Edge effect and structure of mixed-species bird flocks in an Afrotropical lowland forest

Avian mixed-species flocks are a dominant feature of tropical moist forests, yet their cost–benefit balance and habitat dependence in Africa are not fully documented. We recorded the composition of mixed-species bird flocks in a pristine Afrotropical lowland forest site in Salonga NP, DRC. Our data showed that at least four types of flock existed, one of which specialized on edge habitats. We used multivariate analyses to further characterize edge effect on the most documented mixed-species flock type and found a significant effect on flock composition. While neither species guild nor preferential foraging stratum played an important role in flock participation, the nucleus role was played by a different species at the edge and in the interior, and both species abundance and associations were habitat-dependant, suggesting “domino effects” on the structure of mixed-species flocks.     

Composition and structure of a humid montane forest on the Pasochoa volcano, Ecuador

In a one-hectare plot in humid montane forest on Volcán Pasochoa (3,260–3,300 m), one of the last remnants of the inter-Andean forest between the two Ecuadorean cordilleras, there were 1,058 trees ≥ 5 cm dbh in 32 different species with a total basal area of 25.7 m². The most common species were Miconia theaezans, Piper andreanum , and Miconia pustulata with 32%, 14% and 8% of the individuals, respectively. Three species ( Hesperomeles lanuginosa, Boehmeria fallax and Lamiaceae sp.) were represented by only a single individual. Melastomataceae was the most important family with six species, over 50% of the individual trees, and 68% of the basal area. Other important families were Piperaceae and Asteraceae. The abundance of assumed pioneer species suggests that either the forest has been altered by humans in the past, or that their dominance is a part of the dynamic regeneration of a forest growing on steep slopes with inclinations between 36 and 120% in the plot. The diameter class distribution of Miconia corymbiformis and Axinaea macrophylla shows a light-demanding species behavior. Trees with multiple stems characterize this forest perhaps in order to distribute their weight on such steep slopes.     

Breeding systems in a montane tropical cloud forest in Venezuela

The breeding systems, reproductive efficacies and population densities of 75 species of trees, shrubs, perennial herbs and vines in a montane tropical cloud forest in Venezuela are investigated. 56.96% and 44.32% of the trees, versus the other life forms considered, respectively possess obligate outbreeding mechanisms. Two shrubs are non-pseudogamous apomicts. The percentage of dioecy among tree species (31%) is among the highest recorded in tropical forests. Reproductive efficacy is similar under all breeding systems in the forest interior. Obligately outbred taxa are slower recolonizers of a disturbed border as compared with non-obligate outbreeders. Explanations are advanced for the high incidence of dioecy combined with a low level of self-incompatibility among hermaphroditic species. It is concluded that the breeding system spectrum reflects an unpredictable pollination pattern, rather than insufficient pollinator servicing.     

Topographic and spatial controls of palm species distributions in a montane rain forest, southern Ecuador

The northern Andes harbour a flora that is as species-rich or even richer than the 18-times larger lowland Amazon basin. Gaining an understanding of how the high species richness of the Andean region is generated and maintained is therefore of particular interest. Environmental sorting due to elevational gradients in climate has been emphasized as a driver of vegetation distribution and plant community assembly in tropical mountain areas such as the Andes for two centuries, while alternative mechanisms have been little studied. Here, we investigated the importance of topography and spatial location as factors controlling species distributions in a palm community in a montane rain forest landscape in the Andes of southern Ecuador (1900–2150 m above sea level). Eleven species were present: Aiphanes verrucosa, Ceroxylon parvifrons, Chamaedorea pinnatifrons, Dictyocaryum lamarckianum, Euterpe precatoria, Geonoma densa, Geonoma orbignyana, Geonoma paradoxa, Prestoea acuminata and Wettinia aequatorialis. To study their spatial distribution, forty 250 m² (5 × 50 m²) plots were laid out perpendicular to four paths that were categorized into three areas and two topographic units (ridges and gullies). Mantel tests and indicator species analysis showed that both topography and spatial location imposed strong controls on palm species distributions at the study site. Our results suggest that species distributions in the studied montane forest landscape were partly determined by the species’ habitat requirements, but also by unknown spatial effects. Although a number of possible explanations exist for the latter, such as unmeasured environmental variables and historical disturbance events, we believe dispersal limitation is likely to be involved. Furthermore, although the gully- or ridge-association of some species corresponded to their general elevational ranges in southern Ecuador, this was not the case for other species. Based on such considerations, we conclude that elevational climatic gradients are likely to only form part of the explanation for the topographic effects on palm species distributions at the study site. Other factors must also be involved, notably wind-exposure and hydrology, as discussed for lowland palm communities. Our results show that to understand plant community assembly in the tropical montane forests of the Andes it is too simple to focus just on environmental sorting by elevational climatic gradients.     

Disturbance effects on diversity of epiphytes and moths in a montane forest in Ecuador

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.     

Epiphytic and terrestrial mycorrhizas in a lower montane Costa Rican cloud forest

The epiphyte community is the most diverse plant community in neotropical cloud forests and its collective biomass can exceed that of the terrestrial shrubs and herbs. However, little is known about the role of mycorrhizas in this community. We assessed the mycorrhizal status of epiphytic (Araceae, Clusiaceae, Ericaceae, and Piperaceae) and terrestrial (Clusiaceae, Ericaceae) plants in a lower montane cloud forest in Costa Rica. Arbuscular mycorrhizas were observed in taxa from Araceae and Clusiaceae; ericoid mycorrhizas were observed in ericaceous plants. This is the first report of intracellular hyphal coils characteristic of ericoid mycorrhizas in roots of Cavendishia melastomoides, Disterigma humboldtii, and Gaultheria erecta. Ericaceous roots were also covered by an intermittent hyphal mantle that penetrated between epidermal cells. Mantles, observed uniquely on ericaceous roots, were more abundant on terrestrial than on epiphytic roots. Mantle abundance was negatively correlated with gravimetric soil water content for epiphytic samples. Dark septate endophytic (DSE) fungi colonized roots of all four families. For the common epiphyte D. humboldtii, DSE structures were most abundant on samples collected from exposed microsites in the canopy. The presence of mycorrhizas in all epiphytes except Peperomia sp. suggests that inoculum levels and environmental conditions in the canopy of tropical cloud forests are generally conducive to the formation of mycorrhizas. These may impact nutrient and water dynamics in arboreal ecosystems.     

Floristic composition, structure, and dynamics of an upper montane rain forest in Southern Ecuador

Two 1 ha plots of undisturbed upper montane rain forest in southern Ecuador were sampled for all trees with a dbh ≥ 5 cm. An extraordinarily high α-diversity for a forest near treeline is described. The "non-ridge forest" plot at 2900 m elevation has 75 species and 28 families ha^-1 and is believed to represent an advanced stage of succession whereas, the "ridge forest" plot at 2700 m elevation with 90 species and 38 families ha^-1 has a more rapid turn-over rate probably due to a more unstable environment. Downslope forces (soil creep) are discussed as a possible cause for 32–44% of all trees being inclined more than 30° in the moderately steep terrain (average slope 20° in both study plots). The plot of "non-ridge forest" is characterized by a much greater biomass whether expressed as basal area (44 m² versus 15 m²) or stem volume (214 m³ versus 52 m³), while the density is equally high (2310 versus 2090 trees ha^-1) in both plots. Families with Family Importance Value >25 / ha^-1 are Clusiaceae, Cunoniaceae, Melastomataceae, Myrsinaceae, and Ternstroemiaceae.     

Tree species richness complementarity, disturbance and fragmentation in a Mexican tropical montane cloud forest

Tropical montane cloud forest exhibits great heterogeneity in speciescomposition and structure over short geographic distances. In central Veracruz,Mexico, the conservation priority of seven cloud forest fragments was assessedby considering differences in woody plant species richness and complementarityof species among sites, forest structure, tree mortality, and timber andfirewood extraction as indicators of anthropogenic disturbance. Densities oftrees 5 cm dbh (360–1700 trees/ha) weredifferent among the sites, but basal area (35.3–89.3m²/ha) did not differ among fragments. The number of dead trees rangedfrom 10–30 to 170–200 trees/ha. The fragments'species composition was different but complementary. The Morisita–Hornindex indicated low similarity between fragments. A non-parametric estimator ofspecies richness indicated that more sampling effort would be necessary tocomplete the inventory (15 additional trees and two understory shrub species).Unfortunately, most of the fragments are threatened with deforestation. The numberof cut trees was similar among sites (0–15 cut trees/0.1 ha).Sites with immediate need for conservation were close to settlements, with highnumbers of cut trees and no legal protection. The selected sites represent thevariety of situations that exist in the region. Given the high complementarityobserved between fragments, a regional conservation approach will be required topreserve the last repositories of part of the tremendous biodiversity of theonce continuous forest in this region.     

Fern richness,tree species surrogacy,and fragment complementarity in a Mexican tropical montane cloud forest

We related pteridophytes versus tree species composition to identify surrogate measures of diversity, and complementarity of seven cloud forest fragments. Forest structure, and fern and tree composition were determined in 70 (2 × 50 m) transects. Fern density (10,150–25,080 individuals/ha) differed among sites. We recorded 83 fern species in the transects. Nonparametric richness estimators indicated that more sampling effort was needed to complete fern inventories (14 more species). However, ferns recorded outside of the transects increased richness to 103 species (six more species than predicted). Twenty-eight species were unique and rare due to special habitat requirements (Diplazium expansum, Hymenophyllum hirsutum, Melpomene leptostoma, Terpsichore asplenifolia), or were at a geographical distribution edge (Diplazium plantaginifolium, Lycopodium thyoides, Pecluma consimilis, Polypodium puberulum). Correlations between fern richness and tree richness and density were not significant, but were significant between fern richness and fern density, between epiphytic fern density and tree richness and density. Tree richness is not a good surrogate for fern diversity. Only three species were recorded in all fragments (Polypodium lepidotrichum, P. longepinnulatum, P. plebeium); thus fragments pteridophytes compositions are highly complementary, but more similar for ferns than for trees. A regional conservation approach which includes many small reserves needs to focus supplementarity on patterns of tree and fern species richness.     

Change in water quality during the passage through a tropical montane rain forest in Ecuador

We studied five 20-m transects onthe lower slope under tropical lower montanerain forest at 1900–2200 m above sea level. We collectedsamples of soil and of weekly rainfall,throughfall, litter leachate, and stream waterbetween 14 March 1998 and 30 April 1999 anddetermined the concentrations of Al, totalorganic C (TOC), Ca, Cl^–, Cu, K, Mg, Mn,NH₄ ⁺-N, NO₃ ^–-N, total N (TN), Na, P, S, and Zn. The soils were shallowInceptisols; pH ranged 4.4–6.3 in the Ohorizons and 3.9–5.3 in the A horizons, totalCa (6.3–19.3 mg kg^–1) and Mgconcentrations (1.4–5.4) in the O horizon weresignificantly different between the transects.Annual rainfall was 2193 mm; throughfall variedbetween 43 and 91% of rainfall, cloud waterinputs were 3.3 mm a^–1 except forone transect (203). The volume-weighted mean pHwas 5.3 in rainfall and 6.1–6.7 in throughfall.The median of the pH of litter leachate andstream water was 4.8–6.8 and 6.8, respectively.The concentrations of Ca and Mg in litterleachate and throughfall correlatedsignificantly with those in the soil (r =0.76–0.95). Element concentrations inthroughfall were larger than in rainfallbecause of leaching from the leaves (Al, TOC,Ca, K, Mg), particulate dry deposition (TOC,Cu, Cl^–, NH₄ ⁺-N), and gaseousdry deposition (NO₃ ^–-N, total N, S).Net throughfall (= throughfall-rainfalldeposition) was positive for most elementsexcept for Mn, Na, and Zn. High-flow eventswere associated with elevated Al, TOC, Cu, Mn,and Zn concentrations.     

Ficus lacunata (Moraceae), a new species from pluvial montane forest of northwest Ecuador

Ficus lacunata Kvitvik, a new species of subgen.Pharmacosycea sect.Pharmacosycea collected in the provinces of Carchi, Imbabura, and Pichincha in Ecuador, shows affinities to bothFicus macbridei Standl. andFicus tonduzii Standl. but differs distinctly in leaf and syconium characters.     

Photosynthetic parameters,dark respiration and leaf traits in the canopy of a Peruvian tropical montane cloud forest

Few data are available describing the photosynthetic parameters of the leaves of tropical montane cloud forests (TMCF). Here, we present a study of photosynthetic leaf traits (V _cmax and J _max), foliar dark respiration (R _d), foliar nitrogen (N) and phosphorus (P), and leaf mass per area (LMA) throughout the canopy for five different TMCF species at 3025 m a.s.l. in Andean Peru. All leaf traits showed a significant relationship with canopy height when expressed on an area basis, and V _cmax-area and J _max-area almost halved when descending through the TMCF canopy. When corrected to a common temperature, average V _cmax and J _max on a leaf area basis were similar to lowland tropical values, but lower when expressed on a mass basis, because of the higher TMCF LMA values. By contrast, R _d on an area basis was higher than found in tropical lowland forests at a common temperature, and similar to lowland forests on a mass basis. The TMCF J _max–V _cmax relationship was steeper than in other tropical biomes, and we propose that this can be explained by either the light conditions or the relatively low VPD in the studied TMCF. Furthermore, V _cmax had a significant—though relatively weak and shallow—relationship with N on an area basis, but not with P, which is consistent with the general hypothesis that TMCFs are N rather than P limited. Finally, the observed V _cmax–N relationship (i.e., maximum photosynthetic nitrogen use efficiency) was distinctly different from those in tropical and temperate regions, probably because the TMCF leaves compensate for reduced Rubisco activity in cool environments.     

Interactions between epiphytes during canopy soil formation: an experiment in a lower montane cloud forest of southeast Mexico

• In several montane forests around the world, epiphytes coexist in mats, sharing the rhizosphere and forming histosol-type soils rich in nutrients. The role of these epiphytes in the formation of canopy soil and the fitness costs that epiphytes face when cohabiting in these mats are unknown.
• In a lower montane cloud forest in central Veracruz, Mexico, a 2-year factorial experiment was carried out with the presence/absence of ramets of Phlebodium areolatum (Polypodiaceae), Tillandsia kirchhoffiana, T. multicaulis and T. punctulata (Bromeliaceae). We examined (i) which epiphyte species contribute to the formation of canopy soil, (ii) the role of epiphyte composition in the soil nutrient composition, and (iii) the fitness costs faced by epiphytes when cohabiting.
• Canopy soil formation highest when P. areolatum is present. Soil nutrient content does not change with epiphyte composition, is influenced by the microbiota, and P content decreases with the presence of epiphytes. The fitness costs show that the species compete, decreasing their survival and growth, but the competitive capacity differs between the species.
• We conclude that P. areolatum is an ecosystem engineer that promotes the creation of canopy soil but is a poor competitor. The results coincide with the model of succession by facilitation. Canopy soil is a slow-created component whose nutrient content does not depend on the epiphytic flora. In epiphyte mats, the dominant interactions are competitive, but there is also facilitation.

     

Understory spider diversity in two remnants of tropical montane cloud forest in Chiapas, Mexico

We evaluated the spider diversity of a tropical montane cloud forest understory in two nearby sites with different degree of human disturbance at the Biosphere Reserve Volcán Tacaná, Chiapas, Mexico. The study was conducted over a 24 days period distributed in 6 months in 2009, covering dry and rainy seasons. A total of 8,370 spiders (1,208 adults and 7,162 juveniles) were collected. Determined specimens (7,747) represented 112 species and morphospecies, 71 genera and 22 families. The results showed that human disturbance has an influence on spider communities: species richness was significantly higher in the preserved site as regards to the disturbed site. Despite their proximity, the composition of spider communities showed only a moderate similarity between the two sites. No differences in abundance were found among sites when considering the whole sample, but sites differed clearly when seasons were analyzed separately. The rainy season had a negative effect on the abundance of spiders in the preserved site. Although the spider community structure was very similar between sites, there was a trend towards a greater species evenness in the preserved site for the whole sampling period and for the dry season.     

Linking vegetation structure and bird organization: response of mixed-species bird flocks to forest succession in subtropical China

As forests undergo natural succession following artificial afforestation, their bird assemblages also change. However, interspecific avian social organization associated with forest succession has not been fully understood, particularly for mixed-species bird flocks. To disentangle how mixed-species flocks change as a function of local forest structure, we analyzed flock characteristics (particularly species richness, flocking frequency and propensity) and vegetation physiognomies along a presumed successional series (early, middle, and advanced) simultaneously in subtropical forests in southern China. As hypothesized, monthly point counts demonstrated that complexity of flocks increases with the progression of natural forest succession at a local scale. Advanced forests differed significantly from pioneering plantations with respect to vegetation structure, flock characteristics and constituents (especially for understory specialists). Importantly, forest succession affected flock patterns particularly in relation to the flocking propensity of regular species, and the frequency of nuclear species (Huet’s fulvetta Alcippe hueti), which in turn determined flocking occurrence at different successional stands. Canonical correspondence analysis indicated that understory flocking species (mainly Timaliidae babblers) were significantly associated with intact native canopy cover, complex DBH diversity, as well as high densities of dead trees and large trees, representing a maturity level of successional stands. Our study reveals that the effect of natural forest succession on mixed-species bird flocks is species-specific and guild-dependent. From a conservation perspective, despite a high proliferation of pine plantation in southern China, priority should be placed on protecting the advanced forest with a rich collection of understory flocking specialists.     

Leaf demography and leaf traits of temperate-deciduous and tropical evergreen-broadleaved trees in a Mexican montane cloud forest

The spatial variation in epilithic lichen community structure was investigated as part of a larger study of the vegetation and ecology of the tall limestone cliffs within the Niagara Escarpment Biosphere Reserve in southern Ontario, Canada. The cover of all lichen taxa was visually estimated for a total of 199 samples taken from the top, middle, or bottom of the cliff face at five sites. Twelve environmental variables were also measured. Twenty-seven lichen taxa were identified on the samples. Multivariate ordinations of species composition (DCA, CCA, PCCA) revealed variation in community structure on multiple scales, but no groupings of sites that would have suggested the presence of several distinct species assemblages. A gradient in species composition from north to south, most clearly reflected in the decreasing cover of foliose and fruticose species, may reflect a gradient in human disturbance. There was also intermediate-scale patchiness in species composition in a horizontal plane across cliffs, but despite earlier claims made in the literature, no evidence of vertical zonation of the lichens on cliffs was found. Species composition also responded to small-scale factors possibly related to exposure, light, or moisture. Unlike community composition, the total cover of all lichens was homogeneous over large spatial scales and varied only on a small scale, illustrating that scale as well as resolution of a study may influence the ecological patterns seen. More than half of the species found on the Niagara Escarpment are rare on rock substrates elsewhere in southern Ontario, and two are new for North America (Candelariella heidelbergensis (Nyl.) Poelt and Lecanora perpruinosa Fröberg). The result that cliffs support a distinct flora containing many rare species suggests that they are a reservoir for biodiversity not just for vascular plants, but also for lichens.     

Nutrient limitation restricts growth and reproductive output in a tropical montane cloud forest bromeliad: findings from a long-term forest fertilization experiment

From studies in seasonal lowland tropical forests, bromeliad epiphytes appear to be limited mainly by water, and to a lesser extent by nutrient supply, especially phosphorous. Less is understood about the mineral nutrition of tropical montane cloud forest (TMCF) epiphytes, even though their highest diversity is in this habitat. Nutrient limitation is known to be a key factor restricting forest productivity in TMCF, and if epiphytes are nutritionally linked to their host trees, as has been suggested, we would expect that they are also nutrient limited. We studied the effect of a higher nutrient input on reproduction and growth of the tank bromeliad Werauhia sintenisii in experimental plots located in a TMCF in Puerto Rico, where all macro- and micronutrients had been added quarterly starting in 1989 and continuing throughout the duration of this study. We found that bromeliads growing in fertilized plots were receiving litterfall with higher concentrations of N, P, and Zn and had higher concentrations of P, Zn, Fe, Al, and Na in their vegetative body. The N:P ratios found (fertilized = 27.5 and non-fertilized = 33.8) suggest that W. sintenisii may also be phosphorous limited as are lowland epiphytes. Fertilized plants had slightly longer inflorescences, and more flowers per inflorescence, than non-fertilized plants, but their flowers produced nectar in similar concentrations and quantities. Fertilized plants produced more seeds per fruit and per plant. Frequency of flowering in two consecutive years was higher for fertilized plants than for controls, suggesting that fertilized plants overcome the cost of reproduction more readily than non-fertilized plants. These results provide evidence that TMCF epiphytic bromeliads are nutrient limited like their lowland counterparts.