Diversity and phytogeography of vascular epiphytes in a tropical ...

Flora ] (]]]]) ]]]–]]] 

Diversity and phytogeography of vascular epiphytes in a 

tropical–subtropical transition island, Taiwan 

Rebecca Hsu a, ,1 , Jan H.D. Wolf b 

a 

Taiwan Forestry Research Institute, NO. 67, Sanyuan street, 100 Taipei, Taiwan 

b 

Universiteit van Amsterdam, Institute for Biodiversity and Ecosystem Dynamics (IBED), P.O. Box 94062, 1090 GB Amsterdam, 

The Netherlands 

Received 14 May 2008; accepted 20 August 2008 

Abstract 

We present the first checklist of vascular epiphytes in Taiwan, based on herbarium specimens, literature records, and 

field observations. Epiphyte phytogeography was analyzed using Takhtajan’s modified division in floristic regions. We 

ascertain the presence of 336 species of vascular epiphytes (24 families, 105 genera) in Taiwan. Pteridophytes 

contribute most species (171 species), followed by orchids (120 species). Epiphytes contribute 8% to Taiwanese floristic 

diversity and epiphyte endemism is near 21.3%. The extensive mountain system is probably the most effective driver 

for epiphyte diversification and endemicity in Taiwan. Phytogeographically, Taiwanese epiphytes exhibit equal affinity 

to the Malesian region, southern China and Indo-China and Eastern Asiatic regions. However, some species have a 

disjunctive distribution between Taiwan and SW China and/or E Himalaya, presumably related to low habitat 

similarity with adjacent China and/or the legacy of Late Quaternary climate change. Vascular epiphyte distribution 

patterns corroborate the phytogeographical separation of the island of Lanyu from the main island of Taiwan along 

Kanto’s Neo-Wallace Line. 

r 2009 Elsevier GmbH. All rights reserved. 

Keywords: Endemism; Epiphyte-quotient; Floristic affinity; Neo-wallace line; Paleotropics; Late Quaternary climate change 

Introduction 

The conspicuous vascular epiphyte community in the 

canopy of wet tropical forests has attracted botanists as 

early as 1888, especially during the second half of the 

last century (Benzing, 1990; Gentry and Dodson, 1987a; 

Johansson, 1974; Kress, 1986; Madison, 1977; Richards, 

1952). These studies have shown that the epiphytic lifeform 

is a successful adaptation of plants to conditions in 

Corresponding author. 

E-mail addresses: ecogarden@tfri.gov.tw (R. Hsu), 

wolf@science.uva.nl (J.H.D. Wolf). 

1 C.-C. Hsu is the corresponding author’s name in Taiwanese. 

ARTICLE IN PRESS 

0367-2530/$ - see front matter r 2009 Elsevier GmbH. All rights reserved. 

doi:10.1016/j.flora.2008.08.002 

www.elsevier.de/flora 

the canopy, comprising ca. 29,000 species, or approximately 

10% of all vascular plants, in 83 different 

families and 876 genera (Gentry and Dodson, 1987a). 

Whereas the number of epiphyte inventories is gradually 

increasing, inventories from the paleotropics are still 

rare and especially from Asia few inventories are 

available (Wolf and Flamenco-S, 2003). In addition, 

little is known about epiphytes in tropical–subtropical 

transition zones. Consequently, the differences in 

vascular epiphyte diversity and composition between 

temperate and tropical areas and between paleotropics 

and neotropics remain ambiguous and lack generally 

accepted explanations (Benzing, 1987; Gentry and 

Dodson, 1987a; Zotz, 2005). 

Please cite this article as: Hsu, R., Wolf, J.H.D., Diversity and phytogeography of vascular epiphytes in a tropical–subtropical transition island, 

Taiwan. Flora (2009), doi:10.1016/j.flora.2008.08.002

2 

Taiwan (formerly known as Formosa) is a continental 

island, separated from Southeast China by 

the ca. 200 km wide Taiwan Strait, which reaches a 

depth of 70 m. The Tropic of Cancer crosses through 

the middle of the southern half of the island, and 

about 70% of the total area is covered by mountains. 

Taiwan owes its existence to a collision of the 

Philippines Sea plate with the Eurasian continental 

margin some 5 million years ago, which induced 

orogenesis (Ho, 1988). In contrast to many other 

regions at the tropic of Cancer or Capricorn, 

Taiwan has a humid climate thanks to the high 

mountains that induce cloud formation in high-humidity 

oceanic winds. Frequent typhoons in summer and 

NE monsoon in winter provide most precipitation 

throughout the year. 

Taiwan floristic diversity is high, comprising ca. 4077 

species (Hsieh, 2003). Being a mountainous island, 

species diversity is the result of great habitat heterogeneity. 

Furthermore, situated at the transition from 

tropics to subtropics, in Taiwan many tropical plant 

species reach their northern limit (Hsueh and Lee, 2000), 

whereas temperate species are found in the high 

mountains (Hosokawa, 1958). Phytogeographically, 

Taiwan belongs to the Eastern Asiatic region (Takhtajan, 

1986). Yet the south end of Taiwan, Henchun 

Fig. 1. Location of Taiwan, Lanyu, Lutao, and the Neo- 

Wallace Line (Kanto, 1933). 


R. Hsu, J.H.D. Wolf / Flora ] (]]]]) ]]]–]]] 

Fig. 2. Takhtajan’s floristic regions. Numbers indicated: 2, 

Eastern Asiatic region; 2–20, Ryukyu islands; 2–25, SW China; 

2–27, E Himalaya; 12, Sudano-Zambezian region; 15, Madagascan 

regions; 16, Indian region; 17, Indochinese region; 18, 

Malesian region; 18–104, Philippines; 19, Fijian region; 20, 

Polynesian region; 22, Neocaledonian region; 29, NE Australian 

region. Regions that not covered in above map but with 

Taiwanese epiphyte occurrence are: 3, North American 

Atlantic region; 4, Rocky Mountain region; 6, Mediterranean 

region; 8, Iran-Turanian region; 9, Madrean region; 10, 

Guineo-Congolian region; 21, Hawaiian region; 23, Caribbean 

region; 24, Guayana Highlands; 25, Amazonian region; 26, 

Brazilian region; 27, Andean region. The figure was modified 

from Takhtajan (1986). 

Peninsula, and two small volcanic islands, Lanyu and 

Lutao, located in the south-eastern Taiwan, are 

pertained to Malesian region (Figs. 1 and 2). The 

vegetation of Lanyu is characterized by tropical rain 

forests, and its flora and fauna have more in common 

with the Philippines than with Taiwan. On this basis, 

Kanto (1933) proposed the Neo-Wallace Line by 

extending the boundary of Dickerson and Merrill’s Line 

(Dickerson, 1928) from northern Luzon to Lanyu 

through the middle sea of Lanyu and Lutao (Fig. 1). 

Kanto’s proposal was corroborated by several subsequent 

biogeological studies (Hosokawa, 1958; Kanehira, 

1935; Yen et al., 2003). 

In this study we describe the epiphyte flora of 

Taiwan for the first time. Specifically, we address the 

following research questions: (i) Is species richness, 

endemism, and familial makeup similar to that of 

other floristic regions such as tropical and temperate 

areas in the neotropics? (ii) What is the phytogeographical 

affinity of epiphytes and several 

sub-categories? (iii) Do epiphytes corroborate the Neo- 

Wallace Line? 



Materials and methods 

Study site 

Taiwan is situated between 21145 0 N–25156 0 N and 

119118 0 E–124134 0 E with an area of 36,000 km 2 (Fig. 1). 

The Central Ridge of Taiwan comprises over 200 peaks 

higher than 3000 m a.s.l., and Yushan is the highest 

(3952 m) peak in this island. The annual rainfall ranges 

from 1000 to over 6000 mm (data from 1949 to 2004). 

Mean monthly temperature in the lowlands ranges from 

15 to 20 1C, and is about 28 1C in summer. Based on 

bioclimatic analyses, Taiwan can be classified into seven 

climatic regions, and Lanyu is separated independently 

(Su, 1984, 1992). Lanyu (ca. 46 km 2 , also known as 

Botel Tobago, Kotosho, and Orchid I.) and Lutao 

(ca. 16 km 2 , Green I., Kwasyoto I., and Samasana I.) are 

small tropical islands located at 22103 0 N, 121132 0 E and 

22140N, 121129E, respectively. During summer and 

early autumn, typhoons frequently hit Taiwan, which 

have less impact in western Taiwan, sheltered by the 

Central Ridge. 

Epiphyte definition 

We define epiphytes as organisms that grow on plants 

without extracting water or nutrients from hosts’ living 

tissues, following Barkman (1958). In this paper, focus is 

on vascular plants, but many other epiphytic organisms 

are found in the canopy of the forest. In addition, it is 

not rare to find accidental epiphytes growing on other 

plants, which are unable to reproduce in the canopy 

(Moffett, 2000). We excluded accidental epiphytes 

from our checklist and classified vascular epiphytes in 

following sub-categories: 

(i) Holo-epiphytes: epiphytes that complete their entire 

life cycle without contacting the forest floor 

(Benzing, 1990). 

(ii) Hemi-epiphytes: epiphytes that complete part of 

their life cycle as terrestrial plants. Primary hemiepiphytes 

begin their life cycle as epiphytes and 

eventually send their roots to the ground (e.g. 

strangler figs), whereas secondary hemi-epiphyte 

seedlings germinate terrestrially to become epiphytic 

secondarily when their rooting shoots decompose 

(e.g. aroids). 

(iii) Facultative epiphytes: species in which some individuals 

are terrestrial. 

Epiphyte checklist 

Botanically, Taiwan is one of the best explored 

regions in the tropics. The national database houses 

over 200,000 botanical records (ca. 60% of herbarium 


R. Hsu, J.H.D. Wolf / Flora ] (]]]]) ]]]–]]] 3 

collections). We gratefully made use of this wealth of 

information, scrutinizing for epiphytes in well-known 

epiphytic taxonomic groups (Benzing, 1990). In addition, 

we used epiphyte records in published plant 

inventories and floras. Nomenclature follows the 2nd 

edition of the Flora of Taiwan (Boufford et al., 2003). 

To compile this checklist, species listed in Flora of 

Taiwan were examined one by one, and the approximate 

number of epiphytes was ascertained. 

Phytogeography analyses 

We assessed the presence of Taiwanese vascular 

epiphytes in Takhtajan’s floristic regions (Takhtajan, 

1986). The floristic provinces, SW China, E Himalaya, 

Ryukyu and Philippines under Eastern Asiatic and 

Malesian regions of Takhtajan’s system, were recognized 

independently (Fig. 2). Species geographical 

distributions were characterized based on the flora of 

Taiwan and collections in the global biodiversity 

information facility (GBIF) online database. For 

smaller floristic provinces, such as SW China and 

Ryukyu, floras of Japan and China were consulted to 

determine the specific occurrence locations. 

Results 

Species richness, family makeup, and endemism 

There are 336 species of vascular epiphytes in 105 

genera and 24 families in Taiwan and two subsidiary 

isles, Lanyu and Lutao (Appendix A). Obligate holoepiphytes 

comprise 271 (81%) species, 41 (12%) species 

are facultative holo-epiphytes, and 7 (2%) and 17 (5%) 

species are primary and secondary hemi-epiphytes, 

respectively. 

The Taiwanese epiphyte flora is dominated by 

Pteridophytes, i.e. ferns and fern allies, comprising 171 

species (Table 1). The number of orchids is also 

substantial, 120 species (Fig. 3). The 10 most speciesrich 

families contain 89% of all epiphytes and the 

remaining plant families with epiphytic representatives 

only contribute about 11% to total epiphyte richness 

(Fig. 3). At the genus level also, epiphytism is 

concentrated in few taxa. Only 5% of the genera contain 

more than 10 species and 54 (51%) genera are 

represented with one single species only in the region. 

More than a quarter of native Pteridophytes (Table 1) 

and 36% of native orchids are epiphytes. In contrast, the 

Epiphyte-quotient (Ep.-Q, Hosokawa, 1950), i.e. the 

proportion of epiphytic species in the flora, is only 

approximately 8% (Table 1). 

Of the 336 epiphytes, 75 are endemic species. 

Sixty-nine species are confined to Taiwan, and one 



4 

disjunctively occurs in Taiwan and Lanyu. Despite the 

small size of Lanyu and Lutao, five species are confined 

here (four species are endemic to Lanyu, and one species 

is shared by both). The proportion of Taiwan endemic 

epiphytes (21.3%, Table 2) is less than that in the entire 

flora (26.2%, Hsieh, 2003). Most endemic epiphytes are 

orchids (54.2%) despite overall higher number of 

epiphytic pteridophytes in Taiwan. Of all 114 epiphytic 

orchids, 38 species (33.3%) are endemic to Taiwan, 

as opposed to 19 species (11.2%) of pteridophytes 

(Table 2). 

Epiphyte phytogeography 

With respect to phytogeographical region, about 41% 

of epiphytes in Taiwan also occur in the Malesian 

region, including 10% of species shared with only the 

Philippines (Table 2). About 39% of species are shared 

with Indo-China, and about the same proportion is 

shared with Eastern Asiatic regions, which cover 

temperate E Asia, E Himalaya, SW China, and Ryukyu. 

The islands Lutao and Lanyu share most species (over 

70%) with the Malesian region, whilst Lutao has a high 

proportion (40%) of species that also occur in temperate 


Table 1. Contribution of vascular epiphytes to the flora of Taiwan in various taxonomic categories (data Flora of Taiwan, 

Boufford et al., 2003). 

All vascular plants Ferns and allies Angiosperm Dicotyledons Monocotyledons 

Families 24/235(10%) 12/37(32%) 12/190(6%) 10/151(7%) 2/39(5%) 

Genera 105/1419(7%) 48/145(33%) 57/1257(5%) 16/901(2%) 41/356(12%) 

Species 336/4077(8%) a 

171/629(27%) 165/3420(5%) 40/2410(2%) 125/1010(12%) 

a Epiphyte-quotient. 

Moraceae (11) 3% 

Piperaceae (13) 4% 

Vittariaceae (10) 3% 

Lycopodiaceae (10) 3% 

Davalliaceae (11) 3% 

Aspleniaceae (17) 5% 

Grammitidaceae (19) 6% 

Hymenophyllaceae (31) 

9% 


other families (37) 11% 

Polypodiaceae (57) 17% 

E Asia. Only Lanyu shares an exceptional high 

proportion (22%) of species with the Philippines 

(Table 2). 

Overall, epiphytic ferns shared more species with 

other floristic regions than total epiphytic species 

(Table 2). Over 40% of Taiwanese epiphytic ferns also 

occurred in Eastern Asiatic, Malesian, and Indochinese 

regions. Epiphytic orchids exhibited the highest affinity 

(35%) to Indo-China, yet shared no species with 

Neotropical and Holarctic areas, except E. Asia. 

Discussion 

Orchidaceae (120) 36% 

Fig. 3. Ten most species-rich epiphytic families and their contribution to total epiphyte flora in Taiwan. Numbers in parentheses are 

species numbers. Shading indicates Pteridophyta. 

Species richness and taxonomic distribution 

For a paleotropical region, the island of Taiwan is 

with 336 species rich in epiphytes (Table 1). There is no 

distinct dry season in Taiwan and abundant rainfall and 

warm climate promote epiphyte species richness and 

growth. Another reason why epiphyte richness is high 

may be that Taiwan served as a refuge during Late 

Quaternary climate change, which has been used 

to explain the exceptionally high diversity in Taiwan 



(4077 plant species; further discussed below). In view of 

this high floristic diversity, Taiwan may even be 

considered relatively poor in vascular epiphytes. The 

contribution of vascular epiphytes to total vascular flora 

is only 8%, whilst the EP.-Q worldwide is nearly 10%. 

Moreover, about 36% of orchids are epiphytic in 

Taiwan, which is far less than the 70% worldwide level 

(Atwood, 1986). Possibly frequent tropical storms have 

reduced epiphyte diversity in Taiwan. On average, five 

typhoons hit Taiwan each year (data from 1958 to 2007, 

Central Weather Bureau). Typhoons may have a 

dramatic impact on forest canopies and cause understory 

light levels to increase to 30% of outside levels 

(Lin et al., 2003). Similarly, low epiphyte diversity in 

Puerto Rico has been attributed to island isolation 

and large-scale hurricane disturbances (Migenis and 

Ackerman, 1993). 

Epiphyte richness in neotropical areas, moreover, is 

generally higher. For example, Wolf and Flamenco-S 

(2003) report 1173 species for the state of Chiapas, 

Mexico (75,000 km 2 , 161N–181N). Typical for any 

epiphyte flora, the diversity is concentrated in few taxa 

(Fig. 3, Table 1). In contrast to the Neotropics, 

paleotropical areas lack particularly species-rich epiphyte 

families (e.g. Bromeliaceae, Cactaceae, and 

Marcgraviaceae) and genera in the orchids (e.g. Pleurothallis, 

1500 spp.; Epidendrum, 720 spp.; Maxillaria, 

570 spp.; Stelis, 540 spp.) and in the aroids (Anthurium, 

600 spp.; Philodendron, 350 spp. – Benzing, 1990). In 

Taiwan, the most abundant epiphytes are ferns, and in 

this respect Taiwanese epiphyte flora is typical for 

temperate regions. However, in comparison with other 

vegetation types, ecosystems, and floristic regions, the 

relative proportion of epiphytic ferns and orchids of 

Taiwan is not dramatically different, showing a transition 

from tropical to temperate regions (Table 3). A high 



Table 2. Floristic affinity of Taiwan epiphyte flora with phytogeographical regions, following Takhtajan (1986). 

Floristic regions Taiwan (324) Lanyu (69) Lutao (25) Pteridophytes (170) Orchids (114) 

Eastern Asiatic Region 38.9 (126) 50.7 (35) 64.0 (16) 48.8 (83) 25.4 (29) 

China, Japan, Korea 27.2 (88) 21.7 (15) 40.0 (10) 31.2 (53) 20.2 (23) 

E. Himalaya & S.W. China 13.0 (42) 4.4 (3) 0.0 (0) 13.5 (23) 13.2 (15) 

Ryukyu 13.0 (42) 29.0 (20) 24.0 (6) 18.2 (31) 6.1 (7) 

Malesian Region 40.9 (132) 71.0 (49) 72.0 (18) 51.8 (88) 25.4 (29) 

Malay archipelago 31.2 (101) 49.3 (34) 64.0 (16) 42.4 (72) 14.0 (16) 

Philippines 9.6 (31) 21.7 (15) 8.0 (2) 9.4 (16) 11.4 (13) 

Indo-China 39.2 (127) 46.4 (32) 60.0 (15) 43.5 (74) 35.1 (40) 

India and Sirilanka 23.5 (76) 29.0 (20) 52.0 (13) 28.8 (49) 14.9 (17) 

Melanesia and Hawaii 12.0 (39) 26.1 (18) 44.0 (11) 20.0 (34) 1.8 (2) 

Africa 4.9 (16) 8.7 (6) 8.0 (2) 7.1 (12) 0.9 (1) 

Australia 9.0 (29) 18.8 (13) 36.0 (9) 12.4 (21) 1.8 (2) 

Neotropis 2.5 (8) 5.8 (4) 4.0 (1) 3.5 (6) 0.0 (0) 

Holarctis other than E.A. 1.5 (5) 2.9 (2) 0.0 (0) 2.9 (5) 0.0 (0) 

Endemicity 21.3 (69) 5.8 (4) 0.0 (0) 11.2 (19) 33.3 (38) 

Given is the proportion (%) and number of Taiwanese species, in parentheses, of epiphytic Taiwanese species per region. 

proportion of ferns and fern allies is probably due to the 

presence of temperate mountains in Taiwan that favour 

epiphytic ferns over, for example, orchids (Kessler et al., 

2001; Zotz, 2005). In Taiwan, no epiphytic orchids are 

found above approximately 2300 m a.s.l. (Gastrochilus 

hoii, pers. comm.) in contrast to epiphytic ferns with 

ultimate altitudes of ca. 3000 m a.s.l. (e.g. Pyrrosia spp., 

Lepisorus spp., Mecodium wrightii, pers. observ.). 

Epiphyte endemism 

Many islands are considered global biodiversity 

hotspots because of high endemicity of insular biota 

(Kreft et al., 2008). Taiwan is no exception, having 

extraordinary plant endemicity. More than 1000 vascular 

plant species are endemic to the island, comprising 

26% of the entire flora. The strikingly high flora 

endemism can be explained by Taiwan’s extensive 

mountain system. Taiwan was formed from the collision 

between the Philippines Sea plate and the Eurasian 

continental margin and gave rise to the Central Ridge of 

Taiwan in Mid Pliocene (3 Ma) (Ho, 1988). Active 

orogenesis induced a massive earthquake in central 

Taiwan as recent as 1999. Orogenesis results in greater 

microhabitat differentiation of mountainous regions, 

which promotes island-wide biodiversity and endemicity. 

Kreft et al. (2008) concluded that in continental 

islands, geographic isolation from the mainland may 

contribute less to species diversity than mountain 

isolation. Our data are in agreement with this conclusion. 

For example, several epiphytic genera of mountainous 

regions, Bulbophyllum (24 spp.), Gastrochilus 

(9 spp.), and Oberonia (7 spp.), show exceptionally high 

endemicity of nearly 50%. Furthermore, Goodyera, a 

mid-elevation (ca. 1500–2000 m a.s.l.) species, evolved 




Taiwan. Flora (2009), doi:10.1016/j.flora.2008.08.002 

Table 3. Epiphyte number of species (S) and taxonomic distribution among floristic regions and vegetation types. 

Location Vegetation type Latitude Rainfall (mm) S Ferns (%) Orchids 

(%) 

Paleotropics 

Taiwan Tropical lowland 

to montane 

temperate forests 

Cameroon Semi-deciduous 

rain forest 

Congo, upper Tropical lowland 

Katanga to montane forests 

SW China, Wet sub-tropical 

Mt. Ailao montane forests 

Liberia, 

Nimba 

mountains 

Neotropics 

Mexico, 

Chiapas 

Tropical 

submontane forest 

Tropical lowland 

to montane 


Ecuador Tropical lowland 

to montane 


Ecuador, Tropical lowland 

Yasuní 

Amazonia 

Ecuador, Río 

Guajalito 

Ecuador, Río 

Palenque 

Costa Rica, La 

Selva 

Costa Rica, 

Santa Rosa 

Costa Rica, 

Monteverde 

Panama, 

Barro 

Colorado 

Guyana, 

Mabura Hill 

Temperate 

Chile, Fundo 

San Martín, 

Valdivia 

Tropical montane 

forest 


wet forest 


rainforest 

Seasonal forest 

(with 6-month dry 

season) 

Tropical montane 

forest 


moist forest 


moist forest 

EQ (%) Endemism 

(%) 

Source Regional area/ 

sampling effort 

21.9N–25.3N 2467 324 52 35 8 21 This study 36,000 km 2 

4.25N–2.5S ca. 1500–1900 78 25 65 Zapfack et al. 

(1996) 

150 trees 

ca. 7.5S–13.4S 780–1500 127 28 62 Schaijes and 

Malaisse (2001) 

109,000 km 2 

24.53N 2450–2700 32 53 0.9 Xu and Liu (2005) 80 trees 

6N–8N 1500–3100 153 25 66 Johansson (1974) 463 trees 

16N–18N 800–5000 1173 21 48 14 Wolf and 

Flamenco-S (2003) 

75,000 km 2 / 

12,276 coll. 

1.4N–5S 100–4500 4231 ca. 5–20 ca. 30–53 ca. 25 35 Küper et al. (2004) 256,370 km 2 

0.63S 2750 313 22 30 21 a 

10 Kreft et al. (2004) 650 ha 

0.23S 2700 122 22 57 28 Nieder et al. 

(2001), Rauer and 

Rudolph (2001) 

400 ha 

ca. 1S 2980 238 12 34 23 Gentry and 

Dodson (1987a) 

170 ha 

10.43N 4000 368 16 30 23 Gentry and 

Dodson (1987b) 

1536 ha 

10.83N 1550 24 29 33 4 Gentry and 

Dodson (1987a), 

Janzen and 

Liesner (1980) 

37,000 ha 

10.3N 2500 878 22 b 

36 b 

29 Haber (2001) 10,500 ha 

9.15N 2750 216 20 38 16 Croat (1978) 1560 ha 

5.33N 2700 216 18 42 13 Ek (1997) 10,000 ha 

Temperate forests 39.63S 16 63 6 Riveros and 

Ramírez (1978) 

6 


ARTICLE IN PRESS

New Zealand Temperate rain 34S–47S 150–5400 50 70 12 2 Oliver (1930) 268,680 km 

forests 

2 

Japan Temperate forest 24N–45N 800–3600 52 63 35 ca. 1 Zotz (2005) 377,873 km 2 

India, west Montane 

ca. 29N 1600 17 76 24 Gupta (1968), 3422 km 

Himalaya, temperate forest 

Zotz (2005) 

Nainital 

2 

North Korea Temperate forest 38N–43N 560–1500 9 100 0 Kolbeck (1995) 120,540 km 2 

World 

28,200 9 71 ca. 10 Madison (1977) 

23,456 11 59 ca. 10 Kress (1986) 

29,505 9 77 ca. 10 Gentry and 

Dodson (1987a) 

a 

Freiberg and Freiberg (2000). 

b 

Ingram et al. (1996) (composition of 256 epiphyte spp.). 



three epiphytic species, including two endemics. This is 

the first report of epiphytism in this genus. Finally, 

endemicity increases with altitude in Taiwan up to 

nearly 60% above 3500 m a.s.l. 

Yet, vascular epiphytes show lower endemism 

(21.3%) than terrestrial plants (Table 2). This may be 

due to their superior dispersal ability; 89% of vascular 

epiphytes in Taiwan disperse by wind. The arboreal 

habitat and dust-like seeds and diaspores enable longdistance 

dispersal. Overall, ferns show wider ranges and 

lower endemicity than angiosperms (Gentry and Dodson, 

1987a; Kelly et al., 2004) (Table 2). In contrast with 

epiphytic seed plants, most large epiphytic fern genera 

are preponderantly pantropical (Gentry and Dodson, 

1987a). Apart from dispersal ability, historical factors 

may also explain species geographical range (Lester et 

al., 2007). Kelly et al. (2004) reported that in the tropical 

Andes species endemism increased from primitive to 

advanced taxonomic groups (bryophytesopteridophytesoangiosperms). 

Furthermore, taxa with narrow 

geographical range are often considered to have high 

speciation rates (Kelly et al., 2004). In this view, the high 

endemism (33%) in Taiwanese epiphytic orchids relates 

to their highly specific pollination system, which, 

together with the fragmented canopy habitat, promotes 

rapid speciation (Benzing, 1987; Gentry, 1982; Gentry 

and Dodson, 1987a; Gravendeel et al., 2004). 

Epiphyte phytogeography 

Taiwan has a relatively unique vascular epiphyte 

flora. The regions with closest affinity are the Malesian 

region, Indo-China, and Eastern Asiatic regions; ca. 

40% of Taiwanese species are shared with those regions. 

Interestingly, about 13% of vascular epiphytes have a 

disjunctive distribution between Taiwan and SW China 

and/or E Himalayan regions (Table 2). This floristic 

disjunction is consistent with Hosokawa’s (1958) finding 

that Taiwan’s flora, especially of the highland, is more 

closely related to SW China and E Himalaya than to 

adjacent coastal provinces of mainland China. Kuo 

(1985) indicated similar observations on Taiwanese 

pteridophyte flora. He found that the pteridophytes of 

warm-temperate forests (500–1800 m a.s.l.) were closely 

related to SW China and the Himalayan regions, whilst 

lowland species showed higher affinity to Ryukyu, 

south-eastern China and Indo-China. 

The simplest explanation for the lower epiphyte 

affinity of Taiwan with adjacent coastal regions of 

south-eastern China is lack of suitable habitats (Kuo, 

1985). Due to long-term population pressure and 

associated agricultural activities, south-eastern China 

has endured extensive habitat change. Since epiphytes 

are most diverse and abundant in old-growth forests 

(Cascante-Marin et al., 2006; Köhler et al., 2007; Wolf, 



8 

2005), epiphyte diversity is especially affected. Furthermore, 

lowland south-eastern China shows little habitat 

similarity with Taiwan mountain areas. 

Late Quaternary climate change offers another 

explanation. On an evolutionary time-scale, epiphytism 

is relatively recent, occurring in evolutionary advanced 

families of ferns and seed plants. Orchidaceae did not 

evolve until the Quaternary (1.6 Ma ago) (Benzing, 

1990). Zotz (2005) discussed the possibility that the 

Pleistocene extinction was one of the limits of epiphytism 

in temperate zones, whilst few temperate areas 

(e.g. Chile, New Zealand, Himalayas, Japan) have a 

high number of epiphytes for being Tertiary refugia. The 

common feature of the flora in these areas is a high 

proportion of autochthonous and monotypic taxa. 

During the ice age in the Quaternary, the sea level in 

the Taiwan Strait dropped, connecting Taiwan with 

mainland Eurasia. According to the projected vegetation 

map of Last Glacial Maximum (LGM, 18,000 ago), 

Eurasia had relatively scarce tree cover with scattered 

areas of close forests in the uplands across southwestern 

China and along the south-eastern coast of 

Eurasia (Ray and Adams, 2001). Presumably, the 

oceanic climate facilitated Taiwan as a refuge during 

Quaternary glaciations. Moreover, apart from high 

endemicity, more than half of plant genera in Taiwan 

are monotypic (Hsieh, 2003). There is an endemic 

monotypic epiphyte genus Haraella (Orchidaceae) 

in Taiwan. Thus, we propose that Late Quaternary 

climate change helps explain the disjunctive distribution 

of many vascular epiphytes between Taiwan 

and south-western China as well as eastern Himalayan 

regions. 

Table A1. The vascular epiphyte checklist of Taiwan. 



Interestingly, the epiphyte flora of Lanyu and Lutao is 

phytogeographically distinct. Lanyu has more affinity 

with the Philippines (22%) in the Malesian region than 

Lutao (8%), whereas Lutao shares more species with 

China, Japan and Korea in the Eastern Asiatic Region 

(40%) than Lanyu (22%) (Table 2). This pattern is in 

agreement with the proposed Neo-Wallace Line based 

on insect distributions (Kanto, 1933). 

In summary, this one of the few epiphyte inventories 

in Asia shows that the Taiwanese epiphyte flora is rich in 

species and has an extraordinarily high endemicity. 

Regional mountain isolation is probably the most 

effective driver for epiphyte diversification in Taiwan. 

Regarding the proportional contribution of epiphytic 

ferns and orchids, Taiwan is transitional between 

tropical and temperate zones. The disjunctive distribution 

of epiphytes between Taiwan and SW China as well 

as E Himalaya suggests low habitat similarity to 

adjacent China and/or a legacy of Late Quaternary 

climate change. Taiwanese vascular epiphyte distributions 

are in agreement with the Neo-Wallace Line. 

Acknowledgement 

We thank Chung S.-W., Yu S.-K., Lu P.-F., Chang 

Y.-H., for sharing personal observations on Taiwanese 

epiphytes in the field. 

Appendix A 

See Table A1. 

No Family Species/taxon Habit Floristic_Region 

1 

Pteridophytes 

Aspleniaceae Asplenium adiantoides FacuE 15, 18, 22, 29 

2 Aspleniaceae Asplenium antiquum E 2 

3 Aspleniaceae Asplenium australasicum E 18, 22, 29 

4 Aspleniaceae Asplenium bullatum E 16, 17 

5 Aspleniaceae Asplenium cuneatiforme E EndemicF 

6 Aspleniaceae Asplenium ensiforme FacuE 2–25, 17, 16 

7 Aspleniaceae Asplenium griffithianum FacuE 2–20, 16, 17 

8 Aspleniaceae Asplenium incisum FacuE 2 

9 Aspleniaceae Asplenium laciniatum E 2–27 

10 Aspleniaceae Asplenium neolaserpitiifolium E 2–20, 17 

11 Aspleniaceae Asplenium nidus E 2–20, 17, 18, 19, 20, 21, 22, 23, 

29, 15, 12 

12 Aspleniaceae Asplenium normale FacuE 2, 15, 17, 18, 20, 29, 12, 21 

13 Aspleniaceae Asplenium oldhami FacuE 2–20, 17 

14 Aspleniaceae Asplenium planicaule FacuE 2, 17, 18–104 

15 Aspleniaceae Asplenium prolongatum FacuE 16, 17, 2 

16 Aspleniaceae Asplenium 

pseudolaserpitiifolium 

E 17 



Table A1. (continued ) 



17 Aspleniaceae Asplenium ritoense FacuE 2, 17 

18 Davalliaceae Araiostegia parvipinnata E 2–25 

19 Davalliaceae Davallia formosana E 17 

20 Davalliaceae Davallia mariesii E 2 

21 Davalliaceae Davallia solida E 17, 18, 22 

22 Davalliaceae Humata chrysanthemifolia E 18–104 

23 Davalliaceae Humata griffithiana E 2–27, 2–25 

24 Davalliaceae Humata pectinata E 18, 20, 29 

25 Davalliaceae Humata repens E 2, 15, 17, 18, 29 

26 Davalliaceae Humata trifoliata E 2–20, 17, 18 

27 Davalliaceae Humata vestita E 17, 18 

28 Davalliaceae Leucostegia immersa E 2–27, 16, 17, 18 

29 Grammitidaceae Calymmodon cucullatus E 16, 18, 22, 29 

30 Grammitidaceae Calymmodon gracilis E 17, 18 

31 Grammitidaceae Ctenopteris curtisii E 18 

32 Grammitidaceae Ctenopteris merrittii E 18 

33 Grammitidaceae Ctenopteris mollicoma E 18 

34 Grammitidaceae Ctenopteris obliquata E 16, 17, 18 

35 Grammitidaceae Ctenopteris subfalcata E 16, 17, 18 

36 Grammitidaceae Ctenopteris tenuisecta E 18 

37 Grammitidaceae Grammitis adspera E 18, 29 

38 Grammitidaceae Grammitis congener E 17, 18 

39 Grammitidaceae Grammitis fenicis E 18–104 

40 Grammitidaceae Grammitis intromissa E 18 

41 Grammitidaceae Grammitis jagoriana E 18 

42 Grammitidaceae Grammitis nuda E EndemicF 

43 Grammitidaceae Grammitis reinwardtia E 18 

44 Grammitidaceae Prosaptia contigua E 16, 18, 19, 20, 22, 29 

45 Grammitidaceae Prosaptia urceolaris E 17, 18 

46 Grammitidaceae Scleroglossum pusillum E 17, 18 

47 Grammitidaceae Xiphopteris okuboi E 2, 17 

48 Hymenophyllaceae Abrodictyum cumingii E 2, 18 

49 Hymenophyllaceae Crepidomanes bilabiatum E 2–20, 17, 18 

50 Hymenophyllaceae Crepidomanes birmanicum E 2, 17, 16 

51 Hymenophyllaceae Crepidomanes kurzii E 16, 17, 18, 29 

52 Hymenophyllaceae Crepidomanes latealatum FacuE 2, 16, 17, 18 

53 Hymenophyllaceae Crepidomanes latemarginale FacuE 2–20, 16, 17, 18 

54 Hymenophyllaceae Crepidomanes palmifolium E EndemicF 

55 Hymenophyllaceae Crepidomanes schmidtianum FacuE 2–27, 18–104 

var. latifrons 


56 Hymenophyllaceae Gonocormus minutus E 2, 16, 17, 18, 20, 22 

57 Hymenophyllaceae Hymenophyllum barbatum E 2, 16, 17 

58 Hymenophyllaceae Hymenophyllum devolii E EndemicF 

59 Hymenophyllaceae Hymenophyllum fimbriatum E 18–104 

60 Hymenophyllaceae Hymenophyllum productum E 17, 18 

61 Hymenophyllaceae Hymenophyllum simonsianum E 2–27 

62 Hymenophyllaceae Hymenophyllum taiwanense E EndemicF 

63 Hymenophyllaceae Mecodium badium E 2, 16, 17, 18 

64 Hymenophyllaceae Mecodium javanicum E 16, 18, 19 

65 Hymenophyllaceae Mecodium oligosorum E 2 

66 Hymenophyllaceae Mecodium polyanthos E 2, 15, 17, 18 

67 Hymenophyllaceae Mecodium wrightii E 2, 4 

68 Hymenophyllaceae Meringium blandum E 18 

69 Hymenophyllaceae Meringium denticulatum FacuE 2–20, 16, 17, 18, 19 

70 Hymenophyllaceae Meringium holochilum FacuE 18 

71 Hymenophyllaceae Microgonium bimarginatum FacuE 2–20, 16, 17, 18, 20, 29 

72 Hymenophyllaceae Microgonium motleyi FacuE 2–20, 16, 17, 18, 20 



10 




73 Hymenophyllaceae Microgonium omphalodes FacuE 2–20, 18, 20, 29 

74 Hymenophyllaceae Microtrichomanes nitidulum E 16, 17, 18, 29 

75 Hymenophyllaceae Pleuromanes pallidum E 16, 17, 18, 20 

76 Hymenophyllaceae Vandenboschia auriculata E 2, 16, 17, 18, 20 

77 Hymenophyllaceae Vandenboschia maxima FacuE 2–20, 17, 18 

78 Hymenophyllaceae Vandenboschia radicans E 2–27, 2–20, 6, 12, 16, 17, 18, 23, 

24, 25, 27 

79 Lomariopsidaceae Elaphoglossum callifolium E 17, 18 

80 Lomariopsidaceae Elaphoglossum commutatum E 10, 12, 15, 16, 18, 21, 25 

81 Lomariopsidaceae Elaphoglossum luzonicum E 18 

82 Lomariopsidaceae Elaphoglossum marginatum E EndemicF 

83 Lomariopsidaceae Elaphoglossum yoshinagae E 2, 17 

84 Lomariopsidaceae Lomariopsis spectabilis E 2–20, 18 

85 Lycopodiaceae Lycopodium carinatum E 2–20, 17, 18, 20, 29 

86 Lycopodiaceae Lycopodium cryptomerianum E 2 

87 Lycopodiaceae Lycopodium cunninghamioides E 2 

88 Lycopodiaceae Lycopodium fargesii E 2 

89 Lycopodiaceae Lycopodium fordii E 2, 16, 17 

90 Lycopodiaceae Lycopodium phlegmaria E 2, 18, 22, 29, 15, 12 

91 Lycopodiaceae Lycopodium salvinioides E 2–20, 18–104 

92 Lycopodiaceae Lycopodium sieboldii E 2 

93 Lycopodiaceae Lycopodium squarrosum E 2, 20, 18, 22 

94 Lycopodiaceae Lycopodium taiwanense E 2–27, 2–20, 16 

95 Oleandraceae Nephrolepis auriculata FacuE 2–20, 16, 17, 18, 9, 23, 24, 25, 

26, 27, 22, 21, 15, 29 

96 Oleandraceae Nephrolepis biserrata FacuE 2–20, 19, 20, 18, 23, 12, 15, 16, 

10, 29, 27, 25 

97 Oleandraceae Nephrolepis multiflora FacuE 2–20, 16, 17, 18–104 

98 Oleandraceae Oleandra wallichii E 2–25, 2–27, 16, 17 

99 Opioglossaceae Ophioderma pendula E 17, 18, 15, 21, 29 

100 Polypodiaceae Aglaomorpha meyeniana E 18–104 

101 Polypodiaceae Arthromeris lehmanni E 2, 16, 17, 18–104 

102 Polypodiaceae Belvisia mucronata E 16, 18, 20, 22, 19, 29 

103 Polypodiaceae Colysis hemionitidea FacuE 2–27, 16, 17, 18–104 

104 Polypodiaceae Colysis pothifolia FacuE 2, 16, 17, 18–104 

105 Polypodiaceae Colysis shintenensis FacuE 2 

106 Polypodiaceae Colysis wrightii FacuE 2–20, 17 

107 Polypodiaceae Crypsinus echinosporus E EndemicF 

108 Polypodiaceae Crypsinus engleri E 2 

109 Polypodiaceae Crypsinus hastatus FacuE 2, 18–104 

110 Polypodiaceae Crypsinus quasidivaricatus FacuE 2–27, 16 

111 Polypodiaceae Crypsinus taeniatus var. 

FacuE 18, 20 

palmatus 


112 Polypodiaceae Crypsinus taiwanensis FacuE EndemicF 

113 Polypodiaceae Crypsinus yakushimensis FacuE 2–20 

114 Polypodiaceae Drymotaenium miyoshianum E 2 

115 Polypodiaceae Drynaria fortunei E 17 

116 Polypodiaceae Lemmaphyllum diversum E 2 

117 Polypodiaceae Lemmaphyllum microphyllum E 2 

118 Polypodiaceae Lepisorus clathratus E 2,8,16 

119 Polypodiaceae Lepisorus kawakamii E EndemicF 

120 Polypodiaceae Lepisorus kuchenensis E 2–25 

121 Polypodiaceae Lepisorus megasorus E EndemicF 

122 Polypodiaceae Lepisorus monilisorus E EndemicF 

123 Polypodiaceae Lepisorus morrisonensis E 2–25, 2–27 

124 Polypodiaceae Lepisorus obscurevenulosus E 2 

125 Polypodiaceae Lepisorus pseudoussuriensis E EndemicF 






126 Polypodiaceae Lepisorus suboligolepidus E 2 

127 Polypodiaceae Lepisorus thunbergianus E 2, 18–104 

128 Polypodiaceae Lepisorus tosaensis E 2 

129 Polypodiaceae Leptochilus decurrens FacuE 16, 17, 18, 20 

130 Polypodiaceae Loxogramme confertifolia E EndemicF 

131 Polypodiaceae Loxogramme formosana E 2–25 

132 Polypodiaceae Loxogramme grammitoides E 2 

133 Polypodiaceae Loxogramme remotefrondigera E EndemicF 

134 Polypodiaceae Loxogramme salicifolia E 2, 17 

135 Polypodiaceae Microsorium buergerianum E 2, 17 

136 Polypodiaceae Microsorium dilatatum E 2–20, 16, 17 

137 Polypodiaceae Microsorium fortunei FacuE 2–27, 2–20 

138 Polypodiaceae Microsorium membranaceum FacuE 2–25, 2–27, 16, 17, 18–104 

139 Polypodiaceae Microsorium punctatum E 16, 17, 22, 29 

140 Polypodiaceae Microsorium rubidum E 2–20, 16, 17, 18, 20 

141 Polypodiaceae Polypodium amoenum E 2–27, 17 

142 Polypodiaceae Polypodium argutum E 2–25, 2–27, 17, 18–104 

143 Polypodiaceae Polypodium formosanum E 2–20 

144 Polypodiaceae Polypodium microrhizoma E 2–25, 2–27 

145 Polypodiaceae Polypodium raishanense E EndemicF 

146 Polypodiaceae Polypodium transpianense E EndemicF 

147 Polypodiaceae Pseudodrynaria coronans E 2–20, 2–25, 2–27, 17 

148 Polypodiaceae Pyrrosia adnascens E 2–20, 16, 17, 18, 20 

149 Polypodiaceae Pyrrosia gralla E 2–25 

150 Polypodiaceae Pyrrosia linearifolia E 2 

151 Polypodiaceae Pyrrosia lingua E 2, 17 

152 Polypodiaceae Pyrrosia matsudae E EndemicF 

153 Polypodiaceae Pyrrosia polydactylis E EndemicF 

154 Polypodiaceae Pyrrosia sheareri E 17 

155 Polypodiaceae Pyrrosia transmorrisonensis E EndemicF 

156 Polypodiaceae Saxiglossum angustissimum E 2 

157 Psilotaceae Psilotum nudum E 2, 17, 18, 21, 22, 29, 10, 12, 15, 

23, 9, 3, 25, 27, 26 

158 Selaginellaceae Selaginella delicatula E 2, 16, 17, 18, 20 

159 Selaginellaceae Selaginella involvens E 2, 16, 17, 18 

160 Selaginellaceae Selaginella stauntoniana FacuE 2 

161 Selaginellaceae Selaginella tamariscina FacuE 2, 16, 18 

162 Vittariaceae Antrophyum formosanum FacuE 2–20 

163 Vittariaceae Antrophyum obovatum FacuE 2, 16, 17 

164 Vittariaceae Antrophyum parvulum FacuE 2–20, 18 

165 Vittariaceae Antrophyum sessilifolium FacuE 18–104 

166 Vittariaceae Vaginularia paradoxa E 16, 18, 20, 21 

167 Vittariaceae Vaginularia trichoidea E 18, 21 

168 Vittariaceae Vittaria anguste-elongata E 18 

169 Vittariaceae Vittaria flexuosa E 2, 16, 17, 18 

170 Vittariaceae Vittaria taeniophylla E 2–27, 2–25, 17, 18–104 

171 Vittariaceae Vittaria zosterifolia E 2–20, 18, 20 

172 

Dicotyledons 

Araliaceae Schefflera arboricola E 17 

173 Asclepiadaceae Dischidia formosana E EndemicF&L 

174 Asclepiadaceae Hoya carnosa E 2, 16, 17 

175 Ericaceae Rhododendron kawakamii E EndemicF 

176 Ericaceae Vaccinium dunalianum var. E EndemicF 

caudatifolium 


177 Ericaceae Vaccinium emarginatum E EndemicF 

178 Gesneriaceae Aeschynanthus acuminatus E 2–27, 16, 17, 18 



12 





179 Gesneriaceae Lysionotus pauciflorus E 2 

180 Gesneriaceae Lysionotus pauciflorus var. 

ikedae 

E EndemicL 

181 Melastomataceae Medinilla formosana E EndemicF 

182 Melastomataceae Medinilla hayataina E EndemicL 

183 Melastomataceae Pachycentria formosana E EndemicF 

184 Moraceae Ficus benjamina HemiE-P 17, 18, 29 

185 Moraceae Ficus caulocarpa HemiE-P 2–20, 17, 18, 16 

186 Moraceae Ficus heteropleura HemiE-P 2–27, 18, 17 

187 Moraceae Ficus microcarpa var. 

microcarpa 

HemiE-P 2–20, 18, 17, 16, 29 

188 Moraceae Ficus microcarpa var. 

crassifolia 

HemiE-P 18–104 

189 Moraceae Ficus pumila HemiE-S 2, 16 

190 Moraceae Ficus pumila L. var. awkeotsang HemiE-S EndemicF 

191 Moraceae Ficus sarmentosa var. henryi HemiE-S 2 

192 Moraceae Ficus sarmentosa var. nipponica HemiE-S 2 

193 Moraceae Ficus superba var. japonica HemiE-P 2, 16, 17, 18 

194 Moraceae Ficus virgata HemiE-P 2–20, 16, 17, 18, 29, 22 

195 Piperaceae Peperomia japonica E 2 

196 Piperaceae Peperomia nakaharai E EndemicF 

197 Piperaceae Peperomia reflexa E 2, 23, 26, 25, 21, 12, 10, 15, 25, 

29 

198 Piperaceae Peperomia rubrivenosa E 18–104 

199 Piperaceae Peperomia sui E EndemicF 

200 Piperaceae Piper arborescens HemiE-S 18 

201 Piperaceae Piper betle HemiE-S 18 

202 Piperaceae Piper interruptum var. 

multinervum 

HemiE-S 18 

203 Piperaceae Piper kadsura HemiE-S 2 

204 Piperaceae Piper kawakamii HemiE-S EndemicF 

205 Piperaceae Piper kwashoense HemiE-S EndemicL&G 

206 Piperaceae Piper sintenense HemiE-S EndemicF 

207 Piperaceae Piper taiwanense HemiE-S EndemicF 

208 Rubiaceae Psychotria serpens HemiE-S 2, 17 

209 Saxifragaceae Hydrangea integrifolia E 18–104 

210 Saxifragaceae Pileostegia viburnoides E 2–20, 16, 17 

211 Urticaceae Procris laevigata E 2–25, 15, 16, 17, 18 

Monocotyledons 

212 Araceae Epipremnum formosanum HemiE-S EndemicF 

213 Araceae Epipremnum pinnatum HemiE-S 2, 18, 20, 29 

214 Araceae Pothoidium lobbianum HemiE-S 18 

215 Araceae Pothos chinensis HemiE-S 2 

216 Araceae Remusatia vivipara E 2–25, 15, 16, 17, 18, 12, 29, 10, 

25 

217 Orchidaceae Acampe rigida E 2–27, 16, 17, 18 

218 Orchidaceae Appendicula fenixii E EndemicL 

219 Orchidaceae Appendicula reflexa E 17, 18 

220 Orchidaceae Arachnis labrosa E 17 

221 Orchidaceae Ascocentrum pumilum E EndemicF 

222 Orchidaceae Bulbophyllum affine E 2–27, 16, 17 

223 Orchidaceae Bulbophyllum albociliatum E EndemicF 

224 Orchidaceae Bulbophyllum aureolabellum E EndemicF 

225 Orchidaceae Bulbophyllum chitouense E EndemicF 

226 Orchidaceae Bulbophyllum drymoglossum E 2 

227 Orchidaceae Bulbophyllum electrinum E 2–25, 17 







228 Orchidaceae Bulbophyllum hirundinis E 17 

229 Orchidaceae Bulbophyllum insulsum E 17 

230 Orchidaceae Bulbophyllum japonicum E 2 

231 Orchidaceae Bulbophyllum macraei E 2, 16 

232 Orchidaceae Bulbophyllum melanoglossum E EndemicF 

233 Orchidaceae Bulbophyllum omerandrum E 2 

234 Orchidaceae Bulbophyllum pectenveneris E 17 

235 Orchidaceae Bulbophyllum pectinatum E 17 

236 Orchidaceae Bulbophyllum pingtungense E EndemicF 

237 Orchidaceae Bulbophyllum retusiusculum E 2–27, 17, 16 

238 Orchidaceae Bulbophyllum riyanum E 17 

239 Orchidaceae Bulbophyllum rubrolabellum E EndemicF 

240 Orchidaceae Bulbophyllum setaceum E EndemicF 

241 Orchidaceae Bulbophyllum taitungianum E EndemicF 

242 Orchidaceae Bulbophyllum taiwanense E EndemicF 

243 Orchidaceae Bulbophyllum tokioi E EndemicF 

244 Orchidaceae Bulbophyllum umbellatum E 2–27, 16, 17 

245 Orchidaceae Bulbophyllum wightii E 16 

246 Orchidaceae Chiloschista segawai E EndemicF 

247 Orchidaceae Cleisostoma paniculatum E 17 

248 Orchidaceae Cleisostoma uraiensis E 2–20, 18–104 

249 Orchidaceae Cymbidium dayanum E 2, 16, 17, 18 

250 Orchidaceae Dendrobium catenatum E 2 

251 Orchidaceae Dendrobium chameleon E 18–104 

252 Orchidaceae Dendrobium chryseum E 2, 16, 17 

253 Orchidaceae Dendrobium crumenatum E 16, 17, 18 

254 Orchidaceae Dendrobium equitans E 18–104 

255 Orchidaceae Dendrobium falconeri E 2–27, 16, 17 

256 Orchidaceae Dendrobium 

E EndemicF 

furcatopedicellatum 

257 Orchidaceae Dendrobium goldschmidtianum E 18–104 

258 Orchidaceae Dendrobium leptocladum E EndemicF 

259 Orchidaceae Dendrobium linawianum E 2 

260 Orchidaceae Dendrobium moniliforme E 2 

261 Orchidaceae Dendrobium somae E EndemicF 

262 Orchidaceae Dendrochilum uncatum E 18–104 

263 Orchidaceae Diploprora championii E 2–27, 16, 17 

264 Orchidaceae Epigeneium fargesii E 2–27, 17 

265 Orchidaceae Epigeneium nakaharae E EndemicF 

266 Orchidaceae Eria amica E 2–25, 2–27, 17 

267 Orchidaceae Eria corneri E 2–20, 17 

268 Orchidaceae Eria japonica E 2–20, 17 

269 Orchidaceae Eria javanica E 2, 16, 17, 18 

270 Orchidaceae Eria ovata E 2–20, 18 

271 Orchidaceae Eria robusta E 18 

272 Orchidaceae Eria tomentosiflora E 18–104 

273 Orchidaceae Flickingeria comata E 18, 29, 19, 20, 22 

274 Orchidaceae Flickingeria tairukounia E EndemicF 

275 Orchidaceae Gastrochilus ciliaris E 2 

276 Orchidaceae Gastrochilus formosanus E 2 

277 Orchidaceae Gastrochilus fuscopunctatus E EndemicF 

278 Orchidaceae Gastrochilus hoii E EndemicF 

279 Orchidaceae Gastrochilus japonicus E 2 

280 Orchidaceae Gastrochilus linii E EndemicF 

281 Orchidaceae Gastrochilus matsudai E EndemicF 

282 Orchidaceae Gastrochilus rantabunensis E 2 

283 Orchidaceae Gastrochilus raraensis E EndemicF 



14 





284 Orchidaceae Goodyera bilamellata E EndemicF 

285 Orchidaceae Goodyera pendula E 2 

286 Orchidaceae Goodyera nantoensis E EndemicF 

287 Orchidaceae Haraella retrocalla E EndemicF 

288 Orchidaceae Holcoglossum quasipinifolium E 2 

289 Orchidaceae Liparis bootanensis E 2, 17, 18 

290 Orchidaceae Liparis caespitosa E 17, 18, 16, 12, 15, 19, 20 

291 Orchidaceae Liparis condylobulbon E 17, 18 

292 Orchidaceae Liparis cordifolia FacuE 2–27, 2–25, 16 

293 Orchidaceae Liparis elliptica E 2, 16, 17 

294 Orchidaceae Liparis grossa E 17, 18–104 

295 Orchidaceae Liparis nakaharai E EndemicF 

296 Orchidaceae Liparis somai E EndemicF 

297 Orchidaceae Liparis viridiflora E 2–27, 16, 17, 18 

298 Orchidaceae Luisia cordata E EndemicF 

299 Orchidaceae Luisia megasepala E EndemicF 

300 Orchidaceae Luisia teres E 2 

301 Orchidaceae Microtatorchis compacta E 18–104 

302 Orchidaceae Oberonia arisanensis E 2–20 

303 Orchidaceae Oberonia caulescens E 2–25, 2–27, 17 

304 Orchidaceae Oberonia gigantea E EndemicF 

305 Orchidaceae Oberonia japonica E 2 

306 Orchidaceae Oberonia pumila E EndemicF 

307 Orchidaceae Oberonia rosea E 17 

308 Orchidaceae Oberonia seidenfadenii E EndemicF 

309 Orchidaceae Papilionanthe taiwaniana E EndemicF 

310 Orchidaceae Phalaenopsis aphrodite E 18–104 

311 Orchidaceae Phalaenopsis equestris E 18–104 

312 Orchidaceae Pholidota cantonensis E 17 

313 Orchidaceae Phreatia caulescens E 18–104 

314 Orchidaceae Phreatia formosana E 2–25, 17 

315 Orchidaceae Phreatia morii E EndemicF 

316 Orchidaceae Phreatia taiwaniana E EndemicF 

317 Orchidaceae Pleione bulbocodioides FacuE 2 

318 Orchidaceae Pomatocalpa acuminata E EndemicF 

319 Orchidaceae Schoenorchis vanoverberghii E 18–104 

320 Orchidaceae Staurochilus luchuensis E 2–20 

321 Orchidaceae Sunipia andersonii E 2–27, 16, 17 

322 Orchidaceae Taeniophyllum complanatum E EndemicF 

323 Orchidaceae Taeniophyllum glandulosum E 2, 17, 18, 29 

324 Orchidaceae Thelasis pygmaea E 2–27, 16, 17, 18 

325 Orchidaceae Thrixspermum annamense E 17 

326 Orchidaceae Thrixspermum eximium E 18–104 

327 Orchidaceae Thrixspermum fantasticum E 2–20, 18–104 

328 Orchidaceae Thrixspermum formosanum E 17 

329 Orchidaceae Thrixspermum laurisilvaticum E 2 

330 Orchidaceae Thrixspermum merguense E 17, 18 

331 Orchidaceae Thrixspermum pensile E 17, 18 

332 Orchidaceae Thrixspermum saruwatarii E EndemicF 

333 Orchidaceae Thrixspermum subulatum E 17, 18 

334 Orchidaceae Trichoglottis rosea E 18–104 

335 Orchidaceae Tuberolabium kotoense E EndemicL 

336 Orchidaceae Vanda lamellata E 2–20, 18–104 

Abbreviations: E: epiphyte, FacuE: facultative epiphyte, HemiE-P: primary hemi-epiphytes, HemiE-S: secondary hemi-epiphyte, EndemicF: endemic 

species in Taiwan, EndemicL: endemic species in Lanyu, EndemicG: endemic species in Lutao, floristic codes refer to Fig. 2. 



References 

Atwood, J.T., 1986. The size of the Orchidaceae and the 

systematic distribution of epiphytic orchids. Selbyana 9, 

171–186. 

Barkman, J.J., 1958. Phytosociology and Ecology of 

Cryptogamic Epiphytes. Van Gorcum & Comp. N. V., 

Assen. 

Benzing, D.H., 1987. Vascular epiphytism: taxonomic participation 

and adaptive diversity. Ann. Mo. Bot. Gard. 74, 

183–204. 

Benzing, D.H., 1990. Vascular Epiphytes. Cambridge University 

Press, Cambridge. 

Boufford, D.E., Hsieh, C.-F., Huang, T.-C., Kuoh, C.-S., 

Ohashi, H., Peng, C.-I., Tsai, J.-L., Yang, K.-C., 2003. 

Flora of Taiwan, second ed. Department of Botany, NTU, 

Taipei. 

Cascante-Marin, A., Wolf, J.H.D., Oostermeijer, J.G.B., Den 

Nijs, J.C.M., Sanahuja, O., Duran-Apuy, A., 2006. 

Epiphytic bromeliad communities in secondary and mature 

forest in a tropical premontane area. Basic Appl. Ecol. 7, 

520–532. 

Croat, T., 1978. Flora of Barro Colorado Island. Stanford 

University Press, Stanford. 

Dickerson, R.E., 1928. Distribution of Life in the Philippines. 

Bureau of Science Manila, Manila. 

Ek, R.C., 1997. Botanical diversity in the tropical rain forest 

of Guyana. Ph.D. Dissertation, Universiteit Utrecht, 

Utrecht. 

Freiberg, M., Freiberg, E., 2000. Epiphyte diversity and 

biomass in the canopy of lowland and montane forests in 

Ecuador. J. Trop. Ecol. 16, 673–688. 

Gentry, A.H., 1982. Neotropical floristic diversity: phytogeographical 

connections between Central and South America, 

Pleistocene climatic fluctuations, or an accident of the 

Andean orogeny? Ann. Mo. Bot. Gard. 69, 557–593. 

Gentry, A.H., Dodson, C.H., 1987a. Diversity and biogeography 

of neotropical vascular epiphytes. Ann. Mo. Bot. 

Gard. 74, 205–233. 

Gentry, A.H., Dodson, C.H., 1987b. Contribution of nontrees 

to species richness of a tropical rain forest. Biotropica 19, 

149–156. 

Gravendeel, B., Smithson, A., Slik, F.J.W., Schuiteman, A., 

2004. Epiphytism and pollinator specialization: drivers for 

orchid diversity? Philos. Trans. R. Soc. London B 359, 

1523–1535. 

Gupta, R., 1968. Flora Nainitalensis: A Handbook of the 

Flowering Plants of Nainital. Navayug Traders, New 

Delhi. 

Haber, W.A., 2001. Number of species with different plant 

growth forms. In: Nadkarni, N.M., Wheelwright, N.T. 

(Eds.), Monteverde: Ecology and Conservation of a 

Tropical Cloud Forest. Oxford University Press, New 

York, Oxford, pp. 519–522. 

Ho, C.-S., 1988. An Introduction to the Geology of Taiwan, 

Explanatory Text of the Geologic Map of Taiwan, second 

ed. Central Geological Survey, Ministry of Economic 

Affairs, Taipei. 

Hosokawa, T., 1950. Epiphyte-quotient. Bot. Mag. Tokyo 63, 

739–740. 



Hosokawa, T., 1958. On the synchorological and floristic 

trends and discontinuities in regard to the Japan-Liukiu- 

Formosa area. Plant Ecol. 8, 65–92. 

Hsieh, C.-F., 2003. Composition, endemism and phytogeographical 

affinities of the Taiwan flora. In: Boufford, D.E., 

Hsieh, C.-F., Huang, T.-C., Kuoh, C.-S., Ohashi, H., Peng, 

C.-I., Tsai, J.-L., Yang, K.-C. (Eds.), Flora of 

Taiwan, second ed. Department of Botany, NTU, Taipei, 

pp. 1–14. 

Hsueh, M.-L., Lee, H.-H., 2000. Diversity and distribution of 

the mangrove forests in Taiwan. Wetl. Ecol. Manage. 8, 

233–242. 

Ingram, S.W., Ferrell-Ingram, K., Nadkarni, N.M., 1996. 

Floristic composition of vascular epiphytes in a neotropical 

cloud forest, Monteverde, Costa Rica. Selbyana 17, 88–103. 

Janzen, D.H., Liesner, R., 1980. Annotated check-list of plants 

of lowland Guanacaste Province, Costa Rica, exclusive of 

grasses and non-vascular cryptogams. Brenesia 18, 15–90. 

Johansson, D., 1974. Ecology of vascular epiphytes in west 

African rain forest. Acta Phytogeogr. Suec. 59, 1–129. 

Kanehira, R., 1935. The phytogeographical relationship 

between Botel Tabago (Kotosyo) and the Philippines on 

the basis of the ligneous flora. Bull. Biogeogr. Soc. Japan 5, 

209–212. 

Kanto, T., 1933. Zoogeographical studies of Botel Tobago, 

with a consideration of the northern part of Wallace’s line. 

Geogr. Rev. Japan 9, 5–8 (In Japanese). 

Kelly, D.L., O’Donovan, G., Feehan, J., Murphy, S., 

Drangeid, S.O., Marcano-Berti, L., 2004. The epiphyte 

communities of a montane rain forest in the Andes of 

Venezuela: patterns in the distribution of the flora. J. Trop. 

Ecol. 20, 643–666. 

Kessler, M., Parris, B.S., Kessler, E., 2001. A comparison of 

the tropical montane Pteridophyte floras of Mount 

Kinabalu, Borneo, and Parque Nacional Carrasco, Bolivia. 

J. Biogeogr. 28, 611–622. 

Köhler, L., Tobón, C., Frumau, K., Bruijnzeel, L., 2007. 

Biomass and water storage dynamics of epiphytes in oldgrowth 

and secondary montane cloud forest stands in 

Costa Rica. Plant Ecol. 193, 171–184. 

Kolbeck, J., 1995. Notes on epiphytic communities in forests 

of North Korea. Preslia, Praha 67, 41–45. 

Kreft, H., Koster, N., Kuper, W., Nieder, J., Barthlott, W., 

2004. Diversity and biogeography of vascular epiphytes in 

Western Amazonia, Yasuní, Ecuador. J. Biogeogr. 31, 

1463–1476. 

Kreft, H., Jetz, W., Mutke, J., Kier, G., Barthlott, W., 2008. 

Global diversity of island floras from a macroecological 

perspective. Ecol. Lett. 11, 116–127. 

Kress, W.J., 1986. The systematic distribution of vascular 

epiphyte: an update. Selbyana 9, 2–22. 

Kuo, C.-M., 1985. Taxonomy and phytogeography of 

Taiwanese Pteridophytes. Taiwania 30, 5–100. 

Küper, W., Kreft, H., Nieder, J., Koster, N., Barthlott, W., 

2004. Large-scale diversity patterns of vascular epiphytes 

in neotropical montane rain forests. J. Biogeogr. 31, 

1477–1487. 

Lester, S.E., Ruttenberg, B.I., Gaines, S.D., Kinlan, B.P., 

2007. The relationship between dispersal ability and 

geographic range size. Ecol. Lett. 10, 745–758. 



16 

Lin, T.-C., Hamburg, S.P., Hsia, Y.-J., Lin, T.-T., King, H.-B., 

Wang, L.-J., Lin, K.-C., 2003. Influence of typhoon 

disturbances on the understory light regime and stand 

dynamics of a subtropical rain forest in northeastern 

Taiwan. J. For. Res. 8, 139–145. 

Madison, M., 1977. Vascular epiphytes: their systematic 

occurrence and salient features. Selbyana 2, 1–13. 

Migenis, L.E., Ackerman, J.D., 1993. Orchid phorophyte 

relationships in a forest watershed in Puerto-Rico. J. Trop. 

Ecol. 9, 231–240. 

Moffett, M.W., 2000. What’s ‘‘Up’’? A critical look at the 

basic terms of canopy biology. Biotropica 32, 569–596. 

Nieder, J., Prosperí, J., Michaloud, G., 2001. Epiphytes and their 

contribution to canopy diversity. Plant Ecol. 153, 51–63. 

Oliver, W.R.B., 1930. New Zealand epiphytes. J. Ecol. 18, 

1–50. 

Rauer, G., Rudolph, D., 2001. Vaskuläre Epiphyten eines 

Westandinen Bergregenwaldes in Ecuador. In: Nieder, J., 

Barthlott, W. (Eds.), The Flora of the Río Guajalito 

Mountain Rain Forest (Ecuador). Results of the Bonn- 

Quito Epiphyte Project, funded by the Volkswagen 

Foundation. GmbH, Bonn, pp. 323–470. 

Ray, N., Adams, J.M., 2001. A GIS-based vegetation map of 

the world at the last glacial maximum (25,000–15,000 bp). 

Internet Archaeol. 11 /http://intarch.ac.uk/journal/issue11/ 

2/toc.htmlS. 

Richards, P.W., 1952. The Tropical Rain Forest. Cambridge 

University Press, Cambridge. 

Riveros, M., Ramírez, C., 1978. Fitocenoses epífitas de la 

asociación Lapagerio-Aextoxiconetum en el fundo San 

Martín (Valdivia-Chile). Acta Cient. Venez. 29, 163–169. 



Schaijes, M., Malaisse, F., 2001. Diversity of Upper Katanga 

epiphytes (mainly orchids) and distribution in different 

vegetation units. Syst. Geogr. Plants 71, 575–584. 

Su, H.-J., 1984. Studies on the climate and vegetation types of 

natural forests in Taiwan (iii). A scheme of geographical 

climatic regions. Quart. J. Chin. For. 18, 33–44. 

Su, H.-J., 1992. Vegetation of Taiwan: altitudinal vegetation 

zones and geographical climatic regions. Bot. Ins. Acad. 

Sin. Monogr. Ser. 11, 39–53 (in Chinese). 

Takhtajan, A., 1986. Floristic Regions of the World. 

University of California Press, London. 

Wolf, J.H.D., 2005. The response of epiphytes to anthropogenic 

disturbance of pine-oak forests in the highlands of 

Chiapas, Mexico. For. Ecol. Manage. 212, 376–393. 

Wolf, J.H.D., Flamenco-S, A., 2003. Patterns in species 

richness and distribution of vascular epiphytes in Chiapas, 

Mexico. J. Biogeogr. 30, 1689–1707. 

Xu, H., Liu, W., 2005. Species diversity and distribution of 

epiphytes in the montane moist evergreen broad-leaved 

forest in Ailao mountain, Yunnan. Biodiversity Sci. 13, 

137–147 (in Chinese). 

Yen, S.-H., Kitching, I.J., Tzen, C.-S., 2003. A new subspecies 

of hawkmoth from Lanyu island, Taiwan, with a revised 

and annotated checklist of Taiwanese Sphingidae 

(Lepidoptera). Zool. Stud. 42, 292–306. 

Zapfack, L., Nkongmeneck, A., Villiers, J., Lowman, M., 

1996. The importance of pteridophytes in the epiphytic 

flora of some phorophytes of the Cameroonian semideciduous 

rain forest. Selbyana 17, 76–81. 

Zotz, G., 2005. Vascular epiphytes in the temperate zones – a 

review. Plant Ecol. 176, 173–183.

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