Epiphytes and tropical trees relationship counseling

3 EPIPHYTES MODERATE CLIMATIC EXTREMES IN TROPICAL TREE aspects of the relationship between arthropods and epiphytes, and will outline I had fruitful talks and got valuable advice from Don Windsor, Annette Aiello and. Waikato region of the North Island, and the water relations of the shrub You have each been very kind in giving me advice, encouragement, and your time, tropical rainforest tree in Costa Rica to be °C warmer and 15 % less humid. Trees with epiphytes had significantly higher numbers of species and individuals and insects larger than 5 mm of tropical trees resulted in a great number of studies conducted within .. no long-lasting relationship with the plant, but which may be . advice; J.G. Garcıa-Franco, Federico Escobar, S. Philpott,. V. Rico- Gray.

Davalliaceae comprise four to ten genera and 50— species Copeland, ; Kato, ; Kramer, a ; Nooteboom, The Davalliaceae are characterized by the epiphytic life form, long creeping dorsiventral rhizomes, peltate or pseudopeltate scales densely covering the rhizomes, and indusiate sori Kato, ; Kramer, a. Among the genera, Leucostegia is distinct in having basifixed scales and a few other characters described below, although it shares those characters with the other genera.

In a molecular rbcL analysis of the interfamilial relationships of ferns, Hasebe et al. Available data suggest that epiphytism of the group evolved at or near the base of the clade. The precise relationships of the davallioid—polygrammoid group with more non-epiphytic relatives are useful to understand the evolution of the epiphytic Davalliaceae. The aim of our study was to clarify the life forms of Davalliaceae and related species from field observations, and to infer the evolution of epiphytism in Davalliaceae, based on the phylogenetic relationships of the family and related ferns using two chloroplast genes, rbcL and accD.

Plants were carefully observed with a focus on root conditions connected to the soil or notthe direction of rhizome elongation upward or downwardthe length of rhizomes, the height of tree trunks where the rhizomes attach lower or upperand, when available, the place where young plants grow.

Vouchers for the field observations and molecular analysis are deposited in the University of Tokyo Herbarium TI. Paradavallodes, Parasorus, and Trogostolon were excluded from the analysis, because material was not available. Parasorus is a specialized monotypic genus characterized by simple leaves and coenosori Kato, Trogostolon is also such a genus, characterized by the finely dissected leaves and acicular scales Copeland, Therefore, the present analysis excluding those specialized genera may be able to reveal a general systematic structure of the Davalliaceae, and exclusion of those epiphytic genera does not seem to influence seriously the inferred evolution of the life forms of the Davalliaceae.

However, there is no doubt that the phylogenetic relationships of Davalliaceae await analysis with more species representing all recognized genera and other particular groups. In addition we examined species of Colysis, Crypsinus, Gymnogrammitis, Loxogramme, Microsorum, Grammitis, and Pyrrosia, which are members of the polygrammoid group sister to Davalliaceae Hasebe et al.

We also examined four of the six genera of the Lomariopsidaceae, including the epiphytic Elaphoglossum, and some genera of the Dryopteridaceae, which were less closely related species to the davallioid—polygrammoid group. Athyrium and Matteuccia were used for outgroups Hasebe et al. The species examined in this study represent all major subclades of a monophyletic clade that diverges from the other sister clade to which the outgroup species are assigned in all-family analyses Hasebe et al.

The analysed sequences were limited to the obtained coding regions, because the intergenic spacer between rbcL and accD included too many indels to construct objective alignment.

Therefore, sequences downward to the ACG codon were used as accD sequences. Operational taxonomic units with identical sequences were treated as a single unit in phylogenetic analysis.

The base that could not be identified in this study was treated as N. The rbcL and accD sequences of Davallia denticulata Philippine material had one unidentifiable base, although all other sequences were the same as those of Davallia denticulata Sumatra material. Hence, both sequences were examined in the phylogenetic analysis.

In the MP method, all characters were equally weighted and heuristic searches were conducted with random addition replicates involving tree-bisection-reconnection TBR branch swapping. Bootstrap values were calculated with replicates with ten random additions for both rbcL and accD data and with replicates with random addition replicates for the combined data.

Bayesian searches were conducted by mcmc with four chains over 1 generations, sampling every generations. One thousand trees were discarded as burn-in trees and the rest of the trees were used to calculate the majority rule consensus tree. All characters were treated as unordered and plotted on to the topologies recovered in the MP and ML analyses.

In the obligate epiphytic Elaphoglossum callifolium LomariopsidaceaeLoxogramme avenia Polypodiaceaeand Grammitis reinwardtii Grammitidaceaethe rhizomes were short creeping and attached to tree trunks or branches. Many species of the Davalliaceae with long creeping rhizomes were usually obligate epiphytes.

Both young and mature plants of Davallia trichomanoides occurred on tree trunks and their roots were always separated from the soil. Scyphularia pentaphylla was a high epiphyte occurring on tree branches. Humata vestita, often occurred on fallen wood, although they sometimes grew on mossy rocks as lithophytes with their roots unconnected to the soil.

Oleandra and Nephrolepis were secondary hemi-epiphytes. The long creeping rhizomes of most individuals relatively young plants of Oleandra pistillaris climbed tree trunks and may have reached a height of about 10 m on tree trunks with the basal roots connected to the soil.

In some other plants maturethe proximal parts of the climbing rhizomes were dried and free from the ground.

Nephrolepis acuminata and N. Most individuals were terrestrials or climbers with the basal roots connected to the soil, and some individuals were epiphytic with stolons dry at the proximal parts. Young plants usually climbed trees and had roots connected to the soil. Some polygrammoid species were also secondary hemi-epiphytes or hemi-lithophytes. The secondary hemi-epiphytic Microsorum buergerianum, M. Some species of the davallioid—polygrammoid ferns had two life forms, i. In Araiostegia hymenophylloides, Davallia denticulata, and D.

Crypsinus enervis and Goniophlebium persicifolium Polypodiaceae may be obligate epiphytes and secondary hemi-epiphytes, because they were usually low epiphytes and sometimes climbers with rhizomes and roots connected to the soil, although we did not find epiphytic juveniles. Arguably, narrow host tree specificity is found only in exceptional cases, whereas a certain degree of host preference is not uncommon terStege and Cornelissen ; Laube and Zotz, ; Martinez-Melendez et al. Mechanistically, such preferences are probably related to differences in tree architecture, bark structure and chemistry or leaf phenology of the host.

Several studies have investigated the impact of a variety of tree characteristics on epiphyte distribution, such as tree size Zotz and Vollrathbranch diameter Zimmerman and Olmsted or bark water-holding capacity WHC, Callaway et al. Differing tree characteristics create a variety of microclimates within a tree crown that should directly impact epiphyte distributions Pittendrigh ; Benzing, The changing light regime along the vertical axis of the canopy is one important factor.

Lower and more central crown parts are more humid than the outer portions and exposed parts within a tree are usually the driest e. Freiberg ; Wagner et al.

However, gradients are not stable in time but are affected by phenological changes of the tree usually associated with wet and dry seasons. Seasonality may be an important factor for the establishment of epiphytes; especially when the host tree is deciduous Zotz and Winter Indeed, it has been found that differing microenvironments in evergreen and deciduous trees can lead to significant differences in epiphyte cover Cardelus The situation is complex, however, because other traits, e.

The work presented here was initiated to understand the effect of tree leaf phenology on vascular epiphytes at different levels, from leaf traits of individual epiphytes and demographic parameters to community composition, both at a taxonomic and a functional level. To this end, we documented the differences in environmental conditions in the crowns of five tree species on Barro Colorado Island BCIwhich were deciduous, semi-deciduous and evergreen.

This biological reserve in the Gatun Lake of the Panama Canal, which is administered by the Smithsonian Tropical Research Institute STRIis covered by a semi-deciduous lowland forest with varying canopy heights of up to 50 m Leigh et al. The annual precipitation averages mm with a pronounced dry season from January through April.

During these 4 months, both total rainfall and its frequency are strongly reduced.

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Rainless periods regularly expand to around 10 days, the maximum rainless period on record being 31 days based on data from to measured by the Physical Monitoring Program of STRI— http: Five tree species were selected for the study. Selection criteria were emergent crowns, thus deciduousness would have a strong effect on microclimate; more or less even distribution across the island; accessibility with the single rope technique. The following species were selected: Moraceae; both evergreenCeiba pentandra L.

Malvaceae as a semi-deciduous species and Pseudobombax septenatum Jacq. Dugand and Cavanillesia platanifolia Humb. Kunth both deciduous Malvaceae. In the following, species are addressed by their genus names. The deciduous trees are leafless during the entire dry season Fig. Semi-deciduous Ceiba trees usually lose their leaves only for a few weeks, but every 4 to 5 years when flowering and fruiting the leafless phase lasts for up to 20 weeks Windsor, unpublished.