Rediscovery of Gasteranthusextinctus L.E.Skog & L.P.Kvist (Gesneriaceae) at multiple sites in western Ecuador.

We report the rediscovery of the Critically Endangered cloud forest herb Gasteranthusextinctus, not seen since 1985. In 2019 and 2021, G.extinctus was recorded at five sites in the western foothills of the Ecuadorian Andes, 4-25 km from the type locality at the celebrated Centinela ridge. We describe the species' distribution, abundance, habitat and conservation status and offer recommendations for further research and conservation efforts focused on G.extinctus and the small, disjunct forest remnants it occupies.

Mature Andean forests as globally important carbon sinks and future carbon refuges.

It is largely unknown how South America's Andean forests affect the global carbon cycle, and thus regulate climate change. Here, we measure aboveground carbon dynamics over the past two decades in 119 monitoring plots spanning a range of >3000 m elevation across the subtropical and tropical Andes. Our results show that Andean forests act as strong sinks for aboveground carbon (0.67 ± 0.08 Mg C ha-1 y-1) and have a high potential to serve as future carbon refuges. Aboveground carbon dynamics of Andean forests are driven by abiotic and biotic factors, such as climate and size-dependent mortality of trees. The increasing aboveground carbon stocks offset the estimated C emissions due to deforestation between 2003 and 2014, resulting in a net total uptake of 0.027 Pg C y-1. Reducing deforestation will increase Andean aboveground carbon stocks, facilitate upward species migrations, and allow for recovery of biomass losses due to climate change.

The legacy of biogeographic history on the composition and structure of Andean forests.

The biogeographic origin of species may help to explain differences in average tree height and aboveground biomass (AGB) of tropical mountain forests. After the Andean uplift, small-statured trees should have been among the initial colonizers of the highlands (new cold environment) from the lowland tropics, since these species are pre-adapted to cold conditions with narrow vessels that are relatively resistant to freezing. If the descendants of these small-statured clades continue to dominate tropical highland forests, there will be a high co-occurrence of close relatives at high elevations. In other words, this scenario predicts a systematic decline in tree size, AGB, and phylogenetic diversity with elevation. In contrast, the colonization of Andean forests by some large-statured clades that originated in temperate regions may modify this expectation and promote a mixing of tropical and temperate clades, thereby increasing the phylogenetic diversity in tropical highland forests. This latter scenario predicts an increase or no change of tree size, AGB, and phylogenetic diversity with elevation. We assessed how the historical immigration of large-statured temperate-affiliated tree lineages adapted to cold conditions may have influenced the composition and structure of Andean forests. Specifically, we used 92 0.25-ha forest inventory plots distributed in the tropical Andes Mountains of Colombia to assess the relationship between the phylogenetic diversity and AGB along elevational gradients. We classified tree species as being either "tropical affiliated" or "temperate affiliated" and estimated their independent contribution to forest AGB. We used structural equation modeling to separate the direct and indirect effect of elevation on AGB. We found a hump-shaped relationship of phylogenetic diversity, AGB, and tree size with elevation. The high phylogenetic diversity found between 1,800-2,200 m above sea level (asl) was due to the mixing of highland floras containing many temperate-affiliated species, and lowland floras containing mostly tropical-affiliated species. The high AGB in highland forests, which contrasted with the expected decline of AGB with elevation, was likely due to the significant contribution of temperate-affiliated species. Our findings highlight the lasting importance of biogeographic history on the composition and structure of Andean mountain forests.

Elevation and latitude drives structure and tree species composition in Andean forests: Results from a large-scale plot network.

Our knowledge about the structure and function of Andean forests at regional scales remains limited. Current initiatives to study forests over continental or global scales still have important geographical gaps, particularly in regions such as the tropical and subtropical Andes. In this study, we assessed patterns of structure and tree species diversity along ~ 4000 km of latitude and ~ 4000 m of elevation range in Andean forests. We used the Andean Forest Network (Red de Bosques Andinos, https://redbosques.condesan.org/) database which, at present, includes 491 forest plots (totaling 156.3 ha, ranging from 0.01 to 6 ha) representing a total of 86,964 identified tree stems ≥ 10 cm diameter at breast height belonging to 2341 identified species, 584 genera and 133 botanical families. Tree stem density and basal area increases with elevation while species richness decreases. Stem density and species richness both decrease with latitude. Subtropical forests have distinct tree species composition compared to those in the tropical region. In addition, floristic similarity of subtropical plots is between 13 to 16% while similarity between tropical forest plots is between 3% to 9%. Overall, plots ~ 0.5-ha or larger may be preferred for describing patterns at regional scales in order to avoid plot size effects. We highlight the need to promote collaboration and capacity building among researchers in the Andean region (i.e., South-South cooperation) in order to generate and synthesize information at regional scale.

Functional composition of epiphyte communities in the Colombian Andes.

We identify changes in the functional composition of vascular epiphytes along a tropical elevational gradient with the aim of quantifying the role of climate in determining the assembly of epiphyte communities. We measured seven leaf functional traits (leaf area, specific leaf area, leaf dry-matter content, leaf thickness, force to punch, stomatal density, and potential conductance index) in the 163 most abundant epiphyte species recorded across 10 sites located along an elevational gradient between 60 and 2,900 m above sea level in the Colombian Andes. We grouped the epiphyte species into seven hierarchical functional groups according to their most characteristic leaf traits. Along the elevational gradient, the two main independent leaf trait dimensions that distinguished community assemblages were defined primarily by leaf area-photosynthetic (LAPS) and mass-carbon (LMCS) gradients. Mean annual temperature was the main determinant of species position along LAPS. In contrast, local changes in specific leaf area due to variation in the epiphytes' relative height of attachment was the main determinant of their position along the LMCS. Our findings indicate that epiphytic plant leaves have evolved to optimize and enhance photosynthesis through a leaf area-based strategy and carbon acquisition through investments in construction costs of leaf area per unit of biomass that aim to regulate light capture and tissue development. Given that most studies of plant functional traits neglect vascular epiphytes, our quantification of the multiple dimensions of epiphyte leaf traits greatly augments our understanding of vascular plant function and adaptation to changing environments.

Global climate change increases risk of crop yield losses and food insecurity in the tropical Andes.

One of the greatest current challenges to human society is ensuring adequate food production and security for a rapidly growing population under changing climatic conditions. Climate change, and specifically rising temperatures, will alter the suitability of areas for specific crops and cultivation systems. In order to maintain yields, farmers may be forced to change cultivation practices, the timing of cultivation, or even the type of crops grown. Alternatively, farmers can change the location where crops are cultivated (e.g., to higher elevations) to track suitable climates (in which case the plants will have to grow in different soils), as cultivated plants will otherwise have to tolerate warmer temperatures and possibly face novel enemies. We simulated these two last possible scenarios (for temperature increases of 1.3°C and 2.6°C) in the Peruvian Andes through a field experiment in which several traditionally grown varieties of potato and maize were planted at different elevations (and thus temperatures) using either the local soil or soil translocated from higher elevations. Maize production declined by 21%-29% in response to new soil conditions. The production of maize and potatoes declined by >87% when plants were grown under warmer temperatures, mainly as a result of the greater incidence of novel pests. Crop quality and value also declined under simulated migration and warming scenarios. We estimated that local farmers may experience severe economic losses of up to 2,300 US$ ha-1  yr-1 . These findings reveal that climate change is a real and imminent threat to agriculture and that there is a pressing need to develop effective management strategies to reduce yield losses and prevent food insecurity. Importantly, such strategies should take into account the influences of non-climatic and/or biotic factors (e.g., novel pests) on plant development.

Contrasting outcomes of species- and community-level analyses of the temporal consistency of functional composition.

Multiple anthropogenic drivers affect every natural community, and there is broad interest in using functional traits to understand and predict the consequences for future biodiversity. There is, however, no consensus regarding the choice of analytical methods. We contrast species- and community-level analyses of change in the functional composition for four traits related to drought tolerance using three decades of repeat censuses of trees in the 50-ha Forest Dynamics Plot on Barro Colorado Island, Panama. Community trait distributions shifted significantly through time, which may indicate a shift toward more drought tolerant species. However, at the species level, changes in abundance were unrelated to trait values. To reconcile these seemingly contrasting results, we evaluated species-specific contributions to the directional shifts observed at the community level. Abundance changes of just one to six of 312 species were responsible for the community-level shifts observed for each trait. Our results demonstrate that directional changes in community-level functional composition can result from idiosyncratic change in a few species rather than widespread community-wide changes associated with functional traits. Future analyses of directional change in natural communities should combine community-, species-, and possibly individual-level analyses to uncover relationships with function that can improve understanding and enable prediction.

Ancient human disturbances may be skewing our understanding of Amazonian forests.

Although the Amazon rainforest houses much of Earth's biodiversity and plays a major role in the global carbon budget, estimates of tree biodiversity originate from fewer than 1,000 forest inventory plots, and estimates of carbon dynamics are derived from fewer than 200 recensus plots. It is well documented that the pre-European inhabitants of Amazonia actively transformed and modified the forest in many regions before their population collapse around 1491 AD; however, the impacts of these ancient disturbances remain entirely unaccounted for in the many highly influential studies using Amazonian forest plots. Here we examine whether Amazonian forest inventory plot locations are spatially biased toward areas with high probability of ancient human impacts. Our analyses reveal that forest inventory plots, and especially forest recensus plots, in all regions of Amazonia are located disproportionately near archaeological evidence and in areas likely to have ancient human impacts. Furthermore, regions of the Amazon that are relatively oversampled with inventory plots also contain the highest values of predicted ancient human impacts. Given the long lifespan of Amazonian trees, many forest inventory and recensus sites may still be recovering from past disturbances, potentially skewing our interpretations of forest dynamics and our understanding of how these forests are responding to global change. Empirical data on the human history of forest inventory sites are crucial for determining how past disturbances affect modern patterns of forest composition and carbon flux in Amazonian forests.

Thermophilization of adult and juvenile tree communities in the northern tropical Andes.

Climate change is expected to cause shifts in the composition of tropical montane forests towards increased relative abundances of species whose ranges were previously centered at lower, hotter elevations. To investigate this process of "thermophilization," we analyzed patterns of compositional change over the last decade using recensus data from a network of 16 adult and juvenile tree plots in the tropical forests of northern Andes Mountains and adjacent lowlands in northwestern Colombia. Analyses show evidence that tree species composition is strongly linked to temperature and that composition is changing directionally through time, potentially in response to climate change and increasing temperatures. Mean rates of thermophilization [thermal migration rate (TMR), °C ⋅ y(-1)] across all censuses were 0.011 °C ⋅ y(-1) (95% confidence interval = 0.002-0.022 °C ⋅ y(-1)) for adult trees and 0.027 °C ⋅ y(-1) (95% confidence interval = 0.009-0.050 °C ⋅ y(-1)) for juvenile trees. The fact that thermophilization is occurring in both the adult and juvenile trees and at rates consistent with concurrent warming supports the hypothesis that the observed compositional changes are part of a long-term process, such as global warming, and are not a response to any single episodic event. The observed changes in composition were driven primarily by patterns of tree mortality, indicating that the changes in composition are mostly via range retractions, rather than range shifts or expansions. These results all indicate that tropical forests are being strongly affected by climate change and suggest that many species will be at elevated risk for extinction as warming continues.

Large-scale patterns of turnover and Basal area change in Andean forests.

General patterns of forest dynamics and productivity in the Andes Mountains are poorly characterized. Here we present the first large-scale study of Andean forest dynamics using a set of 63 permanent forest plots assembled over the past two decades. In the North-Central Andes tree turnover (mortality and recruitment) and tree growth declined with increasing elevation and decreasing temperature. In addition, basal area increased in Lower Montane Moist Forests but did not change in Higher Montane Humid Forests. However, at higher elevations the lack of net basal area change and excess of mortality over recruitment suggests negative environmental impacts. In North-Western Argentina, forest dynamics appear to be influenced by land use history in addition to environmental variation. Taken together, our results indicate that combinations of abiotic and biotic factors that vary across elevation gradients are important determinants of tree turnover and productivity in the Andes. More extensive and longer-term monitoring and analyses of forest dynamics in permanent plots will be necessary to understand how demographic processes and woody biomass are responding to changing environmental conditions along elevation gradients through this century.

Compositional shifts in Costa Rican forests due to climate-driven species migrations.

Species are predicted to shift their distributions upslope or poleward in response to global warming. This prediction is supported by a growing number of studies documenting species migrations in temperate systems but remains poorly tested for tropical species, and especially for tropical plant species. We analyzed changes in tree species composition in a network of 10 annually censused 1-ha plots spanning an altitudinal gradient of 70-2800 m elevation in Costa Rica. Specifically, we combined plot data with herbarium records (accessed through GBIF) to test if the plots' community temperature scores (CTS, average thermal mean of constituent species weighted by basal area) have increased over the past decade as is predicted by climate-driven species migrations. In addition, we quantified the contributions of stem growth, recruitment, and mortality to the observed patterns. Supporting our a priori hypothesis of upward species migrations, we found that there have been consistent directional shifts in the composition of the plots, such that the relative abundance of lowland species, and hence CTS, increased in 90% of plots. The rate of the observed compositional shifts corresponds to a mean thermal migration rate (TMR) of 0.0065 °C yr(-1) (95% CI = 0.0005-0.0132 °C yr(-1) ). While the overall TMR is slower than predicted based on concurrent regional warming of 0.0167 °C yr(-1) , migrations were on pace with warming in 4 of the 10 plots. The observed shifts in composition were driven primarily by mortality events (i.e., the disproportionate death of highland vs. lowland species), suggesting that individuals of many tropical tree species will not be able to tolerate future warming and thus their persistence in the face of climate change will depend on successful migrations. Unfortunately, in Costa Rica and elsewhere, land area inevitably decreases at higher elevations; hence, even species that are able to migrate successfully will face heightened risks of extinction.

Directional changes in the species composition of a tropical forest.

Long-term studies have revealed that the structure and dynamics of many tropical forests are changing, but the causes and consequences of these changes remain debated. To learn more about the forces driving changes within tropical forests, we investigated shifts in tree species composition over the past 25 years within the 50-ha Forest Dynamics Plot on Barro Colorado Island (BCI), Panama, and examined how observed patterns relate to predictions of (1) random population fluctuations, (2) carbon fertilization, (3) succession from past disturbance, (4) recovery from an extreme El Niño drought at the start of the study period, and (5) long-term climate change. We found that there have been consistent and directional changes in the tree species composition. These shifts have led to increased relative representations of drought-tolerant species as determined by the species' occurrence both across a gradient of soil moisture within BCI and across a wider precipitation gradient from a dry forest near the Pacific coast of Panama to a wet forest near its Caribbean coast. These nonrandom changes cannot be explained by stochastic fluctuations or carbon fertilization. They may be the legacy of the El Niño drought, or alternatively, potentially reflect increased aridity due to long-term climate change. By investigating compositional changes, we increased not only our understanding of the ecology of tropical forests and their responses to large-scale disturbances, but also our ability to predict how future global change will impact some of the critical services provided by these important ecosystems.

Extinction risks of Amazonian plant species.

Estimates of the number, and preferably the identity, of species that will be threatened by land-use change and habitat loss are an invaluable tool for setting conservation priorities. Here, we use collections data and ecoregion maps to generate spatially explicit distributions for more than 40,000 vascular plant species from the Amazon basin (representing more than 80% of the estimated Amazonian plant diversity). Using the distribution maps, we then estimate the rates of habitat loss and associated extinction probabilities due to land-use changes as modeled under 2 disturbance scenarios. We predict that by 2050, human land-use practices will have reduced the habitat available to Amazonian plant species by approximately 12-24%, resulting in 5-9% of species becoming "committed to extinction," significantly fewer than other recent estimates. Contrary to previous studies, we find that the primary determinant of habitat loss and extinction risk is not the size of a species' range, but rather its location. The resulting extinction risk estimates are a valuable conservation tool because they indicate not only the total percentage of Amazonian plant species threatened with extinction but also the degree to which individual species and habitats will be affected by current and future land-use changes.

Assessing evidence for a pervasive alteration in tropical tree communities.

In Amazonian tropical forests, recent studies have reported increases in aboveground biomass and in primary productivity, as well as shifts in plant species composition favouring fast-growing species over slow-growing ones. This pervasive alteration of mature tropical forests was attributed to global environmental change, such as an increase in atmospheric CO2 concentration, nutrient deposition, temperature, drought frequency, and/or irradiance. We used standardized, repeated measurements of over 2 million trees in ten large (16-52 ha each) forest plots on three continents to evaluate the generality of these findings across tropical forests. Aboveground biomass increased at seven of our ten plots, significantly so at four plots, and showed a large decrease at a single plot. Carbon accumulation pooled across sites was significant (+0.24 MgC ha(-1) y(-1), 95% confidence intervals [0.07, 0.39] MgC ha(-1) y(-1)), but lower than reported previously for Amazonia. At three sites for which we had data for multiple census intervals, we found no concerted increase in biomass gain, in conflict with the increased productivity hypothesis. Over all ten plots, the fastest-growing quartile of species gained biomass (+0.33 [0.09, 0.55] % y(-1)) compared with the tree community as a whole (+0.15 % y(-1)); however, this significant trend was due to a single plot. Biomass of slow-growing species increased significantly when calculated over all plots (+0.21 [0.02, 0.37] % y(-1)), and in half of our plots when calculated individually. Our results do not support the hypothesis that fast-growing species are consistently increasing in dominance in tropical tree communities. Instead, they suggest that our plots may be simultaneously recovering from past disturbances and affected by changes in resource availability. More long-term studies are necessary to clarify the contribution of global change to the functioning of tropical forests.

The role of gap phase processes in the biomass dynamics of tropical forests.

The responses of tropical forests to global anthropogenic disturbances remain poorly understood. Above-ground woody biomass in some tropical forest plots has increased over the past several decades, potentially reflecting a widespread response to increased resource availability, for example, due to elevated atmospheric CO2 and/or nutrient deposition. However, previous studies of biomass dynamics have not accounted for natural patterns of disturbance and gap phase regeneration, making it difficult to quantify the importance of environmental changes. Using spatially explicit census data from large (50 ha) inventory plots, we investigated the influence of gap phase processes on the biomass dynamics of four 'old-growth' tropical forests (Barro Colorado Island (BCI), Panama; Pasoh and Lambir, Malaysia; and Huai Kha Khaeng (HKK), Thailand). We show that biomass increases were gradual and concentrated in earlier-phase forest patches, while biomass losses were generally of greater magnitude but concentrated in rarer later-phase patches. We then estimate the rate of biomass change at each site independent of gap phase dynamics using reduced major axis regressions and ANCOVA tests. Above-ground woody biomass increased significantly at Pasoh (+0.72% yr(-1)) and decreased at HKK (-0.56% yr(-1)) independent of changes in gap phase but remained stable at both BCI and Lambir. We conclude that gap phase processes play an important role in the biomass dynamics of tropical forests, and that quantifying the role of gap phase processes will help improve our understanding of the factors driving changes in forest biomass as well as their place in the global carbon budget.

Habitat fragmentation and effects of herbivore (howler monkey) abundances on bird species richness.

Habitat fragmentation can alter herbivore abundances, potentially causing changes in the plant community that can propagate through the food web and eventually influence other important taxonomic groups such as birds. Here we test the relationship between the density of red howler monkeys (Alouatta seniculus) and bird species richness on a large set of recently isolated land-bridge islands in Lago Guri, Venezuela (n = 29 islands). Several of these islands host relict populations of howler monkeys at densities up to more than 30 times greater than those on the mainland. These "hyperabundant" herbivores previously have been shown to have a strong positive influence on aboveground plant productivity. We predicted that this should lead to a positive, indirect effect of howler monkey density on bird species richness. After accounting for passive sampling (the tendency for species richness to be positively associated with island area, regardless of differences in habitat quality) we found a significant positive correlation between howler monkey density and bird species richness. A path analysis incorporating data on tree growth rates from a subset of islands (n = 9) supported the hypothesis that the effect of howler monkeys on the resident bird communities is indirect and is mediated through changes in plant productivity and habitat quality. These results highlight the potential for disparate taxonomic groups to be related through indirect interactions and trophic cascades.