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Reproductive activity patterns of four species of Cinnamomum in Purwodadi Botanic Garden

The aims of this study were to know flowering and fruiting pattern of the four Cinnamomum and their response in relation to seasonal change in rainfall, temperature and air humidity in Purwodadi Botanic Garden. The vegetative and reproductive activities were observed once per week based on the abundance of leaves, flowers and fruits using a scoring method from 0-4, for 6-8 years of observations. The rainfall per month, temperature, and humidity were simultaneously recorded. Data were analysed by Principal Component Analyses to determine the microclimate variables that have correlation to the reproductive activity patterns of all four species. Most species started flower budding during Juni – August, except C. burmanii which produced flowers 2-3 times per year in small quantity. Flowering of C. burmanii showed positively correlations with rainfall intensity, while flowering the other three species did not relate to microclimate in PBG. Six dry months preceding a long dry season in 2015 caused C. burmanii did not produce flowers and fruits.


Introduction
Cinnamomum is a genus of evergreen aromatic trees belonging to the Lauracea family. The species of Cinnamomum have aromatic oils in their leaves and bark. The genus includes a great number of economically important trees. Many Cinnamomum species are well known as spice and medicine. Main characteristic of the Cinnamomum genus is their leaves are mostly triplinerved or sometimes inconspicuously five-nerved, with conspicuous midrib on both surfaces. Mature leaves are dark green, while young leaves are reddish brown to yellowish-red. Cinnamomum is a genus of monoecious species, have hermaphrodite flowers, small, greenish white, white to yellow. The purplish-black fruit is an ovate, ellipsoidal or subglobose drupe. There are about 250 species of the genus Cinnamomum of the family Lauraceae in the world and they are spread out mostly in tropical and subtropical Asia [1].
Phenology can be defined as scientific study of the seasonal timing of life events. In plants, it is related to dates of plant growth phenomenon, such as flowering, leaf flushing or ripening of fruit [2]. Plant phenological patterns are influenced by environmental and biotic processes. In tropical regions, there is less seasonality with regard to temperature and day length; indeed, there is often greater diurnal than seasonal temperature variation. However, many tropical regions experience seasonality of rainfall, and phenological patterns are often related to this seasonality with peaks in fruit production occurring at the beginning of the rainy season [3,4]. Climate elements affect the phenology of plants such as the vegetative phase and reproductive components. This effect can be minor or major. It is suggested that  [5]. The production of flowering trees depends on the regularity of vegetative and reproductive activity. In the tropics, plant phenology is influenced by hydro period variation. The beginning and the end of rainy season cause the changes of plant vegetative and reproductive activities. Flowering, fruiting and seed set are very sensitive to water deficit [6].
Purwodadi Botanic Garden (PBG) is an ex situ conservation institution that has a main task to conserve plant species diversity. The garden has a collection of 2055 plants species including some Cinnamomum species (Registration data of Purwodadi Botanic Garden until December 2018). There are six species of Cinnamomum in the garden, namely Cinnamomum burmanii, C. iners, C. verum, C. sintoc. C. camphora, and C. celebicum. The first four species are periodically flowering and fruiting in the garden, but C. camphora and C. celebicum have never bloomed (preliminary research). Each plant species has a different pattern of flowering and fruiting time as influenced genetic and environmental properties [2]. Purwodadi Botanical Garden is included in a slightly wet tropical climate (Group C) with a dry climate (Group E) which occurs regularly, seasonal variations often occur [7]. The seasonal variations in the Purwodadi Botanic Garden appeared to affect phenological of species plant collections. Some fruit tree species showed different reproductive activities in response to hydro period variation in the Purwodadi Botanic Garden [8]. Environmental factors are related to the flowering of four Annona species that grow on Purwodadi Botanic Garden. Rainfall intensity (0-550 mm) affected to the flowering of Annona muricata, temperature (25,56-28,33°C) and humidity (66,83-85,02%) to A. squamosa , and humidity to A. glabra (71,62-85,02%) and A. montana (71,62 to 82,94 %) as well [9]. The aim of this research are to determine the reproductive activity pattern of all four Cinnamomum species and to know their respon in relation to seasonal change in rainfall, temperature and air humidity in Purwodadi Botanic Garden.

Methods
Four Cinnamomum species growing in PBG i.e: Cinnamomum iners, C. sintoc C. burmanii, and C. verum were observed their vegetative and reproductive activities once per week for 6-8 years of observation. Vegetative activities were the activity of plants in the formation of young leaves, mature leaves and senescence leaves. Whereas reproductive activities consist of flowering and fruiting activity that was found flowers (buds and blooms) and fruits (immature and ripe fruit) on plants [12]. The observation was based on phenological characteristic such as the abundance of leaves (young, mature, senesence), flowers (bud, bloom) and fruits (immature, ripe) by scoring 0-4 in Table 1 [7,11]. Reproductive activities pattern was determined by frequency of flowering and fruiting of plant species in one year. There are continual (flowering or fruiting with sporadic brief break), sub annual (flowering or fruiting in more than one cycle per year), annual (only one major cycle per year), and supraannual (one cycle over more than one year) [12,13]. The rainfall per month (mm), temperature (°C), and humidity (%) were simultaneously recorded. All data were analyzed by Principal Component Analyses (PCA) to determine the microclimate variables that have correlation to the reproductive activity patterns of all four species.

Reproductive activity patterns of four Cinnamomum species
Growth of young leaves of C. iners showed similar pattern to two other species of Cinnamomum i.e. C. burmanii and C. verum (Table 2). This species produced young leaves throughout the year. Whereas C. sintoc, had sub-annual patterns, young leaves grew two/ three times in a year. In 2015, C. burmanii did not produce young leaves for 6 dry months (rainfall <100 mm) (Figure 1). While the mature leaves and old leaves of the four Cinnamomum species showed the same continual patterns, because the four species are known as evergreen trees. Flowering of C. iners and C. sintoc showed the same patterns, their flowering occured once a year (annual). Cinnamomum iners produced flowers in July-August, when rainfall decreased (Table 2). Similarly, C. sintoc flowered in June-July, when rainfall decreased. While C. burmanii and C. verum had different patterns from C. iners and C. sintoc, namely subannuals (Table 2). This trees which categorized as subannual flowered 2-3 times yearly. C. burmanii flowered with minor period in January-March. Whereas C. verum flowered with minor period in April-May and major period in July /August (Figure 2). This condition is possible in the tropics because the sun passes overhead twice each year, influencing insulation rates and weather patterns. Biotic and abiotic factors have entrained a cyclic rhythm in the plants, at over evolutionary time. When conditions are not favorable for flowering, plants will be reduced of flowering intensity until this condition favorable for flowering [14]. This is thought  From four Cinnamomum species observed, C. burmanii showed reproductive activity that was different from the others. Cinnamomum burmanii produced flowers and fruits only slightly abundance (score <1). While fruiting times of C. burmanii occured in June/ July. Cinnamomum verum produced fruits twice yearly. Fruiting major period occurred in August, September, and October. Cinnamomum verum showed more regularity in flowering and fruiting seasons compared to the other three species (Table 2).
Fruiting times of C. iners and C. sintoc showed the same annual patterns, those plants produced fruit once a year. Cinnamomum iners started to fruit in August and the fruiting peak was attained in September/ October (Figure 3). Cinnamomum sintoc onset to fruit in July/ August and the peak of fruiting times attained in September/October. In 2010, like C. iners, C. sintoc produced fruits with minor period.  (Figure 4). Based on the results of the PCA analysis, there were a relationship between climate factors and vegetative activity of C. sintoc, C. burmanii and C. verum. Whereas the vegetative activity of C. iners did not relate to climate ( Figure 5). The abundance of mature leaves of C. burmanii showed a negative relationship to rainfall intensity, while the abundance of deciduous leaves was positively associated with humidity. Abundance of C. sintoc young leaves decreased, but senescence leaves will be more  Flowering of C. iners, C. sintoc, and C. verum was not related to climate, but flowering of C. burmanii showed a positively associated with rainfall intensity. Decreasing rainfall caused C. burmanii produced little flowers. More over a long dry season (6 dry moths) in 2015 caused this species did not produce flower. The low C. burmanii flower produced was related to adaptability of this species. Cinnamomum burmanii grows best at an elevation between 500-1,500 metres, whereas Purwodadi Botanic Garden is located at 350 meter above sea level (masl). In addition, this species prefers a mean annual rainfall in the range 2,000 -2,500 mm, but tolerates 1,800 -3,500 m. During observation, when along dry season, rainfall intensity in the garden was less 2000 mm.
Flowering of C. iners, C. sintoc, and C. verum which were not influenced by microclimate showed that the three species of plants carry out flowering activities regularly, not affected by the increasing and decreasing of rainfall, temperature and humidity. This indicates that these species are more adaptable to the changes of environment, so there is no change in flowering and fruiting patterns [13]. Figure 5 showed that the fruiting process of C. burmanii did not relate to climatic factors. This was because during the 4 years of observation this species did not produce fruit (its flowers were very litlle). However, fruiting of others Cinnamomum were related to air temperature and humidity. The fruiting of C. sintoc was negatively correlated to humidity. The increasing humidity caused decreasing of C. sintoc fruits. The fruiting of C. verum was related to temperature, increasing the temperature would increase its fruiting. The fruiting of C. iners was positively correlated to temperature and negatively to humidity. Figure 5 showed that the fruiting of C. iners would increase with the increasing temperature and decreasing humidity. Cinnamomum iners never produced ripe fruit, when longer wet months or longer dry months.

Conclusions
Flowering of C. iners and C. sintoc occured once a year (annual), while C. burmanii and C. verum flowered twice yearly. Flowering and fruiting time for Cinnamomum species was almost similar, when rainfall decreased until the beginning of the dry season. Flowering of C. iners, C. sintoc, and C. verum was not related to climate, but flowering of C. burmanii showed a positively associated with rainfall intensity. Whereas the fruiting of C. iners, C. sintoc, and C. verum were associated with temperature and humidity.