Accepted Manuscript

The impact of forest conversion on bird Communities in Northern Flank of Knucklesmountain forest range, Sri Lanka

K. Subasinghe, A.P. Sumanapala, S.R. Weerawardhena

PII: S2287-884X(14)00045-4

DOI: 10.1016/j.japb.2014.07.004

Reference: JAPB 29

To appear in: Journal of Asia-Pacific Biodiversity

Received Date: 27 June 2014

Revised Date: 24 July 2014

Accepted Date: 24 July 2014

Please cite this article as: Subasinghe K, Sumanapala AP, Weerawardhena SR, The impact of forestconversion on bird Communities in Northern Flank of Knuckles mountain forest range, Sri Lanka,Journal of Asia-Pacific Biodiversity (2014), doi: 10.1016/j.japb.2014.07.004.

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The impact of Forest Conversion on Bird Communities in Northern Flank

of Knuckles Mountain Forest Range, Sri Lanka

K. Subasinghe*, A. P. Sumanapala , S.R. Weerawardhena,

Department of Zoology, University of Kelaniya, Sri Lanka

Manuscript type: Original Paper

Corresponding author: K. Subasinghe

E-mail: kalya_subasinghe@yahoo.com

Postal address: No. 258, Wedamulla lane, Kelaniya 11600, Sri Lanka.

T.p: +94112910641/ +94779319100

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Abstract

The consequence of forest conversion into human land-use systems on tropical

biodiversity is needed to be assessed in order to initiate proper conservation and management

strategies. In this study the bird species diversity and composition were characterized along

with the vegetation structural components in four land-use types in Northern Flank of

Knuckles mountain fore range, which is a part of one of the world heritage site in Sri Lanka.

This includes three human modified land-uses (cardamom, pinus and abandoned tea

plantations), and an undisturbed forest. A total of 1,023 individuals belonging to 51 bird

species were recorded using fixed radius point count method. The cardamom plantation with

native shade trees, had a bird species richness and composition comparable to undisturbed

forest (One-way ANOVA; p>0.05, Jaccard index Cj= 0.56). Overall species diversity in

terms of Shannon Wiener index was highest in undisturbed forest. Pearson’s correlation

coefficient suggested a strong positive linear relationship between bird species richness with

canopy cover (r = 0.738) and vertical stratification (r = 0.813). This study reveals that the

land-use systems formed with considerable alterations to vegetation structure significantly

reduce the bird diversity and supports a bird community less comparable to undisturbed

forest.

Key words: Bird species richness, Canopy cover, Cardamom, Point count method, Pinus,

Vegetation structure.

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Introduction

Much of the natural forests throughout the world have been destroyed and converted

to human land-uses in order to meet the ever growing demands for resources (FAO, 2012).

Consequently, most of the remaining forest landscapes are with mosaic of human modified

land-areas such as urban, agricultural and plantation lands. The conditions of these land areas

are highly deviated from its previous undisturbed state.

Although the rate of loss of forest cover has slowed compared to the period of 1990–

2000 (FAO, 2011), it is still occurs at an alarming rate (net annual loss of the world’s forests

was 6.4 million ha between 2000 and 2010) which need a concerted effort to slow or reverse

the trend. Most of the tropical forests that contain a large amount of world’s biological

diversity are still being cleared for agricultural purposes and also for timber industry. Today

forest plantations have established all around the world to compensate forest loss through

agricultural and other anthropogenic disturbances which also have became a major

contributor to local and global economies. However, most of these plantations that have

established using tree species with fast growth rates greatly differ from the naturally

regenerated forests in both composition and structure leading to different ecological

processes and thereby different functional outcomes (Davis et al. 2012).

It is clearly evident, that the conversion of natural forests into human land-uses

negatively affects the rich biodiversity associate with forest ecosystems (Holloway, 1996;

Houlahan and Findlay, 2011; Lupatini et al. 2012; Meijer et al. 2011; Pekin and Pijanowski,

2012; Pimm et al. 2006) and this trend is likely to continue with the rapid growth of

population that results in increasing demands for forest resources. According to Birdlife

International (2013), two-thirds of bird species are found in forests and much of them are

restricted to forest ecosystems. Forest conversion has been identified as a factor that poses a

major threat to the substantial number of bird species, as such it can destroy and degrade the

habitats of species via reducing available food sources, roosting and nesting places and as

well as restricting the movements. However, not all bird species are equally vulnerable to

forest conversions (Sekercioglu et al. 2002; Sodhi et al. 2004). Rare and restricted range

birds, rainforest habitat specialists and altitudinal migrants are more susceptible to habitat

alterations than generalists (Raman, 2001). Several studies have shown that some of the

human land-uses hold a considerable biological diversity. Agro-forestry systems such as

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shade coffee (Coffea arabica) and cacao plantations (Theobroma cacao) shown to support

greater number of forest birds species with higher diversity than open agricultural systems

with few or no trees (Estrada et al. 1997; Greenberg et al. 1997; Petit et al. 1999). This has

lead to a doubt how vital are to incorporate human land-uses in conservation planning.

The Knuckles mountain forest range (KMFR) in Sri Lanka, which has been declared

as a World heritage site for its rich biodiversity (UNESCO, 2012), was severely impacted by

the economic trends over the past several decades. Some parts of the forest within the range

had been cleared to cultivate coffee and subsequently to grow tea at the middle of last century

(Gunawardane, 2003). Since the climatic condition in this forest range is favorable for

cardamom plantation, several planters also engaged in under-planting of cardamom where

they remove quarter of the over-story of the natural forest (Abeygunawardena and Vincent,

1993). This under-planting had become a bigger threat to the forest range in 1960 when the

government authorized individuals and groups to grow cardamom. With the initiation of new

reforestation policies in 1960s a number of pinus plantations were established around the

range (Medawatte et al. 2008). Several attempts have been made to analyze the differences in

ecological functions and diversities in these human land-uses and undisturbed forest

(Dharmaparakrama, 2006; Nizam, 2011; Weerakoon et al. 2010).

Present study reports on bird communities in four land-uses in Northern Flank of

KMFR: 1. the undisturbed sub-montane forest (the type of forest formation common to the

selected altitudinal range) 2. cardamom plantation 3. abandoned tea plantation 4. exotic pinus

plantation. Thereby assess differences in bird community structure in terms of bird species

richness, species diversity and species similarity. Also this study investigates the structural

components of vegetation in all selected land-uses in order to examine the influence of each

component to species richness of birds.

Materials and methods

Study area

This study was conducted in Northern region of KMFR. The study area was restricted

to an elevation range of 600 to 1300 m asl which is the range specific to sub-montane forest

vegetation (Bambaradeniya and Ekanayake, 2003). The study area consists of a continuous

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and fragmented sub-montane forest, commercial plantations of pinus, cardamom, abandoned

tea plantation and small patches of eucalyptus plantations. This study focused on the

prominent human land-use types within the study area i.e. (1) cardamom plantations, (2)

pinus plantations, (3) abandoned tea plantations and as well as the (4) sub-montane forest as

unmodified land-use. The map of study location is given in Fig. 1.

Figure 1: Map of the study area

Bird sampling

Bird sampling was carried out during March 2012 to September 2012. A standard

fixed radius point-count method (Ralph et al. 1995) with the radius set at 25 m was used to

sample birds in all modified land-uses and undisturbed sub-montane forest. Sampling was

carried out for three consecutive days in the months of March, April, June and September. In

each visit a similar number of point count stations were sampled from each land-use type. A

total of 28 point count stations were established within each of the four land-uses. All stations

were established with a distance of 100 m from one another and 100 m from the edge of the

each land-use. Sampling stations were visited between 0600 and 0930 in the morning and

1530 and 1700 in the evening. Each station was surveyed for 10 minutes period and all the

birds detected visually were recorded to the species level using Harrison (1999) and

Kotagama and Ratnavira (2010). The detected but doubtful, uncommon or rare bird species

were marked and identification was confirmed subsequently by the acoustical and visual

recordings. Any flying birds that were not seen to take-off from the point count station were

excluded from analysis. No surveys were made on rainy or windy days. Bird counting was

not repeated on any of the point count station (Ralph et al. 1995).

Vegetation sampling

To describe the vegetation structure of sub-montane forest and the human land-uses,

eight 10 × 10 m subplots were established in eight randomly selected point count station of

each plantation type and forest. Vegetation variables were estimated in each subplot and then

averaged for each land-use type. In each subplot, every tree with a diameter at breast height

(dbh) greater than 10 cm was counted for tree density. Tree height was visually estimated

within each subplot. A digital rangefinder was used to improve the estimates of tree height

(Van Bael et al. 2007).

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Vertical stratification (distribution of foliage at different vertical levels) was assessed

in 16 points by recording presence of foliage in height classes (in metres) of 0-1, 1-2, 2-4, 4-

8, 8-16, 16-24, 24-32, and >32 in an imaginary vertical cylinder of 0.5 m radius around the

observer (Sidhu et al. 2010). This was measured using two transects established along the

center of the point count station with an interval of 5 m. Canopy cover was assessed using a

sighting tube with a single central cross-wire (Ganey and Block, 1994; Goodenough and

Goodenough, 2012). At each two meter point along the same transects, the observer looked

directly overhead through the tube and recorded whether or not the crosshair intercepted

overhead foliage (Ganey and Block, 1994). The result of each individual measurement is

recorded as 1 if the crosshair intercepted and 0 otherwise. Canopy cover is then estimated as

the percentage of points intercepted. Percentage of herbaceous cover (non-woody vegetation

with height < 1 m) was measured using the point quadrat method (Sutherland, 2006).

Data analysis

One-way ANOVA followed by the Tukey Test was used to compare species richness

and relative abundance (number of individuals per point count station) among undisturbed

forest and human modified land-uses after testing the normality of data sets using Anderson

Darling test. Statistical analysis was conducted using MINITAB 14 statistical software.

Shannon-Weiner diversity index (H´) was used to characterize bird species diversity in all

land-uses. Jaccard coefficients (Cj) were used to study the similarities of bird species

composition between human modified land-uses and the undisturbed sub-montane forest

(Magurran, 2004).

The average number of strata with foliage across 16 points within each point count

station was obtained to estimate the vertical stratification. Pearson correlation was performed

in order to evaluate the relationship between bird species richness and each of the vegetation

variable assessed. For that, only the bird community data obtained from the eight point count

stations that have selected to determine vegetation structural attributes were considered.

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Results

Bird richness and diversity

A total of 1,023 birds belonging to 51 species arrayed among 27 families were

recorded on all point count stations during the study. From these, 11 species were endemic to

Sri Lanka. Out of the 51 species recorded, 31 bird species were recorded from sub-montane

forest whereas 33 bird species were recorded from the cardamom plantations. Only fewer

number of bird species were recorded from abandoned tea plantations (13) and pinus

plantations (20). The number of bird species recorded per point count station from the

undisturbed forest was not significantly different from the number recorded from cardamom

plantation (One way ANOVA; p>0.05), but was significantly higher than the pinus or

abandoned tea plantations (One way ANOVA; p<0.05). Relative abundance of birds

measured as the number of individuals per point count station in undisturbed forest, was

significantly higher than the abandoned tea plantation, but was lower than the cardamom

plantation (Table 1). The Shannon Weiner diversity was highest in the sub-montane forest out

of all land-use types considered.

Table 1

Cardamom plantation shown the highest similarity to the sub-montane forest in terms

of bird species composition (Cj=0.56; Table 2). However, only fewer number of bird species

were reported in common between the undisturbed forest and other modified land-use

systems; pinus (Cj=0.28) and the abandoned tea plantations (Cj=0.19).

Table 2

Asian black bulbul (Hypsipetes leucocephalus), Yellow-fronted barbet (Megalaima

flavifrons) and Sri Lanka Scimitar-babbler (Pomatorhinus melanurus) were abundant in

undisturbed forest. Yellow-eared bulbul (Pycnonotus penicillatus) and Sri Lanka dull blue

flycatcher (Eumyias sordidus) were dominant in cardamom plantation. Bird species that are

commonly found in more disturbed habitats such as Red-vented bulbul (Pycnonotus cafer)

and the White-bellied drongo (Dicrurus caerulescens) were reported from the pinus and

abandoned tea plantations (Annexure).

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Vegetation structure of land-uses

Vegetation structural attributes showed differences between the undisturbed forest

and the human modified land-uses. Vertical distributions of foliage were distinguishable

between the land-use types. Canopy cover was highest in undisturbed forest, moderate in

cardamom plantation and least in pinus and abandoned tea plantations. Tree density and tree

height were highest in pinus plantation while the vertical stratification was highest in

cardamom plantation (Table 3). The herbaceous cover was highest in abandoned tea

plantations.

Table 3

The relationship of bird species richness to the vegetation structural attributes

All vegetation structural variables, except the herbaceous cover, positively correlated

with the bird species richness (Table 4). Bird species richness showed a strong positive linear

relationship with the canopy cover (Pearson’s correlation coefficient r= 0.813) and with the

vertical stratification (Pearson’s correlation coefficient r= 0.738). However, herbaceous cover

showed a negative linear relationship with the bird species richness (Pearson’s correlation

coefficient r= -0.456).

Table 4

Discussion

Conversion of natural forests into human modified land-uses created noticeable

impacts on bird community parameters and composition, as species tend to attract or avoid

the modifications. From the present study it is clear that the human modified land-uses could

significantly reduce the bird species richness and diversity. However, land-use systems with

native canopy tree species have the potential to support avifaunal community comparable to

the natural forests. In accordance with previous studies (Dawson et al. 2011), highest bird

diversity in terms of Shannon Weiner index was recorded from the undisturbed forest within

the study area. According to the results, the total abundance of birds are seems not to be

affected by the conversions as the modified areas are colonized by few generalist species.

However high visibility associated with the human modified land-uses might have resulted in

greater number of observations than in sub-montane forest. Therefore, noticeable differences

between land-use types should be judged carefully if corresponding tests met the level of

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statistical significance as explained in Schulze and Riedl (2008). Among the suite of variables

that may influence bird community structure in different land-uses, vegetation structural

characteristics seem to have a particularly strong influence.

The native sub-montane trees that had preserved for shade in cardamom plantations

have grown well due to release from competition. These trees have provided a high floristic

diversity and a complex vegetation structure to the monoculture. This might have lead to high

bird species richness in cardamom plantation. The high bird species richness found in rustic

coffee and shaded monoculture was explained by the structural complexity of the system

(Cruz and Sutherland, 2004). Also the proximity to undisturbed forest may also have

enhanced the bird community in the cardamom plantation. Bird species composition of

cardamom plantation is shown to be much more similar to sub-montane forest (Cj= 0.56).

Sub-montane tree species retained for shade provides suitable habitat for forest species and

may have resulted in more similar bird composition between these land-use types (Raman

and Sukumar, 2002).

Current study reported a significantly lower number of bird species from pinus

plantations than from the sub-montane forest; as such exotic monoculture plantations are

unsuitable for many native species. Structural components of native habitats such as

understory vegetation, snags and dead trees are absent in those plantations (Clout and Gaze,

1984). Senanayake (1987) observed 3 species of bird in a pinus monoculture and 5 in

eucalyptus, compared with 25 in natural forest. Studies conducted in New Zealand have also

shown that pinus plantations to be poor habitat for native birds, particularly for those which

feed on fruit and nectar, nest in holes or insectivorous species (Clout, 1984). However, the

presences of few eucalyptus trees and considerable undergrowth in pinus plantation within

the study area have resulted in greater number of bird species than expected.

A dense non woody plant covers with herbs and grasses in abandoned tea plantations

allowed bird species that can utilize herbaceous strata. Bird species such as Ashy Prinia

(Prinia socialis), Blyth's Reed-warbler (Acrocephalus dumetorum) and Tawny-bellied

Babbler (Dumetia hyperythra) with preferred foraging heights of 1 m were common in this

land use type. Though the bird species composition of abandoned tea plantation deviated

from the undisturbed forest, this has contributed to increase the species richness within the

landscape through harboring bird species that dwell in more open habitats. Presence of open

habitat birds such as Common Iora (Aegithina tiphia), Ashy Prinia, Black throated Munia

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(Lonchura kelaarti) and Common Tailorbird (Orthotomus sutorius) in the undisturbed forest

could be due to the existence of more open habitats in close proximity. Large billed leaf

warbler (Phylloscopus magnirostris), Brown capped babbler (Pellorneum fuscocapillus) and

Dark fronted babbler (Rhopocichla atriceps) that are found in undergrowth forests were

reported only from the undisturbed sub-montane forest and the cardamom plantations.

The vegetation structure is considered to be an important determinant of bird diversity

(Diaz, 2006; Jankowski et al. 2012; Karr and Roth, 1971; MacArthur and MacArthur, 1961).

In the present study some of the structural attributes have shown to be directly influence the

bird species richness. This includes vertical stratification and canopy cover. The increase in

number of vertical strata in a habitat allows greater variety of bird species that have adopted

to utilize each of the vertical strata. Tropical forest vertical stratification is known to be

important for bird species (Winkler and Preleuthner, 2001). According to Raman (2001)

vertical stratification mostly related with bird community attributes, only when the habitats

are highly altered. Contrary to the results of this study, Daniels et al. (1992) has shown an

inverse relationship between bird species richness and vertical stratification of different

vegetation formations. Posa and Sodhi (2006) have demonstrated the important for the

having high values of canopy cover (60% or more) for the existence of many forest bird

species. Several studies have shown a strong positive correlation between bird species

richness and tree height (Gillespie and Walter, 2001; Joshi et al. 2012). However, present

study did not show such strong correlation between tree heights and the bird species richness.

Conclusion

This research concurs that the bird community structure and composition of forests

could considerably altered by converting them into human land-uses. But the degree of

change depends on the type of land-use. Human modified land-uses with a complex

vegetation structure would have the potential to sustain high avifaunal diversity, though the

long-term impacts of such land-uses are still unknown. Therefore it is important to consider

some of the human modified land-uses specially shade plantations into conservation

planning. Since many bird species are undoubtedly dependent on undisturbed forest, it is

essential to take actions to prevent further forest destructions.

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Acknowledgment

I would like to thank University of Kelaniya and the Department of Forest

Conservation, Sri Lanka for giving me the opportunity to carry out this research. Further my

sincere gratitude goes to my parent and to Mr. Upali Ekanayake who is an experienced

ornithological tour leader for extensive support given during field surveys.

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Annexure

Relative abundances of bird species in four land-use types.

Family Name Scientific Name SM CA PI AT

National Conservation

Status (MOE, 2012)

Accipitridae Accipiter badius (Gmelin, 1788) 0.07 0.04 - - LC Accipitridae Nisaetus nipalensis Hodgson, 1836 0.07 - - - VU Accipitridae Pernis ptilorhyncus (Temminck, 1821) M - - 0.04 - NT Accipitridae Spilornis cheela (Latham, 1790) - - 0.04 - LC

Aegithinidae Aegithina tiphia (Linnaeus, 1758) 0.11 - - 0.11 LC Campephagidae Pericrocotus flammeus (Forster, 1781) - 0.25 0.04 - LC Cisticolidae Prinia inornata Sykes, 1832 - - - 0.04 LC Cisticolidae Prinia socialis Sykes, 1832 0.18 - - 0.57 LC Columbidae Chalcophaps indica (Linnaeus, 1758) - - 0.07 - LC Columbidae Stigmatopelia chinensis (Scopoli, 1786) - - 0.29 0.21 LC Columbidae Treron pompadora (Gmelin, 1789) - 0.04 - - LC Cuculidae Centropus sinensis (Stephens, 1815) - 0.04 - - LC Cuculidae Clamator coromandus (Linnaeus, 1766) M 0.14 - - - N/E Dicaeidae Dicaeum erythrorhynchos (Latham, 1790) - 0.32 - 0.11 LC Dicruridae Dicrurus caerulescens (Linnaeus, 1758) - - 0.21 0.07 LC

Estrildidae Lonchura kelaarti (Jerdon, 1863) E 0.21 - - - VU Laniidae Lanius cristatus Linnaeus, 1758 M 0.14 - - - N/E Monarchidae Hypothymis azurea (Boddaert, 1783) - 0.07 - - LC

Monarchidae Terpsiphone paradise (Linnaeus, 1758) M 0.07 - 0.11 - LC

Muscicapidae Culicicapa ceylonensis (Swainson, 1820) 0.04 0.50 - - Not given Muscicapidae Cyornis tickelliae Blyth, 1843 0.29 0.14 0.04 - LC

Muscicapidae Eumyias sordidus (Walden, 1870) E 0.11 0.71 - - VU

Nectariniidae Nectarinia lotenia (Linnaeus, 1766) 0.21 0.04 0.07 - LC Paridae Parus major Linnaeus, 1758 0.25 0.11 0.32 0.04 LC Phasianidae Gallus lafayetii Lesson, 1831 E - 0.04 - - LC Picidae Chrysocolaptes lucidus (Scopoli, 1786) 0.14 0.11 - - LC

Picidae Dinopium benghalense (Linnaeus, 1758) - 0.07 0.07 - LC Psittacidae Loriculus beryllinus (Forster, 1781) E 0.07 0.43 - - LC Psittacidae Psittacula calthropae (Blyth, 1849) E - 0.21 - - NT Pycnonotidae Hypsipetes leucocephalus (Gmelin, 1789) 0.68 0.71 0.89 - LC Pycnonotidae Pycnonotus cafer (Linnaeus, 1766) - - 0.04 0.5 LC Pycnonotidae Pycnonotus penicillatus Blyth, 1851 E 0.43 0.82 - - VU Ramphastidae Megalaima flavifrons (Cuvier, 1816) E 0.57 0.64 0.61 - LC

Ramphastidae Megalaima rubricapillus (Gmelin, 1788) 0.07 0.04 0.07 - LC

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Rhipiduridae Rhipidura aureola Lesson, 1830 0.11 - - - LC Sittidae Sitta frontalis Swainson, 1820 - 0.14 0.11 - LC Sturnidae Gracula ptilogenys Blyth, 1846 E 0.04 0.43 0.43 - VU Sturnidae Gracula religiosa Linnaeus, 1758 0.11 0.04 0.36 - LC Sylviidae Acrocephalus dumetorum Blyth, 1849 M 0.11 - - 0.11 N/E

Sylviidae Orthotomus sutorius (Pennant, 1769) 0.39 0.04 0.14 0.29 LC

Sylviidae Phylloscopus magnirostris Blyth, 1843 M 0.14 0.04 - - N/E Sylviidae Phylloscopus trochiloides (Sundevall, 1837)

M - 0.07 - - N/E

Timaliidae Chrysomma sinense (Gmelin, 1789) - - - 0.14 LC Timaliidae Dumetia hyperythra (Franklin, 1831) - - - 0.07 LC Timaliidae Pellorneum fuscocapillus (Blyth, 1849) 0.29 0.14 - - LC

Timaliidae Pomatorhinus melanurus Blyth, 1847 E 0.5 0.36 - - LC Timaliidae Rhopocichla atriceps (Jerdon, 1839) 0.11 0.11 - - LC

Trogonidae Harpactes fasciatus (Pennant, 1769) - 0.04 - - NT Turdidae Zoothera spiloptera (Blyth, 1847) E 0.04 0.11 - - VU

Zosteropidae Zosterops ceylonensis Holdsworth, 1872 E 0.14 0.32 0.21 - NT

Zosteropidae Zosterops palpebrosus (Temminck, 1824) 0.18 0.04 - 0.11 LC

(SM: sub-montane forest, CA: cardamom plantation, AT: abandoned tea plantation, PI: pinus plantation, PP: pitawala patana, E: Bird species endemic to Sri Lanka. M: Migrants)

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Table 1: The results of bird community assessments in undisturbed forest and the three

human modified land-use types. Tabled values are means and standard errors (SE).

(*) denote the human land-uses that have significantly different means (at the 0.05 level of significance) for bird species richness and

relative abundance from that of undisturbed forest (ANOVA, Tukey’s HSD).

Table 2: The Jaccard coefficients of similarity (below diagonal) between the land-use types.

(SM: Sub-montane forest, CA: Cardamom plantation, PI: Pinus plantation, AT: Abandoned

tea plantation).

Land-use types N No. of species per

point (Mean ± SE)

Relative abundance (Mean ± SE)

Shannon Weiner diversity

Sub-montane forest 28 6.2 ± 0.42 8.9 ± 0.67 3.067

Cardamom plantation 28 7.2 ± 0.31 15.4 ± 0.91 * 2.837

Pinus plantation 28 4.2 ± 0.24 * 9.1 ± 0.68 2.343

Abandoned tea plantation

28 2.2 ± 0.18 * 3.3 ± 0.33 * 2.098

SM CA PI SM CA 0.56 PI 0.28 0.29 AT 0.19 0.10 0.18

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Table 3: Vegetations characteristics of the selected land-use types within the study area.

Table 4: The relationship between bird species richness and vegetation structural attributes.

(r = Pearson’s correlation coefficient)

Sub-montane forest

Cardamom plantation

Pinus plantation

Abandoned tea plantation

Tree density (per 100m2)

13.9 ±1.15 7.4 ± 0.77 20.4 ± 1.18 0.9 ± 0.23

Tree height (m) 7.76 ± 0.22 12.2 ± 0.32 19.4 ± 0.09 3.2 ± 0.49

Canopy cover (%) 86.9 ± 2.30 62.5 ± 3.27 23.1 ± 1.32 4.4 ± 2.90

Herbaceous cover (%) 55.9 ± 0.72 28.5 ± 8.11 17.5 ±2.05 88.2 ± 0.73

Vertical stratification 4.1 ± 0.13 4.2 ± 0.21 1.8 ± 0.15 2.2 ± 0.09

Vegetation attributes r p value

Tree density 0.413 0.019

Tree height 0.410 0.020

Canopy cover 0.738 0.001

Herbaceous cover -0.456 0.009

Vertical stratification 0.813 0.001

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