Vascular epiphyte assemblage structure and distribution patterns in the south-temperate zone

<p>Vascular epiphytes, which are specialised to spend their entire life cycle within trees, are significant contributors to local ecosystem services. However, our current understanding of epiphyte distributions, co-occurrences, and general ecology lags far behind that of terrestrial plants. Furthermore, the majority of epiphyte research is undertaken in tropical forests, with comparatively few studies extending into temperate climates. As such, whether epiphytic plant assemblage structure varies geographically, or is influenced by area and isolation effects needs further scrutiny. In addition, how epiphytes are distributed in relation to host tree ontogeny and microclimates specific to south-temperate forests is poorly understood. Here, I attempt to bridge this gap by researching epiphyte distributions and assemblage structure in New Zealand, southern Chile, and Australia. In the first biogeographic study of epiphyte-host interactions, I determined if epiphyte-host network structure (i.e. nestedness, species co-occurrences, species specialisation) varied among New Zealand and Chilean temperate forests (Chapter 2). At the forest stand level, network structure was consistent with stochastic structuring, which suggests that dispersal and disturbances are important drivers of epiphyte distributions at a biogeographic scale. However, deterministic structure was observed in New Zealand networks with regards to nestedness (i.e. when specialists interact with generalists), which suggests that positive species interactions influence epiphyte distributions at a within-tree scale. Second, I determined whether the composition of plant communities residing in epiphytic birds’ nest ferns (Asplenium goudeyi) on Lord Howe Island, Australia, are influenced by fern size, isolation from a major propagule source and resident plant community richness (Chapter 3). Results suggest that plant communities are structured by dispersal. For one, there was a significant isolation effect on resident plant community richness. Additionally, wind-dispersed taxa were well represented in isolated ferns, while animal-dispersed taxa and taxa with no specific dispersal strategies were absent. This is the first study to test the combined effects of area, isolation and resident plant richness on epiphytic plant assemblage structure. Third, using Darwin’s geological theory of island ontogeny as a theoretical construct, I explored changes in epiphyte species richness throughout tree ontogeny (Chapter 4). Theoretical frameworks have helped bridge the gap between our understanding of vascular epiphytes and terrestrial plants, however, none have been implemented to guide investigations on epiphyte assemblage development. Based on the general features of island ontogeny, I found three stages of epiphyte assemblage development: (i) an initial stage where host trees are devoid of epiphytes, (ii) a second stage where trees acquire epiphytes into maturity, and (iii) a hypothetical stage where epiphyte assemblages follow a period of species decline following host tree mortality. In addition to these results, I found interspecific variation in the ontogenetic stage at which host trees become favourable for epiphyte establishment and the rate at which epiphyte assemblages develop. Lastly, I explored the systematic distribution of epiphytes and mistletoes in relation to microclimate gradients around the trunks of trees (Chapter 5). In addition, I tested the physiological responses of epiphytes and mistletoes to reductions in their most limiting resources to determine if the responses were consistent with their distribution patterns. The radial distributions of epiphytes and mistletoes were highly directional, and paralleled gradients of humidity, light and water. Additionally, the photochemical efficiency of epiphytes and CO₂ assimilation in mistletoe leaves decreased in plants growing in environments with lower water and light availability, respectively. However, mistletoe leaves still assimilated CO₂ in lower light conditions, which suggests a high plasticity of mistletoes to growing in a canopy environment. Despite over 120 years of recognising the importance of vertical microclimates on epiphyte distributions, this is the first systematic study of epiphytic plant distributions in relation to microclimate gradients around the trunks of trees. This thesis has increased our understanding of epiphytic plant assemblage structure, and how it is influenced by host tree species, isolation, area and resident plant species richness. In addition, this thesis has increased our understanding of the effect of host tree ontogeny and microclimate on epiphyte distribution patterns. Together, these studies may be built upon more broadly to further elucidate drivers of epiphyte assembly and distribution patterns.</p>

Vascular epiphyte assemblage structure and distribution patterns in the south-temperate zone

<p>Vascular epiphytes, which are specialised to spend their entire life cycle within trees, are significant contributors to local ecosystem services. However, our current understanding of epiphyte distributions, co-occurrences, and general ecology lags far behind that of terrestrial plants. Furthermore, the majority of epiphyte research is undertaken in tropical forests, with comparatively few studies extending into temperate climates. As such, whether epiphytic plant assemblage structure varies geographically, or is influenced by area and isolation effects needs further scrutiny. In addition, how epiphytes are distributed in relation to host tree ontogeny and microclimates specific to south-temperate forests is poorly understood. Here, I attempt to bridge this gap by researching epiphyte distributions and assemblage structure in New Zealand, southern Chile, and Australia. In the first biogeographic study of epiphyte-host interactions, I determined if epiphyte-host network structure (i.e. nestedness, species co-occurrences, species specialisation) varied among New Zealand and Chilean temperate forests (Chapter 2). At the forest stand level, network structure was consistent with stochastic structuring, which suggests that dispersal and disturbances are important drivers of epiphyte distributions at a biogeographic scale. However, deterministic structure was observed in New Zealand networks with regards to nestedness (i.e. when specialists interact with generalists), which suggests that positive species interactions influence epiphyte distributions at a within-tree scale. Second, I determined whether the composition of plant communities residing in epiphytic birds’ nest ferns (Asplenium goudeyi) on Lord Howe Island, Australia, are influenced by fern size, isolation from a major propagule source and resident plant community richness (Chapter 3). Results suggest that plant communities are structured by dispersal. For one, there was a significant isolation effect on resident plant community richness. Additionally, wind-dispersed taxa were well represented in isolated ferns, while animal-dispersed taxa and taxa with no specific dispersal strategies were absent. This is the first study to test the combined effects of area, isolation and resident plant richness on epiphytic plant assemblage structure. Third, using Darwin’s geological theory of island ontogeny as a theoretical construct, I explored changes in epiphyte species richness throughout tree ontogeny (Chapter 4). Theoretical frameworks have helped bridge the gap between our understanding of vascular epiphytes and terrestrial plants, however, none have been implemented to guide investigations on epiphyte assemblage development. Based on the general features of island ontogeny, I found three stages of epiphyte assemblage development: (i) an initial stage where host trees are devoid of epiphytes, (ii) a second stage where trees acquire epiphytes into maturity, and (iii) a hypothetical stage where epiphyte assemblages follow a period of species decline following host tree mortality. In addition to these results, I found interspecific variation in the ontogenetic stage at which host trees become favourable for epiphyte establishment and the rate at which epiphyte assemblages develop. Lastly, I explored the systematic distribution of epiphytes and mistletoes in relation to microclimate gradients around the trunks of trees (Chapter 5). In addition, I tested the physiological responses of epiphytes and mistletoes to reductions in their most limiting resources to determine if the responses were consistent with their distribution patterns. The radial distributions of epiphytes and mistletoes were highly directional, and paralleled gradients of humidity, light and water. Additionally, the photochemical efficiency of epiphytes and CO₂ assimilation in mistletoe leaves decreased in plants growing in environments with lower water and light availability, respectively. However, mistletoe leaves still assimilated CO₂ in lower light conditions, which suggests a high plasticity of mistletoes to growing in a canopy environment. Despite over 120 years of recognising the importance of vertical microclimates on epiphyte distributions, this is the first systematic study of epiphytic plant distributions in relation to microclimate gradients around the trunks of trees. This thesis has increased our understanding of epiphytic plant assemblage structure, and how it is influenced by host tree species, isolation, area and resident plant species richness. In addition, this thesis has increased our understanding of the effect of host tree ontogeny and microclimate on epiphyte distribution patterns. Together, these studies may be built upon more broadly to further elucidate drivers of epiphyte assembly and distribution patterns.</p>

Cedroxylon shakhtnaense (Blokhina 2010) Dolezych, Mantzouka et L.Kunzmann comb. nov.; A fossil Abies wood from the late early Miocene Mastixioideae flora of Wiesa (east Germany)

We describe the first evidence of fossil Abies wood from the late early Miocene fossil plant assemblage of Wiesa in east Germany. The comparatively well-preserved piece of xylitic wood was recovered in the kaolin quarry at Hasenberg hill in Wiesa. The Wiesa assemblage is characterized as being allochthonous and partly parautochthonous mass deposits of diaspores, leaves, and wood. The latter component is rather incompletely studied so far. The described fossil is characterized by high rays, mostly uniseriate bordered pits, generally thick and pitted horizontal and tangential ray cell walls, but also partly smooth horizontal ray cell walls, absence of ray tracheids, the occurrence of traumatic resin canals, and rare occurrence of axial parenchyma of two types. This type of fossil wood has been described as Abietoxylon shakhtnaense Blokhina from the Oligo-Miocene of Sakhalin, Russia. Due to nomenclatural issues of Abietoxylon a recombination to Cedroxylon Kraus emend. Gothan is proposed following common practice for affiliation of abietoid fossil wood of Cenozoic age. Cedroxylon shakhtnaense comb. nov. shares anatomical characteristics with the wood of extant Abies Mill., in particular with sections Abies and Grandis, and is most closely related to section Grandis. The properly preserved fossil wood from Wiesa provides the opportunity of applying qualitative and quantitative analyses for testing and discussing its placement in relationship to intra-tree variability and ontogenetic aspects. The first evidence of fossil wood of Abies from Wiesa confirms again the presence of the genus in mid-latitude subtropical zonal vegetation during the beginning of the Miocene Climatic Optimum.

Dynamics of Silurian Plants as Response to Climate Changes

The most ancient macroscopic plants fossils are Early Silurian cooksonioid sporophytes from the volcanic islands of the peri-Gondwanan palaeoregion (the Barrandian area, Prague Basin, Czech Republic). However, available palynological, phylogenetic and geological evidence indicates that the history of plant terrestrialization is much longer and it is recently accepted that land floras, producing different types of spores, already were established in the Ordovician Period. Here we attempt to correlate Silurian floral development with environmental dynamics based on our data from the Prague Basin, but also to compile known data on a global scale. Spore-assemblage analysis clearly indicates a significant and almost exponential expansion of trilete-spore producing plants starting during the Wenlock Epoch, while cryptospore-producers, which dominated until the Telychian Age, were evolutionarily stagnate. Interestingly cryptospore vs. trilete-spore producers seem to react differentially to Silurian glaciations—trilete-spore producing plants react more sensitively to glacial cooling, showing a reduction in species numbers. Both our own and compiled data indicate highly terrestrialized, advanced Silurian land-plant assemblage/flora types with obviously great ability to resist different dry-land stress conditions. As previously suggested some authors, they seem to evolve on different palaeo continents into quite disjunct specific plant assemblages, certainly reflecting the different geological, geographical and climatic conditions to which they were subject.

Macrofloral and microfloral changes in the Middle Jurassic plant assemblages of the Cianowice 2 borehole (southern Poland)

The flora of the Cianowice 2 borehole (c. 20 km NW of Cracow, Poland), dominated by cycadophytes (mainly bennettitaleans) and conifers, shows high taxonomic diversity relative to the low number of specimens. Twenty species were identified in the 96 determinable plant fragments found in 27 core samples: Cladophlebis sp. (ferns), Pachypteris rhomboidalis (Ettingshausen) Nathorst and Ptilozamites cycadea (Berger) Möller (seed ferns), Anomozamites nilssonii (Phillips) Seward, Nilssoniopteris solitaria (Phillips) Cleal & Rees, Otozamites mimetes Harris, Otozamites parallelus Phillips, Pterophyllum thomasii Harris, Pterophyllum cf. aequale (Brongniart) Nathorst, Ptilophyllum cf. okribense forma ratchiana Doludenko & Svanidze, Ptilophyllum pecten Phillips, Ptilophyllum sirkennethii Watson & Sincock, Cycadolepis sp. (bennettitaleans), Pseudotorellia grojecensis Reymanówna, Pseudotorellia samylinae Nosova & Kiritchkova, Pseudotorellia sp. (Gymnospermae incertae sedis), Bilsdalea dura Harris, Mirovia szaferi Reymanówna, and Brachyphyllum stemonium Kendall (conifers). The floristic composition is supplemented by palynological data. The taxa were connected to five depositional successions distinguished along the core: one, alluvial fans; two, four and five, meandering/anastomosing river depositional systems with fluvial plain deposits; and three, lacustrine/backswamp environment developed on fluvial plain. The composition of the fossil plant assemblage changes with the depositional setting within the same range of taxa, seen mainly in changed combinations of taxa, which are most diverse in the fluvial plain deposits. Some taxa occur in a single depositional succession; some are present in two or three. The sporomorph assemblages of particular depositional environments differ significantly from the composition of the co-occurring macroflora: ferns occur sporadically in the macroflora of each depositional environment but they strongly dominate the sporomorph assemblage. Our proposed reconstruction of the palaeoenvironment is a slight rise descending into a valley with a depositionary basin, with gymnosperms on the slope and ferns at the base. Some species are shared between Cianowice and nearby Middle Jurassic localities in Grojec and Zabierzów, and the majority of taxa are known from the Middle Jurassic, suggesting that the Cianowice deposits are of that age.