates a novel effect of disturbances; the reversion of the adaptive nature of extended phenotypes, which may have strong consequences on population dynamics and assemblage structure through the invert of dominance relationships.

Evidence of indirect biotic resistance: native ants decrease invasive plant fitness by enhancing aphid infestation.

The biotic resistance hypothesis asserts that native species may hinder the invasion of exotic species, which can occur either directly or indirectly by influencing interactions between exotic and local species. Aphid-tending ants may play a key role in the indirect biotic resistance to plant invasion. Ants may protect aphids, thus increasing their negative effect on exotic plants, but may also deter chewing herbivores, thus benefiting exotic plants. We studied native aphid-tending ants (Dorymyrmex tener, Camponotus distinguendus, and Dorymyrmex richteri) on exotic nodding thistles (Carduus thoermeri), which are attacked by thistle aphids (Brachycaudus cardui) and thistle-head weevils (Rhinocyllus conicus). We evaluated the impact of ants, aphids, and weevils on thistle seed set. We compared ant species aggressiveness towards aphid predators and weevils and performed ant-exclusion experiments to determine the effects of ants on aphid predators and weevils. We analysed whether ant species affected thistle seed set through their effects on aphids and/or weevils. The ant D. tener showed the most aggressive behaviour towards aphid predators and weevils. Further, D. tener successfully removed aphid predators from thistles but did not affect weevils. Excluding D. tener from thistles increased seed set. Analyses supported a negative indirect pathway between the aggressive D. tener and thistle seed set through aphid populations, while the other ant species showed no indirect effects on thistle reproduction. Therefore, aggressive aphid-tending ants may enhance biotic resistance by increasing aphid infestation on exotic invasive plants. This study highlights the importance of indirect biotic resistance in modulating the success of invasive species.

Traffic restrictions for heavy vehicles: Leaf-cutting ants avoid extra-large loads when the foraging flow is high.

A better knowledge of the behaviors that reduce traffic congestions is essential to understand the success of the trail system despite of costs. Leaf-cutting ants use a trunk-trail system to transport leaf fragments into their nests. Some ants carry extra-large leaf fragments and walk slower than the rest of laden workers, thus slowing the ant column behind them. Here we experimentally address whether fragment size selection by leaf-cutting ants depends on the foraging ant flow. If ant behavior aims at minimizing delays associated with carrying extra-large loads, we expect that extra-large loads will be selected mostly under low ant flow conditions. In 38 foraging trails from 18 nests of Acromyrmex crassipinus located in Chaco Serrano woodland, Argentina, we recorded the removal of medium and extra-large baits under variable ant flow conditions. Ants selected extra-large loads mainly under low flow conditions; the increment of ant flow caused an exponential decrease in the proportion and in the preference to carry extra-large fragments. Restriction of heavy vehicles during peak hours is a common traffic rule that prevents traffic jams in transport networks. Our results suggest that this rule may also apply in ant societies that use foraging trails. Avoiding delays generated by carrying large loads appear to be another reason to transport leaf fragments below the individual load capacity, which might help to better understand the high variation in load sizes carried by leaf-cutting ants. This work might help to explain how by following simple traffic rules the trail system can be successful despite its costs, and also illustrate how individual ant behavior can be influenced by nestmates, thereby improving resource harvest in the colony as a whole.

Phorid parasitoid attack triggers specific defensive behaviours and collaborative responses in leaf-cutting ants.

Being social adds another level of defence for organisms: social defences. Beside individual defensive behaviours, social organisms can limit parasite infections by using collective and collaborative behaviours. We evaluated whether the social defence of the leaf-cutting ant Atta cephalotes is specific against phorid parasitoids and the occurrence of collaborative responses depends on the context, i.e. ant activity in foraging trails and number of phorid attacks. We exposed workers to freshly dead specimens of phorids, non-phorid flies or a control without flies in different ant flux conditions and number of attacks and evaluated ant response. We found that workers responded more frequently to phorids than to non-phorid flies and controls suggesting that specific chemical or visual cues of phorids are recognized by leaf-cutting ants triggering a behavioural response. Although the probability of collaborative defences was similar in different ant flux conditions and number of attacks, they occurred more frequently when ants were attacked by a phorid than when they were attacked by the other treatments. Therefore, we demonstrated that leaf-cutting ants differentiate parasitoid flies from other flies, showing a collaborative response against them, in contrast to the other flies and the control, for which almost no collaborative responses were displayed.

Effects of Introduced Plants on Diet and Seed Preferences of Pogonomyrmex carbonarius (Hymenoptera: Formicidae) in the Patagonian Steppe.

Invasive species may interact with resident species and disrupt previously established interactions, with effects on the whole community. If introduced seeds are selectively consumed by native granivores, this could limit the establishment or spread of invasive plants (biotic resistance), and reduce the predation pressure upon native seeds. We determined if the presence of introduced plants affects the diet and the seed preferences of the ant Pogonomyrmex carbonarius (Mayr) in a Patagonian steppe. We expected a higher proportion of introduced seeds in the diet of nests located in areas with a high abundance of introduced species (roadsides) than in nests located in low invaded areas. Diet composition was obtained by collecting items retrieved by workers to colonies and compared between areas of contrasting abundance of introduced species. Field-based choice experiments were performed to evaluate whether exotic seeds were preferred to native ones under a paired comparisons design. Native seeds predominated in the diet. A low proportion of introduced species were included only in colonies close to the road. Ants preferred native seeds to introduced ones, and showed a marked preference for seeds of the native grass Pappostipa speciosa (Trin. & Rupr.) Romasch. (Poaceae), typical of the Patagonian steppe. The presence of introduced plants had little influence on interactions of P. carbonarius with seeds in the Patagonian steppe. Therefore, this ant species would not exert a control on the studied introduced plants, illustrating a case of low biotic resistance to invasion by these species.

A Breath of Fresh Air in Foraging Theory: The Importance of Wind for Food Size Selection in a Central-Place Forager.

Empirical data about food size carried by central-place foragers do not often fit with the optimum predicted by classical foraging theory. Traditionally, biotic constraints such as predation risk and competition have been proposed to explain this inconsistency, leaving aside the possible role of abiotic factors. Here we documented how wind affects the load size of a central-place forager (leaf-cutting ants) through a mathematical model including the whole foraging process. The model showed that as wind speed at ground level increased from 0 to 2 km/h, load size decreased from 91 to 30 mm2, a prediction that agreed with empirical data from windy zones, highlighting the relevance of considering abiotic factors to predict foraging behavior. Furthermore, wind reduced the range of load sizes that workers should select to maintain a similar rate of food intake and decreased the foraging rate by ∼70% when wind speed increased 1 km/h. These results suggest that wind could reduce the fitness of colonies and limit the geographic distribution of leaf-cutting ants. The developed model offers a complementary explanation for why load size in central-place foragers may not fit theoretical predictions and could serve as a basis to study the effects of other abiotic factors that influence foraging.

The effects of ant nests on soil fertility and plant performance: a meta-analysis.

Ants are recognized as one of the major sources of soil disturbance world-wide. However, this view is largely based on isolated studies and qualitative reviews. Here, for the first time, we quantitatively determined whether ant nests affect soil fertility and plant performance, and identified the possible sources of variation of these effects. Using Bayesian mixed-models meta-analysis, we tested the hypotheses that ant effects on soil fertility and plant performance depend on the substrate sampled, ant feeding type, latitude, habitat and the plant response variable measured. Ant nests showed higher nutrient and cation content than adjacent non-nest soil samples, but similar pH. Nutrient content was higher in ant refuse materials than in nest soils. The fertilizer effect of ant nests was also higher in dry habitats than in grasslands or savannas. Cation content was higher in nests of plant-feeding ants than in nests of omnivorous species, and lower in nests from agro-ecosystems than in nests from any other habitat. Plants showed higher green/root biomass and fitness on ant nests soils than in adjacent, non-nest sites; but plant density and diversity were unaffected by the presence of ant nests. Root growth was particularly higher in refuse materials than in ant nest soils, in leaf-cutting ant nests and in deserts habitats. Our results confirm the major role of ant nests in influencing soil fertility and vegetation patterns and provide information about the factors that mediate these effects. First, ant nests improve soil fertility mainly through the accumulation of refuse materials. Thus, different refuse dump locations (external or in underground nest chambers) could benefit different vegetation life-forms. Second, ant nests could increase plant diversity at larger spatial scales only if the identity of favoured plants changes along environmental gradients (i.e. enhancing ß-diversity). Third, ant species that feed on plants play a relevant role fertilizing soils, which may balance their known influence as primary consumers. Fourth, the effects of ant nests as fertility islands are larger in arid lands, possibly because fertility is intrinsically lower in these habitats. Overall, this study provide novel and quantitative evidence confirming that ant nests are key soil modifiers, emphasizing their role as ecological engineers.

Collective Response of Leaf-Cutting Ants to the Effects of Wind on Foraging Activity.

One advantage of sociality is to mitigate environmental restrictions through collective behavior. Here we document a colony-level response of leaf-cutting ants to wind, an environmental factor that impedes foraging. Given that larger ants adhere more strongly to the substrate, increasing forager size in windy conditions should reduce the negative effect of wind. We tested this idea for Acromyrmex lobicornis in windy regions of Patagonia. We examined (1) whether the fraction of larger ants versus smaller ants increased in windy conditions and (2) whether the effect of wind on the ants' movement was lower for larger ants. The size-frequency distribution of foragers was skewed more toward larger ants in nature under more windy conditions. Under windy conditions in the field, the mobility of smaller ants was more reduced than that of larger ants. The change toward larger foragers in windy conditions reduced the negative effect of wind by 32%, illustrating how a social organism can collectively mitigate the adverse effects of the environment.

Social life and sanitary risks: evolutionary and current ecological conditions determine waste management in leaf-cutting ants.

Adequate waste management is vital for the success of social life, because waste accumulation increases sanitary risks in dense societies. We explored why different leaf-cutting ants (LCA) species locate their waste in internal nest chambers or external piles, including ecological context and accounting for phylogenetic relations. We propose that waste location depends on whether the environmental conditions enhance or reduce the risk of infection. We obtained the geographical range, habitat and refuse location of LCA from published literature, and experimentally determined whether pathogens on ant waste survived to the high soil temperatures typical of xeric habitats. The habitat of the LCA determined waste location after phylogenetic correction: species with external waste piles mainly occur in xeric environments, whereas those with internal waste chambers mainly inhabit more humid habitats. The ancestral reconstruction suggests that dumping waste externally is less derived than digging waste nest chambers. Empirical results showed that high soil surface temperatures reduce pathogen prevalence from LCA waste. We proposed that LCA living in environments unfavourable for pathogens (i.e. xeric habitats) avoid digging costs by dumping the refuse above ground. Conversely, in environments suitable for pathogens, LCA species prevent the spread of diseases by storing waste underground, presumably, a behaviour that contributed to the colonization of humid habitats. These results highlight the adaptation of organisms to the hygienic challenges of social living, and illustrate how sanitary behaviours can result from a combination of evolutionary history and current environmental conditions.

Correlations between physical and chemical defences in plants: tradeoffs, syndromes, or just many different ways to skin a herbivorous cat?

Most plant species have a range of traits that deter herbivores. However, understanding of how different defences are related to one another is surprisingly weak. Many authors argue that defence traits trade off against one another, while others argue that they form coordinated defence syndromes. We collected a dataset of unprecedented taxonomic and geographic scope (261 species spanning 80 families, from 75 sites across the globe) to investigate relationships among four chemical and six physical defences. Five of the 45 pairwise correlations between defence traits were significant and three of these were tradeoffs. The relationship between species' overall chemical and physical defence levels was marginally nonsignificant (P = 0.08), and remained nonsignificant after accounting for phylogeny, growth form and abundance. Neither categorical principal component analysis (PCA) nor hierarchical cluster analysis supported the idea that species displayed defence syndromes. Our results do not support arguments for tradeoffs or for coordinated defence syndromes. Rather, plants display a range of combinations of defence traits. We suggest this lack of consistent defence syndromes may be adaptive, resulting from selective pressure to deploy a different combination of defences to coexisting species.

Bottom-up effects may not reach the top: the influence of ant-aphid interactions on the spread of soil disturbances through trophic chains.

Soil disturbances that increase nutrient availability may trigger bottom-up cascading effects along trophic chains. However, the strength and sign of these effects may depend on attributes of the interacting species. Here, we studied the effects of nutrient-rich refuse dumps of the leaf-cutting ant, Acromyrmex lobicornis, on the food chain composed of thistles, aphids, tending ants and aphid natural enemies. Using stable isotopes tracers, we show that the nitrogen accumulated in refuse dumps propagates upward through the studied food chain. Thistles growing on refuse dumps had greater biomass and higher aphid density than those growing in adjacent soil. These modifications did not affect the structure of the tending ant assemblage, but were associated with increased ant activity. In contrast to the expectations under the typical bottom-up cascade effect, the increase in aphid abundance did not positively impact on aphid natural enemies. This pattern may be explained by both an increased activity of tending ants, which defend aphids against their natural enemies, and the low capacity of aphid natural enemies to show numerical or functional responses to increased aphid density. Our results illustrate how biotic interactions and the response capacity of top predators could disrupt bottom-up cascades triggered by disturbances that increase resource availability.

Putting plant resistance traits on the map: a test of the idea that plants are better defended at lower latitudes.

• It has long been believed that plant species from the tropics have higher levels of traits associated with resistance to herbivores than do species from higher latitudes. A meta-analysis recently showed that the published literature does not support this theory. However, the idea has never been tested using data gathered with consistent methods from a wide range of latitudes. • We quantified the relationship between latitude and a broad range of chemical and physical traits across 301 species from 75 sites world-wide. • Six putative resistance traits, including tannins, the concentration of lipids (an indicator of oils, waxes and resins), and leaf toughness were greater in high-latitude species. Six traits, including cyanide production and the presence of spines, were unrelated to latitude. Only ash content (an indicator of inorganic substances such as calcium oxalates and phytoliths) and the properties of species with delayed greening were higher in the tropics. • Our results do not support the hypothesis that tropical plants have higher levels of resistance traits than do plants from higher latitudes. If anything, plants have higher resistance toward the poles. The greater resistance traits of high-latitude species might be explained by the greater cost of losing a given amount of leaf tissue in low-productivity environments.

Succession of ground-dwelling beetle assemblages after fire in three habitat types in the Andean forest of NW Patagonia, Argentina.

Wildfires are one of the major disturbances in the dynamics of forests and shrublands. However, little is known about their effects on insect communities that contribute to faunal biodiversity and play key roles in the ecosystem's dynamics. An intense and widespread fire occurred in 1999 in the Nahuel Huapi National Park in the Andean forest in northern Patagonia, Argentina. This fire affected adjacent, but different, habitat types. After the fire, beetle abundance, species richness and assemblage composition were compared among three habitats that were structurally different before the fire. These habitats were: 1) evergreen forest dominated by Nothofagus dombeyi (Mirb.) Oerst. (Fagales: Nothofagaceae), 2) a mixed forest of the evergreen conifer Austrocedrus chilensis (D. Don) Pic. Serm. and Bizzarri (Pinales: Cupressaceae) and N. dombeyi and 3) a shrubland with a diverse community of shrub species. The relationship between beetle diversity and vegetation structure was investigated over three consecutive years. Ground beetles were collected by pitfall traps, and plant species richness, vegetation cover, and height were measured. Beetle communities varied more over years between habitats during the early regeneration after fire. There was a shift in beetle assemblage composition with time after the fire in all habitat types, probably due to similar colonization rates and microclimatic conditions. Therefore, beetle succession was more influenced by recolonization and survivorship, accompanied by climatic conditions and recovery rate of plant communities over time, than it was influenced by pre-fire habitat conditions. These results suggest that in NW Patagonia, wildfire can have a substantial, short-term impact on beetle abundance and species composition. The pre-fire conditions of each habitat type determined the structure of post-fire communities of plants but not beetle assemblages. Wildfires produce simplification and homogenization of habitat types, and this was reflected by beetle diversity.

Ecological engineering by a native leaf-cutting ant increases the performance of exotic plant species.

Numerous mechanisms are proposed to explain why exotic plants successfully invade natural communities. However, the positive effects of native engineers on exotic plant species have received less consideration. We tested whether the nutrient-rich soil patches created by a native ecological engineer (refuse dumps from the leaf-cutting ant Acromyrmex lobicornis) increase the performance of exotic more than native plants. In a greenhouse experiment, individuals from several native and exotic species were planted in pots with refuse dumps (RDs) and non-nest soils (NNSs). Total plant biomass and foliar nutrient content were measured at the end of the experiment. We also estimated the cover of exotic and native plant species in external RDs from 54 field ant nests and adjacent areas. Greenhouse plants showed more biomass and foliar nutrient content in RDs than in NNS pots. Nevertheless, differences in the final mean biomass among RD and NNS plants were especially great in exotics. Accordingly, the cover of exotic plants was higher in field RDs than in adjacent, non-nest soils. Our results demonstrated that plants can benefit from the enhanced nutrient content of ant RDs, and that A. lobicornis acts as an ecosystem engineer, creating a substrate that especially increases the performance of exotics. This supports the fluctuating resource hypothesis as a mechanism to promote biological invasions, and illustrates how this hypothesis may operate in nature. Since ant nests and exotic plants are more common in disturbed than in pristine environments, the role of ant nests in promoting biological invasions might be of particular interest. Proposals including the use of engineer species to restore disturbed habitats should be planned with caution because of their potential role in promoting invasions.

Leaf-cutting ant nests near roads increase fitness of exotic plant species in natural protected areas.

Understanding the mechanisms that promote the invasion of natural protected areas by exotic plants is a central concern for ecology. We demonstrated that nests of the leaf-cutting ant, Acromyrmex lobicornis, near roadsides promote the abundance, growth and reproduction of two exotic plant species, Carduus nutans and Onopordum acanthium, in a national park in northern Patagonia, Argentina and determine the mechanisms that produce these effects. Refuse dumps (RDs) from ant nests have a higher nutrient content than nearby non-nest soils (NNSs); foliar nutrient content and their 15N isotopic signature strongly suggest that plants reach and use these nutrients. Both species of exotic plants in RDs were 50-600% more abundant; seedlings had 100-1000% more foliar area and root and leaf biomass; and adult plants produced 100-300% more seeds than nearby NNS plants. Plants can thus gain access to and benefit from the nutrient content of ant RD, supporting the hypotheses that enhanced resource availability promotes exotic plant performance that could increase the likelihood of biological invasions. The two exotics produce an estimated of 8385000 more seeds ha(-1) in areas with ant nests compared with areas without; this exceptional increase in seed production represents a potential threat to nearby non-invaded communities. We propose several management strategies to mitigate this threat. Removal efforts of exotics should be focused on ant RDs, where plants are denser and represent a higher source of propagules.

La semilla de Campsiandra angustifolia (Fabaceae: Caesalpiniodeae)como un reflejo de las presiones selectivas sobre su dispersión y establecimiento / The seeds of Campsiandra angustifolia (Fabaceae: Caesalpiniodeae) as a reflex of selective pressures on dispersal and establishment

We indirectly evaluated the selective pressures on dispersal and establishment of Campsiandra angustifolia, a common water-dispersed tree from the Peruvian Amazon, analyzing the variation in the relationship between the volume occupied by dispersal and establishment structures in a total of 535 seeds from 13 trees located at three different habitats. The seeds differed one order of magnitude in their total volume. However, independently of their size and the location of the maternal tree, the relationship between the volume occupied by dispersal and establishment structures was relatively constant (~1) and showed a normal distribution with low skewness, indicating stabilizing selection. These results suggest that, in the habitats studied, dispersal and establishment processes may have similar importance to C.angustifolia. In species with seeds confined in pods, and therefore strongly space-limited, the relative volume of their seeds occupied by dispersal and establishment structures could be a better measure of the trade-off between these two processes than the variation in seed size

Evaluamos indirectamente las presiones selectivas sobre la dispersión y el establecimiento en Campsiandra angustifolia, un árbol de la Amazonía Peruana dispersado por agua, analizando variaciones de la relación entre el volumen ocupado por las estructuras de dispersión y de establecimiento en sus semillas. Medimos un total de 535 semillas de 13 árboles ubicados en tres hábitats diferentes, las cuales presentaron una gran variación en su volumen total. Independientemente del tamaño de la semilla y de la ubicación del árbol de origen, la relación entre el volumen asignado a estructuras de establecimiento y a estructuras de dispersión fue relativamente constante (~1) y presentó una distribución normal con baja asimetría, indicando selección estabilizante. Este resultado sugiere que los procesos de dispersión y establecimiento poseen una importancia relativa similar para C.angustifolia en los hábitats estudiados. En especies con semillas empaquetadas, el volumen relativo ocupado por estructuras relacionadas con la dispersión o el establecimiento podría ser una medida más adecuada de la solución de compromiso entre estos dos procesos que la variación del tamaño de la semilla

[The seeds of Campsiandra angustifolia (Fabaceae: Caesalpiniodeae) as a reflex of selective pressures on dispersal and establishment]. / La semilla de Campsiandra angustifolia (Fabaceae: Caesalpiniodeae) como un reflejo de las presiones selectivas sobre su dispersión y establecimiento.

We indirectly evaluated the selective pressures on dispersal and establishment of Campsiandra angustifolia, a common water-dispersed tree from the Peruvian Amazon, analyzing the variation in the relationship between the volume occupied by dispersal and establishment structures in a total of 535 seeds from 13 trees located at three different habitats. The seeds differed one order of magnitude in their total volume. However, independently of their size and the location of the maternal tree, the relationship between the volume occupied by dispersal and establishment structures was relatively constant (approximately 1) and showed a normal distribution with low skewness, indicating stabilizing selection. These results suggest that, in the habitats studied, dispersal and establishment processes may have similar importance to C. agustifolia. In species with seeds confined in pods, and therefore strongly space-limited, the relative volume of their seeds occupied by dispersal and establishment structures could be a better measure of the trade-off between these two processes than the variation in seed size.

[Acumen function in leaves and its vertical distribution in a tropical rain forest of Costa Rica]. / Función del acumen en las hojas y su distribución vertical en un bosque lluvioso tropical de Costa Rica.

Water retention on the leaf surface can be maladaptive to the plant because it increases the colonization of epiphylls and interferes with the physiologic processes of the leaf, diminishing the photosynthetic capacity. To test if leaf driptips facilitate leaf drying after rainfall in a tropical rain forest of Costa Rica, we (1) experimentally measured the capacity to retain water on leaf surfaces of 30 plant species before and after driptip removal, and (2) analyzed the development of driptips along forest strata. We expected leaf driptips to be less developed in the upper strata due to the environmental conditions of the canopy (i.e., high solar radiation, strong winds and low relative humidity), which favor the natural drying of leaves. The presence of driptips increased 100% the water run off capacity of leaves in all the analyzed species. Also, the development of leaf driptips was smaller in canopy species than in understory species. Additionally, they became less developed in canopy species as trees increased in height. These results support the hypothesis that the adaptive role of driptips is to facilitate the drying of leaf surfaces.

Función del acumen en las hojas y su distribución vertical en un bosque lluvioso tropical de Costa Rica / Acumen function in leaves and its vertical distribution in a tropical rain forest of Costa Rica

Water retention on the leaf surface can be maladaptive to the plant because it increases the colonization of epiphylls and interferes with the physiologic processes of the leaf, diminishing the photosynthetic capacity. To test if leaf driptips facilitate leaf drying after rainfall in a tropical rain forest of Costa Rica, we (1) experimentally measured the capacity to retain water on leaf surfaces of 30 plant species before and after driptip removal, and (2) analyzed the development of driptips along forest strata. We expected leaf driptips to be less developed in the upper strata due to the environmental conditions of the canopy (i.e., high solar radiation, strong winds and low relative humidity), which favor the natural drying of leaves. The presence of driptips increased 100 the water run off capacity of leaves in all the analyzed species. Also, the development of leaf driptips was smaller in canopy species than in understory species. Additionally, they became less developed in canopy species as trees increased in height. These results support the hypothesis that the adaptive role of driptips is to facilitate the drying of leaf surfaces.