Raphaël Pélissier, Jean-Pierre Pascal, N. Ayyappan, B. R. Ramesh, S. Aravajy, and S. R. Ramalingam. 2011. Twenty years tree demography in an undisturbed Dipterocarp permanent sample plot at Uppangala, Western Ghats of India. Ecology 92:1376.

INTRODUCTION

Quantifying forest dynamics is fundamental to several aspects of basic and applied forest ecology. Due to the lack of annual rings in many tropical tree species, the study of tropical forest dynamics relies heavily on the use of Permanent Sample Plots (PSP) that have long since been established throughout the world (e.g., Manokaran and Kochummen 1987, Burslem and Whitmore 1999, Sheil et al. 2000). Monitoring tropical forest PSP was often dictated in the past by practical management considerations, with the aim to assess recovering of timber stock after logging, often monitoring "virgin jungles reserves" as controls (Vanclay 1991). In the last decades, however, more emphasis has been put on forest biodiversity, dynamics, and changes in carbon stock in the context of environmental changes (e.g., Phillips and Gentry 1994, Laurance et al. 2004, Lewis et al. 2004, Losos and Leigh 2004). For instance, Phillips and Gentry (1994) concluded from PSP data analysis that tree turnover rates have increased in tropical forests during the latter part of the 20th century, a finding which proved to be controversial (e.g., Phillips 1995; Sheil 1995). More recently, another global issue was put in front from evidences of decelerating tree growth rates in tropical forests over the past two decades (Feeley et al. 2007), a result which contradicts the carbon fertilization hypothesis predicting an increased productivity under rising concentrations of atmospheric CO₂. From these global perspectives, a critical need to share good quality PSP inventory data among ecologists has emerged (e.g., Dallmeier and Comiskey 1998, Losos and Leigh 2004) and we report here, as a contribution to this movement, a data set on 20 years of tree demography monitored in Uppangala PSP, an undisturbed, old-growth wet evergreen Dipterocarp forest in the Western Ghats of India.

Uppangala PSP (UPSP) is located within a 28-ha experimental station established in 1989–1990 by the French Institute of Pondicherry (IFP) in the Kadamakal Reserve Forest, which falls within Pushpagiri Wildlife Sanctuary in Karnataka state, India. The area falls within the world biodiversity hotspot of the Western Ghats (Myers 1990) and is considered as one of the last remnants of undisturbed rainforest in the region (Pascal and Pélissier 1996). A total of 5.07 ha of forest plots have been systematically inventoried in Uppangala, initially in 1990–1993, while re-censuses were conducted every 3–5 years since then, in 1994, 1997–1998, 2001–2002, 2007, and 2010. Throughout these inventories, a total of 3870 trees with a minimum girth at breast height (gbh) of 30 cm have been identified, mapped and fitted with stainless steel dendrometer bands for accurate growth monitoring. We report here the corresponding demographic data recorded over the last 20 years (1990–2010).

METADATA

CLASS I. DATA SET DESCRIPTORS

A. Data set identity: Uppangala Permanent Sample Plot, Western Ghats, India.

B. Data set identification code: IFP_ECODATA_UPSP_v2

C. Data set description

Originators: Raphaël Pélissier, UMR AMAP, TA A51/PS2, 34398 Montpellier cedex 05, France ([email protected])

N. Ayyappan, French Institute of Pondicherry, UMIFRE 21 CNRS-MAEE, 11 St Louis Street, Puducherry 605001, India ([email protected]).

Abstract: We report a data set on demography of trees monitored over 20 years in Uppangala permanent sample plot (UPSP) in undisturbed, old-growth wet evergreen Dipterocarp forest located within the Pushpagiri Wildlife Sanctuary in India's Western Ghats biodiversity hotspot. During 1989–1990, all trees ≥ 30 cm girth at breast height (gbh) were sampled in five north–south transects 20 m wide and 180 to 370 m long covering a total area of 3.12 ha. In 1992–1993, additional rectangular plots were established, bringing the area sampled to 5.07 ha in total. In all, 3870 trees were identified, tagged, mapped, and provided with permanent dendrometer bands. Since then, the sampled area has been regularly censused at 3–5 year intervals, recording tree recruitment, mortality, and growth. We present data from censuses conducted in 1990–1993, 1994, 1997–1998, 2001–2002, 2007, and 2010. These data have been used to study the natural forest dynamics and to calibrate spatially explicit simulation models.

D. Key words: dendrometer bands; Dipterocarp forest; forest dynamics monitoring; India; mortality; recruitment; species demography; tree inventory data; tree growth; tropical rain forest; Western Ghats.

CLASS II. RESEARCH ORIGIN DESCRIPTORS

A. Overall project description

Identity: Long-term monitoring of the forest dynamics in Uppangala Permanent Sample Plot (UPSP), an undisturbed, old-growth dense wet evergreen forest in the Western Ghats of India.

Originator: Jean-Pierre Pascal and Ecology Department, French Institute of Pondicherry, 11 St Louis Street, 605001 Puducherry, India ([email protected]).

Period of Study: Ongoing since 1989.

Objectives: Long-term monitoring of tree growth and demography.

Abstract: In 1989, Jean-Pierre Pascal discovered within the Kadamakal Reserve forest an unlogged compartment that was scheduled for harvesting but spared thanks to the ban on felling promulgated by Karnataka Forest Department (KFD) in all natural forests of Karnataka state in 1988. A cooperative agreement has then been signed with KFD, authorizing conversion of this forest compartment into an experimental station of the French Institute of Pondicherry (IFP). The site is since 1990 strictly protected and shelters Permanent Sample Plots that provide the basic data to the IFP research projects to model the natural forest dynamics, to compare its evolution with the one in neighboring compartments recovering from selective logging, to explore changes in community structure and diversity, to study tree population dynamics and species ecology.

B. Specific subproject description

1. Site description: (Pascal and Pélissier 1996, Elouard et al. 1997a, b).

Site type: Dipterocarpus indicus - Kingiodendron pinnatum – Humboldtia brunonis type of low elevation wet evergreen forest (Pascal 1984, 1988).

Geography: Pushpagiri Wildlife Sanctuary, Kadamakal Reserve Forest, Sampaje Forest

Range, Kodagu District, Karnataka state, Western Ghats of India, ca. 12° 32' 15'' N, 75° 39' 46 E, altitude between 400 and 600 m a.s.l. The nearest town by road is Subrahmanya in Dakshina Kannada District (Fig. 1).

FIG. 1. Location map (Google Earth extraction) of Uppangala Permanent Sample Plot (UPSP) in the Western Ghats of India.

Habitat: UPSP is located within a 28-ha undisturbed forest compartment of the Kadamakal Reserve Forest, a 1200-ha forest area that was inaccessible by road before 1959 nor appeared in official documents as a logged area until it was submitted to a single cycle of selective logging (harvesting of less than 10 trees per ha per annual concession of 28 to 30 ha) from 1974 to 1988 (Loffeier 1989). UPSP was established in one of the concessions, which, according to Karnataka Forest Department, was scheduled for harvesting but spared thanks to the ban on felling promulgated in all natural forests of Karnataka state in 1988. Before the logging track was opened in the 1980s, UPSP was only accessible by foot from the nearest village of Uppangala (about 20 houses, 10 km away from the site). The site has been strictly protected since 1990 and has no known history of major human disturbance and if ever affected by humans, the disturbance probably remained limited to collection of minor forest products by villagers (Salaün 1995). It is moreover surrounded by a large area of continuous quasi undisturbed forest (Fig. 2).

FIG. 2. View of a portion of Pushpagiri Wildlife Sanctuary in the Western Ghats of India in 1992. Picture taken towards northeast from the crest of the Ghats. The white arrow indicates the approximated location of Uppangala Permanent Sample Plot (UPSP).

Geology: In the southern part of the Western Ghats, bedrock is composed of Archean rocks from the Precambrian shield, with a prevalence of Peninsular Gneisses between 11 and 14°N (Pascal 1984, 1988). Ferry (1994) classified the soils in the Kadamakal Reserve Forest as dystric cambisols (FAO/UNESCO 1988). They are generally thick on very old alterites, but may be less evolved in steep rocky boulders-full slopes shaping the talwegs.

Watersheds/hydrology: UPSP is located at mid-slope of the Ghats’ escarpment (see Fig. 2), which crest is at ca. 1,000 m a.s.l. The site is facing north-northwest with a mean slope angle of about 30–35°. Stream valleys running downward from the crest notch the main slope and determine an east–west alternation of more or less flattened interfluve ridges, so that the topographic situation is the main factor of local environmental variation (Gimaret-Carpentier et al. 1998, Pélissier 1998, Robert and Moravie 2003).

Site history: See A.5.Abstract and B.1.Habitat.

Climate: It belongs to the Allepey-Mangalore regime of Pascal (1982), characterized by a peak of heavy rainfall during summer monsoon and a marked winter dry season. Annual precipitation averages 5108 mm with 90% between June and October and an average dry season (monthly rainfall ≤ 45 mm) of 3.8 months (data recorded from 1993 to 2007 at Uppangala village, 200 m a.s.l., 5 km from UPSP; Fig. 3). Temperature has not been recorded at the site, but mean annual temperature in the nearest met station of Sampaji (140 m a.s.l., 15 km from UPSP) is about 27°C, with a mean minimum temperature of about 25°C, which coincides with the July rainfall peak, and a maximum mean temperature of ca. 29°C in April (Pascal and Pélissier 1996).

FIG. 3. Histogram of average monthly rainfall (in mm) recorded between 1993 and 2007 (with years 1998 and 1999 excluded because of some missing data) at 200 m a.s.l., in the village of Uppangala 5 km away from Uppangala Permanent Sample Plot (UPSP) in the Western Ghats of India.

2. Experimental or sampling design.

Design characteristics: Sampling design consists of five north–south oriented transects viz., A, B, C, D, and E, each 20 m wide, 180 to 370 m long, and 100 m apart center to center (Fig. 4).

FIG. 4. Sampling design of Uppangala Permanent Sample Plot (UPSP) in the Western Ghats of India (background extracted from a 2002 Ikonos image). Elevation contour lines are at 3-m intervals.

The 5 transects were installed in 1989–1990 and collectively represent a 3.12-ha systematic sample of the forest compartment, whose main dendrometric and floristic characteristics are given in Pascal and Pélissier (1996). Subsequently, additional rectangular sampling plots viz., H, R and S, which overlap the transects and represent an additional area of 1.95 ha, were established in 1990–1993 to study the dynamics of the forest in particular topographic situations (Pélissier 1997, 1998). A piece of R source code is given in section III-V-D below to extract data according to various sampling configurations. Table 1 gives a summary of plot size and location.

Because of the uneven site topography (see B.1.Watersheds and hydrology), each plot was designed as a grid of 10 × 10 m contiguous elementary quadrats adjusted to account for local slope angle. The purpose of the correction was to ensure that each side of a quadrat was approximately 10 m flat, so that it contains in total a 100 m² area in planar projection. The slope angle (alpha) was measured along each quadrat sides, which length parallel to the ground was adjusted to l = 10/cos(alpha). In order to accurately locate the trees in (x,y) co-ordinates within each quadrat, they were temporarily delineated with a plastic rope. All trees from 30 cm girth at breast height (gbh) were inventoried, measured, tagged, mapped, and identified to species. The point of girth measurement of each tree (pom) was marked with paint in the initial inventory and then fitted with a permanent dendrometer band. For mapping the trees within each quadrat, the flat distance from the centre of the trunk base to the nearest quadrat boundary was measured in two perpendicular directions parallel to the rope. Each quadrat was mapped in the field at a scale of 1:100 with indication of tree locations, main rocks, fallen trees, stream courses, trails, etc. These maps are stored at IFP Botany Lab.

3. Research Methods

Instrumentation: Slope angles were measured with a 1-degree precision using a Suunto hand-held clinometer. Distance measurements were taken with a 1-cm precision thanks to a 20 m fiberglass tape. Initial tree girths were measured at 1.30 m from the ground or above the buttresses with a precision of 0.1 cm using a 3 or 5 m flexible steel tape. A calibrated 1.30 m stick was used to determine pom height. The dendrometer bands were prepared by technical IFP staff using stainless steel bands adjusted to tree girth and equipped with a 1:50 vernier (Hall 1944; Fig. 5) that allows girth to be measured with a theoretical precision of 0.02 cm. The dendrometers were then fitted at pom on the trees and vernier readings taken as initial gbh measurement.

FIG. 5. Close view of a dendrometer band fitted on a tree with details of the 1:50 vernier scale, in Uppangala Permanent Sample Plot (UPSP), Western Ghats of India.

Taxonomy and systematics: All the trees were identified to species level (including 2 morphospecies) and referenced voucher specimens were deposited at IFP Herbarium (HIFP). Botanical identifications were supervised by expert botanists (initially J.-P. Pascal and B. R. Ramesh, with contribution of N. Ayyappan since 2005) and were most often made in the field with the help of a field key based on vegetative characters (Pascal and Ramesh 1987). In doubtful cases, specimens were collected and identified at HIFP. Species were coded in data files (see IV.B.Variable definitions) using a four-letter code (most generally the two first letters of the genus name and two first letters of the specific epithet, except in case of duplication and for the two morphospecies coded as alsp1 and cisp1). A voucher reference number at HIFP is also provided for each species and morphospecies. Synonyms have been checked with reference to July 2010 version of HIFP database (http://www.ifpindia.org/herbarium/). Table 2 gives a plot summary for the 10 most common species.

Recensuses. Recensuses have been conducted every 3–5 years in all plots since May 1990 (census 0), in May 1994 (census 1), November 1997–April 1998 (census 2), December 2001–May 2002 (census 3), April 2007 (census 4), and April 2010 (census 5). Time intervals between censuses are given in Julian day calendar since day 0 on 1st March 1990. However, some plots (H extension, R and S), installed in March 1992 and November 1993 are included with a census 0 date on day 762 and 1341, respectively. Similarly, censuses 2 and 3, which were completed over several field trips, refer to more than one Julian date (see Table 1). At each recensus, all plots were screened for tree mortality, tree growth by girth increments of alive trees reading from dendrometer bands and tree recruitment, i.e., those trees attained the girth threshold of 30 cm gbh since the previous census. The recruited trees were tagged, mapped, identified, and installed with a dendrometer band, following the initial census protocol (see II.B.3. Research methods). Summary data of plots' demography are given in Tables 3 (all plots pooled together) and 4 (tansects A to E pooled together). These summaries have been computed from the raw data file using a piece of R code provided in section III-V-D below.

Tables 3 and 4. Summary tree demography in Uppangala Permanent Sample Plot (UPSP), Western Ghats of India. GBHmin and GBHmax correspond to the range of tree girth considered (GBHmax=3000 includes all the trees); N=number of living trees at each census; D=number of trees that died since previous census (NA at the first census); S=number of surviving trees since previous census (NA at the first census); R=number of trees that were recruited since previous census (NA at the first census); Sp=number of species present at each census; NA=number of missing gbh values at each census; G=total tree basal area (in m²) at each census (to compute G, missing gbh values have been replaced by gbh value at first census when possible; NA indicate the number of missing gbh values that we could not correct); T=mean census date in Julian calendar days since 1st March 1990 (day 0).

Permit history: Permit for conducting a research program at Uppangala PSP was delivered by the Government of India through a Memorendum Of Understanding between IFP and Karnataka Forest Department (KFD) sited in Bangalore, Karnataka state, India.

Legal/organizational requirements: Any visit to the site requires an authorization delivered to IFP staff by KFD.

Project personnel: The project P.I.s were Jean-Pierre Pascal (project initiation, plot selection, definition of measurement protocol, supervision of initial fieldwork and botanical identification, and project supervision during 1990–2000,), B. R. Ramesh (botanical identification), Raphaël Pélissier (database conception and management, contribution to fieldwork in 1992–1994 and project supervision during 2007–2010) and N. Ayyappan (contribution to fieldwork and database management, project supervision since 2010). C. Elourad, L. Henry, and H. Laborde also contributed to various extents in fieldwork. IFP project technicians, who have assisted in fieldwork through time, are S. Aravajy, N. Barathan, G. Jayapalan, K. Kichenassamy, G. Orukaimani, and S. R. Ramalingam. Staff of the IFP Botany Lab is in charge of the reference herbarium specimens. J. Le Bec contributed to plots' georeferencing and provided Fig. 4.

CLASS III. DATA SET STATUS AND ACCESSIBILITY

A. Status

Latest update: August 2010.

Latest Archive date: August 2010.

Metadata status: The metadata are complete and up-to-date.

Data verification: The database has been screened for missing, erroneous and out of bounds entries, which have been checked by read-back from the original field data sheets. In addition, crosschecking was performed in the field at recensus times: tree locations, botanical identification and girth values were systematically checked against maps and data printouts prepared in advance from the database. Similarly, when a tree was uncertainly identified, it has been regularly surveyed till an unambiguous observation could be made such as the appearance of flowers, fruits, or accessible sprouts. This process led to few modifications of initial species list.

B. Accessibility

Storage location and medium: Ecological data archives of the French Institute of Pondicherry (http://www.ifpindia.org/biodiversityportal). Original field data sheets are stored at IFP Botany Lab and digital copies of the database are stored in several locations at IFP and at UMR AMAP in Montpellier.

Contact person: N. Ayyappan, French Institute of Pondicherry, UMIFRE 21 CNRS-MAEE, 11 St. Louis Street, 605001 Puducherry, India, tel. +91 413 2334168 Extn. 128; fax +91 413 233 9534; [email protected]

Copyright restrictions: None.

Proprietary restrictions: None, the data are free to use for further analyses, with due citations to this data paper and, when appropriate, to Pascal and Pélissier (1996), which is the oldest paper on UPSP.

CLASS IV. DATA STRUCTURAL DESCRIPTORS

A. Data Set File

Identity: Data set is downloadable as a single archive, Data.zip (102 KB), which contains the following data files:

Size: The number of rows and columns includes headers; size is given for uncompressed files.

Format and storage mode: ASCII text, tab delimited.

Header information: Headers corresponding to variable names (see IV.B.Variable definitions) are included as first row in the data file.

Alphanumeric attributes: Mixed.

Special characters/fields: NA in data file stands for Non Available information (missing data).

Authentication procedures:

B. Variable information

Variable names are headers included as first row in the data file. For each of the census variables are included with # corresponding to census number.

CLASS V. SUPPLEMENTAL DESCRIPTORS

A. Data acquisition

Data forms: Initial census data (tree numbers, identifications and gbh measurements) were taken on empty sheets, while tree maps were directly drawn on graph paper from (x,y) tree co-ordinates measured within each 100 m² quadrat. Slope angles taken along each quadrat sides were also reported on the maps and later used to generate elevation contour lines as displayed in Fig. 4. Subsequent measurements were taken on sheets prepared in advance from the previous census database, including tree number, species identification and last dendrometer reading, while maps generated from the database were used for field navigation, so that it is likely that most field measurements and data entry errors have been tracked back and corrected (see III.A.4. Data verification).

Location of completed data forms: See E.1. Archiving

Data entry/verification procedures: Data were manually digitized by the people who took the field measurements, one person reading the field sheet to another one entering the data in the computer while repeating what he heard. Further crosschecking procedures are detailed in sections III.A.4. Data verification and V.A.1. Data forms.

B. Quality assurance/quality control procedures: See A.3. Data entry/verification procedures.

C. Related material: Data on tree dimensions (height, crown size, etc.), architecture, phenology, regeneration, growth and competition have also been collected in UPSP, but, since not systematic, have not been included in the present demography database. Many related studies have been carried out by IFP staff and collaborators at UPSP or in the nearby area of the Kadamakal Reserve forest: Loffeier (1988a, 1988b, 1989), De Franceschi and Tissot (1991), Sinha and Davidar (1992), Basu (1994), Ferry (1994), Pascal (1995), Pélissier (1995), Salaün (1995), Pascal and Pélissier (1996), Robert and Salaün (1996), Durand (1997), Elouard et al. (1997a, 1997b), Houllier et al. (1997), Moravie et al. (1997, 1999a, 1999b), Pélissier (1997, 1998), Gimaret-Carpentier et al. (1998), Pascal et al. (1998), Pélissier et al. (1998), Durand (1999), Elouard and Krishnan (1999), Moravie (1999), Pélissier and Goreaud (2001), Robert (2001), Moravie and Robert (2003), Robert (2003), Robert and Moravie (2003), Magnussen et al. (2006), Madelaine-Antin (2009).

D. Computer programs and data processing algorithms: Data format has been made compatible with CTFS 1.00 package for the R platform (http://cran.r-project.org/web/packages/CTFS/). The following R function allows reading UPSP data file for a direct usage with this package [To download the R code please click here for Rcode.txt]:

Example of computation of mortality using CTFS package:

The function below allows demographic data to be extracted from the main data file according to various sampling configurations using functions in ads package:

Example of data extraction from upsp data frame using the above function:

The function below allows computing summary demographic data from demographic data file and functions in CTFS package:

Example of computation of summary demographic data:

E. Archiving: Original data sheets are stored at IFP Botany Lab. One master copy of the complete database and an extraction of the present data set are stored on the IFP server with several copies in computers and CD at IFP and at UMR AMAP, Montpellier, France.

F. Publications and results:

Many published results have used part of this dataset: Pélissier (1995), Pascal and Pélissier (1996), Elouard et al. (1997a, 1997b), Moravie et al. (1997, 1999a, 1999b), Pélissier (1997, 1998), Gimaret-Carpentier et al. (1998), Pascal et al. (1998), Pélissier et al. (1998), Moravie (1999), Pélissier and Goreaud (2001), Robert (2001), Moravie and Robert (2003), Robert (2003), Robert and Moravie (2003), Magnussen et al. (2006), Madelaine-Antin (2009), Ploton (2010).

G. History of data set usage:

1. Data request history: Part of the data set was requested in 2005 by Steen Magnussen, Canadian Forest Service, 506 West Burnside Road, Victoria, British Colombia, V8Z 1M5 Canada, and in 2009 by I. Parmentier, post-doctoral fellow at Laboratoire d'éco-éthologie évolutive, Université Libre de Bruxelles, CP160/12, Av. F. D. Roosevelt 50, 1050 Brussels, Belgium.

2. Data set update history: The database has been regularly updated since 1990. A preliminary version of this datapaper has been posted in 2008 at (http://www.ifpindia.org/biodiversityportal). The papers cited in V.F.Publications and results used different earlier versions of this data set.

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