Co2

Buffering capacity

Small

Large

Ecophysiological functions H2O-saving; preventing Effectively preventing photoinhibition to some photoinhibition extent

Ecophysiological functions H2O-saving; preventing Effectively preventing photoinhibition to some photoinhibition extent

stance. This may sustain the very high nocturnal vacuolar acid accumulation observed in Clusia with day-night changes of titratable proton levels of more than 1 M (Table 6.8).

The diversity of the hemi-epiphyte and strangler Clusia in the tropics has presented us with many unexpected observations and stimulating new reflections on the nature of ecophysiological adaptations. Even individual species can occur as the life forms of free standing terrestrial trees, stranglers, hemi-epiphytes and epiphytes. There are ca. 350-400 species of Clusia occupying a wide range of habitats, e.g. coastal rocks and sand dunes, savannas, gallery forests, open shrub land, dry low land forest, secondary shrub forest, dry montane karstic limestone forest, montane rain forest, upper montane rain forest, cloud and elfin forest, rock outcrops (inselbergs; see Sect. 11.3.2; Fig. 6.31, see also Fig. 3.6D), and again individual species may be found in several of these types of sites (Table 6.10). Perhaps Clusia is so successful because of the high degree of physiological plasticity. This also makes it suitable for reclamation of tropical land by afforestation and as an ornamental tree even in the center of cities.

Fig. 6.31A-E A A Habitat diversity of Clusia. A Clusia fluminensis on sand dunes in the restinga formation on the Atlantic coast near Rio de Janeiro, Brazil. B,C Clusia rosea on granite rocks on the British Virgin Island Virgin Gorda (Lesser Antilles), with aerial adventitious root systems in the rock furrows in C. D Clusia sp. Gran Sabana, Venezuela, with an epiphytic bromeliad Catop-sis berteroniana. E Clusia rosea in montane rainforest on the US Virgin Island St. John (Lesser Antilles)

Fig. 6.31A-E A A Habitat diversity of Clusia. A Clusia fluminensis on sand dunes in the restinga formation on the Atlantic coast near Rio de Janeiro, Brazil. B,C Clusia rosea on granite rocks on the British Virgin Island Virgin Gorda (Lesser Antilles), with aerial adventitious root systems in the rock furrows in C. D Clusia sp. Gran Sabana, Venezuela, with an epiphytic bromeliad Catop-sis berteroniana. E Clusia rosea in montane rainforest on the US Virgin Island St. John (Lesser Antilles)

Table 6.10 Ecological amplitude of individual species of Clusia, C. multiflora (C. mu.), C. parv-iflora (C.p.), C. rosea (C.r.), C. fluminensis (C.f.), C. minor (C.mi.), C. criuva (C.c.) with their modes of photosynthesis indicated (From Lüttge 2007b)

C.mu. C.p. C.r. C.f. C.mi. C.c. C3 C3 CAM CAM C3/CAM weak

CAM inducible

Coastal rocks •

cerrado ecotone

limestone forest

Upper montane •

rain forest

Cloud forest/fog forest/ • elfin forest

6.6.3 Mineral Nutrients

Some special adaptations to the poor nutrient supply in the epiphytic habitat have already been mentioned above in the discussion of life forms of epiphytes (Sects. 6.3 and 6.4) and in relation to water stress (Sect. 6.6.2). They include the use of humus accumulation in trees (Fig. 6.10A) and morphological features of the plants for collecting humus such as the formation of baskets and nests (Fig. 6.10B) as well as tanks (Figs. 6.15 and 6.10C,D). Scales (epidermal trichomes) of bromeliads (Fig. 6.14) and the velamen radicum of aerial roots of aroids and orchids serving atmospheric nutrition also belong to the specialised plant structures formed for nutrient and water uptake (Goh and Kluge 1989; Reinert 1998). The velamen is a mul-tilayered peripheral structure, which is readily infiltrated by water from throughfall or stemflow (Fig. 6.32).

It has even been argued that the successful trapping of rain and throughfall, enriched by leachates from leaves and stems, is nutritional piracy, depriving

Fig. 6.32A-F Velamen radicum in epiphytic orchids. A Velamen of a Dendrobium species. V velamen; Ex exodermis; C cortex. B Detail from A showing the dead velamen cells with the typically perforated walls. C Detail from A showing the exodermis with an aeration cell (AC). D Surface view of an aerial root of Vanda tricolor with dry velamen. The air-filled velamen cells appear homogeneously whitish. E The same detail as in D; however, after wetting the velamen. With the exception of the pneumatothodes (PN), the velamen cells are filled with water and thus appear dark. F Cross-Sect. through the water-imbibed velamen of Vanda tricolor in the region of the pneu-matothode (PN) and aeration cells (AC). The air-filled cells of the pneumatothode appear white. (Goh and Kluge 1989)

Fig. 6.32A-F Velamen radicum in epiphytic orchids. A Velamen of a Dendrobium species. V velamen; Ex exodermis; C cortex. B Detail from A showing the dead velamen cells with the typically perforated walls. C Detail from A showing the exodermis with an aeration cell (AC). D Surface view of an aerial root of Vanda tricolor with dry velamen. The air-filled velamen cells appear homogeneously whitish. E The same detail as in D; however, after wetting the velamen. With the exception of the pneumatothodes (PN), the velamen cells are filled with water and thus appear dark. F Cross-Sect. through the water-imbibed velamen of Vanda tricolor in the region of the pneu-matothode (PN) and aeration cells (AC). The air-filled cells of the pneumatothode appear white. (Goh and Kluge 1989)

host trees of resources which otherwise would reach their rooting medium (Ben-zing 1989a,b, 1990). A comparison of the nitrogen content in leaves of facultative epiphytes at adjacent sites, showed significantly lower N-levels in two aroid species growing epiphytically as compared to their terrestrial counterparts. N-content was similar in two tank-forming bromeliads, whereas in seedlings of Clu-sia low N-content was also related to growth directly on the phorophyte and not to growth inside bromeliad tanks (Fig. 6.33). Epiphytes, especially in the Orchidaceae, may form mycorrhizas (Reinert 1998). The fungal hyphae penetrate the epiphytes as well as the decaying bark of the host phorophyte. This is evidently piracy of a more overt kind and the tree effectively becomes the pedosphere of the epiphyte (Ruinen 1953; Johannson 1977; Benzing 1982; Benzing and Atwood 1984). An interesting example of counter-piracy is found when phorophytes produce adventitious canopy roots which exploit the nutrient debris collected within the epiphyte cover (Nadkarni 1981).

Analyses of the stable isotope 15N have been used to trace several possible sources of nitrogen and processes of nitrogen acquisition in tropical epiphytes as compared to associated soil rooted trees (Stewart et al. 1995; Reinert 1998; Hietz et al. 1999; Wania et al. 2002). The relations are complex. N-isotope signatures of epiphytes vary with canopy position and the related water supply. The variations

Fig. 6.33 Comparison of nitrogen levels in cohabitant epiphytic and terrestrial-life-forms of two aroid and two bromeliad species and of Clusia rosea. Data show N content in leaves of epiphytic minus leaves of terrestrial plants of the same species (numbers are p values of a t test for statistically significant differences or n.s. = non-significant). (Ball et al. 1991)

Fig. 6.33 Comparison of nitrogen levels in cohabitant epiphytic and terrestrial-life-forms of two aroid and two bromeliad species and of Clusia rosea. Data show N content in leaves of epiphytic minus leaves of terrestrial plants of the same species (numbers are p values of a t test for statistically significant differences or n.s. = non-significant). (Ball et al. 1991)

are attributable in part to altered 15N-discrimination during N-acquisition, i.e. atmospheric deposition, leachates and biological dinitrogen fixation, and to changes in partitioning of N isotopes within the plant. Epiphytes may make considerable use of biological N2-fixation by cyanobacteria and free living N2-fixing bacteria of the phyllosphere (see Sect. 6.1) including their own leaves. Brighigna et al. (1992) described the trichome layers of bromeliad leaves as a favourable habitat for microbes including N2 fixing bacteria.

Two additional strategies involve interactions with animals, one of which is predatory and the other one symbiotic, namely:

• mutualism with ants.

Carnivory by plants is quite frequent in the tropics and subtropics (Fig. 6.34). In the plant kingdom, carnivory is generally assumed to be a mechanism for the acquisition of mineral nutrients, especially N, P and S, by photosynthetically autotrophic plants living in nutrient-poor habitats such as peat bogs (Schmucker and Linnemann 1959; Luttge 1983). Hence, one would assume that carnivorous plants would be rather frequent in the canopy habitat. However, this is not the case. Carnivorous plants have developed special organs for the capture of prey, glands for digestion and absorp-

Ce.= Cephalotus Da.= Darling ton ia Di.= Dionaea Dr= Drosophyllum He.= Heliamphora Ro.=Roridula

Fig. 6.34 Global distribution of carnivorous plants. The genus Drosera is almost ubiquitous and occurs between the two lines drawn in the north and in the south respectively, except in the areas left white. The exclusively tropical and subtropical distribution of the genera Nepenthes and Byblis is clearly seen and the islands of endemic Heliamphora in Northern South America should be noted. Distribution of carnivorous genera of the family Lentibulariaceae is not shown on this map, namely Pinguicula: Northern temperate zone; Genlisea: Tropical South America, South Africa; Utricu-laria: cosmopolitan; Biovularia: Cuba, Tropical South America; Polypompholyx: South America, Australia. (After Schmucker and Linnemann 1959)

Ce.= Cephalotus Da.= Darling ton ia Di.= Dionaea Dr= Drosophyllum He.= Heliamphora Ro.=Roridula

Fig. 6.34 Global distribution of carnivorous plants. The genus Drosera is almost ubiquitous and occurs between the two lines drawn in the north and in the south respectively, except in the areas left white. The exclusively tropical and subtropical distribution of the genera Nepenthes and Byblis is clearly seen and the islands of endemic Heliamphora in Northern South America should be noted. Distribution of carnivorous genera of the family Lentibulariaceae is not shown on this map, namely Pinguicula: Northern temperate zone; Genlisea: Tropical South America, South Africa; Utricu-laria: cosmopolitan; Biovularia: Cuba, Tropical South America; Polypompholyx: South America, Australia. (After Schmucker and Linnemann 1959)

tion of low molecular compounds obtained from the prey, and mechanisms for the attraction of small animals such as showy and colourful appendages and production of scent and nectar (see also Sect. 10.2.3.3). Thus, in cost/benefit analyses the rarity of carnivorous plants in epiphytic habitats can be explained by the high costs for investment and maintenance of these complex attraction and capture mechanisms (Givnish et al. 1984; Ellison 2006). Since other factors, particularly water and often light, are equally limiting, a cost-benefit analysis suggested carnivory would not be effective under these circumstances.

On the other hand, there are a few examples of carnivorous plants among climbers and epiphytes. The pitcher plant genus of Nepenthes is native in the Mala-ian archipelago and an exclusively tropical genus (Fig. 6.34). There are 71 species of Nepenthes altogether, among which a few are purely terrestrial, but 6 are epiphytic and many are climbers (Fig. 6.35). The pitchers attract prey by their shiny and often

Fig. 6.35 Nepenthes gracilis (Malaysia)

Fig. 6.36 ► Nepenthes. A Pitcher cut open showing the velvety covering with loose wax particles in the upper part and the gland zone below. B Part of the upper pitcher region with wax particles and protrusions. C Scale-covered gland

colourful rim, which also bears nectaries towards the inside of the pitcher opening. Small animals, predominantly insects, having fallen over the slippery collar into the pitcher lumen, rapidly drown in the digestive fluid produced by glands on the bottom, which contains a protease secreted by the plant and other enzymes provided by microorganisms participating in prey digestion. Escape via the pitcher walls is prevented by downward pointing scale-like tissue over the glands (Fig. 6.36C), modified stomata with protrusions towards the pitcher lumen and a lubrication with small and loose wax particles in the upper part of the pitcher (Gorb and Gorb 2006; Fig. 6.36A,B). Substances obtained from the digested prey are absorbed via the gland cells.

Some of these traits are also shared by tanks of bromeliads. They often contain dead and putrefying insects and may absorb substances like amino acids from such prey via their scales. In some cases, such as the epiphytic bromeliad Catopsis bert-eroniana, there is also wax at the adaxial leaf surfaces lubricating the tank interior (Benzing 1989b). However, these plants have no glands and do not secrete digestive enzymes, so that at most there is only the initial development towards the carnivorous syndrome (see also Sect. 10.2.3.3).

A more sophisticated example is Utricularia. Many species in this genus are aquatic, forming small bladders from modified leaves along the stems. The bladders are tightly closed by a trap-door, and actively transport ions across the trap wall into the outer medium to drive the osmotic loss of water from the trap lumen. This creates tension in the bladder-walls, which sets the bladder trap. Small animals trigger the opening of the trap door by touching the antennae-like protuberances and are swept into the trap as the tension in the trap wall is released (Fig. 6.37). The animals are then digested inside the bladders. In the tropics, Utricularia species often live

Fig. 6.37 Schematic drawing of a longitudinal section of a trap of Utricularia. i Bladder lumen; o outer medium; D trap door; M trap mouth; SH sensitive hairs of the trap door = trigger hairs; A, antenna. Dotted arrow opening and closing of the trap door; the tissue beneath the trap door prevents opening from the inside. H1, H2, H3 various types of gland hairs serving prey digestion and transport functions. (After Schmucker and Linnemann 1959)

Fig. 6.37 Schematic drawing of a longitudinal section of a trap of Utricularia. i Bladder lumen; o outer medium; D trap door; M trap mouth; SH sensitive hairs of the trap door = trigger hairs; A, antenna. Dotted arrow opening and closing of the trap door; the tissue beneath the trap door prevents opening from the inside. H1, H2, H3 various types of gland hairs serving prey digestion and transport functions. (After Schmucker and Linnemann 1959)

Fig. 6.38A-C Utricularia humboldtii in the tanks of Brocchinia tatei. A Inflorescence of U. hum-boldtii emerging from a tank of B. tatei. B Outer tank leaves removed to show the basal system of U. humboldtii bladders on stems and leaves with petioles and lamina emerging from the tank. C Larger and smaller U. humboldtii-bladders

Fig. 6.38A-C Utricularia humboldtii in the tanks of Brocchinia tatei. A Inflorescence of U. hum-boldtii emerging from a tank of B. tatei. B Outer tank leaves removed to show the basal system of U. humboldtii bladders on stems and leaves with petioles and lamina emerging from the tank. C Larger and smaller U. humboldtii-bladders

Fig. 6.39A-D Ant-house epiphytes. A, B The orchid Schomburgkia humboldtiana with pseudob-ulbs cut open in B to show the ants nest. C The bromeliad Tillandsia flexuosa epiphytic on the cactus Pilosocereus ottonis, where ants are nesting in the inflated basal part of the tank. D The Asclepiadeaceae Dischidia rafflesiana with adventitious roots in modified leaves

Fig. 6.39A-D Ant-house epiphytes. A, B The orchid Schomburgkia humboldtiana with pseudob-ulbs cut open in B to show the ants nest. C The bromeliad Tillandsia flexuosa epiphytic on the cactus Pilosocereus ottonis, where ants are nesting in the inflated basal part of the tank. D The Asclepiadeaceae Dischidia rafflesiana with adventitious roots in modified leaves epiphytically between mosses on stems of trees. Most cunning is Utricularia hum-boldtii, which lives inside the tanks of the bromeliad Brocchinia tatei (Fig. 6.38).

Another tropical habitat which is often nutrient limited are the savannas. We will refer to carnivorous plants again below, when we discuss their role and their contribution to nutrient turnover in this habitat (Sect. 10.2.3.3).

In symbiotic mutualisms of epiphytes with ants (Huxley 1980) we may distinguish two forms, which among other benefits, provide mineral nutrition to plants, namely (Davidson and Epstein 1989; Benzing 1989b, 1990):

• ant garden epiphytes,

• ant house epiphytes.

Ants frequently construct nests in trees using various materials which are rich in nutrients (forming an ant-nest "carton"). Seeds of plants may germinate directly from such a nutritive carton. Since the plants offer various goods in return, such as nectar, fruits and seeds, the ants often disperse and plant the seeds of their epiphytes in ant gardens. Conversely, plants themselves may also provide nesting facilities for ants, e.g. cavities in various parts of the plant body or hollow stems (Figs. 3.23 and 6.39; Sect. 3.3.4.4). Among the epiphytes there are many myrmecophytic species with such ant houses, e.g. orchids with ant nests in pseudobulbs and bromeliads with inflated tank leaves (Fig. 6.39A-C). The ants carry soil and other decaying material into the nests and add their faeces, which gives a debris from which the host plants may absorb nutrients. The pitchers of some species of the Asclepiadaceae Dischidia are most sophisticated, since adventitious roots grow into the soil and debris accumulated by ants inside these containers. In effect, epiphytic Dischidias literally construct their own flower pots (Fig. 6.39D). A study with the Malaysian Dischichia major using stable isotopes to trace sources of N and C (Sect. 2.5) has shown that 29% of the host nitrogen is derived from debris deposited into the leaf pitchers by ants and that 39% of the carbon assimilated by the host is derived from ant-related respiration (Treseder et al. 1995). Ant respiration may increase the CO2 concentration in the pitcher lumen above atmospheric levels. Since the host has stomata on the inner pitcher-wall surface this can be directly used for CO2-fixation even in the dark in this obligate CAM-plant.

In many cases these plant-ant interactions are true symbioses with obligate mutualism, since the partners are no longer successful individually. The epiphytic plants benefit nutritionally and may be protected from herbivores, while the ants obtain nest-sites and various items of food.

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