Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India

CHNAES-00684; No of Pages 26 Acta Ecologica Sinica xxx (2020) xxx

Contents lists available at ScienceDirect

Acta Ecologica Sinica journal homepage: www.elsevier.com/locate/chnaes

Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India Kothandaraman Subashree a, Javid Ahmad Dar a, Subbiah Karuppusamy b, Somaiah Sundarapandian a,⁎ a b

Department of Ecology and Environmental Sciences, Pondicherry University, Puducherry 605014, India Department of Botany, Centre for Botanical Research, The Madura College, Madurai 625011, India

a r t i c l e

i n f o

Article history: Received 1 July 2019 Received in revised form 29 December 2019 Accepted 6 February 2020 Available online xxxx Keywords: Phytodiversity Stand structure Regeneration status Tropical forests Western Ghats

a b s t r a c t Diversity, stand structure and regeneration potential are the key elements of any forest ecosystem. For the present study, seven sites were selected with the aims of assessing plant diversity, structure and regeneration potential in tropical forests across Kanyakumari Wildlife Sanctuary (KWLS), Western Ghats, India. The sites were classified based on the similarity: tropical dry deciduous sites (TDDs I and II), tropical semi-evergreen sites (TSEs I and II) and tropical evergreen sites (TEFs I-III). The phytosociological survey was done by laying a total of 70 plots (10 plots in each study site). Standard methods were followed for the assessment of diversity, structure and regeneration patterns. A total of 267 species (205 genera, 70 families) were recorded. The tree species richness ranged 24 (TDD II) – 76 (TEF III). Of the vegetation spectrum, trees, vines and understorey accounted 56.5, 15.3 and 28.2% respectively to the total flora documented. A total of 66 species were endemic. The total tree density and tree basal area (seedlings, saplings, juveniles and adults) were 18,790 individuals (mean 2684) and 137.6 m2 (mean 19.7 m2) in 70 plots respectively. The mean tree adult density and basal area ranged 370 (TDD I) – 900 (TEF I) individuals/ha and 24.2 (TDD I) – 75.3 (TEF III) m2/ha respectively. The overall species richness was highest in TDD I, but TEF III had the highest tree species richness. The diameter class-wise distribution showed the characteristic ‘reverse J-shaped’ curve. Most tree species were ‘newly recruited’. The dominant species had ‘fair’ to ‘good’ regeneration potential. However, 12 tree species showed ‘no’ regeneration. The overall regeneration pattern of trees was ‘good’, but ‘no’ or ‘poor’ regeneration patterns in some tree species, especially endemics is a point of concern. Since a majority of tree species were ‘new recruits’, species composition may likely change in the future. The results obtained would help in understanding diversity patterns, structural attributes and regeneration potential in tropical forests of protected areas for better forest management and conservation. © 2020 Ecological Society of China. Published by Elsevier B.V. All rights reserved.

1. Introduction Plants that grow together in a community have mutual relationships with each other and also with the environment [1,2]. Phytosociological analysis forms the basic foundation for understanding the ecology of any piece of vegetated land [3,4]. Diversity and structure are the most important characteristics of a forest community [5]. Diversity is an attribute of a community that determines the stability, productivity and trophic structure of a forest ecosystem and an area with high diversity tends to be a very productive and stable ecosystem [6]. On the other hand, a forest’s structure arises as a result of self-organization, which

involves competition and many other interactions among the tree individuals as well as environmental factors [7,8]. Regeneration and competition for resources produce particular patterns in forest structure, which in turn generates characteristic processes of growth and regeneration, ultimately resulting in specific patterns of diversity [5,9]. Studies on population structure reveal the efficiency of species cohabitation and endurance capacities to various environmental processes [10]. The quantitative relationship which exists between rampant and rare species is of key significance in a community [3]. It is the population densities of seedlings, saplings, juveniles and adults which hint on the population structure and regeneration

Abbreviations: A/F, Abundance to Frequency ratio; APG, Angiosperm Phylogeny Group; BD, Beta diversity; BIOTIK, Biodiversity Informatics and co-Operation in Taxonomy for Interactive shared Knowledge base; cm, Centimetre; °C, Degree Celsius; DBH, Diameter at breast height; FSI, Forest Survey of India; GRIN, Germplasm Resources Information Network; ha, Hectare; IVI, Importance Value Index; km, Kilometre; KWLS, Kanyakumari Wildlife Sanctuary; m, Metre; m a.s.l, Metres above sea level; mm, Millimetre; PCoA, Prinicipal coordinate analysis; Repl, Replacement; RichDiff, Richness Difference; TDD, Tropical dry deciduous forest; TEF, Tropical evergreen forest; TSE, Tropical semi-evergreen forest; WG, Western Ghats. ⁎ Corresponding author at: Department of Ecology and Environmental Sciences, School of Life Sciences, Pondicherry University, Puducherry 605014, India. E-mail address: [email protected] (S. Sundarapandian).

https://doi.org/10.1016/j.chnaes.2020.02.004 1872-2032/© 2020 Ecological Society of China. Published by Elsevier B.V. All rights reserved.

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

2

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

potential of a forest community [10,11]. Regeneration potential could be defined as “the ability of a species to complete the life-cycle” and a successful regeneration is one that has adequate growth and survival of seedlings and saplings [12]. In contrast, inadequate numbers of seedlings and saplings point towards a poor regeneration and lack of seedlings and saplings of a tree species indicates no regeneration [13]. The regeneration potential of forests not only reveals the current and future composition of the forests but also depicts their long-term sustainability [14]. Assessing the population structure of forests is very crucial because as per the alarming report of Rai and Saxena [15], ~72.1% of the Indian forests have lost the regeneration potential. Regular assessments of regeneration status of forest communities (especially from protected areas) are of extreme significance as it provides the key to successfully maintain and manage natural populations and enhance their productivity. Tropical forests are globally rich centres of biodiversity due to their diverse biomes [16]. Tropical forests are known to harbour more than half of the world’s species, despite occupying only 7% of the land surface [17]. Slik et al. [18] estimated that tropical forests may contain 40,000– 53,000 tree species. However, in part, due to their richness in biodiversity, tropical forests are under continual anthropogenic pressures, which led to the existence of tropical forests as fragments in many parts of the world [19]. Tropical forests that are pristine and undisturbed have nowadays become very rare [20]. Due to their rich biological wealth and also the multiple threats they face, about 15% of the tropical forests are prioritized for conservation [21]. India is one of the 17 mega-diverse countries in the world and it ranks 10th among the countries with a large forest area (329 million ha). It harbours a total of 47,513 plant species [22], of which 28% are endemic and it represents 11.4% of the world’s flora [23]. The tropical forests of Western Ghats are ecologically rich, being endowed with immense biodiversity and are one of the 36 biodiversity hotspots of the world. Of the entire stretch of Western Ghats, the southern Western Ghats are considered to be the richest in terms of floristic composition and has a high concentration of endemic species [24]. Like the other tropical forests worldwide, the tropical forests of Western Ghats are also being fragmented at a greater pace, especially due to their conversion to land-uses [19]. Several regions in the Western Ghats have been demarcated as ‘protected areas’ for conservation as national parks, wildlife sanctuaries and biosphere reserves. Quantitative information on species composition, forest structure and regeneration potential is important to plan and implement conservation strategies in these protected areas, which is currently greatly lacking, in particular, from the southern Western Ghats. In this context, the present study has been framed with the aims of assessing plant diversity, structure and regeneration potential in tropical forests of Kanyakumari Wildlife Sanctuary (KWLS), Western Ghats, India. 2. Materials and methods 2.1. Study area The Western Ghats is well-known for its structural and floristic diversity. The study was carried out in Kanyakumari Wildlife Sanctuary (8°03′–8°35′ N and 77°05′–77°36′ E), which is a part of Agasthyamalai Biosphere Reserve and is located at the southern tip of Western Ghats. It shares its boundaries with Neyyar Wildlife Sanctuary and KalakkadMundanthurai Tiger Reserve. The forests of Kanyakumari district are virgin and about 75 million years old. According to Forest Survey of India’s report, Kanyakumari has 994 km2 of forest cover, which is subdivided into 146 km2 of very dense forest, 554 km2 of moderately dense forest and 294 km2 of open forest [25]. This wildlife sanctuary was declared in 2002 with an original extent of 45777.57 ha. However, in 2007, its boundaries have been redefined to an area of 40239.55 ha. The study area experiences a tropical climate and is greatly influenced by south-west and north-east monsoon winds, with a rainy season

during October and November and a dry season from March to May. The sanctuary gets an average annual rainfall of 1369.5 mm with 79 rainy days. The mean annual maximum and minimum temperatures are 32.7°C and 19°C respectively [26]. The entire sanctuary is hilly, steep and rugged, and the elevation ranges from 50 to 1650 m a.s.l. However, the highest point is located at 1829.4 m at the tri-junction of Mahendragiri, Veerapuli and Kalakad Reserve Forests. The soil is mostly red loamy, but at some places sandy or clayey as well. The sanctuary is traversed by many rivers that run south-westerly such as Manimuttar, Pachiar, Kodayar, etc. Due to variations in temperature, rainfall, elevation, terrain and edaphic characteristics, different vegetation types occur in this sanctuary. For the present study, seven sites were selected from distinct forest ranges across different locations in KWLS (Pechipaarai, Kutriyar, Sangilippaalam, Paayaasappaarai; Table 1; Fig. 1). On the basis of similarity (using species richness and density as input variables) among the study sites (Fig. 2), it has been found that the study sites I and II conform to the tropical dry deciduous forest type (henceforth abbreviated as TDD I and TDD II), sites III and IV belong to the tropical semievergreen forest type (henceforth abbreviated as TSE I and TSE II), and sites V, VI and VII were of tropical evergreen forest type (henceforth abbreviated as TEF I, TEF II and TEF III). 2.2. Field methods The phytosociological analysis was done following Misra [27] by laying ten square plots of 20 m × 20 m in each study site to document the woody plant diversity. These plots were further sub-gridded into four 10 m × 10 m grids for ease in sampling. The plants encountered in 10 m × 10 m grids were then classified into six categories: tree seedlings (tree species individuals with b1 m height), tree saplings (tree individuals with b3 cm diameter at breast height (DBH)), tree juveniles (tree individuals with 3–9.9 cm DBH), tree adults (tree individuals with ≥ 10 cm DBH), climbers (climbing individuals with girth b2.5 cm at 1.3 m from the rooting point), woody climbers (climbing individuals with girth ≥2.5 cm at 1.3 m from the rooting point). In each of these grids, four sub-quadrats of 1 m × 1 m were laid to study the understorey vegetation (herbs and shrubs). Density, basal area, frequency, importance value index (IVI) were calculated by following Misra [27] and diameter class-wise distribution were calculated for each study site. The study period was from 2015 to 2017. The woody plants were enumerated during March-May, while the understorey was documented during its peak growing period, November. The plant species were identified using Flora of the Presidency of Madras [28], A field key to the trees and lianas of the evergreen forests of the Western Ghats (India) [29] and Biodiversity Informatics and co-Operation in Taxonomy for Interactive shared Knowledge base software [30]. The assignment of a species to the family was done as per Angiosperm Phylogeny Group IV (APG IV) system of classification [31]. The plant names and the corresponding author citations were given following GRIN (Germplasm Resources Information Network) Taxonomy and Global Biodiversity Information Facility [32]. 2.3. Data analysis Diversity indices (Shannon’s, Dominance, Evenness and Fisher’s alpha) and cluster analysis were computed using the Past 3.1 program (version 3.1; Øyvind Hammer, Natural History Museum, University of Oslo). The abundance to frequency (A/F) ratio for each species was determined as per Whitford [33]. The ratio indicates the distribution pattern as follows: “regular (b0.025), random (0.025–0.050) and contiguous (N0.050)”. Beta diversity was calculated using R [34]. The regeneration potential of every tree species in each study site was assessed following Pradhan et al. [14] and Shankar [35]: (a) ‘good’, if saplings N juveniles N adults; (b) ‘fair’, if saplings N juveniles ≤ adults; (c) ‘poor’ if a species survives only in juvenile stage, but not as saplings

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

3

Table 1 Information on the seven study sites in KWLS, Western Ghats, India. Variable

Site-I (TDD I)

Site-II (TDD II)

Site-III (TSE I)

Site-IV (TSE II)

Site-V (TEF I)

Site-VI (TEF II)

Site-VII (TEF III)

Latitude Longitude Elevation (m asl) Annual precipitation (mm) Annual mean temperature (°C) Maximum temperature (°C) Minimum temperature (°C)

8° 26' 58.123" N 77° 19' 13.317" E 115–229 1183 27.4 32.7 22.1

8° 31' 25.372" N 77° 17' 28.348" E 317–392 1310 25.6 31.1 20.4

8° 32' 22.400" N 77° 18' 46.856" E 532–630 1347 24.8 30.2 19.5

8° 31' 28.522" N 77° 17' 19.202" E 306–431 1310 25.6 31.1 20.4

8° 32' 10.207" N 77° 19' 15.784" E 521–677 1379 24.3 29.8 19

8° 32' 23.766" N 77° 19' 03.610" E 508–655 1379 24.3 29.8 19

8° 32' 26.574" N 77° 18' 38.541" E 477–544 1347 24.8 30.2 19.5

(although juveniles may be less or more than or equal to adults); (d) ‘none’, if a species is represented by only adults and has no saplings and juveniles; and (e) ‘new’, if a species has no adults and has only saplings and/or juveniles. Regeneration status is finally expressed as a percentage by dividing the number of tree species in each category of regeneration by the total number of tree species in each study site.

3. Results 3.1. Plant species diversity A total of 267 species were enumerated from 205 genera and 70 families from the seven study sites of KWLS, Western Ghats, India (Table 2). Of these, 151 species (56.5%) occur as trees, 41 species are vines (climbers and woody climbers; 15.3%) and 75 species (28.2%) constitute the understorey (shrubs and herbs). Tree species richness ranged from 24 (TDD II) to 76 (TEF III). The overall species richness ranged from 29 (TDD II) to 139 (TDD I). The highest tree species richness was recorded in TEF III, while TDD I was the richest site, with respect to the climber, woody climber and understorey categories. The species composition varied markedly amongst the seven study sites. Aporosa cardiosperma and Terminalia paniculata were the only two species common in all the study sites. A total of 64 tree species, 28 vine species (climbers and woody climbers) and 58 understorey species were documented from only one of the study sites. Of all the documented species, 66 species were endemic to India and of these, 46 species (17.2% of the total flora documented) were endemic to Western Ghats (Table A.1). Besides, a total of 18 species are exotic and of these, seven species are invasive. With respect to threatened categories, two species, viz. Nothopegia aureofulva and Vateria indica are critically endangered, six species (Cycas circinalis, Dipterocarpus indicus, Dysoxylum malabaricum, Hopea parviflora, Kingiodendron pinnatum and Memecylon subramanii) are endangered and nine species (Aglaia elaeagnoidea, Dalbergia latifolia, Diospyros paniculata, Dysoxylum ficiforme, Hydnocarpus pentandrus, Mallotus atrovirens, Orophea uniflora, Psydrax dicoccos and Pterocarpus marsupium) are vulnerable to extinction in the future. Besides, four species (Aglaia edulis, Aglaia simplicifolia, Dimocarpus longan and Gluta travancorica) are near threatened and nine species (Alstonia scholaris, Calophyllum inophyllum, Commelina benghalensis, Dichrostachys cinerea, Knema attenuata, Spatholobus parviflorus, Tetrameles nudiflora, Vanda tessellata and Ziziphus jujuba) are of least concern, as per The IUCN Red List of Threatened Species. A total of 39 tree adult species, 5 climber species, 8 woody climber species and 7 understorey species were of rare occurrence (≤2 individuals in the total studied area (2.8 ha)). Amongst the diversity indices, Shannon’s index ranged from 1.99 (TDD II) to 2.93 (TEF III), 0 (TDD II, TEFs II and III) to 2.43 (TDD I), 0 (TSE I) to 1.64 (TEF III) and 0.001 (TDD II) to 2.30 (TDD I) for tree adults, climbers, woody climbers and understorey respectively (Table 2). The values of the dominance index ranged 0.09 (TDD I) – 0.3 (TEF I) for adult trees, 0.21 (TEF III) –1 (TSE I) for woody climbers and 0.19 (TSE II) – 0.99 (TDD II) for understorey. The highest values of Fisher’s alpha were observed in TEF III (15.7), TSE II (3.92) and TDD I (5.89) for tree

adults, woody climbers and understorey respectively and the lowest values were in TDD II (2.59), TSE I (0) and TDD II (0.22). The total beta diversity among the seven sites was 0.396 and 0.334 based on Jaccard (J) and Sørensen (S) dissimilarities respectively (Table 3). In this study, replacement accounted for around 59% of the total beta diversity measured, compared with ~41% for richness difference. The principal coordinate (PCoA) ordinations of the Podanifamily replacement (ReplJ and ReplS) and richness difference matrices (RichDiffJ and RichDiffS) for species presence-absence data are presented in Fig. 3. The ordinations of replacement matrices were similar, while those of richness difference matrices were distinct. Of these matrices, only the RichDiffS matrix had the mathematical property of being Euclidean. The square-root transformation resulted in RichDiffJ becoming Euclidean. However, both ReplJ and ReplS matrices remained nonEuclidean even after square-root transformation. 3.2. Family composition Of the 70 families recorded, Euphorbiaceae (14 genera and 24 species), Rubiaceae (16 genera and 23 species) and Fabaceae (16 genera and 17 species) were the most speciose families (Table A.2). A total of eight families were represented in all the seven study sites. The number of species in a family varied from 1 to 24. Twenty-seven families were monospecific and thirteen families were bispecific. In terms of density, Poaceae (2,24,500 individuals/2.8 ha), Fabaceae (63,664 individuals/ 2.8 ha) and Zingiberaceae (15,236 individuals/2.8 ha) were the dominant families. Only one family, viz. Loranthaceae was represented by a singleton (Dendrophthoe falcata). 3.3. Density In toto, 18,790 tree individuals were enumerated from the studied 2.8 ha (Table A.1). The tree adult density ranged from 370 individuals/ ha (TDD I) to 900 individuals/ha (TEF I), with a mean of 676 ± 83.6 individuals/ha. The climber and woody climber densities were highest in TDD I (49,768 and 358 individuals/ha respectively). The climbers were absent or least (3 individuals/ha) in the evergreen forest sites (TEFs IIII), while woody climber representation was fairly high in these sites. The understorey density was highest in TDD II (3,95,023 individuals/ ha) and lowest in TEF I (413 individuals/ha). Among the tree seedling population, Vitex altissima and Cipadessa baccifera in TDD I, Cipadessa baccifera and Tabernaemontana gamblei in TSE I were the abundant species. There is no occurrence of tree seedlings in the sites TDD II, TSE II, TEFs I-III. In the case of tree saplings, Helicteres isora and Melia dubia in TDD I, Aporosa cardiosperma and Helicteres isora in TDD II, Aporosa cardiosperma and Chionanthus ramiflorus in TSE I, Ficus tinctoria and Xanthophyllum flavescens in TSE II, Mallotus philippensis and Holigarna arnottiana in TEF I, Diospyros affinis and Aporosa cardiosperma in TEF II and Agrostisachys borneensis and Hopea parviflora in TEF III were the predominant species. The species most prevalent among the tree juveniles are Helicteres isora and Terminalia paniculata in TDD I, Terminalia paniculata and Aporosa cardiosperma in TDD II, Aporosa cardiosperma and Croton malabaricus in TSE I, Ixora pavetta and Xanthophyllum flavescens in TSE II, Xanthophyllum flavescens and Hopea parviflora in

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

4

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

Fig. 1. Location of the seven study sites in KWLS, Western Ghats, India.

TEF I, Diospyros affinis followed by Aporosa cardiosperma and Hopea parviflora in TEF II, and Diospyros affinis and Chionanthus ramiflorus in TEF III. Amongst the tree adults, Terminalia paniculata and Pterocarpus marsupium in TDD I, Terminalia paniculata and Dillenia pentagyna in TDD II, Pterocarpus marsupium and Ficus beddomei in TSE I, Aporosa cardiosperma and Ixora pavetta in TSE II, Hopea parviflora and Vitex altissima in TEF I, Hopea parviflora followed by Aporosa cardiosperma

and Diospyros affinis in TEF II, and Hopea parviflora and Diospyros affinis in TEF III were the abundant species. With respect to climbers, Hemidesmus indicus and Rhamnus oenoplia were more rampant in TDD I, while the climbers were less in TDD II. Thunbergia fragrans and Smilax zeylanica were the dominant species in TSE I, while again, the climbers were very minimal with the occasional occurrence of Salacia chinensis in TSE II. The climbers were absent in

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

Fig. 2. Bray-Curtis cluster dendrogram (single-linkage) for tree adults in the seven study sites.

TEF I and least in TEF II and TEF III. The common woody climbers in TDD I were Calycopteris floribunda and Rhamnus oenoplia, while TDD II had the occurrence of only two species, viz., Rhamnus oenoplia and Aganosma cymosa. On the other hand, TSE I just had only one individual of Rhamnus oenoplia, while Salacia chinensis and Combretum albidum were prevalent in TSE II. Spatholobus parviflorus was the most dominant species in TEFs I-III, followed by Artabotrys zeylanicus in TEF I, Tetrastigma nilagiricum in TEF II, and Tetrastigma nilagiricum and Dioscorea alata in TEF III. With regards to understorey vegetation, TDD I was mainly dominated by dicot species, although, Themeda cymbaria and Stylosanthes fruticosa showed more abundance, while the herbaceous layer of TDD II was monodominant with the monocot, Themeda cymbaria. Stylosanthes fruticosa and Cymbopogon citratus in TSE I, Sauropus androgynus and Ixora alba in TSE II, Psychotria glandulosa and Benkara malabarica in TEFs I and II, and Psychotria nudiflora and Glycosmis mauritiana in TEF III were the much prevalent understorey species. 3.4. Stand basal area and IVI The basal area of tree adults ranged from 24.2 (TDD I) to 75.3 (TEF III) m2/ha (Table 2). The basal area values of woody climbers were highest in TDD I and TEF I (0.7 m2/ha each) and lowest in TSE I (0.0002 m2/ha). The site TSE I had the highest basal area in understorey (3.4 m2/ha), while TSE II and TEF III had the least (0.3 m2/ha each). Greater basal area values were observed in Terminalia paniculata and Terminalia elliptica in TDDs I and II, Pterocarpus marsupium and Ficus beddomei in TSE I, Aporosa cardiosperma and Terminalia paniculata in TSE II, Hopea parviflora in all the three evergreen forest sites (TEFs IIII), followed by Vitex altissima (TEF I), Tetrameles nudiflora (TEF II) and Vateria indica (TEF III; Table A.1). Calycopteris floribunda and Rhamnus oenoplia in TDD I had high basal area values amongst woody climbers, while in TSE II, Combretum albidum and Salacia chinensis had the maximum basal areas. However, Spatholobus parviflorus had the largest basal area in the tropical evergreen forest sites (TEFs I and II), followed by Artabotrys zeylanicus in TEF I, Tetrastigma nilagiricum in TEF II and Tetrastigma nilagiricum followed by Spatholobus parviflorus in TEF III. With respect to tree saplings, Helicteres isora and Melia dubia in TDD I and Aporosa cardiosperma and Helicteres isora in TDD II; Aporosa

5

cardiosperma and Chionanthus ramiflorus in TSE I, and Ficus tinctoria and Xanthophyllum flavescens in TSE II; Mallotus philippensis and Aporosa cardiosperma in TEF I, Diospyros affinis and Aporosa cardiosperma in TEF II and Agrostisachys borneensis and Hopea parviflora in TEF III had the high IVI values (Table A.1). With reference to tree juvenile population, the species with the maximal IVI values were Helicteres isora and Terminalia paniculata in TDD I, Terminalia paniculata and Aporosa cardiosperma in TDD II, Aporosa cardiosperma and Croton malabaricus in TSE I, Ixora pavetta and Xanthophyllum flavescens in TSE II, Hopea parviflora and Xanthophyllum flavescens in TEF I, and Diospyros affinis and Hopea parviflora in TEFs II and III. In the case of tree adults, Terminalia paniculata and Pterocarpus marsupium in TDD I, Terminalia paniculata and Terminalia elliptica in TDD II, Pterocarpus marsupium and Ficus beddomei in TSE I, Aporosa cardiosperma and Ixora pavetta in TSE II, Hopea parviflora and Vitex altissima in TEFs I and III, and Hopea parviflora and Aporosa cardiosperma in TEF II had large IVI values. Regarding climbers, Rhamnus oenoplia had the largest IVI value in the tropical dry deciduous sites TDDs I and II. Calamus rotang and Piper hookeri in TSE I, Salacia chinensis and Piper hookeri in TSE II had the maximum IVI values in their respective sites. Climbers were absent in the evergreen site TEF I, while Rivea hypocrateriformis (TEF II) and Tetracera akara (TEF III) had the largest IVI values. In the woody climber category, Calycopteris floribunda and Rhamnus oenoplia in TDD I had the maximum IVI values. The site TSE I had only one climber species, viz. Rhamnus oenoplia, while Salacia chinensis and Combretum albidum had the most IVI values in TSE II. Spatholobus parviflorus had the largest IVI value in TEFs I-III. Pertaining to understorey, Lantana camara and Themeda cymbaria in TDD I, and Themeda cymbaria in TDD II; Cymbopogon citratus and Psychotria glandulosa in TSE I, and Sauropus androgynus and Ixora alba in TSE II; Benkara malabarica and Psychotria glandulosa in TEFs I and II, and Psychotria nudiflora and Glycosmis mauritiana in TEF III had high IVI values. 3.5. Species distribution pattern Contiguousness was the principal pattern of distribution in all the life-forms. This is followed by the random distribution that is exhibited to a considerable extent among tree saplings, juveniles and adults. Regular distribution was very rarely observed. 3.6. Size class distribution The diameter class-wise distribution of tree juveniles and tree adults showed the characteristic ‘reverse J-shaped’ curve in all the study sites (Fig. 4). Both abundance and species richness declined with an increase in diameter class. In all the study sites, small-sized trees (10–29.9 cm DBH) occupied a greater proportion (68–81.3%) than other size classes. Medium-sized trees (30–59.9 cm DBH) contributed 14.8–29% to density, whereas large-sized trees (60-100 cm DBH) comprised of only 1.7–8.9%. 3.7. Population structure and regeneration potential Among tree species, 12 species occur in all the four stages of development, viz. seedlings, saplings, juveniles and adults. Maesa indica is the only tree species that is not recorded as an adult but found in all the other three stages. A majority of 72 species occur as saplings, juveniles and adults, while 20 species occur as saplings and juveniles, 7 species as saplings and adults, and 5 species as juveniles and adults. A total of 16 species occur only as saplings, six species only as juveniles and 12 species only as adults. A total of 13 species in TDD I, nine species in TDD II, 14 species in TSE I, 21 species in TSE II, 25 species in TEF I, 28 species in TEF II, 32 species in TEF III occur in three stages, viz. saplings, juveniles and adults. A total of two species in TDD I, four species in TSE I, two species in TSE II, three species each in TEFs I and II, six species each in TEF III occur as juveniles and adults. Also, seven species in TDD I, five species in TDD II, 11 species in TSE I, six species in TSE II, 15 species in TEF I, four

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

6

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

Table 2 Consolidated details of phytosociological analyses of the seven study sites. Parameter

TDD I

TDD II

TSE I

TSE II

TEF I

TEF II

TEF III

Total

Species richness Tree seedlings (b1 m height) Tree saplings (b3cm DBH) Tree juveniles (3–9.9 cm DBH) Tree adults (≥10 cm DBH) Total tree species Climbers (b2.5 cm girth) Woody climbers (≥2.5 cm girth) Total climber species Understorey Total No. of families

12 37 35 32 55 22 10 29 55 139 51

23 14 11 24 1 2 2 3 29 20

5 33 38 24 45 5 1 6 11 62 36

53 32 40 67 4 9 9 12 88 39

43 46 35 56 3 3 8 66 33

36 37 34 44 1 3 3 4 51 31

57 55 49 76 1 6 7 10 93 37

13 128 116 108 151 28 21 41 75 267 70

Density (No./ha) Tree seedlings Tree saplings Tree juveniles Tree adults Climbers Woody climbers Understorey

13500 770 828 370 49768 358 167838

948 253 448 5 30 395023

10000 1470 978 525 23750 3 275150

3468 448 775 38 88 1665

998 1218 900 198 413

1400 1133 863 3 88 545

3558 1280 850 3 110 553

11750 1802 877 676 12261 125 120170

Basal area (m2/ha) Tree seedlings Tree saplings Tree juveniles Tree adults Climbers Woody climbers Understorey

0.14 0.23 2.09 24.23 0.35 0.70 2.25

0.19 0.79 30.83 0.00 0.01 2.80

0.08 0.43 2.49 35.04 4.24 0.0002 3.37

0.31 1.53 45.77 0.00 0.20 0.33

0.29 3.34 49.05 0.70 0.44

0.36 3.00 64.64 0.00 0.37 0.60

0.74 3.17 75.34 0.00 0.20 0.29

0.11 0.36 2.34 46.41 0.77 0.31 1.44

Shannon index Tree seedlings Tree saplings Tree juveniles Tree adults Climbers Woody climbers Understorey

1.99 2.74 2.51 2.86 2.43 0.75 2.30

2.21 2.15 1.99 0 0.56 0.001

1.42 2.66 2.74 2.35 1.53 0 1.28

3.03 2.35 2.68 0.72 1.55 1.89

2.77 3.23 2.08 0.64 1.13

2.96 3.02 2.51 0 0.58 0.88

3.04 3.43 2.93 0 1.64 1.79

1.71 2.77 2.78 2.49 0.78 0.82 1.32

Dominance index Tree seedlings Tree saplings Tree juveniles Tree adults Climbers Woody climbers Understorey

0.20 0.12 0.16 0.09 0.12 0.70 0.20

0.19 0.16 0.16 1 0.63 0.99

0.28 0.12 0.12 0.15 0.23 1 0.33

0.08 0.21 0.13 0.65 0.32 0.19

0.14 0.06 0.30 0.65 0.43

0.08 0.07 0.17 1 0.67 0.47

0.08 0.05 0.15 1 0.21 0.22

0.24 0.12 0.12 0.16 0.67 0.60 0.40

Evenness index Tree seedlings Tree saplings Tree juveniles Tree adults Climbers Woody climbers Understorey

0.61 0.42 0.35 0.54 0.52 0.21 0.18

0.40 0.61 0.67 1 0.88 0.33

0.83 0.43 0.41 0.44 0.92 1 0.33

0.39 0.33 0.37 0.51 0.52 0.55

0.37 0.55 0.23 0.63 0.39

0.53 0.55 0.36 1 0.60 0.60

0.36 0.56 0.38 1 0.86 0.60

0.72 0.41 0.48 0.43 0.83 0.67 0.43

Fisher's alpha Tree seedlings Tree saplings Tree juveniles Tree adults Climbers Woody climbers Understorey

1.46 10.98 9.89 12.56 2.44 2.45 5.89

5.39 4.41 2.59 0.80 0.69 0.22

0.56 7.56 10.4 6.99 0.51 0 0.94

10.92 11.35 12.23 1.79 3.92 2.08

12.23 12.46 9.58 0.62 1.76

8.59 9.53 9.36 0 0.79 0.70

11.89 15.63 15.7 0 1.88 2.16

1.01 9.65 10.52 9.86 0.92 1.48 1.96

species in TEF II and 13 species in TEF III occur as saplings and juveniles, whereas two species in TDD I, one species in TDD II, nine species in TSE II and two species in TEF III were recorded in sapling and adult stages. Of the enumerated flora, two species in TDD I occur as only seedlings; seven species in TDD I, eight species in TDD II, four species in TSE I, 17 species in TSE II, three species in TEF I, four species in TEF II and 10 species in TEF III occur as only saplings; five species in TDD I, four species in TSE I, three species each in TSE II and TEF

I, two species in TEF II and four species in TEF III occur as only juveniles and seven species each in TDD I and TEF I, one species in TDD II, three species each in TSE I and TEF II and nine species each in TSE II and TEF III occur as only adults. The assessment of regeneration potential indicated that most of the species were newly recruited (34.2–54.2%) in all the sites, except TEF II (Table 4). A substantial proportion of the species in the site TEF II displayed poor regeneration (38.1%). A considerable number of species

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx Table 3 Decomposition of beta diversity among the seven study sites for presence-absence data (Podani family) Parameter

Jaccard

Sørensen

BDTotal BDmax = 0.5 ReplTotal / BDTotal RichDiffTotal / BDTotal

0.396 79% of BDmax 0.591 0.408

0.334 66% of BDmax 0.585 0.414

showing poor regeneration was also recorded from the study sites TEF I, TDD I and TEF III. Good regeneration of tree species was observed to be high in the sites TEF II, followed by TEF III and TSE I. Fair regeneration was largely seen in the sites TSE II and TDD II. About 4.2–15.1% of the species exhibited no regeneration. It is worth mentioning that different species showed different patterns of regeneration potential in different sites. In general, the dominant adult tree species had adequate representation in both sapling and juvenile stages of development and therefore had fair or good regeneration potential in their respective sites of dominance. However, 12 species occur as only adults, with no younger stage of development and among these are five species (Cycas circinalis, Dipterocarpus indicus, Gluta travancorica, Litsea salicifolia and Palaquium ellipticum) that are endemic to only the Western Ghats, implying that these endemic species displayed no regeneration. 4. Discussion In the present inventory, 267 species were recorded from 205 genera and 70 families from the seven study sites of KWLS, which is a

7

part of the Agasthyamalai Biosphere Reserve, Western Ghats, India. Agasthyamalai hills are known to shelter N 2000 of the ~4000 known plant species of the Western Ghats [36,37] and the flora documented in this study represents 13.3% of Agasthyamalai flora. Very often, the differences in plot dimensions, plot shapes, girth classes, locations make it difficult to compare the results of our study with other studies [4], although a comparison was made to understand the differences in the patterns of structure and species composition with other forests worldwide (Supplementary table 1). The overall species richness observed (267 species/2.8 ha) is comparable with those reported from Caatinga Biogeographic Domain, Brazil (258 species/4.32 ha [38]), Wadi Turbah Zahran, Saudi Arabia (266 species/3.6 ha [39]); lower than Sulawesi mountains, Indonesia (331 species/3.12 ha [40]) and higher than Kodayar (228 species/2 ha [37]), Veerapuli and Kalamalai reserve forests (244 species/0.9 ha [41]), Nelliampathy hills, Kerala of Western Ghats (152 species/3 ha [42]). Out of all the 267 species documented, 151 are tree species, of which, 108 species occur as tree adults (DBH ≥ 10 cm), which very well fits in the range obtained in tropical forests (individuals with DBH ≥ 10 cm) viz. 30 (Nelliampathy, Western Ghats [43]) to 307 species (Amazonian Ecuador [44]). On a broader scale, this represents only 0.20–0.27% of the ~40,000–N53,000 estimated tree species across the pantropical part of the globe and 0.9% of the pantropical species richness (11,371 tree species) and 1.9% of the Indo-Pacific region (5,672 tree species) as per Slik et al. [18]. The plant species composition and forest structure varied greatly amongst the different study sites. The tropical dry deciduous forest site, TDD I had the overall highest species richness with 139 species. However, when considering only tree species richness, the evergreen forest site, TEF III was the most speciose, harbouring 76 tree species. This site is located alongside a perennial

Fig. 3. Principal coordinate (PCoA) ordinations of the replacement and richness difference indices based on Jaccard and Sørensen dissimilarities (Podani family): (a) square-root transformed replacement (ReplJ) (b) square-root transformed richness difference (RichDiffJ) (c) square-root transformed replacement (ReplS) and (d) untransformed richness difference (RichDiffS).

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

40

80

35

70

30

250

25

200

20

150

15

100

10

50

5 4-5

5-6

6-7

7-8

8-9

9-10

20

60 50

15

40 10

30 20

5

10 0

0

0

Diameter class (cm)

Diameter class (cm)

20

300 250

15

200 10

150 100

5

50 0

<3

3-4

4-5

5-6

6-7

7-8

8-9

9-10

80

25

70

Stems/4000 m2

TDD II

350

Stems/4000 m2

Species richness

Species richness/4000 m2

Stems per 4000 square m

400

35 30 25

400

20

300

15

200

10

100

5 <3

3-4

4-5

5-6

6-7

40

7-8

8-9

9-10

Stems per 4000 square m

30

800 600

20

400

10

200 5-6

6-7

Diameter class (cm)

7-8

8-9

9-10

0

Stems/4000 m2

Stems/4000 m2

40

4-5

Species richness

TSE I

80

20 15

60 10 40 5 0

200

60

1000

3-4

0

0

0

50

<3

2

20

Species richness

1200

0

4

20

Diameter class (cm)

TSE II

1400

6

30

100

Species richness/4000 m2

Stems per 4000 square m

8

50

Diameter class (cm)

1600

12 10

0

Stems/4000 m2

Stems/4000 m2

Species richness

500

0

TDD II

Diameter class (cm)

TSE I

600

Species richness

10

0

Species richness/4000 m2

Stems per 4000 square m

Stems per 4000 square m

60

Diameter class (cm)

700

25

Species richness/4000 m2

3-4

TDD I

Species richness/4000 m2

<3

Species richness

Stems per 4000 square m

Species richness

TSE II

150

35 30 25 20

100

15 10

50

5

Species richness/4000 m2

0

Stems per 4000 square m

Species richness/4000 m2

TDD I

300

Stems/4000 m2

Species richness

Stems/4000 m2

Stems per 4000 square m

350

Species richness/4000 m2

8

0

0

Diameter class (cm)

Fig. 4. Diameter class-wise distribution of juveniles and adults in the seven study sites.

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

350 300 250 200 150 100 50 3-4

4-5

5-6

6-7

7-8

8-9

9-10

20

100

15

30

500

25

400

20

300

15

200

10

100

5 5-6

6-7

7-8

8-9

Stems/4000 m2

Stems/4000 m2

Species richness/4000 m2

40

600

4-5

0

9-10

200 180 160 140 120 100 80 60 40 20 0

Diameter class (cm)

50

1200

40

1000 800

30

600

20

400

10

200 0

<3

3-4

4-5

5-6

6-7

200

60

7-8

8-9

9-10

0

Stems/4000 m2

Stems/4000 m2

Species richness

TEF III

1400

Species richness

TEF II

35 30 25 20 15 10 5 0

Diameter class (cm)

Species richness/4000 m2

Stems per 4000 square m 1600

0

Stems per 4000 square m 35

3-4

5

0

Species richness

TEF II

<3

10

50

Diameter class (cm)

Stems per 4000 square m

0

25

150

Diameter class (cm)

700

30

Species richness/4000 m2

<3

35

TEF I

Stems per 4000 square m

Species richness

TEF III

50 40

150

30

100

20

50

10

0

0

Species richness/4000 m2

0

Species richness

200

Stems/4000 m2

Stems/4000 m2

400

Stems per 4000 square m 250

50 45 40 35 30 25 20 15 10 5 0

Species richness/4000 m2

Species richness

TEF I

Species richness/4000 m2

Stems per 4000 square m 450

9

Diameter class (cm)

Diameter class (cm) Fig. 4 (continued).

stream. High species richness in the riverine vegetation was also observed by Tripathi and Singh [45] and Sunil et al. [46] and according to the former, this high species richness is possibly due to the site being more nutrient-rich, with greater soil moisture alongside favourable micro-climatic conditions. Woody climbers play a very important role in maintaining biodiversity and regulating ecosystem functions. In the present study, 21 species of woody climbers were documented, which constitutes only 13.5% of the total woody species (juveniles + adults + woody climbers). Schnitzer and Bongers [47] and Parthasarathy et al. [48] reported that the tropical forests harbour 25% of the woody climber diversity of all woody plant diversity. However, the value reported in the present study roughly equals only half of their findings. Also, the cumulative

woody climber species richness values observed for the tropical dry deciduous forest sites (TDD I and TDD II, 11 species/0.8 ha), tropical semievergreen forest sites (TSE I and TSE II, 9 species/0.8 ha) and tropical evergreen forest sites (8 species/1.2 ha) were very much lower than those observed for seasonal dry forest (70 species/31.5 ha), semi-evergreen forest (126 species/21.5 ha) and wet evergreen forest (126 species/ 38.4 ha) respectively by Parthasarathy et al. [48]. One obvious reason could be because of the vast differences in the sampled area. The site TDD I was observed to have the highest understorey plant diversity. Overstorey tree species influence the composition of understorey species by altering resource levels, such as light penetration, water availability, soil properties, litter composition and depth [49,50]. The presence of pioneer species such as Macaranga and Mallotus

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

10

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

Table 4 Regeneration status of tree species(%) in the seven study sites Status

TDD I

TDD II

TSE I

TSE II

TEF I

TEF II

TEF III

Good regeneration Fair regeneration Poor regeneration Newly recruited No regeneration Total

7.5 17.0 22.6 37.7 15.1 100

4.2 25.0 12.5 54.2 4.2 100

20.0 11.1 15.6 46.7 6.7 100

9.0 34.3 3.0 40.3 13.4 100

7.1 8.9 33.9 37.5 12.5 100

28.6 11.9 38.1 14.3 7.1 100

21.1 11.8 21.1 34.2 11.8 100

species indicates the existence of disturbance in the forest, which might, in turn, indicate the presence of canopy openings [43,51]. Although the disturbances were not given primary focus in this study, based on the occurrence of the above-said species and visual observations, the site TDD I could be perceived as a relatively disturbed site. It is also very important to note that seven invasive species, viz. Chromolaena odorata (3.3%), Lantana camara (1.8%), Clidemia hirta, Mesosphaerum suaveolens, Solanum viarum, Synedrella nodiflora (0.1% each) and Mimosa diplotricha (0.001%) constitute the understorey of TDD I and they are absent in all other natural forest sites. Although their percentages are quite low as of now, it might not be very long before they predominate the entire site. Chromolaena odorata is an indicator of past disturbance [37,41]. Invasive species successfully colonize disturbed ecosystems as they are easy targets to plant invasion as per disturbance hypothesis [52]. Presence of invasive species in TDD I reasserts that this is a relatively disturbed site than all the other natural forest sites. Nevertheless, canopy openness is a feature of a deciduous forest when leaf fall occurs during the dry season [50]. The site TDD I, when compared with the semievergreen and evergreen forest sites had a relatively open canopy and was disturbed which might have favoured the establishment of diverse understorey. Water absorption and transpiration rates of a leafless tree (a deciduous tree) are much lesser than that of an evergreen tree and the lack of a covered canopy results in more water seepage into the soil. Further, the leaf fall leads to litter accumulation, which in turn, maintains soil humidity [50,53]. Thus, a deciduous overstorey facilitates the establishment of a diverse understorey beneath it by altering environmental conditions. On the other hand, in TDD II, the diversity of understorey is very low, but the density is much higher, as it is monodominant with Themeda cymbaria. Based on local enquiry, this region was subjected to the recurrence of annual fires before this forest was declared as a wildlife sanctuary in 2002. At the same time, ground fires occur frequently during summer in this site, which could be the reason for its low diversity. Even though TDD II contains more canopy openings, the same provides a conducive environment for luxuriant booming of this perennial, tall grass species. The tropical semievergreen forest sites (TSEs I and II) had a relatively lesser diverse understorey than the tropical dry deciduous forest site (TDD I), but greater than the tropical evergreen forest sites (TEFs I-III). This could be because these sites might be undergoing a gradual transition from deciduous to evergreen forest type. Also, the understorey diversity is very less in the tropical evergreen forest sites, especially TEFs I and II which might be attributed to the suppression of understorey growth by the robust and fast growth of tree saplings and tree juveniles that outcompete the former for light and other resources, also leading to canopy closure. Western Ghats is the only hotspot with the highest human population density, yet with only 9% of the forests under protected area network, which poses a great concern [54]. Agasthyamalai Biosphere Reserve harbours 7.5% of endemics [55,56]. In the present study, 66 species were found to be endemic to India and of these, 46 species were endemic to only Western Ghats. This is within the range 2-55% of endemics reported for Western Ghats by Utkarsh et al. [57]. Of these 46 endemics, Gluta travancorica is a localized endemic of only Agasthyamalai region [56]. According to Pascal [58], regions of high

endemism and species richness occur where there is high rainfall and a short dry season. The studied forest sites are located in the tip of the peninsular India and hence receive copious rainfall in both the monsoon seasons, which could have therefore supported rich diversity with more endemics. Tropical forests are usually very diverse, which is often indicated by the diversity indices. The Shannon index depicts the richness of an ecosystem which in the present study varied between 1.99 (TDD II) and 2.93 (TEF III) for tree adults. The Shannon index normally ranges from 1.5 to 3.5 and rarely surpasses 4.5 [59] and the observed values conform to this range and also the range prescribed for tropical forests, viz., 0.83– 4.1 [60]. The highest value observed in TEF III is backed by the finding of Joseph et al. [61] who reasoned that the aggregation and co-existence of many species along the riverine forest is possibly due to water availability, coupled with nutrient enrichment. The dominance index is a measure of the abundance of a species in the sample [62], which in this study was from 0.09 (TDD I) to 0.3 (TEF I) for tree adult species. The average value of dominance index for tropical forests is 0.06 and it ranges between 0.21 and 0.92 for Indian tropical forests [63,64]. The values obtained in the present study slightly deviates the lower limit of this range. It is known that lower the dominance index, higher is the diversity of the ecosystem. Beta diversity, which refers to variation in species composition among sites is partitioned into two main components, namely replacement and richness difference [34]. Replacement refers to turnover, where species tend to replace each other along spatial or environmental gradients. On the other hand, richness difference refers to the fact that one community usually has a greater number of species than another [34]. In the present study, the total beta diversity values were 0.396 and 0.334 based on Jaccard (J) and Sørensen (S) dissimilarities respectively (Podani family) of species presence-absence data, while replacement and richness difference values were 0.234 and 0.162, and 0.196 and 0.138 respectively. Presence-absence data are preferred when the plant communities among the sites vary in species composition and partly harbour different species [34]. The PCoA ordinations showed that there exists a variation in species composition among the study sites. Similar results were reported by Sullivan et al. [16] in Asian tropical forests, where the tree communities were the least similar. In this study, a total of 70 families were documented from all the seven study sites and of these, Euphorbiaceae, Rubiaceae and Fabaceae were the most speciose families. These families were listed to be dominant in other studies as well: Ganesh et al. [36]; Ayyappan and Parthasarathy [65]; Panda et al. [66]; Sahoo et al. [67] and Sosef et al. [68]. Out of the 20 most speciose families in tropical forests of Africa [68], 18 were represented in this study, which cumulatively contributes to 49.1% of the total species richness. Gentry [69] had listed Rubiaceae and Euphorbiaceae as one of the ten most speciose families in both palaeo- and neotropics. Large families are often the most represented ones and it is well-known that all the afore-mentioned families are large. A family’s dominance is possibly due to the adaptability of its species (such as the ability to form symbiosis with nitrogen-fixing bacteria as in Fabaceae) coupled with conducive environmental factors that facilitate pollination, seed dispersal and successful ecesis [66]. In the present study, a total of 18,790 tree individuals were enumerated and the tree adult density ranged from 370 individuals/ha (TDD I) to 900 individuals/ha (TEF I), with a mean value of 676 ± 83.6 individuals/ha. This is well-within the range given for tropical forests (245–859 stems/ha [70]) and also within the range 352–1173/2 ha given by Sundarapandian and Swamy [37] from Kodayar, Kanyakumari, Western Ghats. The mean tree adult density (676 individuals/ha), which when compared on a continental scale is 25.1, 8.4 and 7.4% greater than the average tree densities of the tropical forests of Africa [71], Amazonia [72] and Borneo in Asia [73] respectively. These differences may be due to variations in the sampled plot size, methods adopted and

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

geographical features. The composition of a forest is often defined by the species that is most abundant [44]. Although different species dominated different categories of populations in different forest types, the evergreen forest sites (TEFs) displayed a remarkable dominance of a single family, Dipterocarpaceae in their adult population. Dipterocarpaceae was represented by only three species, viz. Hopea parviflora, Vateria indica and Dipterocarpus indicus, against a total of 70 species that occur as adults in TEFs, yet it forms the major bulk of the population, constituting 45.6% (54.4% in TEF I, 40.6% in TEF II and 41.2% in TEF III) and of this, Hopea parviflora alone constitutes 42.7% (53.6% in TEF I, 37.4% in TEF II and 37.1% in TEF III). This family, particularly the above-listed three species contributed to 6.8% of the total tree adult density in tropical evergreen forest at Varagalaiar, Western Ghats [74]. Dipterocarpaceae is a family that is well-known to dominate the Asian tropical evergreen forests and this dominance serves as a classical distinction between the forests of palaeotropics and neotropics [69]. Also, Dipterocarpaceae is a signatory family of all the rainforests in Borneo, Malaysia and Sumatra as well as in many seasonal forests from India to Philippines [75]. Dipterocarpaceae is regarded as the most speciose woody family in the evergreen forests of southeast Asia [69]. As stated by Torti et al. [76], the regional diversity could possibly influence the dominance as a potentially dominant species could achieve the state of dominance, when there occurs a reduction in the number of competing species and they have a tendency to replace itself by recruiting and establishing under its own canopy. A characteristic feature of Dipterocarpaceae is mast fruiting [77] and these fruits, with their accrescent, purplish calyx could be found strewn along the evergreen forest trails in the evergreen study sites. On the other hand, Terminalia paniculata dominated the adult tree population in the tropical dry deciduous forest sites (23.6% in TDD I and 27.4% in TDD II). On the family level, Combretaceae contributed 31.8 and 48.04% in TDD I and TDD II respectively. Combretaceae in tree adult population is represented by only Terminalia species (Terminalia bellirica, Terminalia chebula, Terminalia elliptica and Terminalia paniculata) in this study and these usually exhibit mass flowering in the dry season [78], which attract a wide range of insect pollinators (bees and flies for Terminalia paniculata [79]) that enhance its reproductive success. Also, they produce samarae, which are winged fruits that are easily wind-dispersed. In the case of tropical semi-evergreen forest sites, Pterocarpus marsupium (27.1% in TSE I) and Aporosa cardiosperma (24.8% in TSE II) were dominant. Fabaceae contributed 28.1% in TSE I, while Phyllanthaceae contributed 25.8% in TSE II. Fabaceae is considered to be a pantropical family [80]. Besides forming symbiotic associations with bacterial nitrogen-fixers, Fabaceae species possess rapid seed germination rates than the non-legumes and with the onset of the monsoon, the seedlings exhibit high growth rates, leading to their dominance [81]. This growth strategy enables Fabaceae to combat dry periods and thrive successfully in seasonally dry tropical forests. Phyllanthaceae was the second largest segregated family from Euphorbiaceae sensu lato as per APG and it is pantropical with about 2000 species from 59 genera, comprising several semi-succulent and/or phyllocladous species [82]. These unique family traits might have resulted in greater tree adult densities of species belonging to the above-discussed families in the study sites. Woody climbers were known to contribute 10–45% of the total woody stem density in the tropics [47,48], although in the present study, they constituted only 7.4% of the total woody stems (juveniles + adults + woody climbers). Overall, 349 individuals of woody climbers were enumerated from 2.8 ha with the mean value of 125 individuals/ha. This is within the pantropical range for woody climber density (105–1414 individuals/ha) given by DeWalt et al. [83]. However, it is 9.8, 24.9, 22.6 and 33.7% lesser than the average woody climber densities given by DeWalt et al. [83] for Asian (223 individuals/ha), African (374 individuals/ha), Central American (351

11

individuals/ha) and South American (462 individuals/ha) tropical forests respectively. The mean woody climber densities observed in the present study for the tropical dry deciduous forest sites (194 individuals/ha), tropical semi-evergreen forest sites (46 individuals/ha) and tropical evergreen forest sites (132 individuals/ha) were very much lower than those observed for seasonal dry forest (594.8 individuals/ ha), semi-evergreen forest (647.7 individuals/ha) and wet evergreen forest (345.6 individuals/ha) respectively of peninsular India by Parthasarathy et al. [48]. Such variations in woody climber distribution could be probably due to geographic topography, light penetration, soil properties, host tree sizes, disturbances, etc. [48]. The basal area is a function of net primary productivity and reflects the growing stock and the biomass of forest [72]. In this study, the mean tree adult basal area was 46.4 m2/ha. This is 14.42% higher than the pantropical average basal area given by Dawkins [84], 32 m2/ha. This is also 14.92, 17.42 and 9.42% greater than the average basal area values computed for the tropical forests of Africa [71], Amazonia [85] and Borneo in Asia [73] respectively. However, it is within the range reported in other studies from Western Ghats (28–81.4/2 ha [37], 28.35– 67.4/0.3 ha [41], 25.91–47.75/30 ha [65]). As pointed out by Ramachandran and Swarupanandan [42], although tree saplings and juveniles dominated in terms of density, tree adults dominated in the case of basal area due to their large girths. Species that contributed most to the basal area is often regarded as the most important woody species of the forest [86]. In this study, Terminalia paniculata and Terminalia elliptica in tropical dry deciduous forest sites (TDDs I and II), Pterocarpus marsupium and Ficus beddomei (TSE I), and Aporosa cardiosperma and Terminalia paniculata (TSE II) in tropical semi-evergreen forest sites and Hopea parviflora in the tropical evergreen forest sites (TEFs I–III) were the most important dominant woody tree adult species. Nonetheless, of all the species, Hopea parviflora contributed to the largest basal area (9.6 m2/ha). Dipterocarp species are well-recognized to dominate the basal area due to their massive tree size [74]. Also, the buttressed trees were noted to have large basal area values by Dutta and Devi [87]. The highest basal area in TEF III (75.3 m2/ha) could be due to its close proximity to the perennial stream which provides perpetual water availability, which further enhances the tree size of the already voluminous dipterocarps. In the case of woody climbers, the basal area ranged from 0.0002 m2/ha (TSE I) to 0.7 m2/ha (TDD I and TEF I) and the mean basal area was 0.31 m2/ha. This is less than the pantropical average basal area for woody climbers (0.71 m2/ha) but within the range (0.18–2.1 m2/ha) given by DeWalt et al. [88]. The mean basal area values of woody climbers observed for the tropical dry deciduous forest sites (0.35 m2/ha), tropical semi-evergreen forest sites (0.10 m2/ha) and tropical evergreen forest sites (0.42 m2/ha) were higher, lower and comparable with those values observed for seasonal dry forest (0.21 m2/ha), semi-evergreen forest (0.69 m2/ha) and wet evergreen forest (0.43 m2/ha) respectively by Parthasarathy et al. [48]. In the present study, contiguousness was the most prevalent form of distribution, followed by a considerable extent of random distribution among tree saplings, juveniles and adults, while regular distribution was very rare. According to Odum [89], contiguousness is the commonest form of distribution in natural forests, random distribution is observed in stable habitats and regular distribution indicates severe competition among the species. Contiguous distribution was reported in other studies too: Swamy et al. [41], Gandhi and Sundarapandian [90] and Das et al. [91]. Contiguousness or clumping patterns may be due to the synergistic effect of several factors such as modes of seed dispersal, topography, soil composition, gap formations, climate, etc. [89]. Plotting a size class distribution reveals the species composition and population structure of forest [92]. In this study, the diameter class-wise distribution of tree juveniles and tree adults showed the characteristic ‘reverse J-shaped’ curve in all the study sites. This indicated a decline

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

12

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

in species richness and abundance with increasing diameter class. Although this means that large-sized trees are less in number, they are considered to be ecologically very significant as they contribute to biomass production [93]. Even so, this trend mirrors a good regenerative capacity. However, this could not be assumed for granted as cautioned by Chaturvedi et al. [94] that small-sized trees are much sensitive to changes in the environment than the large-sized trees and even upon the prevalence of ideal environmental conditions, the mortality rates of the former are way higher than the latter [95,96]. Thus, the regeneration of tree species in tropical forests is an important, yet a complex and dynamic process that needs to be periodically assessed. Population structure is an important indicator of regeneration [12]. In the present study, only 12 tree species (7.9%) occur in all the growth phases, viz. seedlings, saplings, juveniles and adults. Moreover, dominant adult tree species were found to have a good representation in sapling and juvenile stages, which indicates a good regeneration potential of these species in their respective sites of dominance. However, a point of concern is that 12 species occur as only adults, with no younger stage of development and among these are five species that are endemic to only the Western Ghats, although it is possible that they may occur in younger phases elsewhere along the entire hill chain of Western Ghats. This calls for attention to understand the phenophases, reproductive potential, germination potential and population dynamics of these endemic species, in order to sustain their population in the future and take prompt conservation measures. Assessment of regeneration potential in the present study revealed that a large proportion of tree species were either newly recruited or with poor regenerative capacity. Also, the presence of some tree species with no regeneration was recorded from all the study sites. Such poor or no regeneration could be the result of less or uncertain seed supply, absence of conducive microsites for germination and growth and/or presence of factors that affect survival [97]. The regenerative capacity of a species depends on several factors such as the ability of the seedlings and saplings to sprout, survive and grow, light penetration, soil properties, nutrients, nature of the landscape (continuous or fragmented), seed dispersal modes, disturbances, etc. [19,98,99]. Species that show poor or no regeneration potential could be at a great risk in the future [100]. Also, as a majority of the tree species are newly recruited, the species composition of these forest sites is highly likely to change over time, which might in turn, affect future micro-environmental conditions and therefore the regenerative potential of other species.

5. Conclusion The present study yielded information on plant diversity, stand structural attributes and regeneration potential of different sites (tropical dry deciduous, tropical semi-evergreen and tropical evergreen) of KWLS, Western Ghats, India. It indicates that the study area has unique plant species richness and regeneration potential. Although the tropical dry deciduous site (TDD I) was the most speciose site, in terms of overall plant species richness, it was the evergreen site (TEF III) which had the

highest tree species richness. The site conditions play a crucial role in determining the patterns of diversity and structure. Trees form the architecture of any forest ecosystem. The overall regeneration pattern of trees was good as could be elucidated from the ‘reverse J-shaped curve’ of the diameter-class distribution and the dominant species had ‘fair’ to ‘good’ regeneration potential in their respective sites of dominance. Also, it was found that a large proportion of tree species in the study sites were ‘newly recruited’. Even though the overall regeneration status may appear good, there were also some tree species that showed ‘poor’ or ‘no’ regeneration. This might alter the species composition in future. Therefore, it is suggested that alongside continuing the existing protection measures, it is also essential to take steps to periodically monitor the population structure and curb plant invasions through community participation. The data generated in this study would help forest managers and key officials to prioritize sites within the sanctuary that need more attention for better conservation. Funding This research did not receive any specific grant from funding agencies in the public, commercial, or not-for-profit sectors. Authors' contributions SS was the Research Supervisor who designed the research. KS and JAD collected and analyzed the data. SK helped in the identification of plant specimens. The manuscript was written by KS and refined by JAD and SS. All authors read and approved the final version of the manuscript. Finally, we thank the anonymous reviewers for their valuable comments that greatly improved this manuscript. Declaration of Competing Interest None Acknowledgement The authors thank the Tamil Nadu Forest Department for granting permission to conduct this study. KS is grateful to the University Grants Commission for granting fellowship during the study period. Our sincere thanks to Dr. W. Arisdason, Scientist ‘D’, Botanical Survey of India and Dr. N. Ayyappan, French Institute of Pondicherry for helping us with plant identification. We thank the anonymous reviewers for their valuable comments that greatly improved the manuscript. Appendix A. Supplementary data Supplementary data to this article can be found online at https://doi. org/10.1016/j.chnaes.2020.02.004. Appendices

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

TDD I Life-form Seedlings Aporosa cardiosperma (Gaertn.) Merr. Breynia retusa (Dennst.) Alston Calophyllum inophyllum L. Ceriscoides turgida (Roxb.) Tirveng. Cipadessa baccifera (Roth) Miq. Dillenia pentagyna Roxb. Helicteres isora L. Maesa indica (Roxb.) A. DC. Morinda coreia Buch.-Ham. Tabernaemontana alternifolia L.* Tabernaemontana gamblei Subr. & A.N. Henry** Terminalia bellirica (Gaertn.) Roxb. Vitex altissima L.f. Saplings Aglaia edulis (Roxb.) Wall. Aglaia elaeagnoidea (A. Juss.) Benth.** Aglaia simplicifolia (Bedd.) Harms** Agrostistachys borneensis Becc. Alstonia scholaris (L.) R. Br. Antidesma ghaesembilla Gaertn. Antidesma montanum Blume Aporosa cardiosperma (Gaertn.) Merr. Aporosa indo-acuminata Chakrab. & N.P. Balakr.* Ardisia elliptica Thunb. Artocarpus heterophyllus Lam. Artocarpus hirsutus Lam.** Atalantia wightii Tanaka** Baccaurea courtallensis (Wight) Müll.Arg.** Blachia umbellata (Willd.) Baill.** Boswellia serrata Roxb.* Breynia retusa (Dennst.) Alston Bridelia retusa (L.) A. Juss. Buchanania cochinchinensis (Lour.) Almeida Calophyllum inophyllum L. Calophyllum polyanthum Wall. ex Choisy Canarium strictum Roxb. Careya arborea Roxb. Cassia fistula L. Cassine glauca (Rottb.) Kuntze Catunaregam spinosa (Thunb.) Tirveng. Ceriscoides turgida (Roxb.) Tirveng. Chassalia curviflora (Wall.) Thwaites Chionanthus ramiflorus Roxb. Cinnamomum malabathrum (Burm.f.) J. Presl** Cinnamomum verum J. Presl Cipadessa baccifera (Roth) Miq. Croton klotzschianus (Wight) Thwaites Croton malabaricus Bedd.**

TDD II

D

BA

IVI

100 100 100 300 700 200 400

0.002 0.002 0.001 0.002 0.008 0.003 0.002

10.22 10.22 7.91 18.83 37.93 13.37 19.82

100 200 600 500 2100

0.001 0.003 0.019 0.002 0.011

7.19 14.24 64.94 17.63 77.71

1

0.000

2.23

3

0.001

3.24

9

0.001

5.62

8

0.002

5.86

1

0.000

1.76

4

0.001

5.39

1

0.000

2.20

D

TSE I BA

IVI

1

0.000

2.22

146 27 6

0.033 0.005 0.001

94.85 21.39 4.24

1

0.000

1.79

TSE II

D

BA

IVI

600 300

0.006 0.002

61.52 32.43

1700

0.005

86.77

400

0.011

53.59

1000

0.007

65.68

10

0.002

6.50

160

0.046

61.55

2

0.001

1.53

2 2

0.001 0.001

2.56 2.49

8

0.002

4.39

39

0.010

19.11

18 73 4

0.005 0.024 0.001

8.76 33.45 3.97

D

4

0.000

4.58

2 20

0.001 0.006

4.30 19.22

1

0.001

2.50

6

0.002

6.59

1

0.001

2.33

1

0.003

0.000

17.38

1.79 59

0.018

28.44

IVI

17 6 3 1 26 16

0.002 0.001 0.000 0.000 0.003 0.004

6.24 1.90 1.13 0.73 9.42 7.27

4 1

0.001 0.000

1.87 0.62

13

0.001

2.47

3

0.000

1.45

1 24

TEF I BA

117

0.000

0.007

D

TEF II BA

IVI

D

TEF III BA

IVI

D

BA

IVI

3 2

0.000 0.000

0.84 0.81

3 8

0.001 0.002

3.97 6.15

12

0.003

7.48

247

0.043

36.18

24 6

0.008 0.002

18.10 5.67

65 25

0.016 0.006

28.33 10.04

19 34

0.005 0.007

5.68 8.53

1 46

0.000 0.013

1.06 21.33

24

0.005

13.43

12 15 27

0.004 0.004 0.008

3.98 4.03 9.40

2

0.001

0.95

2

0.000

2.09

1

0.000

1.64

1

0.000

1.68

1

0.000

1.04

2 9

0.001 0.002

1.65 3.42

3

0.001

5.32

20

0.005

12.80

22

0.007

6.58

5.38

34 10 6

0.010 0.002 0.002

19.36 7.06 2.97

1 90 4

0.000 0.025 0.002

0.67 17.54 1.92

4

0.000

0.90

5

0.001

2.89

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

0.94

15.76

6

0.001

3.61

8 34

0.001 0.004

2.23 6.54

5

4

0.002

0.001

5.99

10

0.002

6.13

(continued on next page)

13

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

Table A.1 Density, basal area and Importance Value Index of different life-forms in the seven study sites.

14

Life-form

TDD I D

Cullenia exarillata A. Robyns* Dalbergia latifolia Roxb. Dillenia pentagyna Roxb. Dimocarpus longan Lour. Diospyros affinis Thwaites Diospyros bourdillonii Brandis** Diospyros buxifolia (Blume) Hiern Diospyros melanoxylon Roxb. Diospyros paniculata Dalzell** Dolichandrone atrovirens (B. Heyne ex Roth) Sprague* Dysoxylum ficiforme (Wight) Gamble** Dysoxylum malabaricum Bedd. ex Hiern** Elaeocarpus variabilis Zmarzty* Euonymus dichotomus B. Heyne ex Roxb.* Ficus beddomei King** Ficus benghalensis L. Ficus hispida L.f. Ficus tinctoria (Blume) Corner Glycosmis pentaphylla (Retz.) DC. Gomphandra tetrandra (Wall.) Sleum. Gossypium hirsutum L.# Helicteres isora L. Holigarna arnottiana Hook.f.** Hopea parviflora Bedd.** Hydnocarpus alpina Wight Hydnocarpus pentandrus (Buch.-Ham.) Oken** Isonandra lanceolata Wight Isonandra perrottetiana A. DC.** Ixora brachiata Roxb. ex DC.** Ixora pavetta Andr. Knema attenuata (Wall. ex Hook.f. & Thomson) Warb.** Lagerstroemia microcarpa Wight* Lepisanthes tetraphylla (Vahl) Radlk. Ligustrum perrottetii A. DC.* Macaranga peltata (Roxb.) Müll.Arg. Maesa indica (Roxb.) A. DC. Mallotus aureopunctatus (Dalzell) Müll.Arg.** Mallotus ferrugineus (Roxb.) Müll.Arg. Mallotus philippensis (Lam.) Müll.Arg. Mangifera indica L. Manihot carthaginensis (Müll.Arg.) Allem# Meiogyne pannosa (Dalzell) J. Sinclair** Melia dubia Hiern Memecylon heyneanum Benth. ex Wight & Arn.** Mesua ferrea L. Miliusa indica Lesch. ex A. DC. Millingtonia hortensis L.f. Morinda coreia Buch.-Ham. Myristica beddomei King

TDD II BA

IVI

D

13

TSE I BA

0.004

IVI

D

0.000

IVI

2

0.001

1.91

5 20

0.001 0.006

4.20 11.32

26 1

0.009 0.000

12.93 1.27

TEF I

D

BA

IVI

5

0.001

2.12

53

0.005

12.95

TEF II BA

IVI

D

BA

IVI

D

BA

IVI

1

0.000

1.93

3 104

0.001 0.030

2.19 45.25

24 82 1 4

0.004 0.023 0.000 0.001

5.34 16.16 0.66 1.00

3

0.001

2.72

17 18

0.004 0.004

11.40 8.06

3

0.001

1.51

4

0.002

3.76

3.56

9

0.000 0.003

4

0.000

1.18

4

0.001

1.64

20

0.002

3.75

2.47

1

0.001

2.03

5.67

3 257 14

84

0.030

70.87

9 52

0.001 0.011

6.07 35.87

17

0.004

7.61

1

0.000

1.07

28 3 20

0.001 0.006

0.001

0.004

0.002

43

4

0.000

0.013

0.001

0.000

0.001

33.48 3.98

40

0.004

10.35

4 1

0.001 0.001

1.81 1.15

85

0.009

18.76

1 31 16

0.000 0.002 0.000

0.63 7.05 3.15

9 20

0.001 0.001

4.01 3.83

5 39 2

0.000 0.003 0.000

1.77 9.68 1.39

0.001 0.001 0.002

1.24 1.52 3.58

26 1 4 10

0.006 0.001 0.001 0.003

5.62 0.84 1.08 3.19

12

0.002

5.98

13 11

0.003 0.002

2.92 4.13

3 32 28 3

0.000 0.005 0.007 0.000

1.41 14.37 16.23 2.20

33 203 51 22 8 7 1 41

0.006 0.041 0.007 0.005 0.002 0.002 0.001 0.008

9.60 32.53 7.02 5.32 3.43 2.12 0.82 8.19

16 4

0.003 0.001

4.99 1.23

11 191

0.002 0.027

3.54 25.93

4 3

0.002 0.001

1.35 1.61

3

0.000

0.91

1 2

0.000 0.001

1.90 2.26

28 18 8 7

0.005 0.005 0.002 0.002

15.06 14.02 6.29 10.21

10 3

0.003 0.000

10.35 2.10

6

0.001

4.71

1

0.001

2.10

2

0.001

2.30

139

0.035

67.70

1.38

1.04

8.99

18.67 6 5 2

1

16.59

4

0.013 0.002

1 2 4

2.50 12

18

0.007

4.42 14.17 1

1

TEF III

D

14.38

3 2

TSE II BA

2.08

30.53

5

0.001

0.002 0.001 0.000

3.11 2.32 1.32

4.51

4

0.000

0.96

4

0.001

1.32

1 2

0.000 0.001

1.58 2.35

4

0.001

3.42

4.72 6

0.002

4.57

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

Table A.1 (continued)

Juveniles Aglaia edulis (Roxb.) Wall. Agrostistachys borneensis Becc. Albizia lebbeck (L.) Benth.

9

0.001

9.85

3

3 7

0.001

4.53

16

0.002

12.62

2

0.000

3.53

5

0.001

0.001

0.002

2.06

81 52 1

0.000 0.001 0.003 0.000

1.84 2.50 12.28 2.28

2 2

0.000 0.001

2.51 2.75

2

0.000

2 5

0.000 0.001

1.42 2.34

5

0.001

2.12

6.37

3.90

3

0.001

0.000

1.81

2 2 30 134 9

1 5 6 2

0.001 0.002 0.002 0.000

2.36 5.67 7.03 2.42

1 1

0.000 0.000

2.28 2.02

23

0.006

0.001 0.005

5.13 17.58

14

0.004

13.02

2

0.010

3.19

3

0.013

5.16

0.001

0.000 0.000 0.005 0.014 0.001

0.001

7

0.003

5.29

1 1

0.000 0.000

1.64 1.70

2

0.001

1.79

6

0.001

1.81

9

0.002

3.60

3

0.000

1.36

1

0.000

0.74

2

0.000

1.27

11 3

0.002 0.001

3.74 1.56

1.99 4.55

1

0.001

2.13

15

0.004

10.71

15

0.006

9.82

9 1

0.003 0.001

6.09 2.08

3

0.001

3.06

19

0.005

10.87

54

0.012

10.69

1 19

0.000 0.004

1.06 11.38

12

0.003

4.72

1

0.000

1.05

13

0.005

9.48

10

0.004

3.56

1

0.001

1.29

30 1

0.006 0.000

6.88 0.66

7

0.020

5.44

14

0.030

9.01

2.94

3

0.001

3.18

0.90 0.78 7.15 26.82 4.15

5.20

6.38 6.49 30.12

4

0.001

3.93

8

0.003

8.37

16

0.004

10.32

1

0.000

1.88

30

0.010

17.77

0.002 0.000 0.006

5.09 1.72 12.52

0.009

17.48

3.95

3.85

0.72

0.001 0.001

0.001 0.001 0.007

0.012

0.001

0.000

1 3

5 5 34

3

6

2

3.51

11.87 20

4 15

0.001

2.75 11

1

14.45 11.23 0.83

3.14

7 1 1 10 1

0.005 0.005 0.000

1

0.000

0.84

8 1 15

150

0.014

27.76

20

1

0.000

0.70

3

0.006

5.71

4

0.009

3.26

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

(continued on next page)

15

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

Neolitsea pallens (D. Don) Momiy. & H. Hara* Nothapodytes nimmoniana (J. Graham) Mabb. Nothopegia aureofulva Bedd. ex Hook.f.** Nothopegia beddomei Gamble** Nothopegia castaneifolia (Roth) Ding Hou* Olea dioica Roxb. Pavetta indica L. Persea macrantha Wight Phyllanthus emblica L. Pinanga dicksonii (Roxb.) Blume** Pleurostylia opposita (Wall.) Alston Poeciloneuron indicum Bedd.** Polyalthia fragrans (Dalzell) Hook. f. & Thomson* Polyalthia korintii Hook. f. & Thomson Polyalthia longifolia (Sonn.) Thwaites Psydrax dicoccos Gaertn. Pterocarpus marsupium Roxb. Pterospermum diversifolium Blume Pterospermum rubiginosum Heyne** Pterospermum suberifolium (L.) Lam. Pterygota alata (Roxb.) R. Br. Rhodomyrtus tomentosa (Aiton) Hassk. Scolopia crenata (Wight & Arn.) Clos Sterculia villosa Roxb. ex Sm. Strombosia ceylanica Gardner Strychnos potatorum L.f. Symplocos cochinchinensis (Lour.) S. Moore Syzygium laetum (Buch.-Ham.) Gandhi** Syzygium mundagam (Bourd.) Chithra** Syzygium munronii (Wight) N.P.Balakr.* Tabernaemontana alternifolia L.* Tabernaemontana gamblei Subr. & A.N. Henry** Tamilnadia uliginosa (Retz.) Tirveng. & Sastre Tarenna asiatica (L.) Kuntze ex K. Schum. Tectona grandis L.f. Terminalia bellirica (Gaertn.) Roxb. Terminalia chebula Retz. Terminalia elliptica Willd. Terminalia paniculata Roth* Vateria indica L.** Vitex altissima L.f. Wrightia tinctoria (Roxb.) R. Br. Xanthophyllum flavescens Roxb.* Zanthoxylum rhetsa (Roxb.) DC. Ziziphus mauritiana Lam.

16

Life-form

TDD I D

Alstonia scholaris (L.) R. Br. Antidesma ghaesembilla Gaertn. Antidesma montanum Blume Aporosa cardiosperma (Gaertn.) Merr. Aporosa indo-acuminata Chakrab. & N.P. Balakr.* Ardisia elliptica Thunb. Artocarpus heterophyllus Lam. Artocarpus hirsutus Lam.** Atalantia wightii Tanaka** Baccaurea courtallensis (Wight) Müll.Arg.** Boswellia serrata Roxb.* Breynia retusa (Dennst.) Alston Bridelia retusa (L.) A. Juss. Buchanania cochinchinensis (Lour.) Almeida Calophyllum inophyllum L. Calophyllum polyanthum Wall. ex Choisy Canarium strictum Roxb. Careya arborea Roxb. Cassia fistula L. Cassine glauca (Rottb.) Kuntze Ceriscoides turgida (Roxb.) Tirveng. Chassalia curviflora (Wall.) Thwaites Chionanthus ramiflorus Roxb. Cinnamomum malabathrum (Burm.f.) J. Presl** Cipadessa baccifera (Roth) Miq. Croton malabaricus Bedd.** Cullenia exarillata A. Robyns* Dalbergia latifolia Roxb. Dillenia pentagyna Roxb. Dimocarpus longan Lour. Diospyros affinis Thwaites Diospyros melanoxylon Roxb. Diospyros paniculata Dalzell** Dolichandrone atrovirens (B. Heyne ex Roth) Sprague* Dysoxylum ficiforme (Wight) Gamble** Elaeocarpus variabilis Zmarzty* Euonymus dichotomus B. Heyne ex Roxb.* Ficus beddomei King** Ficus benghalensis L. Ficus callosa Willd. Ficus hispida L.f. Ficus tinctoria (Blume) Corner Filicium decipiens (Wight & Arn.) Thwaites Flacourtia montana J. Graham* Glycosmis pentaphylla (Retz.) DC. Gomphandra tetrandra (Wall.) Sleum. Gossypium hirsutum L.# Helicteres isora L. Holigarna arnottiana Hook.f.** Hopea parviflora Bedd.**

TDD II BA

IVI

D

25 4

0.013

4.07

2

0.008

4.28

3

0.006

4.33

1 1

0.004 0.001

2.13 1.75

6

TSE I BA

0.040

0.037

IVI

50.55

22.88

D

114

1 2 7 2

TSE II BA

0.281

0.001 0.005 0.023 0.011

IVI

66.22

D

1 7 3

6 0.013

6.76

32

0.128

31.76

11 1

0.035 0.003

1

0.015

0.004

24.54

12.90 1.96

7

0.013

5.07

12

0.016

9.56

26 23

0.054 0.035

15.04 14.28

2 48

5

3

0.026

0.016

0.239

2.26 13.97 5.20

0.003

0.009

2.94

18 1 1

0.050 0.004 0.003

13.59 1.59 1.53

BA

IVI

11

0.036

6.74

45 3

0.134 0.003

25.46 2.64

18

0.035

11.60

2

0.002

1.47

D

TEF III BA

IVI

D

BA

IVI

1 1 23 12

0.007 0.002 0.065 0.015

1.43 1.01 12.89 6.14

18 8 25

0.047 0.017 0.056

10.48 4.86 13.82

2 1

0.004 0.003

1.32 1.06

41 11

0.126 0.031

26.26 6.52

1 28

0.004 0.066

1.33 16.90

25

0.058

16.30

2

0.004

1.50

2 5 1 2

0.003 0.007 0.001 0.002

1.28 3.44 0.95 1.17

25

0.073

17.61

9

0.012

5.30

25 4

0.053 0.013

15.15 3.43

45 4

0.097 0.004

19.67 2.40

13

0.033

10.86

10

0.024

6.40

4 76 10 9

0.012 0.223 0.017 0.014

2.62 41.35 7.32 6.13

5 57

0.011 0.157

3.76 26.79

3

0.000

1.26

2

0.004

1.39

2.63

22

0.067

15.72

2

0.008

5.49

3

0.005

1.88

4

0.010

5.35

1 2 5

0.001 0.003 0.009

1.15 1.50 3.43

3

0.018

7.63

5

0.013

9.58

5

0.014

3.86

3

0.010

2.28

4

0.011

4.29

2 2

0.002 0.009

1.46 2.88

2.72 31.15

4

TEF II

D

15.25

4.11

3

0.016

3.39

5

0.015

6.69

6

0.017

5.19

1

0.001

2.29

3

0.012

6.65

3

0.003

5.30

2 3

0.006 0.004

1.67 2.47

14 2 1

0.040 0.007 0.001

8.50 1.55 0.92

5

0.020

4.27

3

0.005

2.61

1

0.003

1.08

1 2 1

0.001 0.003 0.001

1.19 1.53 1.16

5

0.004

2.20

5 2

0.006 0.003

2.78 1.88

7 54

0.008 0.203

4.67 35.15

4 41

0.006 0.150

4.37 28.28

2 35

0.002 0.090

1.84 20.37

2.18

5 114

0.002 0.090

0.001 0.024 0.003

IVI

1.37 1.99 7.08 3.55 1

4

TEF I BA

71.21

2 10

0.008 0.018

0.020

4.25

7.16 26.13 17

0.058

17.03

2

0.005

3.55

9

0.025

16.86

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

Table A.1 (continued)

15

0.035

11.48

2 1 4

1

3

15

1

9

2

0.002

0.006

0.033

0.001

0.012

0.009

0.001

4.63

13 3

0.030 0.003

7.54 1.70

1

0.001

1.09

3

0.007

3.45

7 4 5 8 1

0.013 0.011 0.020 0.024 0.004

3.68 3.56 4.52 5.36 1.18

0.311

109.31

24 1

0.072 0.002

13.59 0.97

2

0.004

1.90

3

0.003

5.29

7

0.020

5.62

7 1

0.011 0.001

3.83 1.34 2 1

0.008 0.005

5.55 2.86

29

0.036

10.40

2

0.009

4.16

1

0.003

1.23

14 1

0.048 0.001

10.37 0.92

4 13

0.010 0.018

2.87 7.18

1 5

0.001 0.016

0.92 2.88

7

0.016

3.89

8

0.027

5.65

1

0.002

1.00

3

0.012

2.20

9

0.031

6.77

6

0.015

3.64

1.79

6.81 7

0.012

4.10

2

0.006

1.75

4 2 4

0.007 0.006 0.008

2.23 1.76 3.17

2.98 0.001

3.89

3

0.010

2.72

1

0.004

1.63

5 2

0.014 0.005

5.67 3.02

1 1

0.004 0.002

2.68 2.50

6

0.008

3.51

1.82 0.001 0.041

0.015

0.005 0.023

2.55 9.57

1

0.001

1.74

3

0.006

7.56

6

1.77

1

0.028

0.001

0.001

0.004

5

0.011

4.24

8

0.012

4.29

2

0.009

2.72

2.29 5

1

2.13

0.001

13.01 1

2 9

0.001

5.63 17.12

15.19

3

1

0.015 0.089

77

1

0.004

9 25

2.96

1 9

1

3.65 2.75

1.86

1

1

0.016

0.006 0.004

0.009

3.50

2.79

9.29

1

1

0.004

2.69

1

0.003

2.59

0.001

1.14

18

0.047

10.79

20

0.037

7.78

2

0.008

1.89

2

0.004

2.29

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

1.33

2

0.003

3.12

10

0.031

16.82

1.33

8 5 2

0.008 0.008

0.007

3.09

3.40 1.91 (continued on next page)

17

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

Hydnocarpus alpina Wight Hydnocarpus pentandrus (Buch.-Ham.) Oken** Isonandra lanceolata Wight Isonandra perrottetiana A. DC.** Ixora brachiata Roxb. ex DC.** Ixora notoniana Wall. ex G. Don** Ixora pavetta Andr. Kingiodendron pinnatum (Roxb. ex DC.) Harms** Knema attenuata (Wall. ex Hook.f. & Thomson) Warb.** Lepisanthes tetraphylla (Vahl) Radlk. Ligustrum perrottetii A. DC.* Macaranga peltata (Roxb.) Müll.Arg. Maesa indica (Roxb.) A. DC. Mallotus aureopunctatus (Dalzell) Müll.Arg.** Mallotus ferrugineus (Roxb.) Müll.Arg. Mallotus philippensis (Lam.) Müll.Arg. Mangifera indica L. Manihot carthaginensis (Müll.Arg.) Allem# Meiogyne pannosa (Dalzell) J. Sinclair** Melia dubia Hiern Memecylon heyneanum Benth. ex Wight & Arn.** Mesua ferrea L. Morinda coreia Buch.-Ham. Myristica beddomei King Nothapodytes nimmoniana (J. Graham) Mabb. Nothopegia aureofulva Bedd. ex Hook.f.** Nothopegia castaneifolia (Roth) Ding Hou* Olea dioica Roxb. Orophea uniflora Hook.f. & Thomson** Pavetta indica L. Phyllanthus emblica L. Pinanga dicksonii (Roxb.) Blume** Pleurostylia opposita (Wall.) Alston Polyalthia fragrans (Dalzell) Hook. f. & Thomson* Polyalthia korintii Hook. f. & Thomson Psydrax dicoccos Gaertn. Pterocarpus marsupium Roxb. Pterospermum rubiginosum Heyne** Pterospermum suberifolium (L.) Lam. Rhodomyrtus tomentosa (Aiton) Hassk. Scolopia crenata (Wight & Arn.) Clos Sterculia villosa Roxb. ex Sm. Stereospermum chelonoides (L.f.) DC. Strombosia ceylanica Gardner Strychnos potatorum L.f. Symplocos cochinchinensis (Lour.) S. Moore Syzygium cumini (L.) Skeels Syzygium laetum (Buch.-Ham.) Gandhi** Syzygium mundagam (Bourd.) Chithra** Syzygium munronii (Wight) N.P.Balakr.*

18

Life-form

Tabernaemontana alternifolia L.* Tabernaemontana gamblei Subr. & A.N. Henry** Tamarindus indica L.# Tectona grandis L.f. Terminalia bellirica (Gaertn.) Roxb. Terminalia chebula Retz. Terminalia elliptica Willd. Terminalia paniculata Roth* Tetrameles nudiflora R. Br. Vateria indica L.** Vitex altissima L.f. Wrightia tinctoria (Roxb.) R. Br. Xanthophyllum flavescens Roxb.* Adults Aglaia edulis (Roxb.) Wall. Aglaia simplicifolia (Bedd.) Harms** Agrostistachys borneensis Becc. Albizia lebbeck (L.) Benth. Alstonia scholaris (L.) R. Br. Antidesma ghaesembilla Gaertn. Aporosa cardiosperma (Gaertn.) Merr. Aporosa indo-acuminata Chakrab. & N.P. Balakr.* Ardisia elliptica Thunb. Artocarpus heterophyllus Lam. Artocarpus hirsutus Lam.** Atalantia wightii Tanaka** Baccaurea courtallensis (Wight) Müll.Arg.** Blachia umbellata (Willd.) Baill.** Bombax ceiba L. Boswellia serrata Roxb.* Breynia retusa (Dennst.) Alston Bridelia retusa (L.) A. Juss. Buchanania cochinchinensis (Lour.) Almeida Butea monosperma (Lam.) Taub. Calophyllum inophyllum L. Canarium strictum Roxb. Careya arborea Roxb. Cassia fistula L. Cassine glauca (Rottb.) Kuntze Ceriscoides turgida (Roxb.) Tirveng. Chassalia curviflora (Wall.) Thwaites Chionanthus ramiflorus Roxb. Cinnamomum malabathrum (Burm.f.) J. Presl** Cipadessa baccifera (Roth) Miq. Croton malabaricus Bedd.** Cullenia exarillata A. Robyns* Cycas circinalis L.** Dillenia pentagyna Roxb. Dimocarpus longan Lour.

TDD I

TDD II

D

BA

IVI

4 19

0.009 0.038

3.58 14.40

1 2

0.001 0.004

1.75 3.73

6 41

0.019 0.108

8.19 34.81

1 14

0.002 0.030

1.85 13.20

1

0.070

2.91

5

0.561

13.71

D

1 3 26

TSE I BA

0.004 0.017 0.087

IVI

4.99 13.58 74.31

BA

IVI

11 1

0.021 0.002

6.93 1.44

1 1

0.001 0.003

1.32 1.57

16

0.060

16.04

21

0.076

17.86

17 1

8

1 1

0.163

0.122 0.009

0.113

0.329

21.40

D

1 1

17 5.50

8

0.087

10.85

1

0.028

2.48

0.090 0.166

IVI

D

BA

IVI

D

BA

IVI

8

0.023

6.01

36

0.101

20.85

29

0.066

14.07

8

0.019

4.87

5 10

0.011 0.033

3.55 7.98

2 6

0.004 0.022

2.01 5.46

16 2 3 34

0.032 0.004 0.003 0.085

7.46 1.62 1.74 17.75

3 2

0.004 0.010

2.45 2.75

1

0.002

0.99

20

0.047

14.30

13

0.034

7.17

27

0.074

14.37

2.69

2

0.025

2.27

2 2 5 4

0.353 0.044 0.388 0.095

2.56 1.53 5.96 3.89

6 2

1.022 0.047

8.36 2.34

2 2 1

0.207 0.020 0.065

2.08 1.46 1.31

0.078

33.92

58

0.240

33.41

2 1 3

0.272 0.018 0.039

3.93 1.32 3.89

1

0.010

1.60

7

0.241

5.70

16

0.279

13.46

1

0.023

1.66

2

0.028

9

0.102

77

3.380

52.31

1

0.017

1.32

1

0.010

1.28

3

0.538

5.71

TEF III

BA

24

0.014 0.548

2.28 24.11

1 2

0.071 0.058

2.35 2.74

8

0.321

12.95

0.771

25.43

4

0.228

6.27

1

0.069

2.34

2 1

0.040 0.009

2.60 1.91

2

1

5 2

2.33 2.23

TEF II

D

2

0.018

36 5

1.009 0.058

21.48 2.69

3.23

3 9

0.237 0.996

3.83 10.54

6.82

2

0.043

2.79

2

0.031

1.72

3.35

0.043

0.299 0.119

0.001 0.001

IVI

3.45 2.29

3

3 1

TEF I BA

2.33

10.12

1 15 1

0.019

TSE II

D

9.66 3.41 10.36 6.10

2

32

1.358

45.02

0.032

0.042

0.023

2.68 3

0.040

4.83

5

0.092

4.86

1

0.045

1.77

8 1

0.141 0.014

5.92 1.37

10

0.167

7.50

1 6

0.013 0.090

1.61 7.19

3

0.091

3.26

6

0.181

4.76

4

0.047

3.88

4

0.457

4.29

1.35

3.92 3

0.039

3.88

2 4

0.219 0.224

2.74 6.12

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

Table A.1 (continued)

1 1

9 7

3

0.384 0.431

0.032

43 4 2

3.767 0.218 0.098

59.69 7.57 4.39

1

0.011

1.93

17

0.687

18.48

0.014 0.008

1.30 1.27

3

0.102

2.62

36 1

0.905 0.043

21.08 1.47

3

0.057

2.39

6

0.355

5.15

3

0.043

2.32

2

0.026

3.22 1 5 5

0.081 0.266 0.904

1.62 4.52 6.99

2

0.026

2.72

3.87

1

7

0.304

15.44

1

0.019

2.39

1 1

0.241 0.033

4.68 2.53

2

0.029

3.17

1

0.453

0.037

0.014

12

1.252

14.31

1 1

0.032 0.012

1.40 1.29

6.03

2.11

9 70

0.118 1.355

4.45 38.99

4 10

1.355 0.417

10.49 10.01

1 1 3

0.092 0.444 0.063

1.73 3.65 3.11

2

0.031

1.72

0.866

1 8 5

0.156 0.850 0.203

1.61 8.37 4.54

9 3

0.261 0.067

6.71 1.91

9 1 5

0.517 0.011 0.767

8.36 1.13 4.82

126 2

17.175 0.042

102.05 1.53

193 1 14

14.458 0.016 0.241

139.95 1.63 11.44

129

14.473

103.57

1

0.045

1.77

3

0.106

4.34

8 6

0.694 0.159

7.06 4.69

9 1 1 9

0.194 0.013 0.261 0.112

6.49 1.14 1.96 4.62

2 5

0.042 0.072

1.53 3.31

2

0.085

1.67

2 1

0.022 0.038

1.46 1.22

1

0.042

1.23

5

0.215

4.58

1

0.033

1.20

2 2

1

2.33

23.61

6 2 15

0.396

21.27

1

0.035

2.46

11

1.688

27.90

9

57

0.431

5.289

0.020 0.031

0.009

1.92 1.98

2

0.066

1.86

1

0.017

1.38

1.59

0.352 0.411

2 1

0.044 0.023

2.05 1.66

2

0.026

1.95

0.277 0.017

5

0.324

6.79

5.02 9.01 1

15

9.13

1.95

4 7 0.013

0.305

14.58 10.69

3

1

14

0.041

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

1.75

6.15 2.54

5

0.443

7.24

1 2

0.016 0.077

1.37 2.92

12.84

78.58

3 1

0.032 0.122

2.95 1.89

6

0.747

9.27 (continued on next page)

19

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

Diospyros affinis Thwaites Diospyros melanoxylon Roxb. Diospyros montana Roxb. Diospyros paniculata Dalzell** Dipterocarpus indicus Bedd.** Dolichandrone atrovirens (B. Heyne ex Roth) Sprague* Dysoxylum ficiforme (Wight) Gamble** Euonymus dichotomus B. Heyne ex Roxb.* Ficus beddomei King** Ficus benghalensis L. Ficus hispida L.f. Gluta travancorica Bedd.** Gomphandra tetrandra (Wall.) Sleum. Grewia tiliifolia Vahl Helicteres isora L. Hevea brasiliensis (Willd. ex A. Juss.) Müll.Arg.# Hopea parviflora Bedd.** Hydnocarpus alpina Wight Hydnocarpus pentandrus (Buch.-Ham.) Oken** Isonandra lanceolata Wight Isonandra perrottetiana A. DC.** Ixora brachiata Roxb. ex DC.** Ixora pavetta Andr. Kingiodendron pinnatum (Roxb. ex DC.) Harms** Lagerstroemia microcarpa Wight* Lepisanthes tetraphylla (Vahl) Radlk. Ligustrum perrottetii A. DC.* Litsea salicifolia (Nees) Hook. f.** Macaranga peltata (Roxb.) Müll.Arg. Mallotus ferrugineus (Roxb.) Müll.Arg. Mallotus philippensis (Lam.) Müll.Arg. Manihot carthaginensis (Müll.Arg.) Allem# Meiogyne pannosa (Dalzell) J. Sinclair** Melia dubia Hiern Mesua ferrea L. Miliusa indica Lesch. ex A. DC. Morinda coreia Buch.-Ham. Myristica beddomei King Neolamarckia cadamba (Roxb.) Bosser Neolitsea pallens (D. Don) Momiy. & H. Hara* Nothapodytes nimmoniana (J. Graham) Mabb. Nothopegia aureofulva Bedd. ex Hook.f.** Palaquium ellipticum (Dalzell) Baill.** Pavetta indica L. Persea macrantha Wight Phyllanthus emblica L. Pinanga dicksonii (Roxb.) Blume** Polyalthia fragrans (Dalzell) Hook. f. & Thomson* Polyalthia korintii Hook. f. & Thomson Pterocarpus marsupium Roxb.

20

Life-form

TDD I D

Pterospermum rubiginosum Heyne** Scolopia crenata (Wight & Arn.) Clos Sterculia villosa Roxb. ex Sm Stereospermum colais (Buch.-Ham. ex Dillw.) D. L. Mabberley Strombosia ceylanica Gardner Symplocos cochinchinensis (Lour.) S. Moore Syzygium cumini (L.) Skeels Syzygium laetum (Buch.-Ham.) Gandhi** Syzygium mundagam (Bourd.) Chithra** Tabernaemontana alternifolia L.* Tabernaemontana gamblei Subr. & A.N. Henry** Tamarindus indica L.# Tarenna asiatica (L.) Kuntze ex K. Schum. Tectona grandis L.f. Terminalia bellirica (Gaertn.) Roxb. Terminalia chebula Retz. Terminalia elliptica Willd. Terminalia paniculata Roth* Tetrameles nudiflora R. Br. Vateria indica L.** Vitex altissima L.f. Wrightia tinctoria (Roxb.) R. Br. Xanthophyllum flavescens Roxb.* Climbers Anamirta cocculus (L.) Wight & Arn. Asparagus racemosus Willd. Bridelia stipularis (L.) Blume Calamus rotang L. Calycopteris floribunda (Roxb.) Lam. ex Poir. Centrosema molle Benth.# Cissampelos pareira (Buch.-Ham. ex DC.) Forman# Cissus latifolia Lam. Clematis zeylanica (L.) Poir. Cyclea peltata (Lam.) Hook.f. & Thomson Gymnema sylvestre (Retz.) R. Br. ex Schult. Hemidesmus indicus (L.) R. Br. Jasminum flexile Vahl Jasminum multiflorum (Burm.f.) Andrews Parthenocissus semicordata (Wall.) Planch. Piper hookeri Miq. Piper nigrum L. Pothos scandens L. Rhamnus oenoplia L. Rivea hypocrateriformis (Desr.) Choisy Salacia chinensis L. Sida cordata (Burm.f.) Borss. Waalk. Smilax zeylanica L. Stephania japonica (Thunb.) Miers Tetracera akara (Burm.f.) Merr.

TDD II BA

IVI

D

BA

TSE I IVI

D

TSE II BA

IVI

D

1

TEF I BA

0.058

IVI

D

TEF II BA

IVI

BA

IVI

1

0.040

1.47

0.453 0.035

6.86 2.55

2 2

0.018 0.029

3.05 4.68

1 1

0.179 0.303

4.04 5.31

11 35

0.890 3.417

22.68 72.54

7

0.146

10.78

1 1200 1000

0.000 0.010 0.011

1.62 26.05 16.09

1400 600 200

0.014 0.003 0.000

21.91 8.69 2.71

400 1500 200 5100 1401 500 2 2 400 100 3101

0.002 0.003 0.002 0.010 0.004 0.006 0.000 0.000 0.002 0.000 0.048

5.36 17.74 5.32 48.82 13.43 8.36 1.60 1.60 8.36 2.15 61.75

1 2

8 29 49

0.331 3.467 3.704

15.22 60.45 73.54

2

0.000

0.029

2

0.023

1.67

11

0.316

9.78

0.003 0.006 0.000

18.58 12.65 2.32

2.38 3.91

2.72

2

0.020

2.56

1

0.326

3.01

2.331 3.236

23.55 29.14

3.90

0.024

1

0.014

1.14

9 2

0.324 0.023

6.92 2.27

1.40

0.011

1.60

6

0.249

5.76

4

0.068

3.80

3

0.073

3.74

7

0.181

6.81

2

0.067

2.41

0.019 0.698 0.031 1.527

3.18 8.71 2.26 25.13

10 14 11 15

1.183 1.962 0.360 1.024

9.51 17.77 7.64 15.45

6 5 9 12

0.461 0.946 1.381 1.091

5.69 7.01 10.43 13.55

0.380

11.29

3

0.037

3.40

1

0.000

300.00

32.56

7

0.160

7.22

3

0.337

4.61

2 14 3 26

21

0.437

16.37

20

1

0.000

22.89

1

0.000

27.97

1

0.000

26.34

12

0.000

222.80

91.59

2100

0.308

68.86

1900

0.006

39.40

300.00

0.008

0.313

1

1.560

2300

3

4.82

1 1700 700 100

IVI

0.037

22

1.365

BA

3

14 16

600

2

0.075 0.031

D

1.54 1

1 1

TEF III

D

0.000

300.00

53.25

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

Table A.1 (continued)

Woody climbers Aganosma cymosa (Roxb.) G. Don Artabotrys zeylanicus Hook.f. & Thomson Calamus vattayila Renuka* Calycopteris floribunda (Roxb.) Lam. ex Poir. Cissus latifolia Lam. Combretum albidum G. Don Connarus sclerocarpus (Wight & Arn.) G. Schellenb.* Dendrophthoe falcata (L.f.) Ettingsh. Dioscorea alata L. Mucuna pruriens (L.) DC. Piper hookeri Miq. Rhamnus oenoplia L. Rivea hypocrateriformis (Desr.) Choisy Salacia chinensis L. Smilax wightii A. DC. Spatholobus parviflorus (Roxb. ex DC.) Kuntze Tetracera akara (Burm.f.) Merr. Tetrastigma nilagiricum (Miq.) B.V. Shetty Vallaris solanacea (Roth) Kuntze Ventilago maderaspatana Gaertn. Ziziphus rugosa Lam. Understorey Alpinia galanga (L.) Willd. Amomum muricatum Bedd.** Anaphyllum beddomei Engl.** Aristolochia indica L. Benkara malabarica (Lam.) Tirveng. Biophytum intermedium Wight Breynia vitis-idaea (Burm.f.) C.E.C. Fisch. Canthium angustifolium Roxb. Capparis divaricata Lam. Carissa spinarum L. Chromolaena odorata (L.) R.M. King & H. Rob.## Clausena austroindica B.C. Stone & K. Narayanan* Clidemia hirta (L.) D. Don## Commelina benghalensis L. Croton bonplandianus Baill.# Croton caudatus Geiseler Curculigo orchioides Gaertn. Cyanthillium cinereum (L.) H. Rob. Cymbopogon citratus (DC.) Stapf# Desmodium gangeticum (L.) DC. Desmodium triflorum (L.) DC. Dichrostachys cinerea (L.) Wight & Arn.

2600 200 100

0.001 0.011

3

119

0.265

0.008

46.90

4.63 10.27

0.001

86.83 1 1

0.000 0.005

8.27 13.84

13

0.046

57.82

2 7

0.012 0.047

30.83 99.06

1

0.006

20.16

3 1 1 1

0.001 0.000 0.000 0.000

14.80 8.39 8.29 8.69

8

0.002

40.46

18

0.016

107.82

9

0.003

43.96

223.12

2 1

0.001 0.000

6.73 5.81

2

0.002

7.15

9

0.006

23.53

9

0.001

213.17

1

0.000

300.00

1 1

0.000

5.73

6

0.004

10.65

1 1 1

0.000 0.000 0.000

5.73 5.81 5.73

4103

0.081

19.62

100

0.000

1.27

800 1 7

0.002 0.001 0.004

2.48 1.28 1.60

2200

0.032

12.45

100 800 2 4 200 600

0.001 0.004 0.003 0.003 0.001 0.001

1.34 3.83 1.44 3.63 2.61 3.22

400 300 1

0.001 0.003 0.000

1.77 1.92 1.15

10000 200

0.194 0.003

40.46 4.14

21

0.125

18.73

1

0.000

1.90

40400

0.285

33

17

0.011

0.003

4

0.003

14.36

62

0.233

227.82

0.000

10.11

28

0.140

246.65

13

0.021

85.65

6

0.007

43.24

8

0.045

84.50

5

0.003

25.28

2

0.000

4.31

12

0.040

49.24

3

0.007

14.20

16.69

8.14

4

0.001

4.53

3

0.000

4.05

40

0.105

127.84

1

0.000

6.99

85

0.187

164.35

6

0.005

23.78

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

76.72

(continued on next page)

21

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

Thunbergia fragrans Roxb. Tylophora indica (Burm.f.) Merr. Ziziphus rugosa Lam.

22

Life-form

TDD I

Digitaria ciliaris (Retz.) Koeler Dracaena terniflora Roxb. Ecbolium viride (Forssk.) Alston Flueggea leucopyrus Willd. Glycosmis mauritiana (Lam.) Tanaka Hybanthus enneaspermus (L.) F. Muell. Ixora alba L. Ixora coccinea L. Justicia japonica Thunb. Lantana camara L.## Leea asiatica (L.) Ridsdale Leucas aspera (Willd.) Link Leucas biflora (Vahl) R. Br. ex Sm. Lygodium flexuosum (L.) Sw. Mallotus atrovirens Müll.Arg.** Memecylon randerianum S.M. Almeida & M.R. Almeida* Memecylon subramanii A.N. Henry** Mesosphaerum suaveolens (L.) Kuntze## Mimosa diplotricha C. Wright ex Sauvalle## Murraya koenigii (L.) Spreng. Myrovernix scaber (L.f.) Koek. Naregamia alata Wight & Arn. Oldenlandia corymbosa L. Ophiorrhiza mungos L. Osbeckia zeylanica L.f. Pavonia zeylanica (L.) Cav. Phyllanthus reticulatus Poir. Pseudarthria viscida (L.) Wight & Arn. Psychotria anamallayana Bedd.** Psychotria glandulosa (Dennst.) Suresh Psychotria nilgiriensis Deb & M. Gangop.** Psychotria nudiflora Wight & Arn.** Pteris biaurita L. Rauvolfia verticillata (Lour.) Baill. Ruellia tuberosa L.# Sauropus androgynus (L.) Merr. Selaginella sp. Sida cordifolia L. Solanum viarum Dunal## Spermacoce hispida L. Stachytarpheta jamaicensis (L.) Vahl# Streblus asper Lour. Stylosanthes fruticosa (Retz.) Alston Synedrella nodiflora (L.) Gaertn.## Themeda cymbaria Hack. Thottea siliquosa (Lam.) Ding Hou Triumfetta rhomboidea Jacq. Vanda tessellata (Roxb.) Hook. ex G. Don Vanda testacea (Lindl.) Rchb.f. Vigna indica T.M.Dixit, K.V.Bhat & S.R.Yadav Viscum heyneanum DC. Zingiber montanum (J. König) Link ex A. Dietr. Ziziphus jujuba Mill.# 2

2

2

TDD II

D

BA

IVI

1500

0.023

5.88

100 102

0.001 0.001

1.40 1.37

400

0.001

1.77

400 400 1200 1 100 300 100 4

0.005 0.001 0.480 0.002 0.000 0.001 0.000 0.002

3.37 1.79 58.59 1.38 1.27 2.75 1.27 1.37

D

7 100 1 100 300 7100 300 300 700 100 400 600

0.001 0.000 0.002 0.003 0.040 0.000 0.002 0.003 0.001 0.000 0.007

TSE I BA

0.001

IVI

D

TSE II BA

IVI

15.52

D

TEF I BA

IVI

TEF II

D

BA

IVI

1

0.000

6.86

D

TEF III BA

IVI

D

BA

IVI

2

0.000

2.22

54

0.014

53.43

134 42

0.032 0.003

61.02 18.52

23 14

0.029 0.009

52.18 23.65

24

0.031

37.02

4

0.000

5.48

78

0.009

30.40 12

0.017

23.41

1 99 10 1

0.002 0.043 0.008 0.000

7.96 103.14 16.85 6.95

1.40 1.11 1.48 3.00 18.32 2.72 2.86 2.53 1.34 1.74 2.77 10536

700

0.003

2.53

1000

0.003

2.98

300 300 100 200 200 200 14300 100 24600

0.001 0.000 0.000 0.000 0.008 0.000 0.016 0.000 0.044

1.71 2.72 1.27 1.43 2.27 1.45 30.84 1.27 48.22

100 100 500 200 2

0.000 0.085 0.013 0.002 0.001

1.30 10.78 3.25 1.66 2.37

7

0.006

2.87

158000

2

1.116

0.002

0.557

67.81

600

0.001

4.38

47000

0.037

62.42

400

0.006

4.56

2 900

0.000 0.142

1.90 16.97

276.65

7.83

Density (No./4000 m ); basal area (m /4000 m ); * - endemic to India, ** - endemic to Western Ghats, # - exotic, ## - exotic and invasive

101

0.008

33.14

9

0.003

10.55

219

0.034

73.72

122

0.047

99.42

17

0.004

21.34

5

0.002

12.45

82

0.009

61.87

10

0.004

14.29

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

Table A.1 (continued)

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

23

Table A.2 Family-wise contribution in the seven study sites. Family

Rubiaceae Fabaceae Euphorbiaceae Malvaceae Phyllanthaceae Apocynaceae Anacardiaceae Meliaceae Moraceae Annonaceae Lamiaceae Rutaceae Acanthaceae Combretaceae Ebenaceae Arecaceae Lauraceae Melastomataceae Myrtaceae Oleaceae Rhamnaceae Asteraceae Bignoniaceae Calophyllaceae Celastraceae Menispermaceae Sapotaceae Vitaceae Dipterocarpaceae Poaceae Sapindaceae Zingiberaceae Achariaceae Araceae Aristolochiaceae Asparagaceae Burseraceae Dilleniaceae Myristicaceae Orchidaceae Piperaceae Primulaceae Pteridaceae Salicaceae Smilacaceae Solanaceae Stemonuraceae Verbenaceae Violaceae Capparaceae Commelinaceae Connaraceae Convolvulaceae Cycadaceae Dioscoreaceae Elaeocarpaceae Hypoxidaceae Lecythidaceae Loganiaceae Loranthaceae Lygodiaceae Lythraceae Oxalidaceae Polygalaceae Ranunculaceae Santalaceae Selaginellaceae Strombosiaceae Symplocaceae Tetramelaceae

TDD I

TDD II

TSE I

TSE II

TEF I

TEF II

TEF III

G

S

D

G

S

D

G

S

D

G

S

D

G

S

D

G

S

D

G

S

D

11 12 4 7 5 6 2 4 2 2 4 1 3 2

12 13 7 9 7 7 2 5 2 2 5 1 3 4

1691 16458 61 2814 1714 6374 3 7880 211 2 2604 100 1500 2135

1 1

1 1

17 16

6 2 3

10 2 4

249 15 304

3 2 2 1 2 3 1 1

4 2 3 1 2 3 1 1

126 38 49 3 7 22 75 3

228 28 47 6 234 61 36

58

684 2 113 32 359 19 61 18 267 55 4 22

4 1 2 2 3 1 1

1

9 2 7 4 6 3 2 6 4 3 1 3

3 1 2 2 2 1 1

1

6 2 6 4 4 3 1 4 2 3 1 2

161 70 192

74 227 3 27 1 2 9 1

10608 47075 130 2 1211 1035 9 1703 72

8 1 3

3 5 1 2 1 1 1 1

4 4 4 2 5 1 1 2 3

5 1 3

3 4 1 2 1 1 1 1

4 4 3 2 4 1 1 2 1

2 1 1 1

5 1 1 1

114 4 48 1

5 3 3 3 2 3 1 3

7 3 3 4 4 3 1 4

158 99 50 15 67 216 35 98

1

3

160

1 1 1 1 1

1 3 3 1 1

2600 62 67 600 4

2

4

40

1 1

2 2

45 12

29 173

1 2 1

1 2 1

1 101 1

1 18 78 165 16 3

1 1 1 3

1 2 2 3

6 13 20 8

56 277 11 22

3 5

1 2 1 3 1 2

3 3 1 3

1 1

1 2 1 2 1 1

1 1 1 2 1 1

1 1

8 67

2 1 2 2 1

2 1 3 2 1

11 1 32 150 5

1 1 1

1 1 1

1 12 52

1 1 3

1 1 3

4 1 57

1 3 3

1 3 3

11 13 36

1 1 1

1 1 1

1 5 50

2 4

3 4

31 93

1

1

8

25

62 40400 8 11100

3 27 61

2

1 1 2 3

1 2 1

2

1 1 2 3

1 2 1

2

2

272

1 1 2

1 1 2

12 6 209

1 3 3

2 3 3

42 13 378

2 1 1

2 1 1

106 33 7

2

2

20

2

2

12

2

2

72

1

2

64

1

2

4

1 1

1 1

1900 400

1 1

2 1

86 2

1

1

1

1 1 1

1 1 1

1 2 4

1 1 1

3 1 3

1

1

2

2 1 2 2 4

2 3 3 4 4

800 5 1903 3219 3200

2

2

101

4

4

1801

2

2

3

2 1 1 1 1 1 1 1 2 1 1 1 1 1

2 1 1 2 1 1 1 1 2 1 2 2 1 1

26100 1 4103 41 100 100 1200 8 208 4 600 402 12 700

1 1

2 1

701 100

2 1

2 1

1400 400

1 1

1 1

800 2

1

1

5

1 1

1 1

3 200

1

1

1

1

1

1

1

1

1

1

1

1

7

11

158000

50

6

10

1 1

1 1

2100 413

1

1

2300

2

2

28

1

1

1 1 1

1 1

1 100

1

1

800

1 1 1

1 1 1

400 2 300

1

1

1

1

1

47

1

1

6

1

1

19

12

2

1 1

1 1

1

1

1

1

86

1

1

17

1 1 1

34

1

1

5

1

1

3

3

1

1

19

1

1

18

1

1

6

1

1

2 1

1

8

1

1

3

1

1

1

1

1

60

1

1

12

1

1

5

1

1

8

1

1

1

2

2

1

1

3 1

1

3

1

1

35

2

1

1

5

1

1

195

1 1

1 1

2 34

1

1

98

2 600

1

1

16

1

1

16

2

G – Genus/Genera, S – Species, D – Density in No./4000 m .

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

24

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx

References [1] D. Mishra, T.K. Mishra, S.K. Banerjee, Comparative phytosociological and soil physico-chemical aspects between managed and unmanaged lateritic land, Ann. Forest. 5 (1997) 16–25. [2] P. Saikia, J. Deka, S. Bharali, A. Kumar, O.P. Tripathi, L.B. Singha, S. Dayanandan, M.L. Khan, Plant diversity patterns and conservation status of eastern Himalayan forests in Arunachal Pradesh, Northeast India, For. Ecosyst. 4 (2017) 28, https://doi.org/10. 1186/s40663-017-0117-8. [3] G. Mandal, S.P. Joshi, Analysis of vegetation dynamics and phytodiversity from three dry deciduous forests of Doon Valley, Western Himalaya, India, J. Asia Pac. Biodivers. 7 (2014) 292–304, https://doi.org/10.1016/j.japb.2014.07.006. [4] J.A. Dar, S. Sundarapandian, Patterns of plant diversity in seven temperate forest types of Western Himalaya, India, J. Asia Pac. Biodivers 9 (2016) 280–292, https://doi.org/10.1016/j.japb.2016.03.018. [5] K.V. Gadow, G. Zhang, G. Durrheim, D. Drew, A. Seydack, Diversity and production in an afromontane forest, For. Ecosyst. 3 (2016) 15, https://doi.org/10.1186/ s40663-016-0074-7. [6] E.P. Lillo, E.S. Fernando, M.J.R. Lillo, Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, J. Asia Pac. Biodivers. 12 (2019) 83–105, https://doi.org/10.1016/j.japb.2018.07.003. [7] K.V. Gadow, C.Y. Zhang, C. Wehenkel, A. Pommerening, J. Corral-Rivas, M. Korol, S. Myklush, G.Y. Hui, A. Kiviste, X.H. Zhao, Forest structure and diversity, in: T. Pukkala, K. Gadow (Eds.), Continuous Cover Forestry, Book Series Managing Forest Ecosystems, Vol 24, Springer Science + Business Media B. V. 2011, pp. 29–84, https://doi.org/10.1007/978-94-007-2202-6. [8] S. Bano, S.M. Khan, J. Alam, A.A. Alqarawi, E.F. AbdAllah, Z. Ahmad, I.U. Rahman, H. Ahmad, A. Aldubise, A. Hashem, Eco-Floristic studies of native plants of the Beer Hills along the Indus River in the districts Haripur and Abbottabad, Pakistan, Saud. J. Biol. Sci. 254 (2018) 801–810, https://doi.org/10.1016/j.sjbs.2017.02.009. [9] S. Sahney, M.J. Benton, P.A. Ferry, Links between global taxonomic diversity, ecological diversity and the expansion of vertebrates on land, Biol. Lett. 64 (2010) 544–547, https://doi.org/10.1098/rsbl.2009.1024. [10] A. Paul, M.L. Khan, A.K. Das, Population structure and regeneration status of rhododendrons in temperate mixed broad-leaved forests of western Arunachal Pradesh, India, Geol. Landscape Ecol (2018) https://doi.org/10.1080/24749508.2018. 1525671. [11] A.M. Mane, N. Prabakaran, S.S. Manchi, Floral diversity, composition, and recruitment on the karstland of Baratang Island, India, Ecol. Complex. 37 (2019) 47–54, https://doi.org/10.1016/j.ecocom.2018.11.002. [12] S. Singh, Z.A. Malik, C.M. Sharma, Tree species richness, diversity, and regeneration status in different oak Quercus spp. dominated forests of Garhwal Himalaya, India, J. Asia Pac. Biodivers. 9 (2016) 293–300, https://doi.org/10.1016/j.japb.2016.06. 002. [13] M.L. Khan, J.P. Rai, R.S. Tripathi, Population structure of some tree species in disturbed and protected subtropical forests of north-east India, Acta Oecol. 8 (1987) 247–255. [14] A. Pradhan, A. Ormsby, N. Behera, Diversity, population structure, and regeneration potential of tree species in five sacred forests of western Odisha, India, Ecoscience (2018) https://doi.org/10.1080/11956860.2018.1522148. [15] S.N. Rai, A. Saxena, The extent of forest fire, grazing and regeneration status in inventoried forest areas of India, Indian For. 123 (1997) 689–702. [16] M.J. Sullivan, J. Talbot, S.L. Lewis, O.L. Phillips, L. Qie, S.K. Begne, J. Chave, A. CuniSanchez, W. Hubau, G. Lopez-Gonzalez, L. Miles, A. Monteagudo-Mendoza, B. Sonke, T. Sunderland, H. ter Steege, L.J. White, K. Affum-Baffoe, S.I. Aiba, E.C. de Almeida, E.A. de Oliveira, P. Alvarez-Loayza, E.A. Davila, A. Andrade, L.E. Aragao, P. Ashton, C.G. Aymard, T.R. Baker, M. Balinga, L.F. Banin, C. Baraloto, J.F. Bastin, N. Berry, J. Bogaert, D. Bonal, F. Bongers, R. Brienen, J.L. Camargo, C. Cerón, V.C. Moscoso, E. Chezeaux, C.J. Clark, A.C. Pacheco, J.A. Comiskey, F.C. Valverde, E.N. Coronado, G. Dargie, S.J. Davies, C. De Canniere, K.M. Djuikouo, J.L. Doucet, T.L. Erwin, J.S. Espejo, C.E. Ewango, S. Fauset, T.R. Feldpausch, R. Herrera, M. Gilpin, E. Gloor, J.S. Hall, D.J. Harris, T.B. Hart, K. Kartawinata, L.K. Kho, K. Kitayama, S.G. Laurance, W.F. Laurance, M.E. Leal, T. Lovejoy, J.C. Lovett, F.M. Lukasu, J.R. Makana, Y. Malhi, L. Maracahipes, B.S. Marimon, B.H. Junior, A.R. Marshall, P.S. Morandi, J.T. Mukendi, J. Mukinzi, R. Nilus, P.N. Vargas, N.C. Camacho, G. Pardo, M. Peña-Claros, P. Pétronelli, G.C. Pickavance, A.D. Poulsen, J.R. Poulsen, R.B. Primack, H. Priyadi, C.A. Quesada, J. Reitsma, M. Réjou-Méchain, Z. Restrepo, E. Rutishauser, K.A. Salim, R.P. Salomão, I. Samsoedin, D. Sheil, R. Sierra, M. Silveira, J.W. Slik, L. Steel, H. Taedoumg, S. Tan, J.W. Terborgh, S.C. Thomas, M. Toledo, P.M. Umunay, L.V. Gamarra, I.C. Vieira, V.A. Vos, O. Wang, S. Willcock, L. Zemagho, Diversity and carbon storage across the tropical forest biome, Sci. Rep. 7 (2017) https://doi.org/10.1038/srep3910239102. [17] D.F. Cusack, J. Karpman, D. Ashdown, Q. Cao, M. Ciochina, S. Halterman, S. Lydon, A. Neupane, Global change effects on humid tropical forests, Evidence for biogeochemical and biodiversity shifts at an ecosystem scale, Rev. Geophys. 54 (2016) 523–610, https://doi.org/10.1002/2015RG000510. [18] J.W.F. Slik, V. Arroyo-Rodríguez, S.I. Aiba, P. Alvarez-Loayza, L.F. Alves, P. Ashton, P. Balvanera, M.L. Bastian, P.J. Bellingham, E. Van Den Berg, L. Bernacci, P. Da Conceição Bispo, L. Blanc, K. Böhning-Gaese, P. Boeckx, F. Bongers, B. Boyle, M. Bradford, F.Q. Brearley, M.B.N. Hockemba, S. Bunyavejchewin, D.C.L. Matos, M. Castillo-Santiago, E.L.M. Catharino, S.L. Chai, Y. Chen, R.K. Colwell, R.L. Chazdon, C. Clark, D.B. Clark, D.A. Clark, H. Culmsee, K. Damas, H.S. Dattaraja, G. Dauby, P. Davidar, S.J. DeWalt, J.L. Doucet, A. Duque, G. Durigan, K.A.O. Eichhorn, P.V. Eisenlohr, E. Eler, C. Ewango, N. Farwig, K.J. Feeley, L. Ferreira, R. Field, A.T. De Oliveira Filho, C. Fletcher, O. Forshed, G. Franco, G. Fredriksson, T. Gillespie, J.F. Gillet, G. Amarnath, D.M. Griffith, J. Grogan, N. Gunatilleke, D. Harris, R. Harrison,

[19]

[20]

[21]

[22] [23]

[24] [25] [26] [27] [28] [29] [30]

[31] [32]

[33]

[34]

[35]

[36]

[37]

[38]

[39]

[40]

[41]

A. Hector, J. Homeier, N. Imai, A. Itoh, P.A. Jansen, C.A. Joly, B.H.J. De Jong, K. Kartawinata, E. Kearsley, D.L. Kelly, D. Kenfack, M. Kessler, K. Kitayama, R. Kooyman, E. Larney, Y. Laumonier, S. Laurance, W.F. Laurance, M.J. Lawes, I.L. Do Amaral, S.G. Letcher, J. Lindsell, X. Lu, A. Mansor, A. Marjokorpi, E.H. Martin, H. Meilby, F.P.L. Melo, D.J. Metcalfe, V.P. Medjibe, J.P. Metzger, J. Millet, D. Mohandass, J.C. Montero, M. De Morisson Valeriano, B. Mugerwa, H. Nagamasu, R. Nilus, S. Ochoa-Gaona, N. Page Onrizal, P. Parolin, M. Parren, N. Parthasarathy, E. Paudel, A. Permana, M.T.F. Piedade, N.C.A. Pitman, L. Poorter, A.D. Poulsen, J. Poulsen, J. Powers, R.C. Prasad, J.P. Puyravaud, J.C. Razafimahaimodison, J. Reitsma, J.R. Dos Santos, W.R. Spironello, H. Romero-Saltos, F. Rovero, A.H. Rozak, K. Ruokolainen, E. Rutishauser, F. Saiter, P. Saner, B.A. Santos, F. Santos, S.K. Sarker, M. Satdichanh, C.B. Schmitt, J. Schöngart, M. Schulze, M.S. Suganuma, D. Sheil, E. Da Silva Pinheiro, P. Sist, T. Stevart, R. Sukumar, I.F. Sun, T. Sunderland, H.S. Suresh, E. Suzuki, M. Tabarelli, J. Tang, N. Targhetta, I. Theilade, D.W. Thomas, P. Tchouto, J. Hurtado, R. Valencia, J.L.C.H. Van Valkenburg, T. Van Do, R. Vasquez, H. Verbeeck, V. Adekunle, S.A. Vieira, C.O. Webb, T. Whitfeld, S.A. Wich, J. Williams, F. Wittmann, H. Wöll, X. Yang, C.Y.A. Yao, S.L. Yap, T. Yoneda, R.A. Zahawi, R. Zakaria, R. Zang, R.L. De Assis, B.G. Luize, E.M. Venticinque, An estimate of the number of tropical tree species, Proc. Natl. Acad. Sci. 112 (2015) E4628–E4629, https://doi.org/10.1073/pnas. 1423147112. A.M. Osuri, D. Chakravarthy, D. Mudappa, T.S. Raman, N. Ayyappan, S. Muthuramkumar, N. Parthasarathy, Successional status, seed dispersal mode and overstorey species influence tree regeneration in tropical rain-forest fragments in Western Ghats, India, J. Trop. Ecol. 33 (2017) 270–284, https://doi.org/10. 1017/S0266467417000219. T.A. Gardner, J. Barlow, R. Chazdon, R.M. Ewers, C.A. Harvey, C.A. Peres, N.S. Sodhi, Prospects for tropical forest biodiversity in a human-modified world, Ecol. Lett. 12 (2009) 561–582, https://doi.org/10.1111/j.1461-0248.2009.01294.x. J. Poker, K. MacDicken, Tropical forest resources, facts and tables, in: L. Pancel, M. Kohl (Eds.), Tropical Forestry Handbook, Springer-Verlag, Berlin Heidelberg 2016, pp. 3–45, https://doi.org/10.1007/978-3-642-54601-3_7. P. Singh, S.S. Dash, Plant Discoveries 2013 – New Genera, Species and New Records, Botanical Survey of India, Kolkata, India, 2014. W. Arisdason, P. Lakshminarasimhan, http://www.bsienvis.nic.in/Database/Status_ of_Plant_Diversity_in_India_17566.aspx. 2017, Last accessed on 16/05/2019. https://doi.org/10.1007/978-3-642-54601-3_7 M.P. Nayar, Hotspots of Endemic Plants of India, Nepal and Bhutan, Tropical Botanic Garden and Research Institute, Thiruvananthapuram, India, 1996. Forest Survey of India (FSI), India State of Forest Report 2017, http://fsi.nic. in/details.php?pgID=sb:64 2017 Last accessed on 16/05/2019. Worldclim, http://www.worldclim.org/ 2019 Last accessed on 16/05/2019. R. Misra, Ecology Work Book, Oxford and IBH Publications, New Delhi, 1968 244. J.S. Gamble, Flora of the Presidency of Madras, 11 Parts. (Parts 1 – 7 by J.S. Gamble and 8 – 11 by C.E.C. Fischer), Adlard & Sons, London, 1915 – 1936. J.P. Pascal, B.R. Ramesh, A field key to the trees and lianas of the evergreen forests of the Western Ghats (India), Institut Français De Pondichéry, Puducherry, 1997. Biodiversity Informatics and Co–Operation in Taxonomy for Interactive Shared Knowledge Base (BIOTIK), http://www.biotik.org/ 2019 Last accessed on 16/06/ 2019. Angiosperm Phylogeny Group IV (APG IV), http://www.mobot.org/ MOBOT/research/APweb/ 2019 Last accessed on 16/05/2019. Wiersema, GRIN Germplasm Resources Information Network Taxonomy and Global Biodiversity Information Facility, https://www.gbif.org/dataset/66dd09602d7d-46ee-a491-87b9adcfe7b1 2018 Last accessed on 16/05/2019. P.B. Whitford, Distribution of woodland plants in relation to succession and clonal growth, Ecology 30 (1949) 199–208. https://www.jstor.org/stable/ 1931186. P. Legendre, Interpreting the replacement and richness difference components of beta diversity, Glob. Ecol. Biogeogr. 23 (2014) 1324–1334, https://doi.org/10. 1111/geb.12207. U. Shankar, A case of high tree diversity in a Sal (Shorea robusta) dominated lowland forest of Eastern Himalaya, floristic composition, regeneration and conservation, Curr. Sci. 81 (2001) 776–786. https://www.jstor.org/stable/24106397. T. Ganesh, R. Ganesan, M.S. Devy, P. Davidar, K.S. Bawa, Assessment of plant biodiversity at a mid-elevation evergreen forest of Kalakad–Mundanthurai Tiger Reserve, Western Ghats, India, Curr. Sci. 71 (1996) 379–392. https://www.jstor.org/ stable/24099615. S.M. Sundarapandian, P.S. Swamy, Forest ecosystem structure and composition along an altitudinal gradient in the Western Ghats, South India, J. Trop. For. Sci. 12 (2000) 104–123. https://www.jstor.org/stable/23616407. D.M. Apgaua, D.G. Pereira, R.M. Santos, G.C. Menino, G.G. Pires, M.A. Fontes, D.Y. Tng, Floristic variation within seasonally dry tropical forests of the Caatinga Biogeographic Domain, Brazil, and its conservation implications, Int. For. Rev. 17 (2015) 33–44, https://doi.org/10.1505/146554815815834840. S.A. Al-Robai, H.A. Mohamed, S.M. Howladar, A.A. Ahmed, Vegetation structure and species diversity of Wadi Turbah Zahran, Albaha area, southwestern Saudi Arabia, Ann. Agric. Sci. 62 (2017) 61–69, https://doi.org/10.1016/j. aoas.2017.04.001. F. Brambach, C. Leuschner, A. Tjoa, H. Culmsee, Diversity, endemism, and composition of tropical mountain forest communities in Sulawesi, Indonesia, in relation to elevation and soil properties, Perspect. Plant. Ecol. Evol. Syst. 27 (2017) 68–79, https://doi.org/10.1016/j.ppees.2017.06.003. P.S. Swamy, S.M. Sundarapandian, P. Chandrasekar, S. Chandrasekaran, Plant species diversity and tree population structure of a humid tropical forest in Tamil Nadu, India, Biodivers. Conserv. 9 (2000) 1643–1669, https://doi.org/10.1023/A: 1026511812878.

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx [42] V.S. Ramachandran, K. Swarupanandan, Structure and floristic composition of oldgrowth wet evergreen forests of Nelliampathy Hills, Southern Western Ghats, J. For. Res. 24 (2013) 37–46, https://doi.org/10.1007/s11676-013-0323-3. [43] U.M. Chandrashekara, P.S. Ramakrishnan, Vegetation and gap dynamics of a tropical wet evergreen forest in the Western Ghats of Kerala, India, J. Trop. Ecol. 10 (1994) 337–354, https://doi.org/10.1017/S0266467400008014. [44] R. Valencia, R.B. Foster, G. Villa, R. Condit, J.C. Svenning, C. Hernández, K. Romoleroux, E. Losos, E. Magård, H. Balslev, Tree species distributions and local habitat variation in the Amazon, large forest plot in eastern Ecuador, J. Ecol. 92 (2004) 214–229, https://doi.org/10.1111/j.0022-0477.2004.00876.x. [45] K.P. Tripathi, B. Singh, Species diversity and vegetation structure across various strata in natural and plantation forests in Katerniaghat Wildlife Sanctuary, North India, Trop. Ecol. 50 (2009) 191–200. [46] C. Sunil, R.K. Somashekar, B.C. Nagaraja, Riparian vegetation assessment of Cauvery River basin of South India, Environ. Monit. Assess. 170 (2010) 545–553, https://doi. org/10.1007/s10661-009-1256-3. [47] S.A. Schnitzer, F. Bongers, The ecology of lianas and their role in forests, Trends Ecol. Evol. 17 (2002) 223–230, https://doi.org/10.1016/S0169-5347 (02)02491-6. [48] N. Parthasarathy, S. Muthuramkumar, C. Muthumperumal, P. Vivek, N. Ayyappan, M.S. Reddy, Liana composition and diversity among tropical forest types on peninsular India, in: S.A. Schnitzer, F. Bongers, R.J. Burnham, F.E. Putz (Eds.), The Ecology of Lianas, Wiley-Blackwell Publishing, Oxford, UK 2015, pp. 36–49, https://doi.org/ 10.1002/9781118392409.ch4. [49] A.K. Osman, F. Al-Ghamdi, A. Bawadekji, Floristic diversity and vegetation analysis of Wadi Arar, A typical desert Wadi of the Northern Border region of Saudi Arabia, Saud. J. Biol. Sci. 216 (2014) 554–565, https://doi.org/10.1016/j.sjbs.2014.02.001. [50] F.M. Souza, S. Gandolfi, R.R. Rodrigues, Deciduousness influences the understory community in a semideciduous tropical forest, Biotropica 46 (2014) 512–515, https://doi.org/10.1111/btp.12137. [51] J.F. Slik, P.J. Kebler, P.C. van Welzen, Macaranga and Mallotus species Euphorbiaceae as indicators for disturbance in the mixed lowland dipterocarp forest of East Kalimantan Indonesia, Ecol. Indic. 2 (2003) 311–324, https://doi.org/10.1016/S1470160X(02)00057-2. [52] R.L. Myers, Site susceptibility to invasion by the exotic tree Melaleuca quinquenervia in southern Florida, J. Appl. Ecol. 20 (1983) 645–658. https://www.jstor.org/stable/ 2403532. [53] J.M. Facelli, S.T. Pickett, Plant litter, its dynamics and effects on plant community structure, Bot. Rev. 57 (1991) 1–32, https://doi.org/10.1007/BF02858763. [54] V.S. Chitale, M.D. Behera, P.S. Roy, Future of endemic flora of biodiversity hotspots in India, PLoS One 9 (2014), e115264. https://doi.org/10.1371/journal.pone. 0115264. [55] A.N. Henry, M. Chandrabose, M.S. Swaminathan, N. Nair, Agasthyamalai and its environs, a potential area for Biosphere Reserve, J. Bombay Nat. Hist. Soc. 81 (1984) 282–290. [56] K. Dutta, C.S. Reddy, S. Sharma, C.S. Jha, Quantification and monitoring of forest cover changes in Agasthyamalai Biosphere Reserve, Western Ghats, India 1920–2012, Curr. Sci. 110 (2016) 508–520. [57] G. Utkarsh, N.V. Joshi, M. Gadgil, On the patterns of tree diversity in the Western Ghats of India, Curr. Sci. 75 (1998) 594–603. https://www.jstor.org/stable/ 24100563. [58] J.P. Pascal, Wet Evergreen Forests of the Western Ghats of India, French Institute of Pondicherry, Pondicherry, India, 1988. [59] M. Kent, P. Coker, Vegetation Description and Analysis, Belhaven Press, London, UK, 1992. [60] J.S. Singh, Y.S. Rawat, O.P. Chaturvedi, Replacement of oak forest with pine in the Himalaya affects the nitrogen cycle, Nature 311 (1984) 54, https://doi.org/10. 1038/311054a0. [61] S. Joseph, S.C. Reddy, C. Pattanaik, S. Sudhakar, Distribution of plant communities along climatic and topographic gradients in Mudumalai Wildlife Sanctuary, Southern India, Biol. Lett. 45 (2008) 29–41. [62] A.E. Magurran, Why diversity, Ecological Diversity and Its Measurement, Springer, Netherlands 1988, pp. 1–5. [63] D.H. Knight, A phytosociological analysis of species-rich tropical forest on Barro Colorado Island, Panama, Ecol. Monogr. 45 (1975) 259–284, https://doi.org/10. 2307/1942424. [64] L.S. Devi, P.S. Yadava, Floristic diversity assessment and vegetation analysis of tropical semievergreen forest of Manipur, north east India, Trop. Ecol. 47 (2006) 89–98. [65] N. Ayyappan, N. Parthasarathy, Biodiversity inventory of trees in a large-scale permanent plot of tropical evergreen forest at Varagalaiar, Anamalais, Western Ghats, India, Biodivers. Conserv. 8 (1999) 1533–1554 https://doi.org/10.1023/A: 1008940803073. [66] P.C. Panda, A.K. Mahapatra, P.K. Acharya, A.K. Debata, Plant diversity in tropical deciduous forests of Eastern Ghats, India, A landscape level assessment, Int. J. Biodivers. Conserv. 5 (2013) 625–639, https://doi.org/10.5897/IJBC2013.0581x. [67] T. Sahoo, P.C. Panda, L. Acharya, Structure, composition and diversity of tree species in tropical moist deciduous forests of Eastern India, a case study of Nayagarh Forest Division, Odisha, J. For. Res. 28 (2017) 1219–1230, https://doi.org/10.1007/ s11676-017-0408-5. [68] M.S. Sosef, G. Dauby, A. Blach-Overgaard, X. Van Der Burgt, L. Catarino, T. Damen, V. Deblauwe, S. Dessein, J. Dransfield, V. Droissart, M.C. Duarte, H. Engledow, G. Fadeur, R. Figueira, R.E. Gereau, O.J. Hardy, D.J. Harris, J. de Heij, S. Janssens, Y. Klomberg, A.C. Ley, B.A. Mackinder, P. Meerts, J.L. van de Poel, B. Sonké, T. Stévart, P. Stoffelen, J.C. Svenning, P. Sepulchre, R. Zaiss, J.J. Wieringa, T.L.P. Couvreur, Exploring the floristic diversity of tropical Africa, BMC Biol. 15 (2017) 15, https://doi.org/10.1186/s12915-017-0356-8.

25

[69] A.H. Gentry, Changes in plant community diversity and floristic composition on environmental and geographical gradients, Ann. Missouri. Bot. Gard. 75 (1988) 1–34. https://www.jstor.org/stable/2399464. [70] P.W. Richards, The Tropical Rain Forest; An Ecological Study, Cambridge University Press, Cambridge, UK, 1952 450. [71] S.L. Lewis, B. Sonke, T. Sunderland, S.K. Begne, G. Lopez-Gonzalez, G.M.F. van der Heijden, O.L. Phillips, K. Affum-Baffoe, T.R. Baker, L. Banin, J.F. Bastin, H. Beeckman, P. Boeckx, J. Bogaert, C. De Canniere, E. Chezeaux, C.J. Clark, M. Collins, G. Djagbletey, M.N.K. Djuikouo, V. Droissart, J.L. Doucet, C.E.N. Ewango, S. Fauset, T.R. Feldpausch, E.G. Foli, J.F. Gillet, A.C. Hamilton, D.J. Harris, T.B. Hart, T. de Haulleville, A. Hladik, K. Hufkens, D. Huygens, P. Jeanmart, K.J. Jeffery, E. Kearsley, M.E. Leal, J. Lloyd, J.C. Lovett, J.R. Makana, Y. Malhi, A.R. Marshall, L. Ojo, K.S.H. Peh, G. Pickavance, J.R. Poulsen, J.M. Reitsma, D. Sheil, M. Simo, K. Steppe, H.E. Taedoumg, J. Talbot, J.R.D. Taplin, D. Taylor, S.C. Thomas, B. Toirambe, H. Verbeeck, J. Vleminckx, L.J.T. White, S. Willcock, H. Woell, L. Zemagho, Aboveground biomass and structure of 260 African tropical forests, Philos. Trans. R. Soc. B Biol. Sci. 368 (2013) 20120295, https://doi.org/10.1098/rstb.2012.0295. [72] S.L. Lewis, O.L. Phillips, T.R. Baker, J. Lloyd, Y. Malhi, S. Almeida, N. Higuchi, W.F. Laurance, D.A. Neill, J.N. Silva, J. Terborgh, A.T. Lezama, R.V. Martinez, S. Brown, J. Chave, C. Kuebler, P.N. Vargas, B. Vinceti, Concerted changes in tropical forest structure and dynamics, evidence from 50 South American long-term plots, Philos. Trans. R Soc. B Biol. Sci. 359 (2004) 421–436, https://doi.org/10.1098/rstb.2003. 1431. [73] J.W.F. Slik, S.I. Aiba, F.Q. Brearley, C.H. Cannon, O. Forshed, K. Kitayama, H. Nagamasu, R. Nilus, J. Payne, G. Paoli, A.D. Poulsen, N. Raes, D. Sheil, K. Sidiyasa, E. Suzuki, J.L.C.H. Valkenburg, Environmental correlates of tree biomass, basal area, wood specific gravity and stem density gradients in Borneo's tropical forests, Glob. Ecol. Biogeogr. 19 (2010) 50–60, https://doi.org/10.1111/j.1466-8238.2009. 00489.x. [74] N. Ayyappan, N. Parthasarathy, Composition, population structure and distribution of dipterocarps in a tropical evergreen forest at Varagalaiar, Anamalais, Western Ghats, South India, J. Trop. For. Sci. 13 (2001) 311–321. https://www.jstor.org/stable/43582302. [75] P.S. Ashton, Dipterocarpaceae, Flora Malesiana I 9 (1982) 237–552. [76] S.D. Torti, P.D. Coley, T.A. Kursar, Causes and consequences of monodominance in tropical lowland forests, Am. Nat. 157 (2001) 141–153. https://www.jstor. org/stable/10.1086/318629. [77] S. Sakai, K. Momose, T. Yumoto, T. Nagamitsu, H. Nagamasu, A.A. Karim, T. Nakashizuka, T. Inoue, Plant reproductive phenology and general flowering in a mixed Dipterocarp forest, in: D.W.R.S. Sakai, A.A.H. Karim (Eds.), Pollination Ecology and the Rain Forest, Springer, New York, USA 2005, pp. 35–50, https://doi.org/ 10.1007/0-387-27161-9_4. [78] D.H. Janzen, Synchronization of sexual reproduction of trees within the dry season in Central America, Evolution 21 (1967) 620–637, https://doi.org/10.1111/j.15585646.1967.tb03416.x. [79] A. Thangaraja, V. Ganesan, Studies on the pollen biology of Terminalia paniculata Roth. Combretaceae, African J. Plant Sci. 2 (2008) 140–146. [80] L. Banin, T.R. Feldpausch, O.L. Phillips, T.R. Baker, J. Lloyd, K. Affum-Baffoe, E.J.M.M. Arets, N.J. Berry, M. Bradford, R.J.W. Brienen, S. Davies, M. Drescher, N. Higuchi, D.W. Hilbert, A. Hladik, Y. Iida, K.A. Salim, A.R. Kassim, D.A. King, G. LopezGonzalez, D. Metcalfe, R. Nilus, K.S.H. Peh, J.M. Reitsma, B. Sonké, H. Taedoumg, S. Tan, L. White, H. Wöll, S.L. Lewis, What controls tropical forest architecture? Testing environmental, structural and floristic drivers, Glob. Ecol. Biogeogr. 21 (2012) 1179–1190, https://doi.org/10.1111/j.1466-8238.2012.00778.x. [81] G. Vargas, L.K. Werden, J.S. Powers, Explaining legume success in tropical dry forests based on seed germination niches, a new hypothesis, Biotropica 47 (2015) 277–280 https://doi.org/10.1111/btp.12210. [82] P. Hoffmann, H. Kathriarachchi, K.J. Wurdack, A phylogenetic classification of Phyllanthaceae Malpighiales; Euphorbiaceae sensu lato, Kew Bull. 61 (2006) 37–53. https://www.jstor.org/stable/20443245. [83] S.J. DeWalt, S.A. Schnitzer, L.F. Alves, F. Bongers, R.J. Burnham, Z. Cai, W.P. Carson, J. Chave, G.B. Chuyong, F.R. Costa, C.E.N. Ewango, R.V. Gallagher, J.J. Gerwing, E.G. Amezcua, T. Hart, G. Ibarra-Manríquez, K. Ickes, D. Kenfack, S.G. Letcher, M.J. Macía, J.R. Makana, A. Malizia, M. Martínez-Ramos, J. Mascaro, C. Muthumperumal, S. Muthuramkumar, A. Nogueira, M.P.E. Parren, N. Parthasarathy, D.R. PérezSalicrup, F.E. Putz, H.G. Romero-Saltos, M.S. Reddy, M.N. Sainge, D. Thomas, J. van Melis, Biogeographical patterns of liana abundance and diversity, in: S.A. Schnitzer, F. Bongers, R.J. Burnham, F.E. Putz (Eds.), The Ecology of Lianas, WileyBlackwell Publishing, Oxford, UK 2015, pp. 131–146, https://doi.org/10.1002/ 9781118392409.ch11. [84] H.C. Dawkins, The volume increment of natural tropical high-forest and limitations on its improvements, Empire For. Rev. 38 (1959) 175–180. https://www.jstor.org/ stable/42600614. [85] T.R. Baker, O.L. Phillips, Y. Malhi, S. Almeida, L. Arroyo, A.D. Fiore, T.J. Killeen Erwin, S.G. Laurance, W.F. Laurance, S.L. Lewis, J. Lloyd, A. Monteagudo, D.A. Neill, S. Patino, N.C.A. Pitman, J.N.M. Silva, R.V. Martínez, Variation in wood density determines spatial patterns in Amazonian forest biomass, Glob. Chang. Biol. 10 (2004) 545–562 https://doi.org/10.1111/j.1365-2486.2004.00751.x. [86] F. Gurmessa, T. Soromessa, E. Kelbessa, Structure and regeneration status of Komto Afromontane moist forest, East Wollega Zone, west Ethiopia, J. For. Res. 23 (2012) 205–216, https://doi.org/10.1007/s11676-012-0242-8. [87] G. Dutta, A. Devi, Plant diversity, population structure, and regeneration status in disturbed tropical forests in Assam, northeast India, J. For. Res. 24 (2013) 715–720 https://doi.org/10.1007/s11676-013-0409-y. [88] S.J. DeWalt, S.A. Schnitzer, J. Chave, F. Bongers, R.J. Burnham, Z. Cai, G. Chuyong, D.B. Clark, C.E. Ewango, J.J. Gerwing, E. Gortaire, T. Hart, G. Ibarra-Manríquez, K. Ickes, D.

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004

26

[89] [90]

[91]

[92]

[93]

[94]

[95] [96]

[97]

K. Subashree et al. / Acta Ecologica Sinica xxx (2020) xxx Kenfack, M.J. Macía, J.R. Makana, M. Martínez-Ramos, J. Mascaro, S. Moses, H.C. Muller-Landau, M.P.E. Parren, N. Parthasarathy, D.R. Pérez-Salicrup, F.E. Putz, H. Romero-Saltos, D. Thomas, Annual rainfall and seasonality predict pan-tropical patterns of liana density and basal area, Biotropica 42 (2010) 309–317, https:// doi.org/10.1111/j.1744-7429.2009.00589.x. E.P. Odum, Fundamentals of Ecology, Saunders, Philadelphia, Pennsylvania, 1971. D.S. Gandhi, S.M. Sundarapandian, Diversity and distribution pattern of understory vegetation in tropical dry forests of Sathanur reserve forest in Eastern Ghats, India, Int. J. Sci. Nat. 5 (2014) 452–461. A.K. Das, L.B. Singha, M.L. Khan, Community structure and species diversity of Pinus merkusii Jungh. & de Vriese forest along an altitudinal gradient in Eastern Himalaya, Arunachal Pradesh, India, Trop. Ecol. 58 (2017) 397–408. D.M. Newbery, J.S. Gartlan, A structural analysis of rain forest at Korup and Douala– Edea, Cameroon, Proc. R. Soc. Edinb. Sec. B Biol. Sci. 104 (1996) 177–22410.1017/ S0269727000006138. D.B. Clark, D.A. Clark, Abundance, growth and mortality of very large trees in neotropical lowland rain forest, For. Ecol. Manag. 801-3 (1996) 235–24410.1016/ 0378-1127(95)03607-5. R.K. Chaturvedi, A.S. Raghubanshi, K.W. Tomlinson, J.S. Singh, Impacts of human disturbance in tropical dry forests increase with soil moisture stress, J. Veg. Sci. 28 (2017) 997–100710.1111/jvs.12547. P.G. Jarvis, Scaling processes and problems, Plant Cell Environ. 18 (1995) 1079–108910.1111/j.1365-3040.1995.tb00620.x. J. Grogan, M. Pena-Claros, S. Gunter, 2011. Managing natural populations of BigLeaf Mahogany, in: S. Gunter, M. Weber, B. Stimm, R. Mosandl (Eds.), Silviculture in the tropics, Springer, Berlin, DE 2011, pp. 227–235 , 10.1007/978-3-64219986-8_15. J.S. Clark, B. Beckage, P. Camill, B. Cleveland, J. HilleRisLambers, J. Lichter, J. McLachlan, J. Mohan, P. Wyckoff, Interpreting recruitment limitation in forests, Am. J. Bot. 86 (1999) 1–16.

[98] N.F. Good, R.E. Good, Population dynamics of tree seedlings and saplings in a mature eastern hardwood forest, Bull. Torrey Bot. Club 99 (1972) 172–178. https:// www.jstor.org/stable/2484571. [99] M. Ma, Y. Wu, Y. Zhang, H. Kang, Z. Chen, P. Liu, Sprouting as a survival strategy for non-coniferous trees: Relation to population structure and spatial pattern of Emmenopterys henryi (Rubiales), Acta Ecol. Sin. 39 (2019) 1–810.1016/j. chnaes.2018.07.004. [100] Z.A. Malik, A.B. Bhat, Regeneration status of tree species and survival of their seedlings in Kedarnath Wildlife Sanctuary and its adjoining areas in Western Himalaya, India, Trop. Ecol. 57 (2016) 677–690.

Glossary Plant diversity: Number of different species that are represented in a given community.: Stand structure: Distribution of trees by species and size within a stand.: Regeneration potential: The ability of a species to complete the life-cycle.: Ecosystem: A biological community of interacting organisms and their physical environment.: Phytosociology: A branch of science which deals with plant communities, their composition and development, and the relationships between the species within them.: Endemism: Ecological state of a species being unique to a defined geographic location, such as an island, nation, country or other defined zone, or habitat type.: Tree density: Number of trees per unit area.: Tree basal area: Total cross-sectional area of tree stems in a stand measured at breast height, and expressed as per unit of land area.:

Please cite this article as: K. Subashree, J.A. Dar, S. Karuppusamy, et al., Plant diversity, structure and regeneration potential in tropical forests of Western Ghats, India, Acta Ecologica Sinica, https://doi.org/10.1016/j.chnaes.2020.02.004