Neotropical Forests Research Articles

Chorus Organization in a Neotropical Forest Cicada

In many species of animals, males aggregate in particular locations and produce calls to attract searching females for reproduction. One striking feature of such choruses is synchronization. On a scale of hours, choruses are often concentrated at particular times of day, such as dawn or dusk. On a scale of seconds, males may also synchronize the rhythm of their calls with one another. While synchronized calling at this scale suggests benefits acting at the collective level, competitive interactions between males to attract females can also lead to synchronized calling as an epiphenomenon. Why males in some species synchronize the rhythm of their calls is still debated, and more work is needed to understand the evolution of this behavior. I investigated chorus organization in the Emerald cicada (Zammara smaragdula), a Neotropical forest cicada in southern Belize, to explore these issues. Choruses in this species occurred at dawn and dusk and, occasionally, during daytime. There was no evidence for synchronization in the rhythm of calls among males, as bouts of collective calling occurred after quiet intervals of variable rather than fixed durations. The temporal aggregation of calls in this species thus probably emerged from competitive interactions among males to attract females. The degree of temporal overlap in the calls of males during a chorus varied as a function of chorus phase and time of day, suggesting flexibility in chorus organization, perhaps in relation to temporal variations in factors such as the number of calling cicadas, the number of predators present or ambient temperature during a chorus.

Body size and abundance patterns of male orchid bees in a fragmented Neotropical landscape

Abstract Understanding the relationship between functional traits and fragmentation is crucial for promoting biodiversity and ecosystem function. Orchid bees (Euglossini) are important pollinators in Neotropical forests, but how body size may be associated with fragmentation and its influence on community organisation remains unexplored. We systematically sampled orchid bee communities across 15 plots in a fragmented area in the Ecuadorian Amazon basin to evaluate how patch size and isolation are associated with body size and abundance. Our samples comprised large‐bodied bees (Eulaema, Euglossa and the parasitic Exaerete), with small Euglossa notably absent. The relative representation of Eulaema species of varying sizes changed across the fragmented landscape, leading to shifts in overall mean body size consistent with the prediction that larger‐bodied pollinators are better at long‐distance movement. Euglossa, the most abundant genus (67% of the total sample), was found across all patches regardless of body size. Euglossa intersecta and Euglossa orellana, the largest Euglossa in the region, were the most common species across the fragmented landscape. The abundance of male euglossines experienced a general decline with decreasing patch size and increasing isolation. However, at the species level, the abundance of some species was not affected at all. Because body size often mediates dispersal ability, thermoregulation, and resource use, we reinforce the importance of assessing body size patterns to understand changes in diversity, abundance and the composition of tropical pollinator communities in fragmented landscapes.

A methodological procedure for the estimation of ecological value applied to a neotropical cloud forest

A methodological procedure for the estimation of ecological value applied to a neotropical cloud forest

Floral morphology and pollen placement strategies of bat-pollinated flowers: a comparative analysis within a guild of chiropterophilous plants in a neotropical dry forest

BackgroundSynchronopatric plant species, coexisting and flowering simultaneously, often engage in intense competition for pollen deposition on shared pollinators. This study focuses on the intricate dynamics of chiropterophilous flowers within the neotropical bat-pollination plant guild, specifically investigating the mechanical fit of reproductive structures to pollinating bats at the community level.MethodsUtilizing a diverse guild of bat-pollinated plant species from the Caatinga, the largest dry forest in the Neotropics, our research integrates various key components. Initially, we identified specific contact sites for floral reproductive structures on the bat's body, exploring diverse pollen placement strategies. Subsequent efforts involved characterizing floral traits within the guild and examining their associations with different pollen placement strategies. Precision in the contact of floral reproductive structures was estimated, and findings were integrated by associating pollen placement strategies and precision with the investment in pollen production.ResultsWe found that certain bat body parts, particularly the face and neck, were more frequently contacted by reproductive structures. The three identified categories of pollen placement strategies were evenly distributed among plant species, each linked to specific floral traits. Notably, the absence of oriented herkogamy prevailed in 70% of the species. Morphometric analyses unveiled significant variations in operative distances among species, emphasizing exceptional variability in certain outliers. While precision in pollen transfer was influenced by key factors, surprisingly, investment in pollen production did not differ among plants with distinct pollen placement strategies. Furthermore, it showed no correlation with fundamental accuracy.ConclusionsThe subsequent exploration delves into the intricate associations between distinct floral characteristics and various pollen placement strategies, shaping the complex pollen landscape on bat bodies. This research provides valuable insights into the community-level dynamics of chiropterophilous flowers in the Caatinga Dry Forest, emphasizing the role of different pollen deposition strategies in facilitating the coexistence of multiple plant species within the chiropterophilous guild.

Diversidad y distribución de los helechos y licófitos de un bosque subandino del norte de los Andes

Background and aims: Ferns and lycophytes are highly diverse in Neotropical forests, and their distribution is influenced by various biotic and abiotic factors, including disturbance. This study analyzes the effect of some of these factors on the diversity of these plants in a sub-Andean forest. M&M: Four transects of 400 m2 (80×5 m) were established, two in highly disturbed sites, grassland, and stream edge, and two in the forest. Each transect was divided into 16 plots where all terrestrial and epiphytic individuals were recorded. Clustering analyses were performed to recognize the presence of floristic groups, and a canonical correlation analysis was performed to identify how soil chemistry and vegetation structure affect the composition and diversity of these plants. Results: A total of 4935 individuals were found, representing 42 species. Amauropelta rudis (Kunze) Pic. Serm. (2485 individuals) and Asplenium flabellulatum Kunze (1190 individuals) were the dominant terrestrial species. The clustering and ordination analyses revealed that the plots of open sites are more related to each other than to those of the forest. Species of open sites are more correlated with a lower layer of litter, a lower height and coverage of trees or saplings, a higher concentration of aluminum and a lower concentration of sodium. Conclusions: Ferns and lycophytes are influenced by the environmental, structural and disturbance conditions of the site where they grow. Key words Disturbance, diversity, edaphic factors, multivariate analysis, pteridophytes, richness, structural factors.

Effects of urbanization on orchid bee diversity and orchid pollination: From neotropical cloud forests to urban cores

Urbanization is a significant driver of land use change, profoundly impacting biodiversity and ecosystem services worldwide. However, its effects in the tropics, which host some of the planet's highest biodiversity, remain inadequately understood. Orchid bees (Apidae: Euglossini) are key pollinators in Neotropical ecosystems, playing crucial roles in maintaining floral diversity and reproductive success of orchids and other plant families. Yet, little is known about how urbanization influences their diversity and pollination. In this study, we analyzed the diversity and composition or orchid bee communities along an urbanization gradient which extends from the city center to the surrounding cloud forests, which bear high orchid endemism while being highly threatened. Along the same gradient, we further evaluated pollination of a model native orchid, Gongora galeata, which is exclusively pollinated by the bee Euglossa obrima. As expected, increasing urbanization led to a decrease in orchid bee diversity, as well as a clear separation in species composition between urban and non-urban sites and a reduction in G. galeata pollination (i.e. fruit production). However, contrary to our expectations, orchid pollination also decreased with environmental heterogeneity and the abundance of its specific pollinator. Despite urban areas still hosting orchid bee species, our results reveal clear negative effects of urbanization not only on diversity, but also on the ecosystem function of a highly threatened group of bees. This study highlights the importance of considering local factors of urban landscapes for preserving not only biodiversity, but also fundamental ecological processes in cities.

Cattle exclusion increases encounters of wild herbivores in Neotropical forests

Abstract Ongoing habitat loss and species extinctions require managers to implement and quantify the effectiveness of conservation actions for protecting biodiversity. Fencing, when done properly, is an important management tool for conservation in landscapes where wildlife and domestic animals co‐occur, potentially enhancing habitat use through selective exclusion of domestic species. For instance, the fencing of forest patches in the Neotropics is expected to reduce the degradation of understory vegetation by cattle, releasing these resources for the native community of browsers and fruit consumers. Here, we implemented an ecological experiment using a before‐after control‐impact design to quantify the effect of cattle exclusion on encounter probability of the native community of browsers and fruit consumers, and percent ground cover in multifunctional landscapes of the Colombian Orinoquía. We built 14 km of wildlife‐permeable fences along forest edges in four forest patches (i.e. blocks) containing control and fenced (treatment) sites. We installed 33 camera traps to obtain information about wildlife and cattle encounter probabilities, before and after the fences were constructed. We used Bayesian generalised linear mixed effects models to quantify the effect of fences via the interaction between the time period (before and after the fences were built) and treatment (control or fenced sites). Fencing was effective at reducing encounter probabilities of cattle in the treated sites, and it had a positive impact on relative encounter probabilities of four of seven studied wildlife species (herbivores including the black agouti [dry season only], lowland tapir [dry season only] and spotted paca [both seasons] and an omnivore, the South American coati [rainy season only]). The effect of fencing was negative for the collared peccary but only during the dry season. No statistically significant effect was detected for the white‐lipped peccary or white‐tailed deer. Synthesis and applications: We provide experimental evidence that fences are effective at selectively excluding cattle and increasing encounter rates of wild browsers and fruit consumers in forest patches where these species co‐occur with cattle. Our results highlight an important application of fencing ecology in Neotropical forests, where the implementation of wildlife‐permeable fences is feasible due to smaller body sizes of wildlife compared to domestic animals such as cattle.

From Leaves to Reproductive Organs: Chemodiversity and Chemophenetics of Essential Oils as Important Tools to Evaluate Piper mollicomum Kunth Chemical Ecology Relevance in the Neotropics.

Piper mollicomum Kunth (Piperaceae) plays a vital role in the preservation of the Brazilian Atlantic Forest by contributing to the regeneration of deforested areas. Recent scientific investigations have analyzed the chemical constituents and seasonal dynamics of essential oils (EO) from various Piper L. species, highlighting the need to elucidate their chemical-ecological interactions. This study aims to expand the chemical-ecological knowledge of this important taxon in neotropical forests, using P. mollicomum as a model. The methodologies employed include the collection of plant material, EO extraction by hydrodistillation, analysis of EO by gas chromatography-mass spectrometry (GC-MS) and gas chromatography-flame ionization detector (GC-FID), recording the frequency of visits by potential pollinators and microclimatic variables, and by conducting calculations of chemodiversity and chemophenetic indices. Chemical analyses indicated that the diversity of EO and environmental factors are linked to the activities of potential pollinators. In the Tijuca Forest, P. mollicomum revealed significant interactions between its volatile constituents and microclimatic variables, showing that the chemodiversity of the leaves and reproductive organs correlates with pollinator visitation. Additionally, a notable difference in chemical evenness was observed between these vegetative structures. The chemophenetic indices by Ramos and Moreira also revealed correlations with chemical diversity.

Leaf-cutting ants’ distribution in an elevation gradient in central Argentina

ABSTRACT Leaf-cutting ants are among the most prominent herbivores in neotropical forests and savannas, and Argentina represents the southern limit of their distribution. Despite their importance, few systematic studies analyze the presence and distribution of leaf-cutting ants in Argentina. Our aim was to describe, through an intensive field sampling, the leaf-cutting ant species present in the central Argentine mountains and analyze their distribution along the elevation gradient. We collected ants from 69 nests in 56 out of the 250 plots visited along the study elevation gradient (400–2700 m above sea level) in seven different habitats, from forests to rocky areas. We found six species. The most frequent were Acromyrmex lobicornis and Acromyrmex crassispinus, while Acromyrmex heyeri, Amoimyrmex silvestrii, Amoimyrmex striatus, and Amoimyrmex bruchi were far less frequent. The species distributed at the highest elevation was A. lobicornis (1223 to 2163 m a.s.l.), showing significant differences with the other species. We found leaf-cutting ants in all habitats, with a higher presence in grazing lawns, shrublands, and exotic forests. Our results indicate that leaf-cutting ants are present in most of the mountain range, as the area above their upper limit of distribution represents less than 5% of the total mountain area.

Ecotones as Windows into Organismal-to-Biome Scale Responses across Neotropical Forests.

Tropical forests are incredibly diverse in structure and function. Despite, or perhaps because of, this diversity, tropical biologists often conduct research exclusively in one or perhaps a few forest types. Rarely do we study the ecotone-the interstitial region between forest types. Ecotones are hyper-diverse, dynamic systems that control the flow of energy and organisms between adjacent ecosystems, with their locations determined by species' physiological limits. In this review, we describe how studying ecotones can provide key indicators for monitoring the state of Neotropical forests from organisms to ecosystems. We first describe how ecotones have been studied in the past and summarize our current understanding of tropical ecotones. Next, we provide three example lines of research focusing on the ecological and evolutionary dynamics of the ecotone between tropical dry forests and desert; between tropical dry and rainforests; and between Cerrado and Atlantic rainforests, with the latter being a particularly well-studied ecotone. Lastly, we outline methods and tools for studying ecotones that combine remote sensing, new statistical techniques, and field-based forest dynamics plot data, among others, for understanding these important systems.

To improve estimates of neotropical forest carbon stocks more direct measurements are needed: An example from the Southwestern Amazon

Tropical forests play a critical role in the global carbon cycle, storing 40–55 % of terrestrial plant carbon and significantly contributing to primary productivity. However, uncertainties persist in estimating carbon stocks and fluxes, exhibiting variation across the Neotropics, Africa, and Asia tropical forest regions. Despite hosting some of the most densely sampled forests, significant uncertainties persist in biomass and forest carbon stock estimates in the Neotropics. Although the Southwestern Amazon (SWA) forests span over 20 million hectares, no specific biomass or above- and below-ground carbon model has been calibrated for this region thus far. In our study, we conducted direct forest inventories in the SWA to address the following question: Do the allometric patterns, biomass, and carbon stocks observed in the Southwestern Amazon differ from those found in other regions of the Amazon or Pantropical? Our research reveals substantial differences in water and carbon content, biomass stocks, above- and below-ground oven-dry biomass ratios, and allometric patterns between SWA forests and other Amazonian and Pantropical forests. We have demonstrated that these differences result in overestimations of forest biomass when applying allometric equations developed for other Amazonian and Pantropical regions to the open forests of Southwestern Amazonia. This overestimation can reach up to 37 % when using equations from the eastern Amazon, and between 26 % and 46 % depending on the applied Pantropical equation. The use of an inappropriate factor for the root-to-shoot ratio in the Southwestern Amazon (SWA) can lead to overestimates of belowground oven-dry biomass by up to 20 %. To reduce uncertainties related to estimates of forest carbon stock and flux in Neotropical forests, it is necessary to enhance the density of direct biomass measurements, particularly in southwestern Amazonia.

Seed Dispersal Networks in Neotropical Forest Areas of the Amazon, Cerrado, and Associated Ecotone: Abundance as the Driver of Bat Roles

Seed Dispersal Networks in Neotropical Forest Areas of the Amazon, Cerrado, and Associated Ecotone: Abundance as the Driver of Bat Roles

A low-cost UAV survey can unravel Baird's Tapir (Tapirus bairdii) trail network dynamics in a neotropical highland forest

UAV technology has transformed spatial data collection and high-spatial resolution analysis of diverse environments around the world. Its role within geosciences, ecology, engineering, and other sciences has been increasing withing the last decade. In ecological and geomorphological aspects, it offers a powerful tool for studying landforms, surface processes, vegetation, and animal behavior globally. Our study employs high-resolution UAV surveys to analyze Tapirus bairdii trail networks and their connection to the landscape in a Costa Rican highland paramo-forest national park. Using a DJI Phantom 4 V.2 UAV, we conducted a photogrammetric flight over 3.19 ha, producing datasets with a spatial resolution of 2.99 cm/pixel. These datasets facilitated geostatistical analysis of Tapirus bairdii trails, revealing their complexity and hotspots near lagoons. We investigated lagoon characteristics, finding that finer sediments spatially correspond with higher trail activity. This suggests that sediment types in lagoons influence Tapirus bairdii navigation, impacting trail network composition. Our findings highlight UAV technology's role in advancing ecological and geomorphological research in protected areas, providing insights into Tapirus bairdii behavior within highland ecosystems. Overall, to the date this is the first high-resolution UAV survey of Tapirus bairdii within these types of highland tropical environments.

Long-term monitoring reveals the long lifespans of Neotropical forest landbirds.

Long-term monitoring reveals the long lifespans of Neotropical forest landbirds.

Functional traits and ecological niches as correlates of the interspecific growth–mortality trade‐off among seedlings of 14 tropical tree species

Abstract The interspecific trade‐off between growth versus mortality rates of tree species is thought to be driven by functional biology and to contribute to species ecological niche differentiation. Yet, functional trait variation is often not strongly correlated with growth and mortality, and few studies have investigated the relationships of both traits and niches, specifically encompassing above and belowground resources, to the trade‐off itself. These relationships are particularly relevant for seedlings, which must often survive resource limitation to reach larger size classes. We investigated the functional basis of the interspecific growth–mortality trade‐off and its relationship with ecological niches for seedlings of 14 tree species in a tropical forest in southwest China. We found evidence for an interspecific growth–mortality trade‐off at the seedling stage using 15 functional traits and 15 ecological niche variables. None of the organ‐level traits correlated with growth, mortality, nor the trade‐off, whereas specific stem length (SSL), a biomass allocation trait, was the only trait to have a significant correlation (positive). Moreover, light‐defined niches were not correlated with growth, mortality or the trade‐off, but soil‐defined niches did. Species at the faster growth/higher mortality end of the trade‐off were associated with higher fertility defined by lower soil bulk density and slope, and higher soil organic matter concentration and soil total nitrogen. Our findings indicate the importance of stem elongation and soil fertility for growth, mortality and their trade‐off at the seedling stage in this Asian tropical forest. Our findings contrast with analogous studies in neotropical forests showing the importance of photosynthesis‐related leaf traits related to insolation. Therefore, the functional drivers of demographic rates and trade‐offs, as well as their consequences for ecological niches, can vary among forests, likely owing to differences in biogeography, canopy disturbance rates, topography and soil properties. Moreover, the effects of functional trait variation on demographic rates and trade‐offs may be better revealed when biomass allocation is accounted for in a whole‐plant context. Read the free Plain Language Summary for this article on the Journal blog.

Chicatana ant (Atta mexicana) as a potential source of biologically active substances

Atta mexicana (Smith, 1858) is a leaf-cutting ant from the neotropical forests and an important pest. The abdomens of the reproductive specimens are consumed as food and medicinal remedies in the southwest of Mexico. Because their high-quality proteins remain unexplored, the protein profile and the free amino acid content were analyzed through a metabolomic approach. We identified 2247 proteins in the abdomen and edible part of the Chicatana ant and 2098 proteins in the whole insect, visualizing a core of 1186 proteins. Thus, we carried out a targeted metabolomic approach with a bioinformatic prediction that suggests the occurrence of antimicrobial tryptic peptides in the Atta mexicana proteome. These findings are the first description of the free amino acid and protein profiles of Atta mexicana.

Assessing two decades of landscape greenness in relation to temperature and precipitation in a tropical dry forest of Northwestern Mexico

Canopy greenness is an indicator of ecosystem primary productivity, which is often limited by temperature and precipitation. Changes in vegetation greening have been reported mostly at global scales. However, we still have a poor understanding of vegetation greening patterns and drivers for major vegetation types, such as the tropical dry forest, one of the most extensive vegetation types in Mexico. Here, we analyze two decades of interannual variation in greenness and its relationship to temperature and precipitation in the northmost neotropical dry forest occurring in Northwestern Mexico. We constructed time-series linear regression models using standardized anomalies with z-scores (i.e., standard deviations away from the long-term mean) for the Normalized Difference Vegetation Index (NDVI) and the climate data from 2001 to 2021. Our best models indicate both temperature and precipitation exert positive effects on vegetation greenness, particularly from a lagged effect perspective, as retained predictors were the accumulated precipitation of two monsoon (summer) seasons and previous year mean temperature. The lowest levels of landscape greenness seem connected to prolonged droughts and extreme frost events. In fact, a switch from negative to positive NDVI anomalies was observed in the years following the February 2011 extreme frost that affected much of North America, including northern Mexico. Notable, under the stricter statistical criterion of −1.7 ≥ z score ≥ 1.7, only climatic variables presented very-extreme anomalies and these were not necessarily linked to a very-extreme response in landscape greenness. However, considering the criterion of −1.3 ≥ z score ≥ 1.3, we identified several extreme NDVI anomalies and their corresponding climatic anomaly. Therefore, more flexible statistical criteria might reveal climate extremes of ecological and social relevance. Our overall findings have implications for forest and climate risk management, particularly as extreme climatic anomalies are expected to continue increasing in light of climate change.

Limited drought tolerance in the neotropical seasonally dry forest plants impairs future species richness.

Neotropical seasonal dry forest (NSDF) is one of the most threatened ecosystems according to global climate change predictions. Nonetheless, few studies have evaluated the global climate change impacts on diversity patterns of NSDF plants. The lack of whole biome-scale approaches restricts our understanding of global climate change consequences in the high beta-diverse NSDF. We analysed the impact of global climate change on species distribution ranges, species richness, and assemblage composition (beta diversity) for 1,178 NSDF species. We used five representative plant families (in terms of abundance, dominance, and endemism) within the NSDF: Cactaceae, Capparaceae, Fabaceae, Malvaceae, and Zygophyllaceae. We reconstructed potential species distributions in the present and future (2040-2080), considering an intermediate Shared Socioeconomic Pathway and two dispersal ability assumptions on the taxa. Using a resource use scores index, we related climate-induced range contractions with species' water stress tolerance. Even under a favourable dispersal scenario, species distribution and richness showed future significant declines across those sites where mean temperature and precipitation seasonality are expected to increase. Further, changes in species range distribution in the future correlated positively with potential use of resources in Fabaceae. Results suggest that biotic heterogenization will likely be the short-term outcome at biome scale under dispersal limitations. Nonetheless, by 2080, the prevailing effect under both dispersal assumptions will be homogenization, even within floristic nuclei. This information is critical for further defining new areas worth protecting and future planning of mitigation actions for both species and the whole biome.

The role of changing landscape in the dispersal of a soil‐feeding termite in Suriname and French Guiana

Abstract Neotropical forest ecosystems harbour significant biodiversity. To develop effective insect conservation practices, it is important to understand the factors that influence the diversity and population structure of the species. Dispersal, a key determinant of population structure, is well studied in termites nesting in wood, for which it can be influenced by wood transport, or in termites living in urban environments. However, understanding the dispersal of termites whose mobility remains unaffected by wood transport remains understudied. We investigated the dispersal of Embiratermes neotenicus, a soil‐feeding species with short dispersal distance, in the Neotropical region, where both intact and degraded forests exist. Using mitochondrial and nuclear data, we analysed genetic diversity, structure and factors contributing to population differentiation in Suriname and French Guiana at multiple scales for 70 colonies. The population in French Guiana is the ancestral population in the region that subsequently expanded. Significant genetic differentiation between populations was observed, with distinct patterns identified in Suriname and French Guiana. The Suriname population showed higher genetic diversity and no subpopulation differentiation, whereas the French Guiana population showed substructure into distinct genetic clusters. Analyses at the scale of all colonies suggest the influence of landscape features, such as the Maroni River, on genetic differentiation. At the local scale, genetic differentiation between colonies increases with forest alteration, even when this does not include major changes in forest cover. Our results highlight the sensitivity of soil‐feeding termite populations to habitat change. We argue that multi‐scale studies are needed for a comprehensive understanding of genetic patterns, especially for species with short dispersal distances.

Evolution of Acquired Perfumes and Endogenous Lipid Secretions in Orchid Bees.

Male orchid bees are unique in the animal kingdom for making perfumes that function as sex pheromone. Males collect volatile chemicals from the environment in the neotropical forests, including floral and non-floral sources, creating complex but species-specific blends. Male orchid bees exhibit several adaptations to facilitate perfume collection and storage. When collecting volatile compounds, males apply lipid substances that they secrete from cephalic labial glands onto the fragrant substrate. These lipids help dissolve and retain the volatiles, similar to the process of 'enfleurage' in the traditional perfume industry. We investigated how the chemical composition of acquired perfume and labial gland secretions varied across the phylogeny of orchid bees, including 65 species in five genera from Central and South America. Perfumes showed rapid evolution as revealed by low overall phylogenetic signal, in agreement with the idea that perfume compounds diverge rapidly and substantially among closely related species due to their role in species recognition. A possible exception were perfumes in the genus Eulaema, clustering closely in chemospace, partly mediated by high proportions of carvone and trans-carvone oxide. Labial gland secretions, in contrast, showed a strong phylogenetic signal at the genus level, with secretions of Eufriesea and Exaerete dominated by fatty acids and Eulaema dominated by saturated acetates of chain lengths 12 to 16 C-atoms. Secretions of the majority of Euglossa were heavily dominated by one unsaturated long chain diacetate, (9Z)-Eicosen-1,20-diyldiacetate. However, we also identified few highly divergent species of Euglossa in four subclades (11 species) that appear to have secondarily replaced the diacetate with other compounds. In comparison with environment-derived perfumes, the evolution of labial gland secretion is much slower, likely constrained by the underlying biochemical pathways, but perhaps influenced by perfume-solvent chemical interactions.