13
PRIMARY RESEARCH PAPER | Philippine Journal of Systematic Biology DOI 10.26757/pjsb2020c14004 Volume 14 Issue 3 - 2020 | 1 © Association of Systematic Biologists of the Philippines Rapid assessment of epiphytic pteridophyte biodiversity in Mt. Apo Natural Park, North Cotabato Province, Philippines: a comparison of disturbed and undisturbed forests Abstract Pteridophytes are potential indicators of climate change and environmental disturbances because of their sensitivity to the changes in the environment such as sunlight intensities and humidity levels. The study was conducted to document the species and compare the diversity of epiphytic pteridophytes in two sites–the undisturbed and disturbed areas of Mt. Apo Natural Park, North Cotabato Province in the Philippines. In each site, four stations were established, each with four 20 m × 20 m plots that are at least 20 m apart from each other. Between stations, at least 100 m was maintained. In this study, 102 species of epiphytic pteridophytes belonging to 33 genera and 14 families were identified. Seven species are lycophytes and 95 species are ferns. The number of species recorded represents 10 % of all species of pteridophytes in the Philippines. Polypodiaceae was the most dominant family consisting of 13 genera and 33 species. Among these, Prosaptia was the largest genus represented by nine species followed by Selliguea with five species. Based on the Species Importance Value (SIV), Lindsaea pulchella was the most abundant species in the area studied. Abundance of epiphytic pteridophytes vary between two sites but other parameters such as species richness, evenness, Shannon-Weiner and Simpson diversity indices showed no significant differences. There were three main assemblages of epiphytic pteridophytes based on species composition with 50% overall similarity. Of the 102 species identified, 11 were threatened which represent 6.08% of the total threatened pteridophytes of the Philippines. Noteworthy is the new species record of Asplenium beccarianum for the Philippines. Unique composition of epiphytic pteridophytes was evident and the results showed that Mt. Apo is an important location for the conservation of these communities. Keywords: abundance, Asplenium beccarianum, conservation, phorophtye, epiphyte 1 Department of Biological Sciences, College of Science and Mathematics, University of Southern Mindanao, Kabacan, Cotabato, Philippines 2 Biology Department, College of Arts and Sciences, Central Mindanao University, Maramag, Bukidnon, Philippines *Corresponding email: [email protected] Date Submitted: 29 January 2019 Date Accepted: 11 September 2020 Introduction The Philippines is one of the countries in the tropics which is known to host a variety of flora and fauna with high endemicity, thus, it is recognized as a megadiverse country (Ong et al. 2002). Anthropogenic activities such as logging, land conversion to agricultural uses pose a risk to both floral and faunal components of the remaining forest areas. Despite various researches on the floral and faunal species in the Philippines (Lumbres et al. 2014, Castañares et al. 2017, Replan & Malaki 2017, Vidal et al. 2018), threats due to anthropogenic activities to Philippine biodiversity are still increasing (Amoroso et al. 2019, Zapanta et al. 2019), thereby putting the country as one of the biodiversity hotspots in the world. World forests are endowed with diverse communities of flora and fauna with complex interactions. Ecologically, the plant-plant interactions regulate the structure and composition of plant communities in a forest ecosystem (Tewari et al. 2009, Chomba et al. 2011, Adhikari et al. 2012, Cach-Perez et al. 2013, Zotz et al. 2014). Epiphytic pteridophytes have gained attention as a diverse group with approximately 2,600 species all over the world (Kress 1986), comprising about 29 % of all pteridophyte species. Various factors were identified to influence and shape the occurrence, distribution and diversity of epiphytic pteridophytes including phorophyte microenvironment (Kromer et al. 2007), structure of the forest surrounding the epiphyte, spore dispersal and establishment (Hirata et al. 2009), and presence of phorophyte (Zotz et al. 2014). In addition, Cherie Cano-Mangaoang 1* , Bryan Lloyd P. Bretaña 1 , and Victor B. Amoroso 2

Rapid assessment of epiphytic pteridophyte biodiversity in ...asbp.org.ph/wp-content/uploads/2020/10/PJSB_2020-004.pdf · epiphytes face exceptional challenges because of their specific

  • Upload
    others

  • View
    2

  • Download
    0

Embed Size (px)

Citation preview

Page 1: Rapid assessment of epiphytic pteridophyte biodiversity in ...asbp.org.ph/wp-content/uploads/2020/10/PJSB_2020-004.pdf · epiphytes face exceptional challenges because of their specific

PRIMARY RESEARCH PAPER | Philippine Journal of Systematic Biology

DOI 10.26757/pjsb2020c14004

Volume 14 Issue 3 - 2020 | 1 © Association of Systematic Biologists of the Philippines

Rapid assessment of epiphytic pteridophyte biodiversity in

Mt. Apo Natural Park, North Cotabato Province, Philippines:

a comparison of disturbed and undisturbed forests

Abstract

Pteridophytes are potential indicators of climate change and environmental disturbances because of their sensitivity to the

changes in the environment such as sunlight intensities and humidity levels. The study was conducted to document the

species and compare the diversity of epiphytic pteridophytes in two sites–the undisturbed and disturbed areas of Mt. Apo

Natural Park, North Cotabato Province in the Philippines. In each site, four stations were established, each with four 20 m

× 20 m plots that are at least 20 m apart from each other. Between stations, at least 100 m was maintained. In this study,

102 species of epiphytic pteridophytes belonging to 33 genera and 14 families were identified. Seven species are

lycophytes and 95 species are ferns. The number of species recorded represents 10 % of all species of pteridophytes in

the Philippines. Polypodiaceae was the most dominant family consisting of 13 genera and 33 species. Among these,

Prosaptia was the largest genus represented by nine species followed by Selliguea with five species. Based on the Species

Importance Value (SIV), Lindsaea pulchella was the most abundant species in the area studied. Abundance of epiphytic

pteridophytes vary between two sites but other parameters such as species richness, evenness, Shannon-Weiner and

Simpson diversity indices showed no significant differences. There were three main assemblages of epiphytic

pteridophytes based on species composition with 50% overall similarity. Of the 102 species identified, 11 were threatened

which represent 6.08% of the total threatened pteridophytes of the Philippines. Noteworthy is the new species record of

Asplenium beccarianum for the Philippines. Unique composition of epiphytic pteridophytes was evident and the results

showed that Mt. Apo is an important location for the conservation of these communities.

Keywords: abundance, Asplenium beccarianum, conservation, phorophtye, epiphyte

1Department of Biological Sciences, College of Science and

Mathematics, University of Southern Mindanao, Kabacan, Cotabato,

Philippines 2Biology Department, College of Arts and Sciences, Central Mindanao

University, Maramag, Bukidnon, Philippines

*Corresponding email: [email protected]

Date Submitted: 29 January 2019

Date Accepted: 11 September 2020

Introduction

The Philippines is one of the countries in the tropics which

is known to host a variety of flora and fauna with high

endemicity, thus, it is recognized as a megadiverse country

(Ong et al. 2002). Anthropogenic activities such as logging,

land conversion to agricultural uses pose a risk to both floral

and faunal components of the remaining forest areas. Despite

various researches on the floral and faunal species in the

Philippines (Lumbres et al. 2014, Castañares et al. 2017, Replan

& Malaki 2017, Vidal et al. 2018), threats due to anthropogenic

activities to Philippine biodiversity are still increasing (Amoroso

et al. 2019, Zapanta et al. 2019), thereby putting the country as

one of the biodiversity hotspots in the world.

World forests are endowed with diverse communities of

flora and fauna with complex interactions. Ecologically, the

plant-plant interactions regulate the structure and composition of

plant communities in a forest ecosystem (Tewari et al. 2009,

Chomba et al. 2011, Adhikari et al. 2012, Cach-Perez et al.

2013, Zotz et al. 2014). Epiphytic pteridophytes have gained

attention as a diverse group with approximately 2,600 species all

over the world (Kress 1986), comprising about 29 % of all

pteridophyte species. Various factors were identified to

influence and shape the occurrence, distribution and diversity of

epiphytic pteridophytes including phorophyte microenvironment

(Kromer et al. 2007), structure of the forest surrounding the

epiphyte, spore dispersal and establishment (Hirata et al. 2009),

and presence of phorophyte (Zotz et al. 2014). In addition,

Cherie Cano-Mangaoang1*, Bryan Lloyd P. Bretaña1, and Victor B. Amoroso2

Page 2: Rapid assessment of epiphytic pteridophyte biodiversity in ...asbp.org.ph/wp-content/uploads/2020/10/PJSB_2020-004.pdf · epiphytes face exceptional challenges because of their specific

Volume 14 Issue 3 - 2020 | 2 Philippine Journal of Systematic Biology Online ISSN: 2508-0342

Cano-Mangaoang et al.: Epiphytic pteridophyte biodiversity in Mt. Apo Natural Park

epiphytes face exceptional challenges because of their specific

adaptations on non-soil substrate as well as sensitivity to

ecological disturbances and microclimatic changes (Martin et

al. 2007, Chomba et al. 2011, Gowland et al. 2011, Quaresma

& Jardim 2014, Zhao et al. 2015). In view of this sensitivity to

disturbances and microclimates, and importance in tropical

forest ecology, epiphytes serve as good indicators or guides for

careful and effective management of forest ecosystems (Heitz

1999). On the other hand, Schmitt & Windisch (2009) pointed

out the importance of host plants with respect to the existence of

the epiphytic pteridophytes given that majority of them are

habitual and that some occur solely on preferred tree ferns.

Epiphytic pteridophytes also plays an important role in

catching rainfall (Chomba et al. 2011) and in holding

atmospheric nutrients. According to several studies (Arévalo &

Betancur 2006, Aguirre et al. 2010, Hirata et al. 2009, Zhao et

al. 2015) epiphytic plants are important components of tropical

forests, not just for the biomass they accumulate but also in

providing nesting sites for many species of arboreal fauna

(Andama et al. 2003, Zhang et al. 2010). Epiphytic ferns such

as Asplenium nidus L. trap nutrients by collecting falling

detritus on its leaf bases and this ability to intercept detritus is

nutritionally significant (Reich et al 2003, Karasawa & Hijii

2006). Haro-Carrion et al. (2009) confirmed that epiphyte

diversity, composition, and vertical distributions are useful

indicators of human disturbance in agroforest ecosystem.

Understanding their distribution and diversity is not only of use

in conserving the forest ecosystem but also in the conservation

and protection of biodiversity. Hence, this study aims to

compare the diversity of epiphytic pteridophytes in two

different forest habitats - the disturbed and undisturbed forests

of Mt. Apo Natural Park.

Materials and Methods

Description of sampling sites

The study was conducted in Mt. Apo Natural Park, the

Philippines’ highest peak (2,954 masl) and declared as protected

area under the Republic Act 9237 as a Natural Park. It is one of

the most important and protected areas in the Philippines

because of its diverse biological resources. Despite being

known as a biodiversity-rich area, Mt. Apo is continually

exposed to forest degradation from illegal logging, land

conversion, upland settlers and wildlife hunting/poaching. Two

sampling sites were established, one within the Energy

Development Corporation- Long Term Ecological Research

(EDC-LTER) area in Brgy Ilomavis, Kidapawan City and the

other one is the Makalangit Site in Brgy New Israel, Makilala.

These two sites are situated at 06° 59.728’N; 125° 15.196’E with

an elevation of 1827 masl and 06°56.742’N; 125°13.839’E with

an elevation of 1933 masl, respectively (Fig 1).

With the establishment of EDC since 1996, and its

intervention in protecting the remaining forest of Mt. Apo

within its jurisdiction, extraction of natural resources within the

protected area such as poaching, hunting and other

anthropogenic disturbances have been minimized. It was also

observed during field sampling the abundance of young trees

outside the sampling plots in the EDC-LTER site which might

be an indication that the forest is undergoing early regeneration.

The trail usually used by climbers and hikers to the Mt. Apo

peak has been permanently closed since 1996 thus minimizing

entry of people in this area. With this, the EDC-LTER site is

characterized in this study as an undisturbed site. Meanwhile,

the Makalangit site is found along the Maag trail. In here,

frequent ground clearing is being done for planting of

agricultural commodities by the members of certain religious

group. Cutting of trees was also evident, with trees used in the

construction of temporary shelter whenever they are in the area.

In addition, the trail is also used as one of the entry and exit

points of visitors while poaching and hunting are still evident on

the site. Thus this area is tagged in this study as the disturbed

site.

The type of vegetation present in each sampling site is

characterized. The tagged undisturbed site has more numerous

and larger trees compared with the disturbed site. The Diameter

at Breast Height (DBH) of trees in the undisturbed site ranges

from 12 -150 cm while the DBH in disturbed site ranges from

10 -101 cm. Abundance of smaller trees and tree ferns was

observed in the disturbed site. The undisturbed site on the other

hand has an intact forest with 100% canopy cover for almost all

Figure 1. Sampling sites in Mt. Apo Natural Park, North Cotabato

Province, Philippines.

Page 3: Rapid assessment of epiphytic pteridophyte biodiversity in ...asbp.org.ph/wp-content/uploads/2020/10/PJSB_2020-004.pdf · epiphytes face exceptional challenges because of their specific

Volume 14 Issue 3 - 2020 | 3 © Association of Systematic Biologists of the Philippines

Cano-Mangaoang et al.: Epiphytic pteridophyte biodiversity in Mt. Apo Natural Park

established plots compared to the disturbed site with less

canopy cover. There were plots in the disturbed site that are

situated in abandoned clearing areas.

The study was conducted from March to September, 2016

with an average temperature of 16°C and 21°C in disturbed and

undisturbed site, respectively. The average humidity was 97%

and 85% in disturbed and undisturbed site, respectively. The

average rainfall for the both sampling sites was 7.57 mm during

sampling period.

Sampling strategy

A modification of the protocol by Karger et al. (2014) was

employed in the establishment of the sampling plots within each

site. In each site, four stations were established, each with four

20 m x 20 m plots which are at least 20 m from each other. At

least 100 m between any two stations were maintained in the

survey. Overall, there were 16 plots in each site with a total of

6,400 m2 area covered.

Within each 20 m x 20 m plot, all phorophytes (trees and

tree ferns with epiphytic pteridophytes attached on them

regardless of number) were sampled. Transect walks between

the sampling plots were also done to supplement the data on

species richness. The species of epiphytic pteridophytes in the

sampling sites (both in the sampling plots and outside the

sampling plots) were surveyed by a combination of methods

such as climbing of trees using a rope, cutting selected branches

and using binoculars (Karger et al. 2014).

Five individuals of each species were collected as

herbarium specimens and deposited at Central Mindanao

University Museum (CMUH) and University of Southern

Mindanao (USM). Based on preliminary identification on site,

if any particular species was under endangered or critically

endangered status, no collection was made and only

photographs were taken. All specimens were processed

following the method employed by Amoroso et al. (2016).

Identification of collected specimens was done at the

Botany Division, CMUH following monographs, floras such as

Copeland (1958 – 1961), and other publications such as

Hoovenkamp et al. (1998); Pelser et al. (2011 onwards) and

digitized images of specimens available in Global Plants on

JSTOR. Annotated specimen of epiphytic pteridophytes

deposited at the CMUH were used as reference in the

identification of collected specimens. PPG I classification

scheme was followed in this study.

Data evaluation

PRIMER software was used to calculate the diversity

indices such as the Shannon-Weiner index and Simpson index.

For multivariate analysis of the assemblages, Analysis of

Similarity (ANOSIM) was applied to analyze the difference in

pteridophyte assemblage based on Bray-Curtis similarities.

ANOSIM has been widely used to test the hypotheses on the

impacts of environmental factors on spatial differences and

temporal changes in assemblages (Chapman and Underwood

1999). In addition, Similarity Percentage (SIMPER) was used to

assess the average percent contribution of the species to the

dissimilarity between sites in a Bray-Curtis dissimilarity matrix.

This identified the species that are likely to be major

contributors (90% cumulative percentage) to any difference

between sites (Clarke, 1993). Result of assemblage analysis was

plotted using the nonmetric multidimensional scaling (nMDS).

Species Importance Value was computed using the formula: SIV

= RDen + RFreq + RDom, where RDen = Relative Density,

RFreq = Relative Frequency, Rdom = Relative Dominance

(Amoroso et al. 2019). Conservation and distribution statuses of

the species identified were based on Fernando et al. (2008),

Amoroso et al. (2015) and DENR Administrative Order 2017-11

(DAO 2017-11). Ecological notes were supplemented from

Copeland (1958-1961). Economically important species were

noted based on Amoroso et al. (2015).

Results and Discussion

Species Richness

This study identified 102 species representing 33 genera

and 14 families of epiphytic pteridophytes from 32 sampling

plots and 24 transect walks in Mt. Apo Natural Park (Table 1).

Of these, seven species were lycophytes and 95 species of ferns.

A total of 82 and 78 species were identified from the

undisturbed and disturbed sites, respectively. Most of the

species identified (33 species representing 13 genera) were

under the family Polypodiaceae. These species of pteridophytes

were found attached on 556 individual phorophytes in the

undisturbed site while 918 were in the disturbed area.

The results showed that the species richness of epiphytic

pteridophytes in Mt. Apo represents 10% of all species of

pteridophytes in the Philippines. Moreover, it comprises 23 % of

all species of pteridophytes identified at Mt. Kitanglad Mountain

Range and 36 % of the total species in Mt. Malindang, both

ecosystems found in Mindanao (Amoroso et al. 2011). Results

of the study showed that Mt. Apo is also an important area for

epiphytic pteridophytes. Further, this indicated that research

attention on epiphytic pteridophytes be given including their

various ecological functions which might benefit the ecosystem

as well as the humans (Heitz 1999, Arévalo & Betancur 2006,

Aguirre et al. 2010, Haro-Carrion et al. 2009, Hirata et al. 2009,

Zhao et al. 2015). Kress (1986) mentioned that pteridophytes

are the second group of vascular plants in terms of epiphyte

Page 4: Rapid assessment of epiphytic pteridophyte biodiversity in ...asbp.org.ph/wp-content/uploads/2020/10/PJSB_2020-004.pdf · epiphytes face exceptional challenges because of their specific

Volume 14 Issue 3 - 2020 | 4 Philippine Journal of Systematic Biology Online ISSN: 2508-0342

Cano-Mangaoang et al.: Epiphytic pteridophyte biodiversity in Mt. Apo Natural Park

Species of Epiphytic Pteridophytes Undisturbed Site Disturbed Site

A Lycophytes

Lycopodiaceae

1 Phlegmariurus nummularifolius (Blume) Ching /

2 P. salvinioides (Herter) Ching / /

3 P. banayanicus (Herter) A.R. Field & Bostock /

4 P. verticillatus (L.f.) A.R. Field & Testo /

Selaginellaceae

5 Selaginella aristata Spring /

6 S. eschscholzii Hieron. /

7 S. remotifolia Spring /

B Ferns

Aspleniaceae

8 Asplenium affine Sw. /

9 A. apoense Copel. / /

10 A. beccarianum Ces. /

11 A. callipteris (Fée) E. Fourn. /

12 A. colubrinum Christ / /

13 A. crinicaule Hance / /

14 A. lobulatum Mett. /

15 A. longissimum Blume /

16 A. nigrescens Blume / /

17 A. nidus L. / /

18 A. normale D. Don / /

19 A. thunbergii Kunze / /

20 A. tenerum G. Forst. / /

21 A. vittaeforme Cav. /

22 Asplenium sp. 1 / /

23 Asplenium sp. 2 / /

24 Asplenium sp. 3 / /

25 Asplenium sp. 4 / /

Davalliaceae

26 Davallia embolostegia Copel. /

27 D. hymenophylloides (Blume) Kuhn /

28 D. repens (L.f.) Kuhn / /

29 D. trichomanoides Blume / /

30 D. wagneriana Copel. /

Table 1 cont. List of epiphytic pteridophytes identified from the two sampling sites in Mt. Apo Natural Park.

Page 5: Rapid assessment of epiphytic pteridophyte biodiversity in ...asbp.org.ph/wp-content/uploads/2020/10/PJSB_2020-004.pdf · epiphytes face exceptional challenges because of their specific

Volume 14 Issue 3 - 2020 | 5 © Association of Systematic Biologists of the Philippines

Cano-Mangaoang et al.: Epiphytic pteridophyte biodiversity in Mt. Apo Natural Park

Species of Epiphytic Pteridophytes Undisturbed Site Disturbed Site

Dryopteridaceae

31 Dryopteris hirtipes (Blume) Kuntze /

32 Elaphoglossum blumeanum (Fèe) J. Sm. / /

33 E. callifolium (Blume) T. Moore / /

34 Polysticum elmeri Copel. / /

Hymenophyllaceae

35 Abrodictyum cumingii C. Presl /

36 A. apluma (Hook.) Ebihara & K. Iwats. /

37 Callistopteris apiifolia (C. Presl) Copel. / /

38 Crepidomanes brevipes (C. Presl) Copel. /

39 C. grande Ebihara & K. Iwats. /

40 C. minutum (Blume) K. Iwats. /

41 Hymenophyllum acanthoides (Bosch) Rosenst. /

42 H. badium Hook. & Grev. / /

43 H. emarginatum Sw. / /

44 H. holochilum (Bosch) C. Chr. / /

45 H. imbricatum Blume / /

46 H. pallidum Ebihara & K. Iwats. / /

47 H. paniculiflorum C. Presl /

48 H. productum Kunze / /

49 H. ramosii Copel. /

50 H. demissum (G. Forst.) Sw. /

51 Hymenopnyllum sp. 1 / /

52 Hymenophyllum sp. 2 / /

Hypodematiaceae

53 Leucostegia truncata (D.Don) Fraser / /

Lindsaeaceae

54 Lindsaea apoensis Copel. / /

55 L. pulchella (J. Sm.) Mett. ex Kuhn. / /

56 L. repens (Bory) Thwaites /

Oleandraceae

57 Oleandra neriformis Cav. / /

58 O. sebaldii Grev. / /

Ophioglossaceae

59 Ophioglossum pendulum L. / /

Table 1 cont. List of epiphytic pteridophytes identified from the two sampling sites in Mt. Apo Natural Park.

Page 6: Rapid assessment of epiphytic pteridophyte biodiversity in ...asbp.org.ph/wp-content/uploads/2020/10/PJSB_2020-004.pdf · epiphytes face exceptional challenges because of their specific

Volume 14 Issue 3 - 2020 | 6 Philippine Journal of Systematic Biology Online ISSN: 2508-0342

Cano-Mangaoang et al.: Epiphytic pteridophyte biodiversity in Mt. Apo Natural Park

Species of Epiphytic Pteridophytes Undisturbed Site Disturbed Site

Plagiogyraceae

60 Plagiogyria euphlebia (Kunze) Mett. / /

61 P. glauca (Blume) Mett. / /

Polypodiaceae

62 Aglaomorpha cornucopia (Copel.) M.C.Roos / /

63 Aglaomorpha sagitta (Christ) Hovenkamp & S. Linds. /

64 Calymmodon muscoides (Copel.) Copel.

65 C. gracilis (Fée) Copel. / /

66 Goniophlebium persicifolium (Desv.) Bedd. / /

67 G. pseudoconnatum (Copel.) Copel. / /

68 G. subauriculatum (Blume) C. Presl / /

69 Lecanopteris deparioides (Cesati) Baker /

70 Lepisorus accedens (Blume) Hosok. /

71 L. mucronatus (Fée) Li Wang / /

72 L. platyrhynchos (Kunze) Li Wang /

73 L. longifolius (Blume) Holttum /

74 L. spicatus (L.f.) Li Wang /

75 L. validinervis (Kunze) Li Wang / /

76 Loxogramme avenia (Blume) C. Presl / /

77 Oreogrammitis adspersa (Blume) Parris /

78 O. beddomeana (Alderw.) T.C. Hsu /

79 O. dolichosora (Copel.) Parris /

80 O. fasciata (Blume) Parris /

81 O. setigera (Blume) T.C. Hsu /

82 Prosaptia celebica (Blume) Tagawa & K. Iwats. / /

83 P. contigua (G. Forst.) C. Presl / /

84 P. cryptocarpa Copel. /

85 P. davalliacea (F Muell. & Baker) Copel. / /

86 P. multicaudata (Copel.) Mett. / /

87 P. nutans (Blume) Mett. /

88 P. obliquata (Blume) Mett. / /

89 P. venulosa (Blume) M.G. Price /

90 Selliguea albidosquamata (Blume) Parris / /

91 S. elmeri (Copel.) Ching /

92 S. taeniata (Sw.) Parris / /

Table 1 cont. List of epiphytic pteridophytes identified from the two sampling sites in Mt. Apo Natural Park.

Page 7: Rapid assessment of epiphytic pteridophyte biodiversity in ...asbp.org.ph/wp-content/uploads/2020/10/PJSB_2020-004.pdf · epiphytes face exceptional challenges because of their specific

Volume 14 Issue 3 - 2020 | 7 © Association of Systematic Biologists of the Philippines

Cano-Mangaoang et al.: Epiphytic pteridophyte biodiversity in Mt. Apo Natural Park

Diversity Indices and Species Importance Value (SIV)

The species with the highest SIV in Mt. Apo Natural Park

includes Lindsaea pulchella Mettenius (71.01%) followed by

Tmesipteris zamorarum (60.55%) and Hymenophyllum

paniculiflorum Presl (11.68%) (Table 2).

Tree ferns are the most abundant phorophytes in both

disturbed and undisturbed sites studied and the two species,

Lindsaea pulchella and Tmesipteris zamorarum, have a high

affinity to the stems of tree ferns. This may be the main reason

for their higher species importance value than the others. With

this observation, it implies that they are host specific species

because of their dependence for growth on tree fern trunks as

their only habitat. It implies further that the removal of tree

ferns will result to loss of these two host-specific species. The

condition of the tree ferns will inevitably affect their existence

and survival. Colwell et al. (2012) opined supported that the

dependence of affiliates to these tree fern substrates might result

in co-extinction from host population removal.

In terms of diversity parameters, there is a significant

difference in the abundance (Fig. 3B) of epiphytic pteridophytes

in the two sites with more individuals in the disturbed site. Other

measures of diversity including species richness, evenness,

Shannon-Weiner index and Simpson index showed no

significant differences (Figs. 3A, 3C-E and 4). The higher

abundance of epiphytic pteridophytes in the disturbed area can

be attributed to clearing of the other vegetation, therefore

providing more sunlight for light tolerant species. Another factor

is due to the higher availability of host plants or phorophytes for

diversity with 29 % next to monocots but few studies has been

conducted on epiphytic pteridophytes.

The high species richness of epiphytic pteridophytes in

Mt. Apo has implications for improving the structure of the

forest ecosystem. Epiphytic pteridophytes collect masses of

humus, which provides nesting sites for many species of

arboreal ants and other invertebrates (Andama et al. 2003,

Arévalo & Betancur 2006, Zhang et al. 2010). Moreover, they

contribute to species diversity, primary productivity, biomass,

litter fall, water retention, and provide substrate for nitrogen

fixing bacteria. Establishment of epiphyte communities will

also enhance energy, moisture capture and retention (Cummings

& Rogers 2006).

In this study, a few species of epiphytic pteridophytes

were identified with unique ecological and conservation status.

Endangered species are: Phlegmariurus salvinoides (Herter)

Ching (Fig 2A), Tmesipteris zamorarum Gruézo & Amoroso

(Fig 2B), Ophioglossum pendulum L. (Fig 2C), Lecanopteris

deparioides (Cesati) Baker (Fig 2D). The study also recorded

Vaginularia paradoxa (Fée) Mettenius ex Miquel (Fig 2E) a

rare species. The presence of Asplenium beccarianum Cesati

(Fig. 2F) on Mt. Apo, which was reported to be thriving in

Papua New Guinea and Indonesia based on digitized plant

specimens available in Global Plants on JSTOR, is a new

distribution record in the Philippines. For further verification,

specimen of A. beccarianum Cesati collected by the first author

is deposited at the Department of Biological Sciences, College

of Science and Mathematics, University of Southern Mindanao

with accession number, USM-Bio-MANP-2016-0011.

Species of Epiphytic Pteridophytes Undisturbed Site Disturbed Site

93 S. triloba (Houtt.) M.G. Price / /

94 Tomophyllum macrum (Copel.) Parris / /

Psilotaceae

95 Tmesipteris zamorarum Gruezo & Amoroso / /

Pteridaceae

96 Antrophyum reticulatum (Forst.) Kaulf. / /

97 A. sessilifolium (Cav.) Spring / /

98 A. plantagineum (Cav.) Kaulf. /

99 Haplopteris alternans (Copel.) S. Linds. & C.W. Chen / /

100 H. ensiformis (Sw.) E.H. Crane / /

101 H. scolopendrina (Bory) C. Presl / /

102 Vaginularia junghuhnii Mett. / /

Table 1 cont. List of epiphytic pteridophytes identified from the two sampling sites in Mt. Apo Natural Park.

Page 8: Rapid assessment of epiphytic pteridophyte biodiversity in ...asbp.org.ph/wp-content/uploads/2020/10/PJSB_2020-004.pdf · epiphytes face exceptional challenges because of their specific

Volume 14 Issue 3 - 2020 | 8 Philippine Journal of Systematic Biology Online ISSN: 2508-0342

Cano-Mangaoang et al.: Epiphytic pteridophyte biodiversity in Mt. Apo Natural Park

attachment. Zotz et al. (2014) reiterated the importance of the

phorophytes for the existence of epiphytes as the lack of

suitable substrate (trees) rather than climatic reasons are

responsible for the absence of epiphytes immediately above the

treeline. Furthermore, Hirata et al. (2009) stated that

distribution of epiphytes is affected by two major processes:

dispersal and establishment.

The lower values of diversity indices of disturbed site

might be attributed to anthropogenic disturbances although no

significant differences were noted between two sites. The

clearing of ground cover and cutting of trees for agriculture

might be a factor for the lower diversity of epiphytic

pteridophytes seen here. These activities are evidences of how

forest fragmentation and habitat conversion affect not only the

diversity of epiphytic pteridophytes but also the flora and fauna

thriving in the same ecosystem (Jha et al. 2005, Stireman et al.

2014, Zambrano et al. 2019).

Although no significant difference in diversity indices was

observed in the study, over-all result of ANOSIM based on

species similarity showed that there was a clear separation of

the assemblages (R value = 0.653, p-value<0.05). SIMPER

analysis based on species composition and abundance revealed

a 50% overall similarity between the two sites in Mt. Apo

Natural Park. All stations in the disturbed site including two

plots in stations 1 and 2 of the undisturbed site formed the

assemblage A. Assemblage B includes stations 1 and 2 and

assemblage C contains stations 3 and 4 (Fig. 5). The presence

of a shared species, L. pulchella and T. zamorarum contributed

to the similarity of assemblages A and B. SIMPER analysis

further revealed that the dissimilarity of the assemblage was

mainly due to the top three discriminating species,

Hymenophyllum holochilum (Bosch) C. Christensen, H.

paniculiflorum and Lindsaea apoensis Copeland. These are the

species with high abundance in the disturbed site and few or

Figure 2. Noteworthy species of epiphytic pteridophytes in Mt. Apo Natural Park. A. Phlegmariurus salvinioides, B. Tmesipteris zamorarum,

C. Ophioglossum pendulum, D. Lecanopteris deparioides, E. Vaginularia paradoxa, F. Asplenium beccarianum.

Species of Epiphytic Pteridophytes SIV

Lindsaea pulchella 71.01

Tmesipteris zamorarum 60.55

Hymenophyllum paniculiflorum 11.68

Hymenophyllum holochilum 9.88

Selliguea taeniata 8.89

Davallia repens 8.12

Elaphoglossum blumeanum 7.59

Selliguea triloba 7.27

Polystichum elmeri 7.06

Hymenophyllum emarginatum 5.88

Table 2. Species Importance Value (SIV) of epiphytic pteridophytes

in Mt. Apo Natural Park.

Page 9: Rapid assessment of epiphytic pteridophyte biodiversity in ...asbp.org.ph/wp-content/uploads/2020/10/PJSB_2020-004.pdf · epiphytes face exceptional challenges because of their specific

Volume 14 Issue 3 - 2020 | 9 © Association of Systematic Biologists of the Philippines

Cano-Mangaoang et al.: Epiphytic pteridophyte biodiversity in Mt. Apo Natural Park

Species of Epiphytic Pteridophytes Conservation/Ecological Status

Lycopdiaceae

Phlegmariurus. bayanicus Economically Important Species

P. nummularifolius Endemic and Economically Important Species

P. salvinioides Endangered

P. verticillatus Economically Important Species

Aspleniaceae

Asplenium epiphyticum Endemic

A. apoense Endemic

A. decorum Depleted Species and Economically Important Species

A. nidus Depleted Species and Vulnerable

vittaeforme Vulnerable

Davalliaceae

Davallia embolostegia Other threatened species (OTS)

D. trichomanoides Other threatened species (OTS)

Dryopteridaceae

Polysticum elmeri Endemic, Other Wildlife Species (OWS)

Lindsaeaceae

Lindsaea apoensis Endemic

Ophioglossaceae

Ophioglossum pendulum Endangered

Polypodiaceae

Aglaomorpha cornucopia Endemic, Vulnerable

A. sagitta Vulnerable

Lecanopteris deparioides Endangered; Rare species

Lepisorus platyrhynchos Endangered

Tomophyllum macrum Endemic

Psilotaceae

Tmesipteris zamorarum Endangered

Pteridaceae

Antrophyum sessilifolium Endemic

Haplopteris alternans Endemic

Table 3. Conservation/Ecological status of epiphytic pteridophytes from Mt. Apo Natural Park.

Page 10: Rapid assessment of epiphytic pteridophyte biodiversity in ...asbp.org.ph/wp-content/uploads/2020/10/PJSB_2020-004.pdf · epiphytes face exceptional challenges because of their specific

Volume 14 Issue 3 - 2020 | 10 Philippine Journal of Systematic Biology Online ISSN: 2508-0342

Cano-Mangaoang et al.: Epiphytic pteridophyte biodiversity in Mt. Apo Natural Park

absent in the undisturbed site. This result further implied that

the three species mentioned are indicators of disturbance based

on the analysis. Furthermore, the clear separation of the

assemblage C was not only due to the unshared species but to

the kind of phorophytes thriving in the area. The number of tree

ferns as phorophytes in stations 3 and 4 of the undisturbed site

was lower compared to other stations.

Conservation and Ecological Status

There were eleven (11) threatened species of epiphytic

pteridophytes identified in this study from Mt. Apo Natural

Park. Of these, 5 are Endangered, 3 are Vulnerable, 2 Other

Threatened Species (OTS) and 1 Other Wildlife Species (OWS).

Nine species in the list were also recorded as endemic to Mt.

Apo Natural Park. There were 6 species which are Philippine

endemics and 3 species are restricted only to Mindanao. These

Philippine endemic species are Phlegmariurus nummularifolius

(Blume) Ching, Asplenium epiphyticum Copeland in Perkins,

Aglaomorpha cornucopia (Copeland) M.C. Roos, Antrophyum

sessilifolium (Cavanilles) Sprengel, Polystichum elmeri

Copeland, and Haplopteris alternans (Copeland) S. Lindsay &

C.W. Chen. Mindanao endemic species are Asplenium apoense

Copeland in Perkins, Lindsaea apoensis, and Tomophyllum

macrum (Copeland) Parris. Four species, Phlegmariurus

banayanicus (Herter) A.R. Field & Bostock, P.

nummularifolius, P. verticillatus (L. fil A.R. Field & Testo) and

Asplenium decorum Kuntze are characterized as economically

important (Table 3).

Conclusions and Recommendations

The rapid assessment of epiphytic pteridophytes in Mt.

Apo Natural Park revealed difference in abundance but not in

other diversity parameters. However, different assemblages

were noted due to the differences in species composition,

abundance, and their proportion in different sampling stations

with 50% overall similarity. Noteworthy is the presence of

Asplenium beccarianum, a new record of distribution in the

Figure 3. Diversity indices of epiphytic pteridophytes in the two

sampling sites of Mt. Apo Natural Park. A. Species richness, B.

Abundance, C. Evenness, D. Shannon-Weiner Index, E. Simpson’s Index

Figure 5. Assemblage Analysis (Non-metric dimensional scaling) of

epiphytic pteridophytes in the two sampling sites of Mt. Apo Natural

Park. (M-Malangit, disturbed site; E-EDC, undisturbed site, S-station; P

-plot)

Figure 4. K-dominance plot of the two sampling sites in Mt. Apo

Natural Park.

Page 11: Rapid assessment of epiphytic pteridophyte biodiversity in ...asbp.org.ph/wp-content/uploads/2020/10/PJSB_2020-004.pdf · epiphytes face exceptional challenges because of their specific

Volume 14 Issue 3 - 2020 | 11 © Association of Systematic Biologists of the Philippines

Cano-Mangaoang et al.: Epiphytic pteridophyte biodiversity in Mt. Apo Natural Park

Philippines. Furthermore, results of the study showed

uniqueness of the epiphytic pteridophyte species assemblage as

a result of anthropogenic disturbance and phorophyte

composition. This study suggests strengthening of the

implementation of policies for the proper management of Mt.

Apo Natural Park. Further, the study recommends conducting

studies on habitat preference of epiphytic pteridophytes to better

elucidate their abundance and distribution in the area.

Acknowledgement

The researchers would like to thank Department of

Environment and Natural Resources (DENR) Region XII for

the issuance of gratuitous permit – GP No. 13 on December 15,

2015, Local Government of Makilala most especially to the

Engr. Walter Ruizo, Municipal Environment and Natural

Resources Officer, Mr. Jamel Tawantawan, Protected Area

Superintendent for the assistance in securing the gratuitous

permit. Charissa Joy A. Gumban, Vince Abarquez, Joylinber

Tandingan, Joy Flores, Kuya Nono and Delfin Gordonas for the

assistance during field sampling. Thanks are also extended to

Krizler Tanalgo for revising the map. This study was funded by

the Commission on Higher Education (CHED) and Philippine

Tropical Forest Conservation Foundation (PTFCF) under grant

agreement no. II-2016-20.

Literature Cited

Adhikari, Y.P., H.S. Fischer & A. Fischer, 2012. Micro-site

conditions of epiphytic orchids in a human impact gradient

in Kathmandu valley, Nepal. Journal of Mountain Science,

29: 331-342,

Aguirre, A., R. Guevara, M. Garcia & J.C. Lopez, 2010. Fate of

epiphytes on phorophytes with different architectural

characteristics along the perturbation gradient of Sabal

mexicana forests in Veracruz, Mexico. Journal of

Vegetation Science, 21: 6-15.

Amoroso, V.B., F.M. Acma, F.P. Coritico, F.S. Gorme, N.E.

Lagunday, M.C.S. Salolog & R.D. Colong, 2019. Floral

diversity assessment of the buffer zones and vicinity of Mt.

Hamiguitan Range Wildlife Sanctuary (MHRWS), Davao

Oriental: basis for inclusion to protected area zone.

Philippine Journal of Systematic Biology,12 (2): 36-51.

Amoroso, V.B., C.B. Amoroso, & F.P. Coritico, 2011. Diversity

and status of plants in three mountain ecosystems in

Southern Mindanao, Philippines. Asian Journal of

Biodiversity, 3: 50-73.

Amoroso, V.B., W.L. Chiou, Y.C. Liu, J.R.C. Callado, N.N.G.

Briones, & F.P. Coritico, 2015 Threatened Endemic, Rare

and Economic Species of Philippine Pteridophytes.

Instructional Materials and Development Center, Central

Mindanao University, Musuan, Bukidnon. 173pp.

Amoroso, V.B., F.P. Coritico & P.W. Fritsch, 2016. Species

richness and conservation status of ferns and lycophytes in

Mt. Hamiguitan Range Wildlife Sanctuary, Davao Oriental,

Philippines. Philippine Journal of Science, 145 (2): 127-

137.

Andama, E., E. Charles, M. Michira & G.B. Luilo, 2003.

Studies on epiphytic ferns as potential indicators of forest

disturbances. XII World Forestry Congress.

Arévalo, R. & J. Bentacur, 2006. Vertical distribution of

vascular epiphytes in four forest types of the Serranía de

Chribiquete, Colombian Guayana. Selbyana, 27 (2): 175-

185.

Cach-Perez, M.J., J.L. Andrade, N. Chilpa-Galvan, M. Tamayo-

Chim, R. Orellana & C. Reyes-Garcia, 2013. Climatic and

structural factors influencing epiphytic bromeliad

community assemblage along a gradient of water-limited

environments in the Yucatan Peninsula, Mexico. Tropical

Conservation Science, 6 (1): 283-302.

Castañares, E.D., S.A. Vedra, & J.G. Gorospe, 2017.

Biodiversity and habitat assessment of Mt. Malindawag,

Naawan, Misamis Oriental, Philippines. International

Letters of Natural Sciences, 62: 20-27.

Chapman, M. & A. Underwood, 1999. Ecological patterns in

multivariate assemblages: information and interpretation of

negative values in ANOSIM tests. Marine Ecology

Progress Series, 180: 257-265.

Chomba, C., R. Senzota, H. Chabwela & V. Nyirenda, 2011.

The influence of host tree morphology and stem size on

epiphyte biomass distribution in Lusenga Plains National

Park, Zambia. Journal of Ecology and the Natural

Environment, 3 (12):370-380.

Clarke, K.R., 1993. Non-parametric multivariate analyses of

changes in community structure. Australian Journal of

Ecology, 18: 117-143.

Colwell, R.K., R.R. Dunn & N.C. Harris, 2012. Co-extinction

and persistence of dependent species in a changing world.

Annual Review of Ecology, Evolution, and Systematics, 43:

183– 203.

Copeland, E.B, 1958. Fern Flora of the Philippines. Vols. I-III.

National Institute of Science and Technology Bureau of

Printing, Manila. 557pp.

Cummings, J.M. & A. Rogers, 2006. Quantifying the abundance

of four large epiphytic fern species in remnant complex

Page 12: Rapid assessment of epiphytic pteridophyte biodiversity in ...asbp.org.ph/wp-content/uploads/2020/10/PJSB_2020-004.pdf · epiphytes face exceptional challenges because of their specific

Volume 14 Issue 3 - 2020 | 12 Philippine Journal of Systematic Biology Online ISSN: 2508-0342

Cano-Mangaoang et al.: Epiphytic pteridophyte biodiversity in Mt. Apo Natural Park

notophyll vine forest on the Atherton Tableland, North

Queensland, Australia. Cunninghamia, 9 (4): 521-527.

DENR Administrative Order [DAO] No. 2017-11, 2017.

Updated national list of threatened Philippine plants and

their categories. http://www.denr.gov.ph.

Fernando, E., L. Co, D. Lagunzad, W. Gruezo, J. Barcelona, D.

Madulid, A. Lapis, G. Texon, A. Manila, & P. Zamora,

2008. Threatened plants of the Philippines: a preliminary

assessment. Unpublished draft in the possession of the

authors.

Gowland, K.M., J. Wood, M.A. Clements, & A.B. Nicotra,

2011. Significant phorophyte (substrate) bias is not

explained by fitness benefits in three epiphytic orchid

species. American Journal of Botany, 98 (2): 197-206.

Haro-Carrion, X., T. Lozada, H. Navarrete, & G.H.J. De

Koning, 2009. Conservation of vascular epiphyte diversity

in shade cacao plantations in the Chocó Region of Ecuador.

Biotropica, 41 (4): 520-529.

Hietz, P., 1999. Diversity and conservation of epiphytes in a

changing environment. IUPAC. http://www.iupac.org/

symposia/proceedings/phuket97/hietz.html.

Hirata A., T. Kamijo, & S. Saito, 2009. Host trait preferences

and distribution of vascular epiphytes in a warm-temperate

forest. Plant Ecology, 201: 247–254.

Hoovenkamp,P. H., M.T.M. Bosman, E. Hennipman, H.P.

Nooteboom, G. Rodl-Linder & M.C. Roos, 1998.

Polypodiaceae. Flora Malesiana Series 2, 3: 1–234.

Jha,C.S., L. Goparaju, A. Tripathi, B. Gharai, A.S. Raghubanshi

& J.S. Singh, 2005. Forest fragmentation and its impact on

species diversity: an analysis using remote sensing and

GIS. Biodiversity and Conservation, 14: 1681-1698.

Karasawa, S. & N. Hijii, 2006. Determinants of litter

accumulation and the abundance of litter-associated

microarthropods in bird’s nest ferns (Asplenium nidus L.

complex) in the forest of Yambaru on Okinawa Island,

Southern Japan. Journal of Forestry Research, 11: 313-318.

Karger, D.N., P. Weigelt, V.B. Amoroso, D. Darnaedi, A.

Hidayat, H. Kreft & M. Kessler, 2014. Island biogeography

from regional to local scales: evidence for a spatially scaled

echo pattern of fern diversity in the Southeast Asian

archipelago. Journal of Biogeography, 41: 250-260.

Kress, W.J., 1986. A symposium: The biology of tropical

epiphytes. Selbyana, 9: 1-22.

Kromer, W., M. Kessler, & S.R. Gradstein, 2007. Vertical

stratification of vascular epiphytes in submontane and

montane forest of the Bolivian Andes: The importance of

the understory. Plant Ecology, 189: 261-278.

Lumbres, R.I.C., J.A. Palaganas, K.A. Laruan, E.D. Besic, C.W.

Yun & Y.L. Lee, 2014. Floral diversity assessment in Alno

communal mixed forest in Benguet, Philippines. Landscape

and Ecological Engineering, 10: 361-368.

Martin, C.E., T.C. Lin, C.C. Hsu & S.H. Lin, 2007. No effect of

host tree species on the physiology of the epiphytic orchid

Bulbophyllum japonicum in a subtropical rainforest in

Northeastern Taiwan. Taiwan Journal of Forest Science, 22

(3): 241-251.

Ong, P., L.E. Afuang, & R.G. Rosell-Ambal (eds.), 2002.

Philippine Biodiversity Conservation Priorities: A Second

Iteration of the National Biodiversity Strategy and Action

Plan. Philippine Department of the Environmental and

Natural Resources, Quezon City, 18-113 pp.

Pelser, P.B., J.F. Barcelona & D.I. Nickrent (eds), 2011 onwards.

Co’s Digital Flora of the Philippines.

www.philippineplants.org

PPG 1, 2016. A community-derived classification of extant

lycophytes and ferns. Journal of Systematics and Evolution,

54 (6): 563-603.

Quaresma, A.C. & M.A.G. Jardim, 2014. Floristic composition

and spatial distribution of vascular epiphytes in the restingas

of Maracanã, Brazil. Acta Botanica Brasilica, 28: 68-75.

Reich, A., J.J. Ewel, N.M. Nadkarni, T. Dawson & R.D. Evans,

2003. Nitrogen isotope ratios shift with plant size in tropical

bromeliads. Oecologia, 137: 587-590.

Replan, E.L. & A.B.B. Malaki, 2017. Floral diversity and

habitat assessment of Cabantug Forest, Argao, Central

Visayas, Cebu, Philippines. International Journal of

Scientific & Engineering Research, 8 (10): 775-780.

Schmitt, J.L. & P.G. Windisch, 2009. Biodiversity and spatial

distribution of epiphytic ferns on Alsophila setosa Kaulf.

(Cyatheaceae) caudices in Rio Grande do Sul, Brazil.

Brazilian Journal of Biology, 70 (3): 521-528.

Stireman, J.O., H. Devlin & A.L. Doyle, 2014. Habitat

fragmentation, tree diversity, and plant invasion interact to

structure forest caterpillar communities. Oecologia, 176:

201-224.

Tewari, L.M., G. Tewari, T. Nailwal, & Y.P.S. Pangtey, 2009.

Bark factors affecting the distribution of epiphytic ferns

communities. Nature and Science, 7(5): 76-81.

Vidal, K.C., E.D. Macusi, & A.G. Ponce, 2018. Inventory and

mophometrics of anuran species found in Mt. Kilala of the

Mt. Hamiguitan Range Wildlife Sanctuary, Governor

Generoso, Davao Oriental, Philippines. Philippine Journal

of Science, 147 (4): 631-640.

Zambrano, J., C.X.G. Lopez, L. Yeager, C. Fortunel, N.J.

Cordeiro & N.G. Beckman, 2019. The effects of habitat

loss and fragmentation on plant functional traits and

functional diversity: what do we know so far? Oecologia,

191: 505-518.

Page 13: Rapid assessment of epiphytic pteridophyte biodiversity in ...asbp.org.ph/wp-content/uploads/2020/10/PJSB_2020-004.pdf · epiphytes face exceptional challenges because of their specific

Volume 14 Issue 3 - 2020 | 13 © Association of Systematic Biologists of the Philippines

Cano-Mangaoang et al.: Epiphytic pteridophyte biodiversity in Mt. Apo Natural Park

Zapanta, B.R., M.J.M.M. Achondo, A.F.M. Raganas, F.A.,

Camino, A.G.D. Delima, J.A. Mantiquilla, , R.P. Jr.

Puentespina, & F.R.P. Salvaña, , 2019. Species richness of

tress in disturbed habitats within a protected area and its

implications for conservation: The case of Mt. Apo Natural

Park, Mindanao Island, Philippines. Biodiversitas, 20: 2081

-2091

Zhang, L., S. Nurvianto & R. Harrison, 2010. Factors affecting

the distribution and abundance of Asplenium nidus L. in a

tropical lowland rainforest in peninsular Malaysia.

Biotropica, 42 (4): 464-469.

Zhao, M., N. Geekiyanage, J. Xu, M.M. Khin, D.R. Nurdiana,

E. Paudel & R.D. Harrison, 2015. Structure of the epiphyte

community in a tropical montane forest in SW China. PLoS

ONE 10(4): e0122210.

Zotz, G., G. Mendieta-Leiva & K. Wagner, 2014. Vascular

epiphytes at the treeline-composition of species

assemblages and population biology. Flora—Morphology,

Distribution, Functional Ecology of Plants, 209(8): 385-

390.