Tropical montane forests (TMFs) are found on most of Earth's continents along variable
elevation ranges, whose potential upper limits are influenced by cloud condensation
heights and minimum temperatures. They are most widespread in South America and in
(semi-)humid mountain areas (Richter, 2008). According to FAO and UNEP (2020), the
area covered by tropical and subtropical montane forests is around 305 million hectares,
about 13% of the area covered by tropical and subtropical forests. Their elevational
limits are difficult to establish due to the interactions of the different factors
that determine their characteristics. Among these, geomorphology plays a leading role
in regulating TMF structure, and provides useful clues on the contributing mechanisms.
Most TMFs occur under highly variable topography, including steep slopes (Asner et
al., 2014) and landslide-prone terrain (Shreve, 1914; Larsen and Torres-Sánchez, 1998).
Also, the latitudinal gradient, orography, and vertical thermal gradients have a direct
influence on the fauna and flora of TMFs. The latitudinal pattern is not the same
in all TMFs, the temperature and precipitation conditions occur due to seasonality
in the climate and are unambiguously linked to species climatic affinity preferences
(Ohsawa, 1991; Cuesta et al., 2016; Chu et al., 2019). Another factor is the annual
precipitation that generally exceeds c. 1,000–1,200 mm and can be associated with
low level cloud cover or mist, which results in a lower incidence of sunlight and
lower primary productivity, suggesting that NPP for these forests is driven by changes
in photosynthesis. This highlights the importance of variations in solar radiation.
Girardin et al. (2010) estimated that NPP values recorded in TMFs range widely between
4 and 7 Mg C ha−1 yr−1. Despite this variability, TMFs store significant amounts of
carbon in their soils. Malhi et al. (2017) showed that the soil organic layer depth
sharply increased with lower mean annual temperatures. Lower temperatures also result
in low nutrient inputs through slow mineralization of organic matter (Townsend et
al., 1995). Declining temperature appears to be the principal rate-limiting factor
for litter decay with increasing elevation on tropical mountains (Schuur, 2001; Salinas
et al., 2011). The low temperatures have also been linked to biogeochemical limitations,
by reducing nitrogen availability (Nottingham et al., 2015, 2018b) and N2 fixation
(Houlton et al., 2008). However, biogeochemical cycling in TMFs is further affected—returning
to our primary driver—by geomorphology via landslide activity and uplift, which increases
the supply of rock-derived nutrients such as phosphorus (Tanner et al., 1992; van
de Weg et al., 2012; Nottingham et al., 2015). The novel environments TMFs represent
are, thus, a product of interconnected geological and climatic forces.
Current global climate models (GCMs) suggest enhanced warming of the tropical mid
and upper troposphere (Fu et al., 2011). Consequently, rates of temperature rise are
expected to be larger at higher than at lower elevations, as it has already been reported
in mountains around the world (Bradley et al., 2006; Roman-Cuesta et al., 2014). There
is still uncertainty on the effects that temperature and changing moisture conditions
will have on the cloud belt formation in TMFs (Lawton et al., 2001) but upper displacements
of the condensation belt are expected (Foster, 2001; Halladay et al., 2012). Moreover,
mountain regions are more frequently suffering the impacts of oceanic warming such
as El Nino Southern Oscillation ENSO and North Atlantic Oscillation (NAO)/Atlantic
Multidecadal Oscillation (AMO), but the effect of their drought and flooding spells
on TMFs' flora and fauna are yet under-researched (Foster, 2001; Roman-Cuesta et al.,
2014; Oliveras et al., 2017).
Tropical montane forest ecosystems are fragile but exceedingly valuable ecosystems,
due to their important role in the provision of ecosystem services, including the
regulation of water and the regional climate (Bubb et al., 2004), the capture and
storage of carbon (Cuesta et al., 2009; Tejedor Garavito et al., 2012) and—not least—by
harboring a vast store of biodiversity (Myers, 1995). The complex spatial and environmental
gradients typical of TMFs generate a high diversity of habitats. TMFs are considered
among the most biologically diverse and richest ecosystems on Earth (Kessler and Kluge,
2008; Richter, 2008), and recognized as hotspots of species endemism (Gentry, 1993).
These vital ecosystems services are under threat, as climate change is undoubtedly
affecting their species composition and metabolic profiles in a myriad of ways. Along
an elevation gradient, as global average temperatures rise, elevational shifts in
the distribution of species toward regions of lower temperature are to be expected.
A major concern is that the speed of climate change appears to be greater than the
response capacity (adaptation and migration) for a large number of species in the
Andean Amazon. On the one hand, adaptation within species or communities may result
in the increased dominance of individuals more tolerant of change. On the other hand,
species extinction may occur alongside migration and geographic displacement of susceptible
populations toward areas with a more appropriate climate. This implies a high probability
of extinction for plant species without this response capacity, which in turn would
lead to changes in the carbon cycle, in the dynamics of ecosystems and uncertain impacts
on wildlife. But these changes are evidenced not only in animals and plants, soil
bacterial, and fungal communities on tropical mountains are also sensitive to temperature
(Looby et al., 2016; Nottingham et al., 2018a) and may be affected by rapid climate
warming with negative implications for carbon storage (Nottingham et al., 2019) and
for plant species composition (Corrales et al., 2016). At the same time, these ecosystems
are in a state of global threat due to the dynamics of change in land cover and use
(Webster, 1995; Bubb et al., 2004). In many regions, land use patterns have created
a mosaic of habitats transformed through the expansion of human activities. These
fragmented forests should receive more attention when designing conservation policies.
For all these reasons climate change can have severe impacts on montane tropical ecosystems
by generating changes in the life zones, increasing the vulnerability of forests to
fires, pests, invasive species, and greater deforestation pressure due to the establishment
of productive systems with intensive management (Serreze, 2009).
The articles in this special issue aim to fill some of the existing gaps in our knowledge.
These studies were conducted in a wide range of TMFs from pristine forests in protected
areas to those with varying degrees of human disturbance along South America, Africa,
and Asia. These studies have examined TMF responses to environmental cues in forest
plots using a variety of tools which include remote sensing, on-site instrumentation,
biometrics, and allometry, among others, to model field data and provide us with:
Two contributions to this special issue used leaf chemistry and traits analysis to
determine if plant species are sensitive to changes in environmental conditions. Martin
et al. assessed differences in 19 foliar traits in paired sun and shade leaves along
a humid tropical forest elevation gradient in Peru, to determine if foliar chemical
traits, such as photosynthetic pigments, and other leaf traits like LMA differ between
them, and if the sources for these variations are environmental or genetic. They found
that for most traits (i.e., N, foliar nutrients or defense compounds), there was no
significant difference between sun and shade leaves. Other traits for growth, such
as LMA and δ13C concentrations, maintain constant offsets, suggesting that the characteristics
of shade leaves can be derived from those measured in sun leaves. Their findings also
indicate that variation in sunlit canopy foliar traits are controlled primarily by
changes in community composition, and secondarily by environmental factors, like elevation
or substrate. They conclude that there are significant differences in light-sensitive
traits between sun and leaves evaluated, that were maintained across a wide variety
of environmental conditions along a 3,500-m elevation gradient suggesting that plasticity
associated with light availability is an adaptive change. In contrast, they did not
find sun-shade differences in other foliar traits related to defense and metabolism.
Gong et al. evaluated the protective function and phylogenetic relationships of the
transient red coloration of young leaves in some tropical plant species. They investigated
the metabolism, photosynthetic activities, and chemical defenses of leaves from 250
tropical plant species with either red or green young leaves, in a tropical region
of southwest China. They found that the occurrence of transient reddening of juvenile
leaves in the tropics was coupled with increased levels of both anthocyanins and tannins
and that the red coloration protects them from insect herbivory primarily through
chemical defense. Also, the red coloration in young leaves is predominantly a result
of adaptation to special tropical environmental conditions but without a significant
intrinsic phylogenetic relationship between plant species and suggested that the anthocyanins
might not function as light attenuators to protect for effects of high light intensity.
Two studies in this collection illustrate photosynthetic plant function related to
light and leaf nutrients. Feeley et al. assessed the maximum photosynthetic thermal
tolerances of more than 550 individuals of 164 tropical canopy tree species growing
in a steep elevation gradient ascending from near sea level to near tree line in the
northern Andes mountains of Colombia. They analyzed changes in plant thermal tolerances
between elevations at the species and community level and tested the relationship
between species' thermal tolerances and their changes in abundance through time in
10 forest inventory plots. They found a high amount of variation in the maximum thermal
tolerance (T 50) among species co-occurring within each plot and that this tolerance
decreases with plot elevation. However, their results also indicate that the relationship
between T 50 and temperature is weak and extremely shallow. Ziegler et al. investigated
physiological, chemical, and structural properties of leaves in mature individuals
belonging to 12 tree species in a tropical montane rainforest in Rwanda, Central Africa.
In this study, they explored the relative importance of area-based total leaf N content
and within-leaf N allocation to photosynthetic capacity vs. light-harvesting in controlling
the variation in photosynthetic capacity to explore the controls of interspecific
variation in photosynthetic capacity and other leaf gas exchange traits. They found
that photosynthetic capacity at a common leaf temperature of 25°C was higher in early
succession species than in late. However, total leaf N content did not significantly
differ between successional groups and there was no significant trade-off between
relative leaf N investments in compounds maximizing photosynthetic capacity vs. compounds
maximizing light harvesting.
Litton et al. provide information complementary other studies in this issue by examining
how litterfall, live foliar nutrient concentration, foliar nutrient resorption efficiency,
nutrient return via litterfall, and nutrient use e?ciency vary with mean annual temperature
in two dominant tree species in a gradient in Hawaii. Their aim was to understand
how increasing mean annual temperature impacts on the availability and ecological
stoichiometry of macro and micronutrients. Their findings provide strong evidence
that increased mean annual temperature alters the cycling and availability of a broad
suite of nutrients in TMFs, with important implications for nutrient limitation to
ecosystem processes in a warming world.
de la Cruz-Amo et al. explore the role of Andean TMFs as carbon reservoirs. They calculated
the amount of carbon in aboveground and belowground carbon stocks, and in soil organic
matter, along two elevation gradients in the southeast of Ecuador and North-Central
Peru. They assess how carbon stocks vary along elevation gradients and determine the
influence of climate, particularly precipitation seasonality, on the distribution
of these stocks across different forest compartments. They report that the combination
of annual mean temperature and precipitation seasonality explains the differences
in mean total carbon stocks in these three compartments and also show different partitioning
patterns along the elevation gradients both in Ecuador and Peru but that total carbon
stocks do not change with elevation in either site.
Two studies report results analyzing the literature. Soh et al. systematically mapped
all research on the effects of habitat degradation in TMFs globally to identify deficiencies
in current knowledge and to guide future research prioritization. After a comprehensive
review, they show that habitat degradation in TMFs impacts biodiversity at all ecological
levels and is compounded by climate change. However, despite montane species being
perceived as more extinction-prone, there are some indications of biotic resilience
if disturbance in TMFs is less severe. They confirm that TMFs also provide important
ecosystem services being the most important water provision, but that in recent years
these ecosystems have come under human pressure manifested in the form of highest
rates of deforestation. They highlight the poor research representation of Asian and
African TMFs and list the top research priorities which, if addressed, would advance
the goals of biodiversity conservation and sustainable use of resources in TMFs.
Tito et al. present an analysis of the natural variation of abiotic and biotic factors
along mountain elevation gradients based on two papers that used field experiments
conducted along an elevation gradient in the Peruvian tropical Andes. They highlight
the potential for use of field experiments in future studies focused on determining
the direct and indirect e?ects of climate change. They conclude that, despite abundant
research on the effects of global change climate on TMFs, fundamental questions remain
unanswered and it will be necessary to apply more fine-scale experimental approaches
to help better predict future abundance and distribution patterns of species under
altered climate scenarios, and that natural gradients are essential to quickly gain
a more complete understanding on the possible impacts of climate change.
To date, there have been few experimental studies using artificial material to explore
predation conduct. Murray et al. carried out predation experiments in Peninsular Malaysia
at a landscape scale and across a suite of sites of varying disturbance. They used
four di?erent prey items—artificial nests, artificial seeds, caterpillar models, and
frog models—along a disturbance gradient, from pristine forests to tea plantations.
Their purpose was to assess whether predation probability in different habitat types
di?ers between mountain ranges, if this probability consistently varies in di?erent
habitat types and if predation can be explained by vegetation structure. Their results
show that they confirm the first and third hypotheses, but there is no clear trend
in predation probability along a habitat disturbance gradient.
Results from this collection of articles show that responses to Global Change can
vary greatly among species, ecosystems, and even microsites in TMFs. This suggests
that the fate of these forests in response to climate change and greater deforestation
pressure due to the establishment of productive agricultural systems with intensive
management, can have severe impacts on montane tropical ecosystems by generating changes
in vegetation life zones, increasing their vulnerability to fires, pests, invasive
species, and having dramatic consequences on downstream ecosystem services. We hope
that this selection of papers will stimulate interest and research into those wonderful
ecosystems that are TMFs and their elevation gradients, and provide insights into
the future of TMF ecosystems in a century of rapid climate change.
Author Contributions
NS drafted the first version of the editorial. EC, MS, PM, AN, RR-C, and YM made edits,
additions, and revisions. All authors contributed to the article and approved the
submitted version.
Author Disclaimer
Any use of trade, firm, or product names is for descriptive purposes only and does
not imply endorsement by the U.S. Government.
Conflict of Interest
The authors declare that the research was conducted in the absence of any commercial
or financial relationships that could be construed as a potential conflict of interest.
Publisher's Note
All claims expressed in this article are solely those of the authors and do not necessarily
represent those of their affiliated organizations, or those of the publisher, the
editors and the reviewers. Any product that may be evaluated in this article, or claim
that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.