Publications
Found 23 publication(s)
- of type article
Werner, F.A. & Homeier, J. (2024): Diverging elevational patterns of tree vs. epiphyte species density, beta diversity, and biomass in a tropical dry forest . Plants 13(18), 2555.
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DOI: 10.3390/plants13182555
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Abstract:
Abstract:
There is evidence to suggest that vascular epiphytes experience low competition for resources (light, water, and nutrients) compared to terrestrial plants. We tested the hypothesis that low resource competition may lead to higher nestedness among vascular epiphyte assemblages compared to trees. We studied the species composition and biomass of epiphytes and trees along an elevation gradient in a tropical dry forest in SW Ecuador. Both life-forms were inventoried on 25 plots of 400 m2 across five elevation levels (550–1250 m). Tree species density and total species richness increased with elevation, whereas basal area and biomass did not show significant trends. Epiphyte species density and richness both increased strongly with elevation, in parallel to biomass. Plot-level compositional changes were similarly strong for both life-forms. We attribute elevational increases in the species richness of trees and epiphytes to increasing humidity, i.e., more mesic growth conditions. We attribute the more pronounced elevational increase in epiphyte biomass, species density, and richness—the latter coupled with a higher degree of nestedness—to the greater moisture dependency of epiphytes and relatively low direct competition for resources. Our study provides a first comparison of elevational trends in epiphyte and tree diversity and biomass for a tropical dry forest.
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Keywords: |
biomass |
beta diversity |
species turnover |
Alpha diversity |
competition |
biotic interactions |
Graefe, S.; Rodrigo, R.; Cueva, E.; Butz, P.; Werner, F.A. & Homeier, J. (2020): Impact of disturbance on forest structure and tree species composition in a tropical dry forest of South Ecuador. Ecotropica 22, 202002.
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DOI: 10.30427/ecotrop202002
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Abstract:
Abstract:
Tropical dry forests (TDFs) are often degraded and fragmented through human impact, which is also the case in Southern Ecuador, where land-use pressure is high. In this context we studied tree species composition and forest structure in a protected and adjacent disturbed TDF at altitudes between 560-1080 m asl. Fabaceae and Malvaceae were identified as the most important tree families in both forest types. The disturbed forest displayed lower tree species richness than the protected forest, and the gap in species richness between the two forest types increased with increasing altitude. Ten species of the protected forest were not recorded in the disturbed forest, two of them endemic. The disturbed site was further characterized by a lower number of stems but with larger diameters, in comparison with the protected forest. The majority of the most abundant tree species in the disturbed forest had rather low wood densities, but also the combination of high wood density with browsing tolerance and high resprouting capacity was encountered, and seems to be advantageous for getting established in such sites. Although certain tree species were well represented in the disturbed forest, some endemic species with relatively low abundances (e.g. Simira ecuadorensis, Prockia crucis) should receive more conservation attention.
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Keywords: |
species richness |
altitude |
importance value index |
Tumbesian dry forest |
wood density |
Palomeque, X.; Günter, S.; Siddons, D.; Hildebrandt, P.; Stimm, B.; Aguirre, N.; Arias, R. & Weber, M. (2017): Natural or assisted succession as approach of forest recovery on abandoned lands with different land use history in the Andes of Southern Ecuador. New Forests 1(1), 1-10.
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DOI: 10.1007/s11056-017-9590-8
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Abstract:
Forest recovery on disturbed areas is of special significance in the Ecuadorian Andes, where deforestation is a serious problem. Natural diachronic succession was evaluated on three large plots or sites, differing in their land use and vegetation composition, one is dominated by grass species on an abandoned pasture (Pasture site), the other two are post-fire vegetation dominated by bracken (Bracken site) and various shrubs (Shrub site). Additionally, we assessed the effectiveness of manual removal of competitive herbaceous species to accelerate forest recovery. Monitoring was done in 2003, 2005 and 2007 on 48 subplots of 116 m2 each recording species richness and woody-species density. Results showed that the Pasture site demonstrated a competitive effect of exotic grasses on woody species recruitment with much lower species recruitment and density, suggesting serious inhibition of natural forest regeneration and an unclear successional trajectory. The Bracken and Shrub sites became significantly similar floristically and there is evidence for a marked facilitation of woody plant recruitment correlated with light availability on the ground. Both sites showed characteristics of classic succession, with Shrub showing a higher species richness and density while late-successional species are poorly represented on the Bracken site. However, NMDS ordination using species density showed that the two trajectories may not be converging towards a common ‘‘final state’’. Manual weeding was ineffective for accelerating forest recovery. These results suggest that the main limiting factor for the recruitment of woody species on the Pasture site is strong grass competition and must be addressed before seed availability, while seed availability seems to be theconstraining factor for Bracken and Shrub site development, thus direct seeding or planting may be effective in accelerating forest recovery.
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Keywords: |
succession |
abandoned pasture |
bracken fern |
restoration |
Werner, F.A.; Köster, N.; Kessler, M. & Gradstein, S.R. (2011): Is the resilience of epiphyte assemblages to human disturbance a function of local climate?. Ecotropica 17, 15-20.
Kotowska, M. & Werner, F.A. (2013): Environmental controls over methane emissions from bromeliad phytotelmata: The role of phosphorus and nitrogen availability, temperature, and water content. GLOBAL BIOGEOCHEMICAL CYCLES 27, 1-8.
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DOI: 10.1002/2013GB004612
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Abstract:
Abstract:
Tank bromeliads are common epiphytic plants throughout neotropical forests that store
signi?cant amounts of water in phytotelmata (tanks) formed by highly modi?ed leafs.
Methanogenic archaea in these tanks have recently been identi?ed as a signi?cant source of
atmospheric methane. We address the effects of environmental drivers (temperature, tank
water content, sodium phosphate [P], and urea [N] addition) on methane production in
anaerobically incubated bromeliad slurry and emissions from intact bromeliad tanks in
montane Ecuador. N addition ? 1 mg g 1 had a signi?cantly positive effect on headspace
methane concentrations in incubation jars while P addition did not affect methane
production at any dosage (? 1 mg g 1 ). Tank bromeliads (Tillandsia complanata) cultivated
in situ showed signi?cantly increased ef?uxes of methane in response to the addition of
26 mg N addition per tank but not to lower dosage of N or any dosage of P (? 5.2 mg plant 1 ).
There was no signi?cant interaction between N and P addition. The brevity of the
stimulatory effect of N addition on plant methane ef?uxes (1–2 days) points at N
competition by other microorganisms or bromeliads. Methane ef?ux from plants closely
followed within-day temperature ?uctuations over 24 h cycles, yet the dependency of
temperature was not exponential as typical for terrestrial wetlands but instead linear. In
simulated drought, methane emission from bromeliad tanks was maintained with minimum
amounts of water and regained after a short lag phase of approximately 24 h. Our results
suggest that methanogens in bromeliads are primarily limited by N and that direct effects of
global change (increasing temperature and seasonality, remote fertilization) on bromeliad
methane emissions are of moderate scale.
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Keywords: |
nutrients |
gas emission |
NUMEX |
N-cycle |
nitrogen |
Gas exchange |
phosphorus |
nutrient manipulation |
nutrient limitation |
phosphorus availability |
nutrient cycle |
methane |
Werner, F.A. & Homeier, J. (2014): Is tropical montane forest heterogeneity promoted by a resource-driven feedback cycle? Evidence from nutrient relations, herbivory and litter decomposition along a topographical gradient. Functional Ecology x(x), x.
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DOI: 10.1111/1365-2435.12351
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Abstract:
Abstract:
1. Ridges of tropical mountains often differ strikingly from neighbouring ravines in terms of
forest structure, productivity and species composition. This heterogeneity is poorly understood
despite its critical role in biodiversity maintenance, carbon and nutrient budgets.
2. We examined measures of tree biomass and productivity, foliage and litter quality (nutrient
concentrations, specific leaf mass, phenolics), herbivory and leaf litter decomposition in each
six plots laid out in upper and lower slope position in a tropical montane moist forest in southeastern
Ecuador.
3. Productivity, quality of foliage and litter as well as herbivory were significantly lower in
upper slope position, and closely correlated with soil nutrient concentrations and accumulated
humus. The decomposition of upper slope leaf litter (decomposition rate k) was substantially
lower than in litter from lower slope forest, whereas the site of decomposition (slope position)
only had a marginal effect on the decomposition rate.
4. Our results suggest that the differences in stand structure, productivity, foliar quality, herbivory
and decomposition between slope positions are ultimately due to stronger nutrient limitations
in upper slope forest. We propose a general conceptual model that explains origin and
maintenance of contrasting forest types along topographical gradients through down-slope
fluxes of nutrients and water, and a nutrient-driven positive feedback cycle.
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Keywords: |
topographic heterogenity |
Jantz, N.; Homeier, J.; León Yánez, S.; Moscoso, A. & Behling, H. (2013): Trapping pollen in the tropics — Comparing modern pollen rain spectra of different pollen traps and surface samples across Andean vegetation zones. Review of Palaeobotany and Palynology 193, 57-69.
Villota, A.; León Yánez, S. & Behling, H. (2012): Vegetation and environmental dynamics in the Páramo of Jimbura region in the southeastern Ecuadorian Andes during the late Quaternary. Journal of South American Earth Sciences 40, 85-93.
Werner, F.A.; Homeier, J.; Oesker, M. & Boy, J. (2011): Epiphytic biomass of a tropical montane forest varies with topography. Journal of Tropical Ecology 28, 23-31.
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DOI: 10.1017/S0266467411000526
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Abstract:
Abstract:
The spatial heterogeneity of tropical forest epiphytes has rarely been quantified in terms of biomass. In particular, the effect of topographic variation on epiphyte biomass is poorly known, although forests on ridges and ravines can differ drastically in stature and exposure. In an Ecuadorian lower montane forest we quantified epiphytic biomass along two gradients: (1) the twig-branch-trunk trajectory, and (2) the ridge-ravine gradient. Twenty-one trees were sampled in each of three forest types (ridge, slope, ravine positions). Their epiphytic biomass was extrapolated to stand level based on basal area?epiphyte load relationships, with tree basal areas taken from six plots of 400 m 2 each per forest type. Our results document the successional addition and partial replacement of lichens by bryophytes, angiosperms and finally dead organic matter along the twig-branch-trunk trajectory. Despite having the highest tree basal area, total epiphytic biomass (mean ± SD) of ravine forest was significantly lower (2.6 ± 0.7 Mg half 1) than in mid-slope forest (6.3 ± 1.1 Mg half 1) and ridge forest (4.4 ± 1.6 Mg half 1), whereas maximum bryophyte water storage capacity was significantly higher. We attribute this pattern to differences in forest dynamics, stand structure and microclimate. Although our study could not differentiate between direct effects of slope position (nutrient availability, mesoclimate) and indirect effects (stand structure and dynamics), it provides evidence that fine-scale topography needs to be taken into account when extrapolating epiphytic biomass and related matter fluxes from stand-level data to the regional scale.
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Keywords: |
Ecuador |
succession |
epiphyte |
crown humus |
dead organic matter |
carbon storage |
maximum water storage capacity |
topographic heterogenity |
Werner, F.A. (2011): Reduced growth and survival of vascular epiphytes on isolated remnant trees in a recent tropical montane forest clear-cut. Basic and Applied Ecology 12, 172-181.
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DOI: 10.1016/j.baae.2010.11.002
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Abstract:
Abstract:
Forest fragmentation can negatively affect tropical epiphyte diversity, but the processes leading to such impoverishment are insufficiently understood. Due to a lack of experimental studies, the relative influence of dispersal constraints vs. growth conditions remains particularly controversial. This paper addresses the fate of late juvenile and adult vascular epiphytes in response to severe forest disturbance in montane southern Ecuador. Plant growth and survival on trunks and lower branches of isolated remnant trees was studied for the first three years following clear-cutting. Overall epiphyte mortality was substantially increased on remnant trees (72% over 3 years) relative to undisturbed forest (11%). Mortality on remnant trees was higher during the first year (52%) than during the second (20%) and third year (26%). Pteridophytes and dicots suffered higher losses than monocots. Plants surviving on remnant trees generally showed a marked negative growth regarding maximum leaf length, whereas the annual increment in leaf number varied more strongly among taxa (families). The present study provides the first field-experimental evidence for the adverse effects of forest disturbance on the performance of later, well-established life stages of vascular epiphytes. The results suggest that growth conditions may often be a more important predictor of epiphyte diversity in disturbed habitats than dispersal constraints. Similar plant responses can be expected to occur along forest edges. Therefore, the retention of scattered green trees, narrow strips or small fragments of forest are unlikely to be sufficient management tools for the conservation of epiphyte diversity in tropical landscapes.
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Keywords: |
growth |
land-use change |
human disturbance |
diversity |
microclimate |
edge effects |
forest fragmentation |
population dynamics |
scattered trees |
Werner, F.A. & Larrea, M.L. (2010): Response of vascular epiphyte diversity to different land-use intensities in a neotropical montane wet forest. Forest Ecology and Management 260, 1950-1955.
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DOI: 10.1016/j.foreco.2010.08.029
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Abstract:
Abstract:
Although vascular epiphytes contribute substantially to the biodiversity of tropical montane forests, it is
unclear how their diversity and community composition is affected by forest alteration. We studied the
response of vascular epiphyte assemblages to different intensities of land-use in a montane wet forest of
northeastern Ecuador: (1) unmanaged mature forest; (2) mature forest with mid- and understorey opened
for cattle grazing; and (3) isolated remnant trees in cattle pastures. The numbers of individuals and species
of epiphytes per host tree did not differ significantly between land-use types, neither did total epiphyte
species richness (n = 30 trees). However, total species richness of pteridophytes was significantly lower on
isolated remnant trees compared to unmanaged forest, whereas several taxa rich in xerotolerant species
(Bromeliaceae, Orchidaceae, Piperaceae) exhibited the opposite trend. An analysis of floristic composition
using ordination (NMS) and randomisation techniques (MRPP) showed that epiphyte assemblages on
isolated remnant trees were significantly distinct from unmanaged forest while managed forest was
intermediate between those two vegetation types. Ordination analysis further indicated reduced floristic
heterogeneity in disturbed habitats. These results suggest considerable, rapid species turnover since
land-use change 6 years prior to study, with pteridophytes being replaced by more xerotolerant taxa.
We attribute this floristic turnover primarily to changes in microclimate towards higher levels of light
and desiccation stress associated with forest disturbance. Our results support the notion that community
composition offers a more sensitive indicator of human disturbance than species richness.
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Keywords: |
land-use change |
human disturbance |
microclimate |
deforestation |
beta diversity |
fragmentation |
isolated trees |
secondary forest |
species richness |
species turnover |
Martinson, G.; Werner, F.A.; Scherber, C.; Conrad, R.; Corre, M.; Flessa, H.; Wolf, K.; Klose, M.; Gradstein, S.R. & Veldkamp, E. (2010): Methane emissions from tank bromeliads in neotropical forests. Nature Geoscience 2010(3), 766-769.
Ledesma, K.J.; Werner, F.A.; Spotorno, A. & Albuja, L.H. (2009): A new species of Mountain Viscacha (Chinchillidae: Lagidium Meyen) from the Ecuadorean Andes. Zootaxa 2126, 41-57.
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Abstract:
A new species of mountain viscacha, Lagidium ahuacaense, is described based on a specimen and a mitochondrial cytochrome b (cyt b) sequence obtained from a second individual from Cerro El Ahuaca, Loja Province, Ecuador. In several external and craniodental measurements, the new species differed significantly from the three congeneric species (greatest length of skull, basilar, nasal length, palatilar length, length of diastema, least interorbital breadth, breadth of rostrum and skull height). The cyt b sequence of the Ecuadorean viscacha differed by 14 exclusive nucleotide substitutions from all other sequences of Lagidium examined. Kimura 2-parameter (K2P) genetic distances of the Ecuadorean sequence were 8.1?11.0% to L. peruanum, 7.9?9.9% to L. viscacia and 9.7% to L. wolffsohni. The single known population of the newly described species may not comprise more than a few dozen individuals and warrants urgent conservation actions.
Lehnert, M.; Kessler, M.; Salazar, L.I.; Navarrrete, H.; Werner, F.A. & Gradstein, S.R. (2007): Pteridophytes. Checklist Reserva Biológica San Francisco (Prov. Zamora-Chinchipe, S. Ecuador). Ecotropical Monographs 4, 59-68.
Homeier, J. & Werner, F.A. (2007): Spermatophyta. Checklist of the Reserva Biológica San Francisco (Prov. Zamora-Chinchipe, S-Ecuador). Ecotropical Monographs 4, 15-58.
Nöske, N.; Hilt, N.; Werner, F.A.; Brehm, G.; Fiedler, K.; Sipman, H.J. & Gradstein, S.R. (2008): Disturbance effects on diversity of epiphytes and moths in a montane forest in Ecuador. Basic and Applied Ecology 9, 4-12.
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DOI: 10.1016/j.baae.2007.06.014
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Abstract:
Abstract:
We sampled the diversity of epiphytes (lichens, bryophytes, vascular plants) and moths (Geometridae, Arctiidae) in
mature and recovering forest and in open vegetation in the montane belt in Ecuador. No uniform pattern of change in
species richness was detected among the different taxonomic groups with increasing disturbance. Species richness of
epiphytic bryophytes and vascular plants declined significantly from mature forest towards open vegetation. In
contrast, species richness of epiphytic lichens did not change with increasing forest alteration, while that of geometrid
moths was significantly higher in recovering forest compared with mature forest and open habitats. Arctiidae were
significantly more species-rich in recovering forest and open vegetation than mature forest. Hence, for some organisms,
modified habitats may play an important role for biodiversity conservation in the Andes, whereas others suffer from
habitat disturbance. However, trends of changes in species composition following deforestation were surprisingly
concordant across most studied epiphyte and moth taxa.
Werner, F.A.; Ledesma, K.J. & Hidalgo, R. (2006): Mountain viscacha (Lagidium cf. peruanum) in Ecuador - a first record of Chinchillidae for the northern Andes. Mastozoología Neotropical 13, 271-274.
Matt, F. & Werner, F.A. (2007): Mammals. Checklist of the Reserva Biológica San Francisco (Prov. Zamora-Chinchipe, S-Ecuador). Ecotropical Monographs 4, 125-129.
Werner, F.A. & Gradstein, S.R. (2008): Seedling establishment of vascular epiphytes on isolated and enclosed forest trees in an Andean landscape, Ecuador. Biodiversity and Conservation 17(13), 3195-3207.
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DOI: 10.1007/s10531-008-9421-5
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Abstract:
Abstract:
The impact of human disturbance on colonisation dynamics of vascular epiphytes
is poorly known. We studied abundance, diversity and floristic composition of epiphyte
seedling establishing on isolated and adjacent forest trees in a tropical montane landscape. All
vascular epiphytes were removed from plots on the trunk bases of Piptocoma discolor. Newly
established epiphyte seedlingswere recorded after 2 years, and their survival after another year.
Seedling density, total richness at family and genus level, and the number of families and genera
per plot were significantly reduced on isolated trees relative to forest trees. Seedling assemblages
on trunks of forest trees were dominated by hygrophytic understorey ferns, those on
isolated trees by xerotolerant canopy taxa. Colonisation probability on isolated trees was
significantly higher for plots closer to forest but not for plots with greater canopy or bryophyte
cover. Seedling mortality on isolated trees was significantly higher for mesophytic than for
xerotolerant taxa. Our results show that altered recruitment can explain the long-term impoverishment
of post-juvenile epiphyte assemblages on isolated remnant trees.We attribute these
changes to a combination of dispersal constraints and the harshermicroclimate documented by
measurements of temperature and humidity. Although isolated trees in anthropogenic landscapes
are considered key structures for themaintenance of forest biodiversity inmany aspects,
our results show that their value for the conservation of epiphytes can be limited.We suggest
that abiotic seedling requirementswill increasingly constitute a bottleneck for the persistence of
vascular epiphytes in the face of ongoing habitat alteration and atmospheric warming.
Werner, F.A. & Gradstein, S.R. (2009): Diversity of dry forest epiphytes along a gradient of human disturbance in the tropical Andes. Journal of Vegetation Science 20(1), 59-68.
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DOI: 10.3170/2008-8-18466
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Abstract:
Abstract:
Question: Disturbance effects on dry forest epiphytes are poorly known. How are epiphytic assemblages affected by different degrees of human disturbance, and what are the driving forces?
Location: An inter-Andean dry forest landscape at 2300 m elevation in northern Ecuador.
Methods: We sampled epiphytic bryophytes and vascular plants on 100 trees of Acacia macracantha in five habitats: closed-canopy mixed and pure acacia forest (old secondary), forest edge, young semi-closed secondary woodland, and isolated trees in grassland.
Results: Total species richness in forest edge habitats and on isolated trees was significantly lower than in closed forest types. Species density of vascular epiphytes (species per tree) did not differ significantly between habitat types. Species density of bryophytes, in contrast, was significantly lower in edge habitat and on isolated trees than in closed forest. Forest edge showed greater impoverishment than semi-closed woodland and similar floristic affinity to isolated trees and to closed forest types. Assemblages
were significantly nested; habitat types with major disturbance held only subsets of the closed forest assemblages, indicating a gradual reduction in niche availability. Distance to forest had no effect on species density of epiphytes on isolated trees, but species density was closely correlated with crown closure, a measure of canopy integrity.
Main conclusions: Microclimatic changes but not dispersal constraints were key determinants of epiphyte assemblages following disturbance. Epiphytic cryptogams are sensitive indicators of microclimate and human disturbance in montane dry forests. The substantial impoverishment of edge habitat underlines the need for fragmentation studies on epiphytes elsewhere in the Tropics.
Werner, F.A.; Homeier, J. & Gradstein, S.R. (2005): Diversity of vascular epiphytes on isolated remnant trees in the montane forest belt of southern Ecuador. Ecotropica 11, 21-40.