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	_:	@Relation of forest structure and soil properties in natural, rehabilitated and degraded forest 
Ashish K. Mishra a, b *, Soumit K. Behera a, Kripal Singh a, Nayan Sahu a, Omesh Bajpaia, Anoop Kumar a, R.M. Mishra b, Rashmi Arnold b, L.B. Chaudhary a, Bajrang Singh a
a CSIR-National Botanical Research Institute, Rana Pratap Marg, Lucknow 226 001, Uttar Pradesh, India
b Awadesh Pratap Singh University, Rewa 486006, Madhya Pradesh, India 
* Corresponding author at: 
Plant Diversity, Systematics & Herbarium
CSIR-National Botanical Research Institute, Rana Pratap Marg 226001
Tel.: +05222297960; fax: +91 522 2205836, 2205832. 
Email address: HYPERLINK "mailto:ashishmishramlg@gmail.com"ashishmishramlg@gmail.com (A. K. Mishra) 











Abstract
Plant community structure, species diversity and soil properties of natural, rehabilitated and degraded forests were studied in order to assess relationship of different forests structure and soil properties in upper Indo-Gangetic plain of Uttar Pradesh state of India. The composition of species, population abundance and importance value of the woody species were investigated in each forest; the representative soil samples from 30 cm depth of each forest were also analyzed for the soil properties. The species dominance was shared with many species in the natural and rehabilitated forests and remained confined to a few species in degraded forest as indicated by the Simpson index. The�Shannon-Wiener index of general diversity ranged from 1.95 to 3.36 for degraded to natural forest respectively. Tree density (813 stem ha-1) and basal area (62.32 m2 ha-1) both were highest in natural forest and lowest in degraded forest (656 stem ha-1 and 19.27 m2 ha-1). Principal Component Analysis was applied to observe the vegetation soil interactions which reveal that soil properties (bulk density, soil organic carbon SOC, available phosphorus, pH and cations Na+, K+) having significant correlation with forest association as well as their population distribution pattern. The results indicate that each forest shows a close affiliation to variable soil properties. 
Keywords: forests structure, importance value index, microbial biomass, Principal component analysis (PCA), soil organic carbon, phytosociology 










1 Introduction
Tropical forests are exploited with an alarming rate beyond its natural recovery process [7]. These forests are as important as our lungs for vital life, which always support a variety of life forms (biodiversity) in this magnificent nature [32]. Indian forests are being exploited to significant extent by anthropogenic activities [40]. Protection of natural forests becomes today�s need because they are important in environmental conservation, ecological balance, supplying the goods and ecosystem services. The wide-spread degradation in Indian forests has threatened many endemic species [30].
Forest community structure, species composition and their biodiversity studies denote the importance of individual species and their population to assess the status of any forest [18]. Biodiversity has been related with the ecosystem functioning [29] and forest phytodiversity is interdependent on community structures even in the same region [28]. Structure and species diversity have strong correlation in tropical forest [27]. Biodiversity conservation or ecological restoration has been recently given more importance in the management of forest ecosystems due to extinction of many important taxa. Therefore, restoring, enhancing and maintaining the health of forests benefit the environment and creates biodiversity rich ecosystems. Restoration is the process of assisting the recovery of an ecosystem that has been degraded or destroyed [41]. 
Forest structure is developed with the influence of several factors including soil properties [4]. Soil physical properties play an important role in operating the biological processes in the soil [9], whereas chemical and biological properties contribute mainly nutrient requirement of plants [15]. The correlation between tree species association and soil variables has been successfully demonstrated in numerous studies of tropical forests world-wide [1, 11, 26]. When new forest are established on degraded lands, after assessing the status of recovery and correlating the vegetation structure with soil properties we can manipulate the soil ecosystem to enhance the process of restoration [36, 37, 38]. Along with the competitive effect and climatic parameters forest soil also play a leading role in species association [47]; therefore, knowing the soil characteristics and soil conservation strategy may be helpful in maintaining the sustainable land use systems. Tree species association can also be collated with the climatic variables [29]. In this paper we have studied forest structure, species composition and soil properties of three different forests to establish a relation between vegetation structure and soil properties.
2 Materials and methods
2.1 Site and forest description
In this study we have selected natural, rehabilitated and degraded forests in Upper Gangetic plain region of Uttar Pradesh state, India. Natural forest is situated in Katerniaghat wildlife sanctuary, Bahraich, whilst, rehabilitated and degraded forests are located in Lucknow. Detail of each forest is presented below in separate sections. All the three forests are reserved forest since last 30 years without any biotic disturbance and exist on an inherently same type of Gangetic alluvial soil. The atmosphere temperature is almost identical but rainfall at rehabilitated and degraded forests differs significantly from the natural forest. 
2.2 Natural forest
The�Katerniaghat Wildlife Sanctuary�(KWLS) is a protected area in the Upper�Gangetic plain in Uttar Pradesh, India and covers an area of 440 km2 in Bahraich district.�It was declared as a Sanctuary in 1976. KWLS comprises mesmerizing mosaics of Sal and Teak forests, mixed natural forests, lush grasslands, steaming swamps and wetlands [5, 6, 44]. We selected a natural forest in this Sanctuary lies between 28� 6' to 28� 24' N and 81� 24' to 81�19' E. We have already studied plant communities and response of different forest community association to microclimate [3], phenology of dominant species [2] and diversity of Ficus species [17] of different forests in KWLS. Annual temperature of KWLS varied from maximum 40.50C to minimum 3.5 0C with a mean annual rainfall of 1495 mm. 
2.3 Rehabilitated forest
Rehabilitated forest is a 50 year old forest developed on a sodic wasteland at Banthra Research Station of National Botanical Research Institute, Lucknow (26� 40' to 26� 45' N; 80� 45' to 80� 53' E). This forest is a semi natural forest in which a few mixed species were planted initially and later on allowed natural succession over the years [43]. This rehabilitated forest now accommodates several species belong to about 30 families in ground layer, understory and upper story vegetation [37].
2.4 Degraded forest
We selected a degraded forest at Lucknow in Kukrail crocodile rehabilitation and recreation center�s (26� 55.9' N; 80� 57.5' E). It was poorly developed on a barren land and for recreation purpose and also works as a crocodile rehabilitation centre. Before 30 years it was highly degraded forest where a high, biotic pressure animal grazing, vegetation exploitation and nutrient deposition have created disturbance in natural succession and therefore species richness decreased considerably and dominance was concentrated within a few species in the last 30 years when state government acquired and conserved as reserves forest. 
2.5 Phytosociological study
Phytosociological parameters were studied by laying out 30 quadrats of 20 x 20 m2 size for trees along a line transect in each forest. The size and number of quadrat were standardized using the species area curve [20]. All trees e" 5 cm diameter at breast height (dbh = 1.37 m from ground) were recorded in each quadrat. Voucher specimens of plant species were collected from studied forests and identified from herbarium of CSIR-NBRI and regional flora. Quadrat data were processed mathematically for deriving phytosociological indices such as importance value index (IVI), density, frequency, abundance and basal area [8]. The IVI for the tree species was determined as the sum of the relative frequency, relative density and relative basal area [8]. Jaccard similarity index was used to determine the floristic similarity [22]. The tree species diversity was determined by using Shannon Wiener information function H (H = -" pi ln pi; where, pi= ni/N; and ni = IVI of each species, N= total IVI of all species) [31]. Concentration of dominance was measured by Simpson s index through the formula (Cd = " (ni/N)2) [33]. 
2.6 Soil sampling and analysis
In each forest five plots of 5 X 5 m were randomly selected and composite soil samples were collected from these plots. Three soil samples (0-30 cm) were collected at each plot, mixed thoroughly, homogenized and stored as one composite sample. These soil samples were air dried, ground, and then passed through a 2-mm sieve to remove gravel and boulders. Soil pH was measured in a 1: 2 suspension of soil and de-ionized water using a glass electrode. Organic carbon was determined by Walkley and Black method, [46]; total N was determined by macro-Kjeldahl technique, available phosphorus was estimated by phosphomolybdic blue colorimetric method; potassium (K) and sodium were measured by flame photometry [13]. Microbial biomass carbon (MBC) was estimated by chloroform fumigation and incubation method [14]. 
2.7 Statistical analysis
The values are given as means for each parameter along with standard deviation. The 10 soil physic-chemical and biological parameters were condensed in three forest type using Principal Component Analysis (PCA) by PAST 2.00 [12]. Correlation between forest structure attributes and soil variables was analyzed by statistical analysis package SPSS 16.
3 Results
3.1 Community structure and diversity
Number of species, population density, basal area and similarity index across the different forest types are shown in Table 1. The highest number of species (58) was found in the natural forest followed by rehabilitated (26) and degraded forests (17). The greater Simpson index of 0.24 in degraded forests indicates that species dominance is concentrated among the few species in comparison to the other two forests with lower values (0.08-0.09). The highest value (3.36) of Shannon diversity index in natural forest shows a good integration of species and their population in comparison to degraded forest (1.95). Jaccard similarity value 0.26 was higher in rehabilitated - degraded forests and 0.1 was lower in natural- degraded forests (Table 1). Moraceae was found common dominant family whereas Euphorbiaceae and Rubiaceae were co-dominant families. The tree density ranges from 656 � 813 stem ha-1 and basal area ranged from 19.27-62.32 m2 ha-1 from degraded to natural forest, respectively.  Natural forest appears to be older than other two forest types as the tree population was extended up to a relatively high range (5.15-164.65 cm) leading to greatest basal area per hectare. 
On the basis of importance value index (IVI) Mallotus philippensis, Syzygium cumini and Shorea robusta were the dominant tree species of natural forest. Syzygium cumini and Terminalia arjuna were dominant species of rehabilitated forest and Pongamia pinnata was dominant tree species in the degraded forest (Fig. 1). Only five tree species were common among three forest types i.e. HYPERLINK "https://www.google.co.in/search?q=bauhinia%20variegata&start=0&spell=1&biw=1440&bih=605"Bauhinia variegata, Dalbergia sissoo, Ficus glomerata, Pongamia pinnata and Holoptelea integrifolia. Importance value index of total 75 tree species in all three forest sites are presented in Table 2, www hich indicates the diversity and dominance of different species among the three forests.
3.2 Soil characteristics 
Physico-chemical and biological properties of three forest soils were significantly different (Table 3). Natural and degraded forests have slightly acidic to neutral soil pH (6.74 and 6.78), whereas rehabilitated forest soil was alkaline (pH = 7.8). EC of rehabilitated forest soil was significantly higher in comparison to soils of degraded and natural forest. Soil BD was significantly higher at degraded and rehabilitated forests in comparison to natural forest. These forests are characterized by having moderate WHC (42% to 46%) with highest values in natural forest and lowest in degraded forest. AP was significantly higher in degraded forest (133.5�g g-1) in comparison to rehabilitated (47 �g g-1) and natural (28 �g g-1) forests. However, soil organic carbon (SOC) was significantly higher in natural forest than that of rehabilitated and degraded forest soils. Similarly, soil nutrients like N, P and K were also higher in natural forest than degraded and rehabilitated forests. The concentration of exchangeable Na was higher in rehabilitated forest soil. Soil microbial biomass (MBC) was highest in natural and lowest in degraded forest.
3.3 Correlation between vegetation and soil variables 
Principal Component Analysis (PCA) ordination of three different forest on the basis of their soil properties is presented in Fig 2. Eigen values of dominant axis were 6.14, 3.85 and 2.62. Result of variance among all axes shows that axis 1 and axis 2 covered about 95% of the total variation. Pearson�s correlation coefficients between forest structure attributes (species richness, stem density, basal cover and biodiversity indices) and ten soil variables were studied (Table 4). Among the fifteen variables species richness (R) was significantly positively correlated with SOC (p < 0.05). Stem density (D) was significantly positively correlated with WHC (p < 0.05) and significantly negatively correlated with BD (p < 0.05) and AP (p < 0.01), whereas basal area (BA) was significantly negatively correlated with MBC (p < 0.01). In biodiversity indices, C (Simpson index) and H� (Shannon Wiener index) were significantly positively correlated with BD (p < 0.05) and WHC (p < 0.05), respectively; while H� was significantly negatively correlated with soil AP (P< 0.01) and BD (p < 0.05). 
4 Discussion
4.1 Community structure 
Natural forest constitutes more than double species as also basal area being situated on good alluvial soil and receiving higher rain fall than the rehabilitated and degraded forest, which were developed on degraded alluvium. Dominant tree species of the three forests (Mallotus philippensis, Syzygium cumini, Pongamia pinnata, Shorea robusta and Terminalia arjuna) have been classified in the natural vegetation of the northern sub-tropical forest in India [5]. Degraded forest was dominated by Pongamia pinnata, which is established easily on degraded alluvium. It casts much shade which does not favor the growth of many other species. Species structure, population abundance and basal area are almost reasonable inal and rehabilitated forests compared to other tropical forest ecosystems [39, 42]. But degraded forest consisted of relatively low biodiversity indices values. Diversity index of the present study lies within the range reported for tropical forests which was lowest (1.95) for degraded and highest (3.36) in natural forest. A high diversity index in tropical forests was reported in the range of 5.06 to 5.40 [16] but for Indian tropical forests a lesser range (0.83 to 4.1) was observed [24, 34, 45]. In natural and rehabilitated forests species are almost uniformly distributed over the area but degraded forest showed an inequitable distribution and lower richness. Although, environmental factors play important role in the composition and distribution of species [23] and soil is one of them which exerted a profound effect on the organization of plant communities as well.
            Population density and species dominance are the important indicators of phytosociology which describe the structure of a forest, on which multiple functions are operated in the forest ecosystem. Population density of natural forest was relatively high due to relatively high rainfall, as well as soil fertility status whereas lowest population density of degraded may be associated with poor soil structure, low rainfall and some biotic disturbances during early forest formation. The occurrence of dominant tree species such as Mallotus philippensis, Pongamia pinnata, Syzygium cumini, Shorea robusta and Lagerstroemia parviflora and co-dominant species like Ehretia laevis, Diospyros tomentosa, Tectona grandis, Streblus asper develop the tropical deciduous nature of these forests. IVI value of dominant species revealed that natural and rehabilitated both are miscellaneous forest, whereas degraded was dominated by Pongamia pinnata and Holoptelia integrefolia. On the basis of Jaccard similarity index rehabilitated and degraded forests were much similar whereas natural and degraded forests were much dissimilar due to significant variation in soil properties and precipitation. However, due to the introduction of many species in the rehabilitated forest, for the restoration aim [37], species diversity indices (Simpson and Shannon) were higher than degraded forest and lower than natural forest.
As compared to soil properties of dry tropical forest [7] we have found soil physico-chemical and microbial properties of natural forest are indicative of soil with relatively higher values. However, soil properties of rehabilitated forest were quite similar and of degraded forest were lower. Rehabilitated forest was established on sodic land [43] so it has higher value of pH, EC and Na which was quite similar with previous study [37]. Natural forest having higher SOC [21]due to higher litter fall therefore higher SOC value in natural forest may be due to high stem density and species richness of deciduous plants. MBC of any forest may have significantly influenced nutrient availability to forest community [35]. Similarly, we have found natural forest has higher MBC value with high TN and K whereas degraded forest has lowest MBC values with lowest nutrient availability. 
4.2 Relation of soil properties with vegetation pattern
PCA results revealed PCA axis 1 highly correlated with K, TN and SOC whereas PCA axis 2 with pH and Na. (Table 4). In figure 2 PCA ordinations clearly show the correlation between different forest and soil properties. Natural forest regulated by SOC, MBC and K may be due to high species richness and stem density, whereas; vegetation composition of rehabilitated forest was regulated by pH, EC and exchangeable Na. Soil pH, available P and K are the main factors affecting the variation in species richness in degraded forest [19]. In our study degraded forest vegetation was found more closely associated with BD and available P. Soil properties are the major controlling agents for species distribution in tropical forest [15]. Results of Table 5 show the correlation between forest structure and soil variables. Species composition and age of forest are the main factors in difference of SOC in different forest types [25]. In this study SOC significantly positively correlated with species richness. Stem density and Shannon diversity index significantly negatively correlated with BD and positively correlated with WHC which were agreement with study of [10] in which they have studied revegetating sodic wastelands at Lucknow (UP). Thus the forest structure attributes are are  a closely related to many soil parameters. Soil pH, available P and K are the main factors affecting the variation in species richness in degraded forest [19]. Summarized explanation of forest structure and soil properties correlation results could be that soil conditions differs between the forest types. It should be pointed out, that forest composition not only affects soil but that soil properties also govern forest composition and diversity [48]. The relationships between forest vegetation and edaphic factors established herewith may be utilized for the conservation of preferred species in the upper gangetic plain region of India.
5 Conclusions
Soil properties and vegetation although are closely related to each other but different forests had variable association with soil parameters, such as SOC, MBC and K in natural forest, Na, pH and EC in rehabilitated forest and BD and AP in degraded forest. Soil properties and vegetation composition in the 50 year old rehabilitated forest were in between the degraded and natural in forests. Restoration of degraded land to a functioning ecosystem is often assessed from the natural forests to judge the recovery status of restoration in this scale the rehabilitated forest has achieved close affinity with natural forest during 50 years.
Acknowledgements
The authors are grateful to Dr. C. S. Nautiyal, Director, CSIR-National Botanical Research Institute, Lucknow, India for providing necessary facilities and encouragement. Thanks are also due all the Forest Officers of Katerniaghat Wildlife Sanctuary and Kukrail Reserve Forest for their valuable support. The fund was received from CSIR, New Delhi to carry out this work under NWP-020.
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Figure  legends
Fig. 1 
Importance value index [IVI = relative frequency (RF) + relative density (RD) + relative basal cover (RBC)] of 10 dominant species in (a) natural (b) rehabilitated and (c) degraded forest. Tree species,  S.C., Syzygium cumini; T.A., Terminalia arjuna; L.L., Leucaena leucocephala; H.I., Holoptelea integrifolia; B.V., Bauhinia variegata; A.I., Azadirachta indica; P.R., Putranjiva roxburghii;  P.P., Pongamia pinnata; S.H., Syzygium heyneanum; E.O., Emblica officinalis; H.I., Holoptelea integrifolia; H.A., Haplophragma adenophyllum; P.J., Prosopis juliflora; F.H., Ficus hispida;  K.S., Kydia calycina; C.F., Cassia fistula; A.I., Azadirachta indica; T.A., Terminalia arjuna; A.C., Acacia catechu; M.P., Mallotus philippensis; S.R., Shorea robusta; L.P., Lagerstroemia parviflora; E.L., Ehretia laevis; D.T., Diospyros tomentosa;  L.C., Lannea coromandelica; T.G., Tectona grandis; S.A., Streblus asper; S.O., Schleichera oleosa.
Fig. 2
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