Agent‐based modeling of the effects of forest dynamics,selective logging,and fragment size on epiphyte communities

1. Forest canopies play a crucial role in structuring communities of vascular epiphytes by providing substrate for colonization, by locally varying microclimate, and by causing epiphyte mortality due to branch or tree fall. However, as field studies in the three‐dimensional habitat of epiphytes are generally challenging, our understanding of how forest structure and dynamics influence the structure and dynamics of epiphyte communities is scarce.
2. Mechanistic models can improve our understanding of epiphyte community dynamics. We present such a model that couples dispersal, growth, and mortality of individual epiphytes with substrate dynamics, obtained from a three‐dimensional functional–structural forest model, allowing the study of forest–epiphyte interactions. After validating the epiphyte model with independent field data, we performed several theoretical simulation experiments to assess how (a) differences in natural forest dynamics, (b) selective logging, and (c) forest fragmentation could influence the long‐term dynamics of epiphyte communities.
3. The proportion of arboreal substrate occupied by epiphytes (i.e., saturation level) was tightly linked with forest dynamics and increased with decreasing forest turnover rates. While species richness was, in general, negatively correlated with forest turnover rates, low species numbers in forests with very‐low‐turnover rates were due to competitive exclusion when epiphyte communities became saturated. Logging had a negative impact on epiphyte communities, potentially leading to a near‐complete extirpation of epiphytes when the simulated target diameters fell below a threshold. Fragment size had no effect on epiphyte abundance and saturation level but correlated positively with species numbers.
4. Synthesis: The presented model is a first step toward studying the dynamic forest–epiphyte interactions in an agent‐based modeling framework. Our study suggests forest dynamics as key factor in controlling epiphyte communities. Thus, both natural and human‐induced changes in forest dynamics, for example, increased mortality rates or the loss of large trees, pose challenges for epiphyte conservation.

     

Host identity and functional traits determine the community composition of the arbuscular mycorrhizal fungi in facultative epiphytic plant species

The epiphytic vascular mycobiota is scarce and facultative in semi-arid Mediterranean ecosystems. However, unlike in soil conditions, little is known about the factors driving mycorrhizal communities in epiphytic environments. Here, we investigated the arbuscular mycorrhizal fungi (AMF) harboured by 31 plant species occurring on the trunks of Phoenix dactylifera. We wanted to ascertain if host identity and plant functional traits shape mycorrhizal communities. Specifically, we tested the plant life-cycle (perennial versus annual), the plant life-form (herbaceous versus woody), the plant origin (exotic versus native) and the plant species. The plant affiliation to species strongly influenced the AMF community composition. Plant life-form and plant life-cycle also shaped indicator taxa. The AMF structure differed between annual and perennial species and higher AMF richness was detected in perennial plants. The epiphytic plants associated with AMF irrespective of whether they were native or not, probably because here no functional differences derive from plant origin.     

Seasonal variation in a low shore Fucus serratus (Fucales,Phaeophyta) population and its epiphytic fauna

Seasonal variation in a Fucus serratus assemblage was studied on the Isle of Man, UK over a 13-month period. Algal cover varied greatly, related to shedding reproductive fronds and winter storms. Peak reproduction was recorded in the autumn. There was little change in density of plants, the population was dominated by a bank of germlings (<20 cm) and an adult mode between 50–70 cm. An indistinct third component of recruiting juveniles was present when gaps in the canopy formed. Seasonal variation in the algae was reflected in the population dynamics of the mobile epiphytes such as Littorina mariae and Lacuna pallidula, which exhibited annual life histories. Sessile species (such as bryozoans) utilized the algae year round, dominating the perennial stipes and lower fronds, which were not shed. Other species utilized the algae in a transient fashion: some feeding on the algae (e.g. Idotea), or other epiphytic animals (e.g. Nucella). Fucus serratus, therefore, provides a variety of spatially and temporally variable niches that explain the diversity of epiphytic animal species that exploit this alga.     

Disentangling climate change from air pollution effects on epiphytic bryophytes

At the interface between atmosphere and vegetation, epiphytic floras have been largely used as indicators of air quality. The recovery of epiphytes from high levels of SO₂ pollution has resulted in major range changes, whose interpretation has, however, been challenged by concomitant variation in other pollutants as well as climate change. Here, we combine historical and contemporary information on epiphytic bryophyte species distributions, climatic conditions, and pollution loads since the 1980s in southern Belgium to disentangle the relative impact of climate change and air pollution on temporal shifts in species composition. The relationship between the temporal variation of species composition, climatic conditions, SO₂, NO₂, O₃, and fine particle concentrations, was analyzed by variation partitioning. The temporal shift in species composition was such, that it was, on average, more than twice larger than the change in species composition observed today among communities scattered across the study area. The main driver, contributing to 38% of this temporal shift in species composition, was the variation of air quality. Climate change alone did not contribute to the substantial compositional shifts in epiphytic bryophyte communities in the course of the last 40 years. As a consequence of the substantial drop of N and S loads over the last decades, present-day variations of epiphytic floras were, however, better explained by the spatial variation of climatic conditions than by extant pollution loads. The lack of any signature of recolonization delays of formerly polluted areas in the composition of modern floras suggests that epiphytic bryophytes efficiently disperse at the landscape scale. We suggest that a monitoring of epiphyte communities at 10-year intervals would be desirable to assess the impact of raising pollution sources, and especially pesticides, whose impact on bryophytes remains poorly documented.     

Behavioural studies on the peppered moth Biston betularia and a discussion of the role of pollution and lichens in industrial melanism

We examined the resting behaviour during pairing and egg-laying of bred female Biston betularia (L.) after they were set free at wooded sites in Somerset and Cardiff. Moths were not released immediately on emergence but were held for three nights to overcome an initial period of flight activity. Females (N= 257) were placed on trunks, and on large and small branches of trees, especially oaks, over periods of several weeks in 1984, 1985 and 1986. Releases were not made within 50 m of any known survivor. One hundred and thirty-one pairings were observed. We also examined oviposition behaviour in a cage experiment using different types of branches. Our observations support Mikkola's earlier conclusion from cage experiments with male moths that the species rests predominantly on branches and shows an appropriately specialized resting attitude, demonstrated here in a series of photographs. Many moths will rest underneath, or on the side of, narrow branches in the canopy. Once released females had settled and paired they only moved quite short distances, but positioning and egg-laying were profoundly influenced by the presence of foliose lichens. Females of all phenotypes show a strong preference to oviposit beneath a thallus of foliose lichens. Their tendency to rest against or close to the lichen during the day could markedly influence their crypsis. In the absence of foliose lichens, eggs are laid in cracks in the bark. The survivorship of released females both in different pairing combinations and during their whole reproductive life-span is analysed. These preliminary data suggest that bird predation can give rise to differential mortality of different pairing combinations and of the phenotypes. This technique provides a powerful method of measuring the relative crypsis and differential mortality of phenotypes with living females in natural resting sites. The significance of our observations on resting behaviour is discussed with regard to the role of air pollution and epiphytes in industrial melanism. Particular attention is given to the causal mechanisms underlying recent declines in the frequency of carbonaria melanics in parts of Britain. One important factor is the new growth made by trees since the marked reduction in particulate air pollution in the 1960s. It is emphasized that we need to know much more about the interactions between pollution, epiphytes and resting backgrounds, especially in the canopy, before we can be confident of our understanding of the evolution of industrial melanism.     

Grazing effects of a gammaridean Amphipoda,Ampithoe sp., on the seagrass,Syringodium isoetifolium,and epiphytes in a tropical seagrass bed of Fiji

In the pure stand of tropical seagrass,Syringodium isoetifolium, in a small oceanic island, Fiji, grazing effects of the seagrass-associated gammarid,Ampithoe sp., on seagrass and epiphytes were assessed in October 1989, November 1991, November 1992. Density of the gammarid was estimated with two methods, mesh bag method and tuft method. During the three years surveyed the density of the gammarid increased remarkably from 1989 to 1991, with heavy epiphytism. Gut contents of the gammarid were examined. Grazing rates on seagrass leaf with and without epiphytic blue-green algae were measured in a bottle experiment. Litter bag experiments were conducted using different mesh sizes each containing seagrass only and seagrass and gammarids. The seagrass leaf biomass in the litter bag reduced abruptly in both bags. After one week, 78–86% of seagrass biomass disappeared from the bags. Enhancement of decomposition of seagrass leaf by the gammarid grazing was observed. Oxygen consumption and ammonium excretion rates were measured simultaneously in bottle experiments. Carbon budget in the seagrass bed was estimated as follows: 0.9 gC m⁻² day⁻¹ in seagrass growth, gammarid grazing was about a half of it and further assimilated a half of it, about 0.1 gC m⁻² day⁻¹, and more than half of it become CO₂ by respiration. Grazing effects on epiphyte and seagrass growth and production were discussed through the carbon budget and indirect interactions between seagrass, epiphytes and associated gammarids to explain the temporal change of seagrass and epiphyte dynamics.     

The invertebrate fauna of epiphyte mats in the canopy of northern rata (Myrtaceae: Metrosideros robusta A. Cunn.) on the West Coast of the South Island,New Zealand

Abstract

The range of microhabitats and microclimatic conditions provided by epiphytes has been linked to the high diversity of invertebrates in many forest canopies worldwide, but comparably little is known about the invertebrate fauna in this habitat in New Zealand. This study compiled an inventory of the invertebrate fauna of epiphyte mats in the canopy of northern rata (Myrtaceae: Metrosideros robusta A. Cunn.) at two study sites on the West Coast of the South Island. A total of 242 069 invertebrate specimens was collected over one year, representing 4 phyla, 9 classes and more than 160 families, 225 genera and 446 species. At least 10 new species and 3 new genera were identified, while 5 species were recorded outside their known geographical range. Epiphyte mats provided habitat for an invertebrate fauna, highly diverse and abundant both taxonomically and functionally, dominated in terms of abundance by Acari, Collembola and Hymenoptera (largely ants), and in terms of feeding guilds by epiphyte grazers and ants. As the first inventory of this taxonomic depth and breadth compiled for New Zealand forest epiphyte habitats, this study provides important baseline data for the conservation of biodiversity in New Zealand's indigenous forests.     

Disentangling the roles of chance,abiotic factors and biotic interactions among epiphytic bryophyte communities in a tropical rainforest (Yunnan,China)

• Epiphytes offer an appealing framework to disentangle the contributions of chance, biotic and abiotic drivers of species distributions. In the context of the stress-gradient theory, we test the hypotheses that (i) deterministic (i.e., non-random) factors play an increasing role in communities from young to old trees, (ii) negative biotic interactions increase on older trees and towards the tree base, and (iii) positive interactions show the reverse pattern.
• Bryophyte species distributions and abiotic conditions were recorded on a 1.1 ha tropical rainforest canopy crane site. We analysed co-occurrence patterns in a niche modelling framework to disentangle the roles of chance, abiotic factors and putative biotic interactions among species pairs.
• 76% of species pairs resulted from chance. Abiotic factors explained 78% of non-randomly associated species pairs, and co-occurrences prevailed over non-coincidences in the remaining species pairs. Positive and negative interactions mostly involved species pairs from the same versus different communities (mosses versus liverworts) and life forms, respectively. There was an increase in randomly associated pairs from large to small trees. No increase in negative interactions from young to old trees or from the canopy to the base was observed.
• Our results suggest that epiphytic bryophyte community composition is primarily driven by environmental filtering, whose importance increases with niche complexity and diversity. Biotic interactions play a secondary role, with a very marginal contribution of competitive exclusion. Biotic interactions vary among communities (mosses versus liverworts) and life forms, facilitation prevailing among species from the same community and life form, and competition among species from different communities and life forms.