1.

REVIEW OF RELATED LITERATURE

2.1 Diversity and Distribution of Epiphytes

Vascular epiphytes are plants that derive its moisture and nutrients from the air

and rain, usually grow on trees or shrubs. Without directly harming them, they are

mostly limited to tropical and subtropical forests and may be the most diverse life form

in very humid formations (Hietz, 1999).

Johansson (1974) described that “epiphytes occur in many plant orders that it has

wide-ranging geographical distribution among nonvascular plants such as algae,

mosses, and lichens”. The vascular epiphytes, pteridophytes, and phanerogams are

on the other hand, more or less restricted to tropical regions. In tropical rain forests

especially, they form a significant part of the flora, and their presence and abundance

are often used as a characteristic of such forests (Schimper 1903, Richards 1964).

Vascular epiphytes are rare in temperate climates. In humid climates, such as certain

coastal and high mountain areas, a large number of plant species may occur for

varying periods of time as epiphytes although lacking special adaptations to such a

life (Sharp 1957).

2.2 Ecological Role of Epiphytes

According to Coxson and Nadkarni (1995), here are the five reasons why

epiphytes are important canopy components to ecosystem function: (1) Pools of

nutrients and organic matter generated by epiphytes contained in the canopy

can be high, even exceeding host tree foliage, (2)The contribution of epiphytes

to complex canopy structure may enhance wet and dry deposition by

increasing the physical area of canopy surfaces for impaction and sedimentation,

and by increasing the biotic uptake by nutrient-efficient epiphytic plants, (3) Free-

living and symbiotic biota in the canopy can fix atmospheric nitrogen , (4) Certain

microbial activities such as nitrification in the canopy appear to be suppressed,

relative to the forest floor, which tends to foster nutrients in a form that is less

mobile and leachable, and (5) Pools of canopy-held nutrients are by no means

static; nutrients move from the canopy to the other ecosystem members by

four routes: a) as litter fall via abscission and by "riding down" fallen branches

and whole trees, which is subsequently mineralized at variable rates, b) as

crown-wash in the form of throughfall and stemflow, c) via herbivory and

predation of epiphytes by vertebrates and invertebrates, and d) directly to host

trees via uptake by canopy roots. Furthermore, the importance of epiphytes for

tropical biodiversity is increased because of their providing substrate and food for

many of the animals inhabiting tropical rainforest canopies, which may number

millions of still undescribed species (Hietz, 1999).

2.3 Qualification for Indicator Species

There were known indicators which can reflect the effects of various disturbances

and can be of good use to draw efforts to mitigate them (Milledge, Nelson and Palmer

1991). The following are the indicators which were identified: (1) taxon-based

biodiversity indicators – this kind of indicator species can be used as proxies for other

biodiversity components or for changes in the ecosystem; or (2) structure-based

indicators - forest structure features that can be used as proxies for changes in the

ecosystem (Lindenmayer, Margules, & Botkin, 2000). To further allow hypotheses

tests, Castello, Coelho and Cardoso-Leite (2016) comprehensively discussed that

indicator choice should meet the following standard: (1) be sensitive and respond to

changes in the ecosystem caused by human disturbance, (2) allow easy and reliable

identification, (3) be viable and have low implementation costs.

On a study made by Hietz (1999), the performance, survival and distribution of

epiphytes in disturbed forests or secondary vegetation are influenced by stand density

and microclimate, distance from seed source, tree size and sometimes tree species,

type and history of disturbance, population dynamics of epiphytes and trees, and

epiphyte physiology. Being sensitive to disturbance and microclimate, and because

of their importance for tropical forest ecology, epiphytes may serve as indicators or

guides for careful management. The few studies of Andama, Michir and Luilo (2003)

suggest that epiphytes are a vulnerable group and consequently represent a

 good indicator group of biodiversity that can be monitored to assess the effects of

forest disturbances.

2.4 Effect of Humidity to Epiphytes

Fern distribution in the canopy is correlated with the water availability

Hietz and Briones (1998) studied the correlation between water relations within-

canopy distribution of epiphytic ferns in a Mexican cloud forest. They gave focus on

physiological traits associated with water relations of eight common epiphytic which

were investigated in relation to the distribution of these species within the canopy. In

their paper, it was explained that fern distribution was significantly correlated with the

relative water content at which stomata close, leaf thickness, stomatal density and

size. The result showed that Trichomanes bucinatum desiccated completely within

hours in moderately dry air and was confined to the stem bases, and Asplenium

cuspidatum, with no evident adaptations to cope with drought, grew in the second

most shaded zone within the tree crowns. Despite growing in a humid cloud forest, all

other species had xeric adaptations including coriaceous leaves (Pleopeltis mexicana,

Elaphoglossum glaucum), succulent rhizomes (Polypodium puberulum, Phlebodium

areolatum), low rates of uncontrolled water loss (all species except P. puberulum),

leaf scales (Elaphoglossum petiolatum, Polypodium plebeium), and high cell wall

elasticity (all species). P. plebeium and Pl. mexicanum, which grow in the most

exposed locations, tolerated water loss beyond the turgor loss point before the

stomata closed and appear to be poikilohydric or at least to tolerate high water deficits.
 Inadequacy of water is perhaps the most important abiotic restraint in the epiphytic

habitat. Differences in the evenness of water and nutrient availability prompted

Benzing (1990) to define two functional groups - the ‘continuously supplied’ and ‘pulse

supplied’ epiphytes. The former encompass tank bromeliads and taxa with access to

rooting media with a relatively constant supply of moisture. The latter are comprised

of the remaining forms, so‐called bark epiphytes, in which rainless periods of a few

hours may suffice to cause water stress. Although this dichotomy has some heuristic

value, it conceals substantial variability, not only among different taxa within each

group, but also within a given species. The efficiency of the tanks of epiphytic

bromeliads in bridging rainless periods was analyzed as a function of plant size (Zotz

& Hietz, 2001)

Moreover, canopy cover, luminosity and litter cover associates with each other,

influencing species establishment and growth (Denslow, 1980). Higher canopy cover

leads to low light availability and high litter production, favoring shade tolerant species.

Medeiros and Torezan (2013) found a positive relationship between canopy cover and

ecological integrity.
2.5 Various Forest Disturbances and Its Impact

The loss of habitat of many species of plants represents the greatest threat to

epiphyte survival (Hietz, 1999). It is generally acknowledged that species loss and

the erosion of genetic diversity is highest in the tropics which results to high

deforestation rates which becomes alarming due to the fact that tropical forests holds

relevant amounts of species of plants and animals than any other area of the world.

There are three different types of human impact on tropical forests: (1) Complete

conversion of the original forest to some other arboreal vegetation such as secondary

forests, timber plantations or orchards; (2) fragmentation into smaller patches with

deforested areas in between; and (3) various degrees of disturbance, such as forestry

operations (Hietz, 1999). Disturbance of original forests means any operation that

does not lead to a complete removal of the forest cover but affects some or all species

concerned, and most forestry practices in tropical countries will fall into this category.

The degree of disturbance varies widely from more than half of the tree cover

removed, to the collection of inconspicuous non-timber species.

Further studies was done by Padmawathe, et.al (2004) on effect of selective logging

on vascular epiphyte diversity in a moist lowland forest of Eastern Himalaya. They

specifically studied three epiphytic groups- Orchids, pteridophytes and non-orchid

angiosperm epiphytes in closed, selectively logged and in unlogged forests with tree

fall gaps. It demonstrated that logging reduced the structural complexity of the forests
 and altered their microclimate. With logging, there was a general decline in richness

and abundance of epiphytes except orchids. The abundance and species composition

of pteridophytes and non-orchid angiosperm epiphytes were related to

microclimate and substrate features while their richness were correlated only with

canopy cover. In contrast, orchid species composition was related to forest structure.

It was suggested that a combination of management strategies is required for

conservation of all epiphyte groups. A mosaic of logged and unlogged forest patches

with undisturbed forests in proximity would maintain the diversity of pteridophytes and

other angiosperms. Another study was conducted by Castello, et.al (2015), who

studied disturbance type and intensity – vegetation removal for agricultural land use,

selective logging, selective harvesting of economically important species (e.g. Palm

trees), among others – can lead to species substitution and other structural changes

(e.g. decrease in height and diameter of canopy species) and to a return to an earlier

successional stage (Guariguata and Ostertag, 2001).

2.6 Forest Conservation Status Monitoring

According to Castello, et.al (2015), in order to maintain the ecological integrity of

a forest, it is necessary to provide information about status, condition and conservation

value of protected areas. Monitoring conservation status of protected areas (PA) is a

promising tool for assessing whether conservation objectives have been achieved or

not. However, there are few monitoring programs of PA and many of them are focused

on management and infrastructure (ICMBio, 2011) and not on conservation status,

e.g., meeting biodiversity conservation goals (Le Saout et al., 2013). In developed

countries there are guidelines to monitor biodiversity (Gardner, 2010; Parrotta et al.,

2012), mostly focused on certified forests outside protected areas. In developing

countries, however, they lack systematic programs to monitor PA conservation

effectiveness (Terborg and Davenport, 2002; Gaston et al., 2006; Le Saout et al.,

2013). In addition, the complexity of biodiversity makes it virtually impossible to

monitor all species (Lindenmayer et al., 2000). Therefore, the use of indicators to

assess vegetation conservation status can be a useful tool for monitoring these areas

(Noss, 1990).

For Hietz (1999), great efforts have been made to conserve areas of natural

vegetation and give them various degree of protection. This continues to be an

important and urgent task, but it is clear that severe disturbance and destruction will

not stop soon. It is therefore necessary to study not only diversity in pristine

environments but also the impact of alternative uses and management practices

on biodiversity to conserve as much as possible where disturbance and deforestation

cannot be prevented and, where possible, to improve the conservation value of areas

already degraded. In addition, avoiding the unnecessary damage of trees and saving

large trees also of commercial species as seed sources for future generations of trees

and people is also in the long term economic interest of forest managers and owners.
 A few additional measures such as conserving pockets of completely undisturbed

forest where the microclimate of the forest interior is maintained and perhaps paying

attention to tree species that are particularly good hosts for some or all epiphytes will

conserve most or all species at very little additional costs. These and similar

measures will also help to maintain the diversity of birds and other species. If forests

are cleared for whatever reason, even individual isolated remnant trees are much

better than no trees and may form nuclei for the recolonization in a secondary forest.

2.7 Host Tree Preferences of Vascular Epiphytes

Host tree identity believed to have potential role in the structuring of vascular epiphyte

communities. For decades, it has been proposed that each host tree species has a

specific subset of the local species pool according to its own set of properties such as

physicochemical characteristics of the bark, tree architecture, or leaf phenology

patterns (Laube and Zotz, 2006).

In a study done by Laube and Zotz (2006) on vascular epiphytes on three locally

common host-tree species (Socratea exorrhiza, Marila laxiflora, Perebea

xanthochyma) in Panama, a novel, quantitative approach to this question was

presented, taking advantage of a complete census of the vascular epiphyte

community in 0·4 ha of undisturbed lowland forest. In all three tree species,

abundances of the majority of epiphyte species (69–81 %) were indistinguishable from

random, while the remaining species were about equally over- or under-represented
compared with their occurrence in the entire forest plot. Variations based on the

number of colonized trees (reflecting observed spatial patchiness) yielded similar

results. Finally, canonical correspondence analysis also confirmed host-specific

differences in epiphyte assemblages. In spite of distinct preferences of some

epiphytes for particular host trees, no epiphyte species was restricted to a single host.

The epiphytes on a given tree species are not simply a random sample of the local

species pool, but there are no indications of host specificity either.

More research made by Callaway et.al. (2002) where they investigated species-

specific relationships among two species of vascular epiphytes and ten host tree

species in a coastal plain forest in the southeastern United States. The epiphytes

Tillandsia usneoides and Polypodium polypodioides were said to be highly associated

with particular host species in the field, but host traits that favored colonization were

inadequate to fully explain the epiphyte-host associations for either epiphyte. They did

field transplant experiments that bypassed epiphyte colonization which it

demonstrated the growth of epiphytes as expressively higher on host tree species that

naturally bore high epiphyte loads than on host species with few or no epiphytes.

These species-specific relationships were highly correlated with the water-holding

capacity of the host tree's bark. Positive and negative effects of throughfall, light

attenuation by the canopy, and bark stability did not explain the overall patterns of

host specificity, but did correlate with some epiphyte-host species relationships. The

relative importance of particular host traits differed between the "atmospheric

epiphyte" Tillandsia, and the fern Polypodium, which roots in the bark of its hosts.
 Species-specific interactions among plants, such as those described here, suggest

that communities are more than individualistic assemblages of co-occurring species.

Hietz (1999) also complemented that host tree species had a strong effect on the

composition of the epiphyte community, a fact related to differences in bark chemistry

of the hosts. He added that strong differences between host tree species, even of the

same genus, in their communities of epiphytic cryptogams were also found in tropical

rainforests. However, evidence of host specificity or preference among vascular

epiphytes is limited. Some orchids significantly prefer one host over another which

may be related to bark chemistry inhibiting successful germination. In most cases,

the suitability of a host for vascular epiphytes appears to be related to its size, bark-

roughness and branching patterns, affecting all or most species and there appears to

be little true specificity. Probably the physiology of cryptogams and especially their

mode of water uptake makes them more dependent on substrate chemistry than

vascular plants, and orchids are indirectly affected through their obligate symbiosis

with mycorrhizal fungi.

2.8 Economic Importance of Epiphytic Ferns

Ferns are selected for their unique foliage color, upright habit, round, delicate

beauty and excellent garden performance and are cultivated as ornamental

plants. These species are distributed all over the world. According to Zhigila, et.al,

(2015), ferns are mostly located in the wild and in few cities in Nigeria. However, they
are most times neglected, less cultivated, underutilized, unrecognized plants and

there is lack of awareness of the importance of ferns as ornamental, medicinal, food,

dyeing, for environmental protection and management. The ferns as ornamental

plants can be retained in the environments for years producing fronds for various

economic importance.

Moreover, an extension specialist in floriculture at Kentucky, UK, Anderson (2004)

suggested that ferns are admirable plants for interior decoration since in most cases,

they can tolerate filtered to low light conditions and continue to grow. While terrestrial

ferns are often limited by insufficient humidity in the interior environment, epiphytic

ferns are adapted to a drier habitat than most terrestrial types which made them more

suited to the centrally heated, and air conditioned environment. He explained that

cultural techniques are different for epiphytic ferns than for many other houseplants.

Epiphytic ferns naturally occur on the branches of trees in subtropical and tropical

forests. This habitat is much different from most terrestrial habitats and these ferns

have adaptations appropriate to this unusual location. Thus, epiphytic ferns must be

grown under conditions that mimic their natural habitat, or poor growth and plant death

will occur.