Saikia et al.

Forest Ecosystems (2017) 4:28

DOI 10.1186/s40663-017-0117-8

RESEARCH Open Access

Plant diversity patterns and conservation

status of eastern Himalayan forests in
Arunachal Pradesh, Northeast India
P. Saikia1, J. Deka2, S. Bharali3, Amit Kumar1, O. P. Tripathi4, L. B. Singha4, S. Dayanandan5 and M. L. Khan6*

Abstract
Background: The State of Arunachal Pradesh is part of the Himalaya biodiversity hotspots distributed over an area
with the largest elevation gradients in the world, ranging from lowland tropical forests to alpine vegetation.
Methods: The vegetation was surveyed along an elevation gradient ranging from 87 to 4161 m in 354 belt
transects (500 m × 10 m in size) and analyzed using various ecological indices.
Results: A total of 482 (458 identified, 24 unidentified) plants were recorded in the present study, of which 153 are non-
woody herbs and grasses (145 identified) belonging to 55 families, and 329 are woody trees and shrubs (313 identified)
belonging to 74 families. The 458 identified species belong to 117 families and 251 genera, of which 94.10% are
phenerogams (Angiosperms: 421 species [spp.] and Gymnosperms: 10 spp.) and 5.90% cryptogams (Pteridophytes: 27 spp.).
The family Fabaceae contributed the greatest species diversity with a total of 27 plant species (Papilionaceae: 10 spp.,
Mimosaceae: 9 spp. and Caesalpinaceae: 8 spp.) followed by Poaceae (21 spp.), Ericaceae (20 spp.), Asteraceae (18 spp.),
Lauraceae (17 spp.), Euphorbiaceae (16 spp.), Urticaceae (15 spp.) and 49 monotypic families. The most dominant trees were
Castanopsis indica (24 individuals per ha), followed by Quercus semicarpifolia (12 individuals per ha) and Pinus roxburghii (12
individuals per ha) and some bamboos (Dendrocalamus strictus: 69 individuals per ha; Bambusa pallida: 16 individuals per ha).
The studied forest stands were rich in various bamboos; a total of 14 different bamboos were recorded in the present study
including Dendrocalamus strictus, Bambusa tulda, B. pallida etc. Common IUCN red-listed species were Aquilaria malaccensis,
Begonia tessaricarpa, Gledetsia assamica, Gymnocladus assamicus, Livistona jenkinsiana, Rhododendron meddenii, Rhododendron
thomsonii collected from the Dirrang, Tawang, Yamcha, Itanagar, Along and Mebo areas of Arunachal Pradesh.
Conclusion: The results confirm that the Eastern Himalayan forests are very rich in terms of species diversity
which decreases with increasing elevation. Species richness and distribution patterns of plants are largely
regulated by altitude and other environmental factors. Intensive monitoring and management is needed to protect
this fragile ecosystem from the ever-increasing anthropogenic pressure and changing climatic conditions.
Keywords: Species richness, Endemic species, Forest ecosystems, Eastern Himalayan region

Background different regions. Forests are important as they provide

The vegetation is a fundamental component of ecosystems various ecosystem services such as species conservation,
that reflects the effects of the total environment (Billings prevention of soil erosion and preservation of habitat for
1952). Plants grow collectively in a specific ecosystem and plants and animals (Li et al. 2003; Armenteras et al. 2009).
have a mutual relationship among themselves as well as Tree species diversity is the determinant of total forest
within the local environment (Mishra et al. 1997), which biodiversity as trees provide resources as well as habitats
results in the formation of different vegetation types in for almost all other forest species (Haung et al. 2003). In
forest ecosystems, tree species diversity varies greatly from
place to place mainly due to variation in biogeography,
* Correspondence: [email protected]
6
Department of Botany, Dr. Harisingh Gour Central University, Sagar, Madhya
habitat and disturbance (Whitmore 1998). Environmental
Pradesh 470003, India variability in terms of climatic factors, social resources,
Full list of author information is available at the end of the article

© The Author(s). 2017 Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0
International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and
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the Creative Commons license, and indicate if changes were made.
Saikia et al. Forest Ecosystems (2017) 4:28 Page 2 of 12

grazing by herbivores and human interference are the in the Eastern Himalayan forests of Arunachal Pradesh,
critical factors which regulate the spatial and temporal Northeast India over an area of 177 ha spread along
patterns of the vegetation of an ecosystem (Chapin et al. different altitudinal gradients.
1993). However, the composition of vegetation is fluctuating
and continuous modification takes place depending upon Methods
the temperature, moisture, sunlight and available nutrients Study sites
(Heady 1958). Tropical forests constitute the most diverse Arunachal Pradesh is the largest state in the northeastern
plant communities on earth. However, these forests are region of India and situated between 26°28’ N to 29°30’ N
disappearing at the alarming rate of 13.5 million hectares latitude and 91°30′ E to 97°30′ E longitude. The total geo-
per year globally (Kobayashi 2004). It is estimated that graphical area is 83,743 km2, with a population density of
around 7500 species of organisms are becoming extinct 17 individuals per km2 (Census of India 2011). The state is
every year (Chatterjee 1995) and many of them disappear predominantly hilly and mountainous. It is bordered by
even before they are known to the scientific world (Myers et China to the north and northeast (1080 km), separated by
al. 2000; Pimm and Raven 2000). Ecological and environ- the McMohanline, Bhutan (160 km) in the west, and
mental problems such as soil degradation, soil erosion and Myanmar (440 km) in the east, and in the south by the
overexploitation of natural resources are just some of the Indian states of Assam and Nagaland. The state is divided
negative effects resulting from the destruction of natural into 16 districts (Fig. 1). The climate is exceptionally hot
ecosystems (Hundera et al. 2007). Furthermore, floristic and humid at the lower altitudes and in the valleys, while
diversity along with the respective habitat is also rapidly it becomes exceedingly cold in the higher altitudes. The
declining. The loss of any such forest resource will have average mean minimum and maximum temperature
great implications for the environment, the biological ranges between 17.7 °C and 29.5 °C in the subtropical
diversity as well as the socio-economic setup of local humid regions, and 2.4 °C and 21.4 °C in the cold humid
communities. regions. Generally, the winter months (December–February)
The state of Arunachal Pradesh located in the eastern are the coldest months. July and August are the warmest
Himalayan region is part of the Himalaya biodiversity months with the average maximum temperature during
hotspots. It is the largest mountainous state of India and this period being about 30 °C. However, sometimes the
recognized as a globally important eco-region (out of temperature rises as high as 35 °C during summer in
200). It comprises over 5000 species of plants, which are the plains and in the broad valleys. The mean annual
distributed along one of the largest elevation gradients rainfall ranges from 209 to 297 cm. East Siang and the
in the world, ranging from lowland tropical forests to neighbouring areas of Lohit District experience the
alpine vegetation in the Himalayan Mountains. According highest rainfall, whereas, in the West Kameng District
to FSI report (2015), forest cover in Arunachal Pradesh is the rainfall is the lowest. Almost all vegetation types
67,248 km2; it is ranked as second in terms of forested are represented in the state of Arunachal, i.e. tropical,
areas after Madhya Pradesh (77,462 km2). On the other sub-tropical, temperate, and alpine. Bamboos grow along
hand, it ranks 4th in terms of percentage of forest cover the foothills and grasslands cover the riparian tracts. The
(80.30%). In the Himalayan ranges of Arunachal Pradesh, tropical and subtropical vegetation types are characterized
the vegetation ranges from tropical mixed and broadleaf by a great diversity.
evergreen forest to alpine meadow and scrub, across large
elevation gradients. Knowledge of ecological processes Grid generation and vegetation sampling
and biotic interference helps in understanding the stability The area of topographical sheets (1:50,000) as per Survey
of well-established plant communities as ecological, biotic of India was divided into 16 grids with the help of GIS
and anthropogenic disturbances are prevalent all over software. The entire state was covered by a total of 1920
the Himalaya (Singh and Singh 1991). Determining the grids, each grid occupying approximately 6.25 km ×
composition, structure and function of forest ecosystems is 6.25 km area. Details of the grids in different districts of
of great ecological value not only for the understanding of Arunachal Pradesh are presented in Table 1. The vegetation
energy and element flows within the ecosystem, but will also was studied using the belt transect method, and transects
serve as an indicator of ecological impacts (Ramachandra et of 500 m × 10 m were laid in each of the 349 selected grids
al. 2013). It is imperative to have an assessment of the of the 14 districts of Arunachal Pradesh. As a rule, one
current status of the natural stands of plant resources, their transect was to be positioned in each grid, but due to
geographic distribution and population structure so that it difficulties concerning the location identification in the
will be possible to evaluate the various levels of threats and field, five transects were wrongly positioned within the
to develop working plans for the management and conser- already sampled grids. The transects were further
vation of these forest resources. Therefore, the present divided into five separate quadrats and the dimension of
research has attempted to study the plant diversity pattern each quadrat was maintained as per decided standard (i.e.,
Saikia et al. Forest Ecosystems (2017) 4:28 Page 3 of 12

Fig. 1 Map of the study sites (State of Arunachal Pradesh, Northeast India) showing the locations of studied transects at different altitudinal gradients

Table 1 Details of grids per district and percentage covered in the present study
Name of the Districts Total No. of Grids No. of grids sampled No. of transect sampled % of grid coverage
Anjaw 134 21 21 15.67
Changlang 101 39 40 38.61
Dibang Valley 176 – – –
East Kameng 137 16 17 11.68
East Siang 89 18 18 20.22
Kurung Kumey 138 – – –
Lohit 110 36 38 32.73
Lower Dibang valley 142 35 35 24.65
Lower Subansiri 90 21 21 23.33
Papumpare 77 20 21 25.97
Tawang 61 6 6 9.84
Tirap 27 10 10 37.04
Upper Siang 181 20 20 11.05
Upper Subansiri 153 7 7 4.58
West Kameng 121 51 51 42.15
West Siang 183 49 49 26.78
Total 1920 349 354 18.18
Saikia et al. Forest Ecosystems (2017) 4:28 Page 4 of 12

10 m × 100 m), although in some places the standard was 0.025 and 0.05, and contagious distribution if >0.05
adapted to cover the desired transect area due to geomor- (Curtis and Cottam 1956).
phological constraints such as escarpments in the field.
The GBH and visual height of all plant species in the tran-
sects with ≥10 cm girth at breast height were recorded in Results
standard data reporting sheets. Two quadrats measuring The Eastern Himalayan forests in Arunachal Pradesh,
1 m × 1 m were laid out within each transect for enumer- Northeast India, are rich in terms of floristic compos-
ation of herbaceous species as well as seedlings of trees and ition. In the present study, a total of 482 (458 identified,
shrubs. Herbarium specimen of each species were collected 24 unidentified) plant species were recorded, of which
and sufficient numbers of field photographs were taken. 153 were non-woody herbs and grasses (145 identified)
Species were identified by consulting herbaria, existing belonging to 55 families, and 329 were woody trees, and
literature and the respective vernacular names, which were a variety of shrubs (313 identified) belonging to 74 families.
further verified by the Botanical Survey of India, Itanagar, The 458 identified species belong to 117 families and 251
Arunachal Pradesh. genera, of which 94.10% phanerogams (Angiosperms: 421
spp. and Gymnosperms: 10 spp.) and 5.90% cryptogams
Data analysis (Pteridophytes: 27 spp.). Out of the 421 angiosperms, 369
Quantitative analysis of the vegetation was done following spp. were dicots and 52 spp. were monocots. The most
Misra (1968). The Importance Value Index (IVI) was com- diverse Dicot, Monocot, Gymnosperm and Pteridophytic
puted by summing up relative density, relative frequency, families were Fabaceae (27 spp.), Poaceae (21 spp.),
and relative dominance. The species richness was calcu- Pinaceae (6 spp.) and Pteridaceae (10 spp.) respectively.
lated by using the method ‘Margalef’s Index of Richness’ The maximum diversity of species was contributed by
(Dmg) (Magurran 1988). the family Fabaceae with a total of 27 plant species
(Papilionaceae: 10 spp., Mimosaceae: 9 spp. and Caesalpi-
Dmg ¼ ðS–1Þ= ln N naceae: 8 spp.) followed by Poaceae (21 spp.), Ericaceae
(20 spp.), Asteraceae (18 spp.), Lauraceae (17 spp.),
where S is total number of species, N is total number of
Euphorbiaceae (16 spp.), Urticaceae (15 spp.), whereas
individuals.
49 families were monotypic, represented by single species.
The Shannon-Wiener Diversity Index (H, Shannon and
In the case of woody plants, Ericaceae was the most
Wiener 1963) was calculated from the IVI values using
dominant family with 20 different plant species, followed
the formula given by Magurran (1988)
by Fabaceae with 19 species (Papilionaceae: 4 spp.,
X
s Mimosaceae: 7 spp. and Caesalpinaceae: 8 spp.), whereas
H¼– pi ln pi 23 families were monotypic represented by single species.
i¼1 Asteraceae was the most species-rich family (15 spp.) in
where pi is the proportion of the IVI of ith species and the case of herbs and grasses, followed by Pteridaceae with
the IVI of all the species (ni/N). 10 species (a Pteridophytic family), whereas 28 families
The Concentration of Dominance (CD) was assessed were monotypic represented with single species.
by Simpson’s Index (Simpson 1949) The study shows that the majority of transects have a
number of 200–400 individuals followed by transects
X
s with <200 individuals, whereas the majority of transects
CD ¼ ðP i Þ2 have a species diversity ranging from 20 to 30 species
i¼1
followed by transects with 10–20 species (Fig. 2 and
where pi is the same as for the Shannon-Wiener infor- Table 2). The species richness ranges from 0 to 64 with
mation function. a mean of 22 (±0.65 SE) per transect, whereas the tree
The Evenness Index (E) was calculated from the (≥10 cm GBH) density per transect ranges from 0 to
Shannon-Wiener Diversity Index using the formula. 1220 individuals per ha with a mean of 374 (±8.88 SE).
The highest density as well as species richness was
0
E ¼ H 0 =H max recorded in tropical broadleaf semi-evergreen forests in
the Lohit District (Transect: AR 208, Grid No: 92A5SW2
where H′ is the Shannon-Wiener Diversity Index and and Transect: AR 247, Grid No: 83M13SE3 respectively),
H′max = lnS (where S is a total number of species). whereas 16 transects were laid out in grids where the
The ratio of abundance to frequency was used to inter- entire grid was covered by either a river or riverine grass-
pret the distribution pattern of the species (Whitford 1949). land. The maximum diversity was recorded in the Lohit,
The ratio of abundance to frequency indicates regular Changlang and Tirap districts of Arunachal Pradesh.
distribution if below 0.025, random distribution if between However, these forests were exposed to various levels of
Saikia et al. Forest Ecosystems (2017) 4:28 Page 5 of 12

Fig. 2 Map showing the relationship between a total number of individuals and species diversity per transect in the eastern Himalayan forest

disturbances such as tree felling, fire, and encroachment ecological characteristics of the studied forests are pre-
(Fig. 3a-f). sented in Table 4. All 483 plant species (out of a total
A total number of 82,383 individuals of woody species number of 484 spp.) were distributed contagiously in
were recorded in 354 studied transects with a stand all the studied forest stands with an A/F ratio > 0.05,
density of 465 individuals per ha, whereas the density of whereas Phoebe cooperiana showed a regular distribution
herbaceous species was 375,777 individuals per ha, as with an A/F ratio of 0.004. Out of the total number of 329
recorded in 708 quadrants. The most dominant trees woody plants, only 48 species were recorded in the
were Castanopsis indica (24 individuals per ha), followed seedling stage, competing with other herbs and grasses,
by Quercus semicarpifolia (12 individuals per ha) and which shows that only 14.59% of woody species were
Pinus roxburghii (12 individuals per ha), preceded by regenerating in 177 ha plots (354 studied transects).
various bamboos (Dendrocalamus strictus: 69 individuals The study exhibits distinct stratification (five different
per ha; Bambusa pallida: 16 individuals per ha; Table 3). strata) on the basis of mean height of the species such
In addition, Ageratum conyzoides (an invasive weed) was as emergent (>20 m), canopy (>10–20 m), understorey
identified as the most dominant herb with 12,274 individuals (>5–10 m), shrub (<5 m including sapling of the tree
per ha. The Eastern Himalayan forests are also rich in terms species) and ground (<1 m including the saplings of
of invasive species and some dominant invasive include both tree and shrub species). The highest percentage of
Lantana camara, Ageratum conyzoides, Chromolena species was found in the ground strata (31.66%),
odorata, and Eichhornia cressipes (dominant around higher followed by the understorey (28.60%), and the lowest
altitude lakes). percentage was found in the emergent strata (1.31%)
In the present study, the Shannon-Wiener diversity (Fig. 4). In the emergent layer, there were only 6 species
value for trees and herbs was 4.64 and 4.12, whereas the belonging to timber trees (Tetrameles nudiflora, Acacia
concentration of dominance was 0.02 and 0.03, respectively. pycnantha, Gymnocladus assamicus, Dysoxylum hamil-
The different community parameters including quantitative tonii, Acer campbellii and Albizia lebbeck). Similarly,
the canopy layer was occupied primarily by different
Table 2 Correlation between total number of individuals and timber trees and bamboos, whereas the understory was
species diversity in the studied transects
occupied primarily by Rhododendron spp., small timber
Number of individuals∙ Species diversity class and fruit trees.
transect−1
<10 10–20 20–30 >30 Total The forest ecosystems in the Himalaya have been
<200 37 35 12 10 94 severely affected by natural disturbances including
200–400 18 61 83 74 236 landslides, earthquakes, floods, cloudbursts, heavy rain-
400–600 1 4 3 12 20 falls, as well as different biotic interferences. Landslides
>600 0 2 1 1 4
in the hilly areas and floods in the plain areas during
rainy seasons are recurrent ecological disturbances
Total 56 102 99 97 354
experienced throughout the region. Besides this, the
Saikia et al. Forest Ecosystems (2017) 4:28 Page 6 of 12

Fig. 3 Various disturbances (a) tree felling, (b) landslide due to heavy rainfall, (c) landslide, (d) forest clearance – Jhum, (e) forest fire, and (f) felled
timber in natural forest of Arunachal Pradesh, Northeast India

Ranganadi Dam (a hydroelectric project managed by grids, fire and grazing were recurrent phenomena. NTFPs
NEEPCO) is another major threat to its neighbouring collection by the local people was recorded in all the
forests located at the Lower Subansiri and Papumpare sampled grids. Firewood, resins, wild vegetables, fruits,
districts. Different types of biotic interferences such toko leaves, fodder, medicinal plants etc. were the major
as fire, grazing, jhum, NTFPs collection, hunting, timber NTFPs.
felling, etc. were encountered and observed during the field The resources of the forest are the main livelihood
data collection, which are directly or indirectly influencing source for the people living in the Himalayan region,
the composition of the forest. In some of the sampled including Arunachal Pradesh. It must be noted that a

Table 3 List of 10 most important tree species and their density, frequency, dominance and IVI
Name Family Density RD Frequency RF Dominance RDm IVI
Dendrocalamus strictus Poaceae 68.95 14.81 47.18 3.64 55.68 1.72 20.18
Duabanga grandiflora Lythraceae 9.26 1.99 21.75 1.68 296.14 9.15 12.82
Bombax ceiba Malvaceae 6.58 1.41 22.32 1.72 285.51 8.82 11.95
Castanopsis indica Fagaceae 23.81 5.12 24.86 1.92 153.35 4.74 11.77
Terminalia myriocarpa Combretaceae 10.40 2.23 30.79 2.38 224.53 6.94 11.55
Abies alba Pinaceae 6.33 1.36 3.67 0.28 178.06 5.50 7.14
Gmelina arborea Verbenaceae 5.21 1.12 16.10 1.24 136.75 4.22 6.59
Unknown Unknown 9.76 2.10 33.62 2.59 35.83 1.11 5.80
Pinus roxburghii Pinaceae 11.64 2.50 8.19 0.63 85.64 2.65 5.78
Quercus semicarpifolia Fagaceae 12.19 2.62 15.25 1.18 48.76 1.51 5.30
Saikia et al. Forest Ecosystems (2017) 4:28 Page 7 of 12

Table 4 Community characteristics of Eastern Himalayan forests tulda, B. pallida etc. The area is also rich in terms of
of Arunachal Pradesh, NE India various IUCN red-listed species, including Aquilaria
Parameter Woody plants Non-woody malaccensis, Begonia tessaricarpa, Gledetsia assamica,
plants Gymnocladus assamicus, Livistona jenkinsiana, Rhodo-
Total number of plots studied (No.) 354 transects 708 quadrats dendron meddenii, Rhododendron thomsonii, and Piper
Size of sampled area (m2) 5000 1 wallichii, which were collected primarily from the Dirrang,
Actual sampled area (ha) 177 0.0708 Tawang, Yamcha, Itanagar, Along and Mebo areas in
Dominant species Dendrocalamus Ageratum
Arunachal Pradesh. Some threatened species of Arunachal
strictus conyzoides Pradesh are presented in Fig. 5a-f).
Dominant family Ericaceae Asteraceae Out of the total number of 354 studied transects, 15
transects were laid out in alpine vegetation (>2800 m
No. of families (identified species) 74 55
altitude), 35 transects in temperate forests (m altitude),
Species richness 329 153
70 transects in sub-tropical forests (m altitude), and 234
Density (individual∙ha−1) 465 375,777 transects in tropical forests (m altitude) including 2 tran-
Basal area of trees (m2∙ha−1) 3237.58 – sects in a non-vegetation zone (covered by the river Siang).
Diversity (Shannon’s H′) 4.64 4.12 The species richness study exhibits the low variation in the
Concentration of dominance 0.02 0.03 alpine vegetation (the richness ranges from 2 to 21; mean
(Simpson index) 9 ± 1.57 SE), the moderate variation in the temperate
Evenness Index 0.81 0.78 forests (5 to 44; mean 16 ± 1.21 SE), the high variation in
Species richness Index (Margalef 28.01 15.81
the sub-tropical forests (5 to 53; mean 24 ± 1.27 SE), and
index of richness) the very high variation in the tropical forests (0 to 64; mean
23 ± 0.84 SE) (Table 6). The 15 studied transects in the
alpine forests (7.5 ha area) are located in the West Kameng
wide variety of medicinal plants used to be collected by and Tawang districts with a loamy soil covered by mixed
the local people, to cure various diseases ranging from semi-evergreen forests, where tree density ranges from 196
stomach ailments to blood pressure problems, as recorded to 600 individuals per ha (mean 407 ± 28.40 SE). There is
during an informal discussion with the local villagers. occasional biotic interference in the studied forests, mainly
Some of the most important medicinal plants recorded due to collection of fuelwood and grazing. Occasional
were Houttynia cordata, Drymaria cordata, Commelina NTFPs collection, looping, soil removal as well as forest fire
benghalensis, Centella asiatica, Ageratum conyzoides, were also evident in some forest patches, though tree
Scoparia dulcis, Spilanthesp aniculata, etc. All plant felling was not observed in any of the studied forests.
species ranging from medicinal herbs to timber trees are Similarly, the 35 studied temperate forests (17.5 ha
closely linked with the economic life of the local people. area) were located at the West Kameng, West Siang
Details of some important plant species commonly used and Lower Subansiri districts, with loamy to clayey soil
by the local villagers in the day to day necessities and sub- with humus, covered mainly by mixed to coniferous
sistence generation are mentioned in Table 5. semi-evergreen forests as well as grasslands, where the
The Eastern Himalayan forest in Arunachal Pradesh, tree density ranges from 0 to 1220 individuals per ha
Northeast India, is also rich in terms of various bamboos. (mean 413 ± 41.75 SE). The two transects of Mechukha
In the present study, a total of 14 different bamboo species in West Siang were laid out in grasslands used primarily
were recorded, including Dendrocalamus strictus, Bambusa by locals for grazing and with no tree covers. Looping,

35

30
Percentage of species

25

20

15

10

0
Emergent Canopy Understorey Shrub Ground
Strata/ Layers

Fig. 4 Distribution of species in different strata of studied forests according to their mean height
Saikia et al. Forest Ecosystems (2017) 4:28 Page 8 of 12

Table 5 Some of the economically as well as locally used important plants species recorded from the sampled grids
Timber Dipterocarpus macrocarpus, Terminalia myriocarpa, Lagerstroemia speciosa, Morus laevigata, Gmelina arborea,
Pterospermum acerifolium, Toona ciliata, Altingia excelsa, Duabanga grandiflora, Shorea assamica, Shorea robusta,
Schima wallichii, Mesua ferrea, Stereospermum chelonoides, Kyida glabreseeus, Tectona grandis, Pinus roxburghii,
Castonopsis indica, Bombax ceiba, Dulbergia sissoo, Michelia champaca, Anthocephalus chinensis
Medicinal Oroxylum indicum, Terminalia balerica, Terminalia chebula, Phyllathus emblica, Xanthium strumarium,
Commelina benghalensis, Cynodon dactylon, Centella asiatica, Chelerodendrum viscosum, Leucus aspera,
Rhododendron arboreum, Garcinia cowa, Azadirachta indica, Callicarpa arborea, Caryota urens, Cyathea spinulosa,
Dillenia indica, Spilanthes paniculata, Piper wallichii, Piper sylvaticum, Piper mullsua, Valeriana wallichii, Cassia fistula
Fuel wood and fodder Albizia lebbeck, Alnus nepalensis, Rhododendron arboreum, Cassia fistula, Alstonia scholaris, Castonopsis indica,

Pinus roxburghii, Pinus wallichiana, Quercus semicarpifolia, Mallotus phillipensis, Macaranga denticulata,
Lantana camara, Zizyphus mauritiana, Bambusa spp.
Dye yielding Daphne papyracea, Rubia manjith, Juglans regia, Bischofia javonica
Thatch Impereta cylindrica, Livistona jenkinsiana

grazing and fuelwood collection are the common biotic deciduous, broadleaf semi-evergreen, coniferous evergreen
interferences in this region. The 70 studied transects in and coniferous semi-evergreen. On the other hand, most
the sub-tropical forests (35 ha area) were located in 12 of the transects (234 transects covering 116 ha area) were
different districts of Arunachal Pradesh, where tree laid out in tropical forests covering nine different districts
density ranges from 192 to 902 individuals per ha of Arunachal Pradesh, where the tree density ranges from
(mean 419 ± 15.29 SE), with diverse vegetation types 0 to 650 individuals per ha (mean 352 ± 10.63 SE) and
including mixed semi-evergreen, mixed evergreen, mixed the vegetation is mainly broadleaf semi-evergreen, mixed

Fig. 5 Some threatened species of Arunachal Pradesh (a and b) tree, pods and seedling of Gymnocladus assamicus and Gymnocladu schinensis;
(c) Leaves and Pods of Gledetsia assamica; (d) Tree and harvested leaves of Livistona jenkinsiana; (e) Rhododendron fulgens; (f) Rhododendron meddenii
Saikia et al. Forest Ecosystems (2017) 4:28 Page 9 of 12

Table 6 Species richness and stand density of studied forests at different altitudinal gradients of Arunachal Pradesh, Northeast India
Climatic No. of transects Altitude Located districts Species richness (No.) Stem density
zone studied range (m) (≥10 cm GBH)
(individuals∙ha−1)
Alpine 15 2803–4161 Tawang and West Kameng 2 to 21 196–600
(mean 9 ± 1.57 SE) (mean 407 ± 28.40 SE)
Temperate 35 1824–2788 West Kameng, West Siang and Lower Subansiri 5 to 44 0–1220
(mean 16 ± 1.21 SE) (mean 413 ± 41.75 SE)
Sub-Tropical 70 802–1800 West Kameng, East Kameng, West Siang, East Siang, 5 to 53 192–902
Upper Siang, Lower Subansiri, Upper Subansiri, (mean 24 ± 1.27 SE) (mean 419 ± 15.29 SE)
Lower Dibang Valley, Anjaw, Papumpare,
Tirap and Changlang
Tropical 234 87–795 East Kameng, West Siang, East Siang, Upper Siang, 0 to 64 0–650
Upper Subansiri, Lower Dibang Valley, Lohit, Papumpare, (mean 23 ± 0.84 SE) (mean 352 ± 10.63 SE)
Tirap and Changlang

semi-evergreen, broadleaf deciduous, broadleaf evergreen, Despite the high level of disturbance due to tree felling,
and evergreen bamboo to grassland. In some grids, the fire, and encroachment, the higher diversity in the Lohit,
tree density was zero, as 12 transects were laid out in Changlang and Tirap districts may be due to the higher
grassland having no tree cover. Most of the tropical and rainfall as compared to other parts of the state, which
sub-tropical forests were experiencing different degrees promotes soil nutrient and pH to facilitate organic matter
of anthropogenic pressure, including looping, fuelwood decomposition. In general, tree density varies with forest
collection, fire, NTFPs collection, soil removal, grazing, community type, forest age class, tree species and size
litter collection, tree felling etc., which are influencing class, site history, site condition, and other factors. The
the forest diversity of the region. recorded tree density (465 individuals per ha) in the
present study is comparable with the reported value from
other tropical forests (Saxena and Singh 1982; Singh et al.
Discussions 1984; Parthasarathy et al. 1992; Swan Jr, 1988; Sharma et
Understanding species diversity and distribution patterns al. 2014), but it is lower than the recorded density of
is significant for helping managers to evaluate the com- Northeast Indian forests (Bhuyan et al. 2003; Duchok et
plexity as well as the prospects of forest ecosystems. The al. 2005; Kumar et al. 2006). However, the reported
level of species richness in the studied forests can be 3237.58 m2∙ha−1 basal cover of 82,383 individuals of
attributed to microclimatic variations at different altitudinal woody species was found to be significantly higher than
gradients. Studying the microclimate is important because the recorded basal cover of tropical forests (Bhuyan et al.
of its significant effects on species richness and ecosystem 2003; Kumar et al. 2006). Differences in the basal area
functions. Apart from microclimatic factors, plant species may be attributed to altitudinal variations, species com-
may respond to fine-scale variation in pH (Jansen and position, age structure and successional stage of the forest
Oksanen 2013), and soil moisture and nutrients (Fischer et (Swamy et al. 2000). The Shannon-Wiener diversity index
al. 2014). A higher diversity of woody plants was is generally high for tropical forests of the Indian subcon-
observed in the present study, as compared to herbaceous tinent and ranges from 0.81 to 4.1 (Singh et al. 1984;
species; however, reverse data (herbaceous plants > trees Parthasarathy et al. 1992; Bhuyan et al. 2003). The recorded
and shrubs) was presented from Northwest Himalayan Shannon-Wiener diversity values for trees and herbs in the
forests of India (Sharma et al. 2014). The greater richness present study were 4.64 and 4.12, which was slightly higher
of woody plants, particularly observed in the trees, could than the previously recorded values. In general, species
also be the result of the succession process that tends diversity and concentration dominance show an inverse
to increase species diversity in the studied forests. The relationship (Singh and Mishra 1969; Joshi and Behera
present study recorded Fabaceae as the most species-rich 1991). The values of species diversity corresponded to
family with 27 spp. and 49 monotypic families, whereas the general trend i.e., tree > herb, while the concentration
Asteraceae was the dominant family with 49 spp. and 28 dominance displayed the opposite trend i.e., herb > tree in
monotypic families in the Western Himalayan forests of the investigated forests. The high diversity and low con-
India (Sharma et al. 2014). This difference may be due to centration dominance in the analyzed forests may be due
the variations in terms of growth forms, as trees and to different levels of anthropogenic pressure in different
shrubs are dominant in the Eastern Himalayan forests, forests at varied locations. Dispersal limitation is an
whereas herbs are dominant in the Western Himalayan important ecological factor for controlling species’
forests. distribution pattern (Hubbell et al. 1999). Contagious
Saikia et al. Forest Ecosystems (2017) 4:28 Page 10 of 12

distribution has been accepted as a characteristic pattern highest density of trees was recorded in the forests of
of plant occurrence in nature (Odum 1971); it is an indi- temperate zones as temperate zones are dominated mainly
cation of clusteredness. Only 14.59% of woody species by coniferous species with straight stem/bole and can
were recorded in a seedling stage, which signifies a very accommodate more plants. Tree density varied greatly in
poor rate of regeneration of woody species in the 177 ha the middle altitude (1824–2788 m altitude) forest ranges
studied plots (354 transects). This may be due to the from 0 to 1220 individuals per ha (mean 413 ± 41.75 SE) as
uneven timing of sampling (different months during June two transects of Mechukha, West Siang were laid out in
2007 to April 2010). On the other hand, seedling recruit- grasslands used by locals as grazing land and having no
ment was poorer in the higher altitude forests due to the tree cover. Similar findings have also been reported by
thick ground cover of leaf litter (mainly in coniferous Singh et al. (1994) for the Central Himalayan forests.
forests).
Multilayered canopy configuration is characterized by the Conclusion
presence of a large number of heterogeneous microsites, The study exhibits that the Eastern Himalayan forests
which signifies the suitability and ecological efficiency of are very rich in terms of floristic composition (482 plant
the system in terms of sharing, utilization and conservation species). The species richness decreases with the increasing
of the available resources like sunlight, water and soil elevation. Species richness and distribution patterns of
(Wiersum 1982; Torquebiau 1992). A heterogeneous plants are largely regulated by the altitude and environ-
canopy structure is a relevant attribute of tropical forests mental factors. Knowledge of the relative contribution of
in physiognomic terms. In addition, the degree of vertical spatial and environmental variation on the species’ com-
stratification is an important factor for maintaining higher positional patterns along large-scale gradients would help
woody species diversity in natural forests (Feroz et al. forest managers to understand the processes that primarily
2006, 2008). In conformity with the present study, the maintain plant species patterns. The study also confirms
tropical wet evergreen forests of Bangladesh also showed that the Eastern Himalayan forests are extremely rich in
five distinct strata, with the highest species diversity in the endemic as well as non-endemic species. This needs regu-
ground strata (Feroz et al. 2016). Generally, the lower lar intensive monitoring and management to conserve the
strata support the high species richness in a forest fragile ecosystems from the ever-increasing anthropogenic
(Feroz et al. 2006). On the other hand, Feroz et al. pressure in changing climatic conditions.
(2015) recorded four distinct strata in the subtropical
evergreen broadleaf forests of Japan. Abbreviations
Elevation gradients are ideally suited for examining GBH: Girth at Breast Height; IUCN: The International Union for Conservation
of Nature; NEEPCO: North Eastern Electric Power Corporation Limited;
biodiversity drivers, as elevation is correlated with various NTFPs: Non-Timber Forest Products; spp.: Species
environmental variables including competition, mutualism,
ecotone effects, temperature and rainfall, which determine Acknowledgements
energy availability and ecosystem productivity (McCain Financial support from DBT, Govt. of India, through the project ‘Mapping and
quantitative assessment of geographic distribution and population status of
and Grytnes 2010). Two main patterns of species richness- plant resources of Eastern Himalayan region’ is highly acknowledged
elevation relationships are dominant in the relevant (Reference no. BT/PR7928/NDB/52/9/2006 dated 29.09.2006). The authors
literature, i.e. decreasing and hump-shaped (Rahbek 1995). acknowledge the support and assistance of Ziaur Rahman Laskar, Salam
Dilip, Bijit, Bironjoy and Samar during field visits of the study.
Incomplete sampling of environmental gradients can dir-
ectly bias the resulting pattern. The present study confirms Availability of data and materials
that species richness decreases with increase in altitude, in The data is shared with Global Forestry Biodiversity Initiative (GFBI) and
conformity with earlier studies (Bachman et al. 2004; available upon request to the Corresponding Author.

Jacquemyn et al. 2005; Trigas et al. 2013). With the

Authors’ contributions
increase in altitude, species diversity decreases as a result JD and SB participated in the data collection, analysis and interpretation,
of environmental filtering, and because the higher altitude species identification and herbarium processing; AK prepared the maps and
ecosystems have been mainly colonized by plant species also helped in the manuscript preparation and statistical analysis; PS entered,
analysed and formatted data and also drafted the manuscript. OPT and LBS
tolerant of extreme weather conditions. Differences in participated in field design, coordinated during data collection; SD helped in
species diversity among communities generally resulted data presentation and also finalized the manuscript, and MLK conceived the
from variations in site specificity (Denslow 1980). More- study, coordinated among other teams (as it is part of a mega project
conducted in eight Northeastern states) and forest officials of different parts
over, species diversity is often correlated with the rainfall of Arunachal Pradesh and also contributed in finalizing the manuscript.
and nutrient status of the site (Hartshorn 1980). On the All authors read and approved the final manuscript.
other hand, the density of trees increases with altitude
from the tropical to the temperate range, and then it Funding
This study is an output of a project funded by the Department of
decreases in the alpine zone as the vegetation is usually Biotechnology (DBT), Government of India, New Delhi (Reference no. BT/
sparse above an altitude of 3000 m (Singh et al. 1994). The PR7928/NDB/52/9/2006 dated 29.09.2006).
Saikia et al. Forest Ecosystems (2017) 4:28 Page 11 of 12

Authors’ information Feroz SM, Yoshimura K, Hagihara A (2008) Stand stratification and woody species
P. Saikia is Assistant Professor in the Centre for Environmental Sciences, diversity of a subtropical forest in limestone habitat in the northern part of
School of Natural Resource Management, Central University of Jharkhand, Okinawa Island. J Plant Res 121:329–337
Ranchi, Jharkhand, India. J. Deka is Assistant Professor in the Department of Feroz SM, Kabir ME, Hagihara A (2015) Species composition, diversity and
Botany, University of Science and Technology, Meghalaya (USTM), stratification in the subtropical evergreen broadleaf forest along a latitudinal
Meghalaya, India. S. Bharali is working in the Rubber Board of India, Regional thermal gradient in the Ryukyu archipelago. Japan Glob Ecol Cons 4:63–72
Office, Dimapur-797,112, Nagaland, India. A. Kumar is Assistant Professor in Feroz SM, Mamun AA, Kabir ME (2016) Composition, diversity and distribution of
the Centre for Land Resource Management, School of Natural Resource Man- woody species in relation to vertical stratification of a tropical wet evergreen
agement, Central University of Jharkhand, Ranchi, Jharkhand, India. O.P. Tri- forest in Bangladesh. Global Ecol Conserv 8:144–153
pathi and L.B. Singha are Associate Professors in the Department of Forestry, Fischer H, Micler B, Ewald J (2014) Environmental, spatial and structural components in
North Eastern Regional Institute of Science & Technology, Nirjuli, Arunachal the composition of mountain forest in the Bavarian alps. Folia Geobot 49:361–384
Pradesh, India. S. Dayanandan is Professor in the Centre for Structural and Hartshorn GS (1980) Neotropical forest dynamics. Biotropica (Supplement) 12: 23–30
Functional Genomics, Biology Department, Concordia University, Montreal, Heady HF (1958) Vegetational changes in the California annual type. Ecology
Canada. M.L. Khan is Professor in the Department of Botany, Dr. Harisingh 39(3):402–416. https://doi.org/10.2307/1931750
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Ethics approval and consent to participate
Hubbell SP, Foster RB, O'Brien S, Wechsler B, Condit R, Harms K, Wright SJ, Loo de
Not applicable.
Lau S (1999) Light gaps, recruitment limitation and tree diversity in a
Neotropical forest. Science 283:554–557
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Not applicable. a dry afromontane coniferous forest in Bale Mountains, Ethiopia: implication
to biodiversity conservation. SINET: Ethiop J Sci 30:1–12
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Competing interests species diversity, breeding system and floral traits of orchid species on
The authors declare that they have no competing interests. Réunion Island. J Biogeogr 32:1751–1761
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