Data Papers Ecology, 92(6), 2011, p. 1376 Ó 2011 by the Ecological Society of America

Tree demography in an undisturbed Dipterocarp permanent sample plot at Uppangala, Western Ghats of India Ecological Archives E092-115

RAPHAE¨L PE´LISSIER,1,2,4 JEAN-PIERRE PASCAL,1,3 N. AYYAPPAN,1 B. R. RAMESH,1 S. ARAVAJY,1 AND S. R. RAMALINGAM1 1

Institut Franc¸ais de Pondiche´ry, UMIFRE 21 CNRS-MAEE, 11 Saint Louis Street, Puducherry 605001, India 2 IRD, UMR AMAP, TA A51/PS2, 34398 Montpellier cedex 05, France 3 43 Rue Rockfeller, 69003 Lyon, France

Abstract. We provide 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 total area sampled to 5.07 ha. 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. Key words: dendrometer bands; Dipterocarp forest; forest dynamics monitoring; India; mortality; recruitment; species demography; tree inventory data; tree growth; tropical rain forest; Western Ghats.

The complete data sets corresponding to abstracts published in the Data Papers section of the journal are published electronically in Ecological Archives at hhttp://esapubs.org/archivei. (The accession number for each Data Paper is given directly beneath the title.)

Manuscript received 12 October 2010; revised 10 February 2011; accepted 22 February 2011. Corresponding Editor: W. K. Michener. 4 E-mail: [email protected]

1376

Ecological Archives E092-115-D1 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-VD below to extract data according to various sampling configurations. Table 1 gives a summary of plot size and location.

TABLE 1. Plot size (in ha) and location of the corresponding sampling window in Uppangala Permanent Sample Plot (UPSP), Western Ghats of India. A sampling window is defined by a vector of (xmin,ymin,xmax,ymax) coordinates of the two opposite South-West and North-East corners, with reference to a unique arbitrary origin for all plots. Mean date of first census is given in Julian calendar days since 1st March 1990 (day 0). * inclusive of overlap with transects A to E (see Fig.1). Plot

Size

Sampling window

Mean date of first census

A

0.36

(640,56,660,236)

0

B

0.74

(540,40,560,410)

0

C

0.66

(440,68,460,398)

0

D

0.68

(340,54,360,394)

0

E

0.68

(240,83,260,423)

0

H1*

1.1

(470,65,570,175)

0

H2*

0.55

(420,65,470,175)

762

H3*

0.12

(420,175,460,205)

1341

H=(H1+H2+H3)*

1.77

(470,65,570,175)+ (420,65,470,175)+ (420,175,460,205)

170

R

0.24

(360,234,390,314)+

1341

(300,234,340,314) R*

0.72

(300,234,390,314)

1030

S

0.12

(360,94,380,154)

1341

S*

0.24

(340,94,380,154)

720

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.

TABLE 2. Numbers of the 10 most common species along with basal area (BA) at first census (1/3/1990) in all the plots pooled together and in the systematic sample only (transects A to E) in Uppangala Permanent Sample Plot (UPSP, Western Ghats of India. Details about species nomenclature are given in datafile UPSP_Species_list2.txt. Species names

All plots (5.07 ha)

A to E (3.12 ha)

#trees

BA

#trees

BA

Vateria indica

593

57.45

330

36.26

Humboldtia brunonis

476

5.33

294

3.30

Myristica dactyloides

412

21.41

263

14.20

Knema attenuata

198

9.11

117

5.86

Palaquim ellipticum

161

5.82

98

4.25

Reinwardtiodendron anaimalaiense

117

2.62

62

1.70

Dipterocarpus indicus

113

32.58

67

14.37

Drypetes elata

112

7.53

79

5.34

Mesua ferrea

106

8.87

56

4.55

Syzygium gardneri

77

3.97

38

1.78

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).

TABLE 3. Summary tree demography in all the plots pooled together (5.07 ha) in Uppangala Permanent Sample Plot (UPSP), Western Ghats of India. GBHmin

GBHmax

N

D

S

R

Sp

NA

G

T

30

3000

3351

NA

NA

NA

96

2

206.31

270

30

3000

3458

81

3270

188

98

4

212.91

1492

30

3000

3445

93

3365

80

98

1

217.96

2805

30

3000

3379

119

3325

54

96

1

221.12

4295

30

3000

3310

187

3191

119

94

1

225.52

6240

30

3000

3321

67

3242

79

97

2

228.58

7336

30

60

2016

NA

NA

NA

87

2

28.260

271

30

60

2148

56

1960

188

89

4

28.85

1492

30

60

2113

48

2033

80

89

1

28.49

2808

30

60

2017

78

1963

54

87

1

27.41

4297

30

60

1951

119

1832

119

86

1

27.02

6240

30

60

1947

39

1868

79

89

2

26.98

7336

60

90

628

NA

NA

NA

64

2

27.19

301

60

90

803

13

615

188

64

4

28.11

1492

60

90

705

21

625

80

63

1

28.13

2802

60

90

685

19

631

54

60

1

28.08

4293

60

90

738

34

619

119

57

1

26.78

6240

60

90

687

10

608

79

58

2

26.85

7336

90

120

286

NA

NA

NA

36

2

24.39

249

90

120

470

4

282

188

41

4

26.50

1492

90

120

376

12

296

80

39

1

26.79

2802

90

120

362

7

308

54

40

1

28.53

4293

90

120

433

21

314

119

41

1

28.04

6240

90

120

403

7

324

79

40

2

28.20

7336

120

3000

421

NA

NA

NA

42

2

126.47

239

120

3000

601

8

413

188

43

4

129.45

1492

120

3000

491

12

411

80

44

1

134.54

2802

120

3000

477

15

423

54

44

1

137.09

4293

120

3000

545

13

426

119

44

1

143.68

6240

120

3000

521

11

442

79

43

2

146.55

7336

TABLE 4. Summary tree demography in all transects pooled together (3.12 ha) in Uppangala Permanent Sample Plot (UPSP), Western Ghats of India. GBHmin

GBHmax

N

D

S

R

Sp

NA

G

T

30

3000

1981

NA

NA

NA

91

0

123.52

0

30

3000

2055

66

1915

140

93

1

128.16

1492

30

3000

2046

44

2011

35

93

1

131.74

2805

30

3000

2004

78

1967

37

92

1

132.85

4296

30

3000

1962

114

1889

73

90

0

134.99

6240

30

3000

1943

62

1900

43

92

1

135.41

7336

30

60

1178

NA

NA

NA

78

0

16.44

0

30

60

1273

45

1133

140

80

1

16.98

1492

30

60

1238

24

1203

35

80

1

16.64

2806

30

60

1182

47

1145

37

79

1

16.16

4298

30

60

1149

70

1076

73

75

0

15.95

6240

30

60

1131

36

1088

43

76

1

15.72

7336

60

90

367

NA

NA

NA

55

0

15.92

0

60

90

496

11

356

140

57

1

16.40

1492

60

90

398

11

363

35

58

1

16.25

2802

60

90

402

12

365

37

54

1

16.30

4293

60

90

432

19

359

73

50

0

15.56

6240

60

90

391

9

348

43

51

1

15.70

7336

90

120

173

NA

NA

NA

31

0

14.76

0

90

120

310

3

170

140

35

1

16.45

1492

90

120

221

5

186

35

33

1

17.06

2802

90

120

231

7

194

37

33

1

17.61

4293

90

120

263

16

190

73

34

0

16.99

6240

90

120

236

7

193

43

33

1

16.97

7336

120

3000

263

NA

NA

NA

39

0

76.40

0

120

3000

396

7

256

140

40

1

78.32

1492

120

3000

294

4

259

35

41

1

81.79

2802

120

3000

300

12

263

37

41

1

82.78

4293

120

3000

337

9

264

73

41

0

86.49

6240

120

3000

314

10

271

43

40

1

87.01

7336

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 readback 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: Data file name

Description

UPSP_Demo_data.txt

Individual identification, location and multidates girth data for 3,870 trees with gbh ! 30 cm in 5.07 ha of Permanent Sample Plots

UPSP_Species_list2.txt

Species-wise information table

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

Rows

Columns

Size (KB)

UPSP_Demo_data.txt

3,870

28

401

UPSP_Species_list2.txt

101

6

8

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: Data file name

Authentication procedures

UPSP_Demo_data.txt

Sum of numerical columns: Xsite=1656003; Ysite=784495.8; GBH0=236154.9; Date0=1025697; Dendro0=0; GBH1=198087; Date1=5774040; Dendro1=194; GBH2=245017.4; Date2=10863219; Dendro2=411; GBH3=244338.2; Date3=16639127; Dendro3=653; GBH4=243052.3; Date4=24148800; Dendro4=1016; GBH5=243976.5; Date5=28390320; Dendro5=1235. The most common species codes in variable SpCode are: vain=659; hubr=590 and myda=435. The number of observations coded as "A" are: Status0=3353; Status1=3462; Status2=3445; Status3=3379; Status4=3310; Status5=3322.

UPSP_Species_list2.txt Total number of species recorded is 101, among which the most represented families are: Euphorbiaceae=16; Anacardiaceae=7, Lauraceae=7; Ebenaceae=6; Meliaiceae=6; Annonaceae=5; Clusiaceae=5; Flacourtiaceae=5. Sum of numeric variable PotStrata=191.

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. Data file name

Variable name

Variable definition

UPSP_Demo_data.txt

TreeID

Alphanumeric tree identification number. The letter refers to the plot in which a given tree was recorded. For rectangular plots H, R and S, a second letter indicates 10 m north-south transect within each plot, in alphabetic order from east to west. The three following digits constitute a unique tree number within each plot/transect.

SpCode

Alphanumeric species code as given in UPSP_Species_list2.txt

Xsite and Ysite

Tree co-ordinates in a West-East (X), South-North (Y) oriented Cartesian plane. All tree co-ordinates are rounded to the nearest 0.05 meter and given from a unique arbitrary origin, so that distances within and between plots are conserved.

GBH#

Tree girth rounded to the nearest 0.1 centimeter from dendrometer readings. # indicates census number from 0 to 5. NA=missing data

Date#

Time interval in days from 1st March 1990 (day 0) following Julian calendar. # indicates census number from 0 to 5.

Status#

Code for tree status at each census: "A"=alive; "D"=dead; "P"=not yet large enough for measurement, future recruit. # indicates census number from 0 to 5.

Dendro#

Numeric code indicating if GBH# was measured with initial dendrometer band (0) or after 1 to n changes (max=5) of the dendrometer band. # indicates census number from 0 to 5. NA=missing value for trees without dendrometer because they have been damaged, or tree is dead or not yet recruited.

UPSP_Species_list2.txt SpCode

Alphanumeric species code built from the first four letters of the genus name and first four letters of the specific epithet, except in case of duplication and for 2 morphospecies, which are coded as alsp1 and cisp1.

LatinName

Complete latin name of species updated from latest synonyms from HIFP (http://www.ifpindia.org/herbarium/).

Author

Authority name for the species. NA=missing data for morphospecies.

Family

Botanical family to which belong the species.

VoucherRef

Voucher reference number for the species at HIFP (http://www.ifpindia.org/herbarium/).

PotStrata

Numeric code for potential strata reached by the species at adult stage (tree growth form) according to Pascal (1988) updated from Ramesh et al. (2010): 0=Emergent; 1=Canopy species; 2=Subcanopy species; 3=Understorey species. NA=missing value.

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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