Flowering and fruiting phenology of woody species in tropical dry evergreen forests on the Coromandel coast of India
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1 J. Biosci. Res., Vol. 2(2): ISSN Flowering and fruiting phenology of woody species in tropical dry evergreen forests on the Coromandel coast of India D.R. VENKATESH 1, N. PARTHASARATHY 2, K. MUTHUCHELIAN 1 1 Centre for Biodiversity and Forest Studies, School of Energy Environment and Natural Resources, Madurai Kamaraj University, Madurai Department of Ecology and Environmental Sciences, School of Life Sciences, Pondicherry University, Puducherry Abstract Biology and composting ability of three important earthworm species viz., Eudrilus eugeniae Kinberg (Eudrilidae), Eisenia foetida Savigny (Lumbricidae) and Pheratima posthuma Vaillant (Magascolecidae) were studied under laboratory conditions. Among the species tested, E. eugeniae was found superior in terms of compost production compared with other two species. The physico-chemical nature of the compost produced by E. eugeniae was found to be superior in terms of electrical conductivity, P, K, organic matter content and C:N ratio than other two species. The present study indicated that, E. eugeniae could be used for vermin compost production in eco-climatic conditions prevailing in Sivakasi. Key words: Climate change, Long-term monitoring, Rainfall, Reproductive phenology, Temperature. For correspondence: venk8dr@gmail.com Introduction For phenological observations flowering and fruiting phenologies were recorded monthly from March 2000 through February 2003, in two tropical dry evergreen forest sites. In both the forests, 580 individuals of 34 forest woody species were sampled, which include 24 tree species and 10 liana species. Ten of the 24 tree species and two of the 10 liana species were common to both the study sites. All the species sampled were =20 cm gbh, considered by and large to be sexually mature and without vine infestations and deformities of obvious diseases. Flowering and fruiting phenology of woody species in dry evergreen forests showed a general annual flowering and fruiting as the most common pattern and it peaked during the dry periods. Phenological events were constrained mainly by the seasonal climatic variables such as maximum temperature, and rainfall. Although dry evergreen forests are less diverse than lowland rainforests, it supports a rich insect, bird, reptile mammal life, many of which help in the functioning of these forests and hence these forests deserve more attention with respect to research on long-term monitoring of woody species phenology than is currently received. Introduction Phenology is the study of growth of buds, leaf flushing, leaf fall, anthesis and fruiting in relation to seasonality (Okullo et al. 2004). The phenology of tropical woody plants has been shaped by biotic and abiotic factors (Okullo et JOURNAL OF BIOSCIENCES RESEARCH 2(2):
2 al. 2004). The seasonal rhythms of vegetative and reproductive phenologies of tropical plants are often associated with seasonal pattern in climate, or the interaction with animal pollinators and seed dispersers (Smith-Ramirez and Armesto 1994, Chapman et al. 1999, Morellato et al. 2000, & Okullo et al. 2004). Tropical plants often show temporal phenological events that are associated with well-defined wet and dry seasons (Rathcke and Lacey 1985). Understanding phenology in relation to climate is an important first step. Since climatic variables are often highly intercorrelated and testing their influence on phenology requires, firstly finding significant correlations of phenology with climatic variables (Smith-Ramirez and Armesto 1994, Morellato et al. 2000) then, secondly controlling for interactions among these variables, so that the most parsimonious model of the influence of climate on phenology can be derived (Marques et al. 2004). By and large variations in rainfall, photoperiod and temperature have been implicated in governing the timing of flowering (Morellato et al & Marques et al. 2004). Although detailed studies on quantitative ecological inventory of plant diversity and forest production and functional aspects are available from these forests, data concerning tree population changes and woody species phenology are lacking. Hence, an attempt has been made to investigate community-level phenological events in two representative sites in relation to environmental cues such as maximum temperature and rainfall over a period of three years in tropical dry evergreen forests on the Coromandel coast of south India. Study area This study was conducted in two tropical dry evergreen forest sites on the Coromandel coast of India viz. Kuzhanthaikuppam (KK -11º 45' N lat. & 79º 38' E long.) and Puthupet (PP - 12º 03' N lat. & 79º 52' E long.). These are also sacred groves or temple forests, but composed of native species, located within a 50 km radius of Pondicherry town on the Coromandel coast of southern India. The vegetation in this region is tropical dry evergreen forest, due to anthropogenic disturbances, various areas of these forests were degraded to thorny scrub. The tropical dry evergreen forest was once common along the southeast coast of India, and was the predominant vegetation of coastal Tamil Nadu. Now, only remnants of these are found in the area, with most them already cleared for agriculture and settlements and small areas remain as the pockets of dry evergreen forests (for more information please refer Venkateswaran, 2004). Methods For phenological observations flowering and fruiting phenologies were recorded monthly from March 2000 through February 2003, in the two representative sites, Kuzhanthaikuppam (KK) and Puthupet (PP). In both the forests, 580 individuals of 34 forest woody species were sampled, which include 24 tree species and 10 liana species. Ten of the 24 tree species and two of the 10 liana species were common to both the study sites. All the species sampled were 20 cm gbh, considered by and large to be sexually mature and without vine infestations and deformities of obvious diseases. All individuals were marked with yellow colour paint and appended with sequentially numbered aluminium tags to facilitate relocation. Each individual was visited approximately during the middle of every month and the presence of flowers and fruits was noted with the aid of binoculars. If any individual of a species had flowers or fruits that species was considered to be flowering or fruiting that month. Voucher samples of specimens were deposited in the herbarium of Salim Ali School of Ecology and Environmental Sciences, Pondicherry University. To examine the relationship between phenological pattern and climatic variables such as maximum temperature and rainfall, Spearman s rank correlation was used. Monthly lags in climate were introduced so that phenology could be correlated with climate of the previous months varying from the current date up to JOURNAL OF BIOSCIENCES RESEARCH 2(2):
3 9 months before the current date. Both the variables (maximum temperature and rainfall) were correlated with the two phenological phases (flowering and fruiting). Correlation between the number of woody species in each phenophase and monthly maximum temperature and rainfall was made to check if flowering and fruiting occurred more often during dry than wet months. For this purpose dry months were considered as months in which the mean monthly daily maximum temperature was above 32 C and the monthly precipitation total was 62mm (modified after Okullo et al. 2004). Results Climatic information from March 2000 to February 2003, during when the phenological observations were made, reveals that monthly mean of daily maximum temperature was 36 C in May/June at both the sites. Monthly means of daily maximum temperature were relatively constant between 18.1 C to 28.8 C in site KK and in site PP it was 20.2 C to 27.8 C. Rainfall occurred mostly from September to December in both the sites. Annual rainfall (in mm) totaled 1142, 852, 1090, and 1040 at site KK during the four years, 2000 to 2003, and for the same years, at PP it was 1224, 947, 949 and 1359 mm respectively. In all these four years no rainfall was recorded in the month of March at both the sites and the monthly precipitation value of below 62 mm was recorded in January to May except in February 2002 at both the study sites. Rainfall is strongly seasonal with most rains falling from October to December, although southwest monsoon brings occasional rains. Flowering phenology Flowering phenology of woody species in tropical dry evergreen forests was dispersed more evenly throughout the year, with at least 3 species in flower at a given month. Although some species could be found in flower round the year, flowering was annual in both the sites, with a peak in March-April, i.e. during the dry period. Spearman s rank correlation between the number of species in flower each month and the mean monthly rainfall, revealed that flowering occurred more often in dry than wet months, and was significantly inversely related to monthly rainfall (for KK: rs = , P<0.01; for PP: rs = , P<0.01). Thus, flowering peak occurred after rainfall i.e. a 5-month lag period in site KK (rs = 0.539, P<0.01) and a 6-month lag period in site PP (rs = 0.658, P<0.01). Site-wise, the peak season of flowering was January and extended up to May in site KK; while in PP, it started from January and ended in June. The maximum number of species in flower (65% of all species in 2000, 65% in 2001 and 75% in 2002) was recorded in March at site KK and in site PP, except during 2001, the maximum number of species in flower (69% in 2000 and 73% in 2002) was recorded in April. The total number of species that flowered during February to May was more than 85% of the 34 species in the two sites. They included mainly species such as Capparis zeylanica, Combretum albidum, Garcinia spicata, Drypetes sepiaria, Lepisanthes tetraphylla, Pterospermum xylocarpum and Ventilago madraspatana. All species except Diospyros ferrea, in site PP produced fruits during the three years of study. In the threeyear phenological observation, a peak flowering for most of the species occurred in March at site KK and in April at site PP. Most species surveyed in tropical dry evergreen forests flowered and set fruit in each year of the study. These species also tented to flower at about the same time each year and flowered only once per year. In the terms of Newstrom et al. (1994), such species displayed the regular and annual pattern of flowering. The transition between dry and wet seasons was the period in which the majority of woody species were budding, flowering, or fruiting. Such peaks in flowering or fruiting in the late dry season to early wet season have been described for seasonal tropical forests in India. Dry season flowering has been reported elsewhere in the tropics too (Okullo et al. 2004). Fruiting phenology In general, annual fruiting was the most common pattern in both the forest communities, occurring in 22 woody species (85% of species) at site PP and 17 species (85%) at KK; while a small number of species in fruits was encountered year round. A high concentration of fruiting was JOURNAL OF BIOSCIENCES RESEARCH 2(2):
4 seen during the dry season from March to July. The species of greatest abundance in both the forests bore fruits during these months. The peak in fruiting started from March and ended in July at site PP and at site KK it started from February and ended in June. The correlation between the number of species in fruit each month and mean monthly maximum temperature revealed that fruiting also occurred more often in dry than in wet months. Fruiting was strongly positively correlated with mean monthly temperature in both the forests (KK: rs = 0.592, P<0.01; PP: rs = 0.733, P<0.01) and significantly negatively correlated with rainfall at site KK (rs = , P<0.01) and weakly negatively, but not significantly correlated at site PP (rs = , P>0.05). These results implied that fruiting peak occurred after rainfall i.e. during the dry period and time-lag correlation of fruiting, peak fruiting and rainfall showed the highest correlation for a 8-month lag period for site KK (rs = 0.617, P<0.01) and 9-month lag period for site PP (rs = 0.701, P<0.01). The maximum number of species (75%) in fruiting was consistently recorded in March during the three years at site KK and in site PP, it was random during the month of April to July (in 2000; July, 77%, 2001; July, 65% and 2002; June, 81%). The total number of species that bore fruits during March to July represented together more than 88% of the total species in the two sites. They included mainly, Capparis zeylanica, Garcinia spicata, Lepisanthes tetraphylla, Memecylon umbellatum, Pachygone ovata, Pongamia pinnata, Pterospermum xylocarpum, Ventilago madraspatana, Walsura trifolia etc. A peak fruiting for most of the species occurred in March at site KK and in June at PP during the three-year observations. Knowledge of fruiting phenology is of central importance for conservation management generally, but is of special value in tropical forests which support a diversity of fruit-dependent species (Fenner 1998). Fruiting year-round in the dry evergreen forests follows the tendency of many tropical forests. Fruiting during end of the dry season may have evolved to reduced seedling mortality by dispersing seeds when soil moisture conditions are favorable for seed germination and rapid seedling growth (van Schaik et al. 1993). Fruit production at the end of the dry season ensures the seedlings are not immediately exposed to water stress (Hamann 2004). The relationship between phenology and main climatic factors such as maximum temperature and rainfall provided limited evidence concerning possible triggers as few significant correlations between these variables and the onset of flowering and fruiting was identified, either positive or negative. Temperature and irradiance have been reported to influence flowering in many other tropical forests (van Schaik et al. 1993, Chapman et al. 1999). While in the dry evergreen forests, time-lag analysis with the weather data provided the evidence that rainfall also seemed to influence the number of species with 8 to 9-month lag period. To know the periodicity and seasonality, as well as year-to-year variation in phenological events, it requires long-term observation of particular plants, especially in tropical forests with diverse phenology types (Sakai 2001), because long-term monitoring of plant phenology is more important now that global environmental change is a critical issue. Studies from different parts of the world have shown that vegetative and reproductive phenology, both at community and species level, potentially integrate a large number of selective forces caused by climatic factors as well as biotic factors (Hamann 2004). The alternation of dry and wet periods is often associated with tropical phenological cycles. In general, phenological phases were more strongly associated with day length or temperature with a lag of 4-5 months (Marques et al. 2004). These dry evergreen forests are truly seasonal in both the phenological events i.e. flowering and fruiting, and phenology is partially explained by two climatic variables, maximum temperature and rainfall. Conclusion Flowering and fruiting phenology of woody species in dry evergreen forests showed a general annual flowering and fruiting as the most common pattern and it peaked during the dry periods. Phenological events were constrained mainly by the JOURNAL OF BIOSCIENCES RESEARCH 2(2):
5 seasonal climatic variables such as maximum temperature, and rainfall. Although dry evergreen forests are less diverse than lowland rainforests, it supports a rich insect, bird, reptile mammal life, many of which help in the functioning of these forests, and hence these forests deserve more attention with respect to research on long-term monitoring of tree population changes and woody species phenology than is currently received. Acknowledgments The authors are grateful to Mr. Manickam and his family and Mr. Ravi s family for their hospitality and help in the various stages of my field work and also to Mr. Manikandan, and Mr. Velu for their assistance during my initial fieldwork. Dr. D. R.Venkatesh is grateful to the UGC, New Delhi for financial support through Dr. D.S. Kothari Postdoctoral Fellowship [No. F.4-2/2006 (BSR)/13-253/2008 (BSR)] References Chapman C.A., Wrangham R.W., Chapman L.J., Kennard D.K. and Zanne A.E Fruit and flower phenology at two sites in Kibale National Park, Uganda. J. Trop. Ecol. 15: Fenner M The phenology of growth and reproduction in plants. Perspect. Plant Ecol. Evol. Syst. 1: Hamann A Flowering and fruiting phenology of a Philippine submantane rain forest: climate factors as proximate and ultimate causes. J. Ecol. 92: Marques M.C.M., Roper J.J. and Salvalaggio A.P.B Phenological patterns among plant life-forms in a subtropical forest in southern Brazil. Plant Ecol. 173: Morellato L.P.C., Talora D.C., Takahasi A., Bencke, C.C. Romera E.C. and Zipparro V.B Phenology of Atlantic rain forest trees: a comparative study. Biotropica 32: Newstrom L.E., Frankie G.W. and Baker H.G A new classification for plant phenology based on flowering patterns in lowland tropical rainforest trees at La Selva, Costa Rica. Biotropica 26: Okullo J.B.L., Hall J.B. and Obua J Leafing, flowering and fruiting of Vitellaria paradoxa subsp. nilotica in savanna parklands in Uganda. Agroforest. Syst. 60: Rathcke B. J. and Lacey E.P Phenological patterns of terrestrial plants. Annu. Rev. Ecol. Syst. 16: Sakai S. Phenological diversity in tropical forests Popul. Ecol. 43: Smith-Ramirez C. and Armesto J.J Flowering and fruiting patterns in the temperate rainforest of Chiloe, Chile ecologies and climatic constraints. J. Ecol. 82: van Schaik C.P., Terborgh J.W. and Wright S.J The phenology of tropical forests: adaptive significance and consequences for primary consumers. Annu. Rev. Ecol. Syst. 24: Venkateswaran, R Short-term tree population changes, growth and phenology of woody species in tropical dry evergreen forests on the Coromandel coast of India. Ph.D. thesis. Pondicherry University, Pondicherry, India. JOURNAL OF BIOSCIENCES RESEARCH 2(2):
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