Phenological Behavior of Cenostigma pyramidale (Tul.) E. Gagnon & G.P.Lewis in an Area of Caatinga in the Semi-Arid Region of Paraíba

Abstract

This study aimed to evaluate the phenophases of Cenostigma pyramidale (Tul.) Gagnon & G.P.Lewis in an area of Caatinga in the semi-arid region of Paraíba, Brazil. Field research was carried out in the municipality of Sumé, in the Experimental Space reserved for Studies on Ecology and Dynamics of the Caatinga - Area I of the Laboratory of Ecology and Botany - LAEB/CDSA/UFCG (7°39'38.8'' S and 36°53'42.4'' W; 538 m altitude). A total of 96 plots of 10 x 10 m were arranged in the study area, seeking to identify one specimen of C. pyramidale per plot. Monitoring was performed monthly from March 2019 to February 2020, recording budding, senescence, flower bud, flowering, and fruiting data. The Fournier intensity percentage and the activity index were analyzed. The highest budding intensity index was observed when there was a gradual increase in precipitation. Senescence had the highest peaks of intensity in the months with the absence or reduced levels of precipitation. Flower bud and flowering were practically equal throughout the monitoring period. The highest fruiting intensity occurred in the month with the highest precipitation. Regarding the budding activity index, all specimens expressed this phenophase as soon as the precipitation levels increased. In the dry period, all specimens showed senescence. Flower bud and flowering phenophases occurred synchronously. Fruiting had a higher activity index in periods of higher precipitation. Thus, it was observed that precipitation has a great influence on phenological events.

Keywords:
Semi-aridity; Catingueira; Phenophases

HIGHLIGHTS

The budding, flower bud, and flowering phenophases were more intense in months with high precipitation.

Senescence occurs intensely in periods of absence of rainfall.

Specimens of C. pyramidale showed fruiting synchronicity.

INTRODUCTION

Phenology is of great importance in surveying the dynamics of the Caatinga, consisting of studying the periodicity of events (phenophases) such as budding, flowering, fruiting, and seed dispersal [¹]. From the knowledge of these events, reproductive patterns can be characterized, which can be considered relevant elements for assessing the behavior of species in a given environment [²]. The phenophases are related to climatic and meteorological conditions such as rainfall, solar radiation, air temperature, evaporation, and relative air humidity, as well as to local elements, such as soil and photoperiod, and biotic components such as diseases and pests [³]. Thus, detailing the phenological characteristics of forest species is important, as they have different periodicity and behavior, in addition to the differences between them according to the ecosystem in which they occur [⁴].

Cenostigma pyramidale (Tul.) E. Gagnon & G.P.Lewis is popularly known as “catingueira”, a medium-size species belonging to the Fabaceae family, which can be used for different purposes, especially as wood and fodder, being also known as a pioneer species and of great importance for ecological succession [⁵,⁶,⁷]. This plant is considered a good source of energy because its wood contains a high content of cellulose and lignin, in addition to having the potential for the production of fuel alcohol and metallurgical coke [⁸, ⁹]. Its leaves, flowers, and bark are used in folk medicine to treat diarrhea, catarrhal infections, anemia, and hepatitis, as well as in the treatment of worms in domestic animals [¹⁰,⁹,⁶]. C. pyramidale is one of the most studied species and its phenology is directly related to precipitation pulses, due to its rapid response to the occurrence of rainfall [¹¹].

Therefore, this study aimed to evaluate the phenophases of Cenostigma pyramidale (Tul.) Gagnon & G.P.Lewis in an area of Caatinga in the semi-arid region of Paraíba, Brazil.

MATERIAL AND METHODS

Study Area

This study was carried out in an area of Caatinga in the municipality of Sumé (Paraíba, Brazil), located in the western Cariri microregion. The rainfall in the region is concentrated in 3 - 4 months, with annual averages between 250 and 900 mm, being irregular and poorly distributed in time and space, with average annual temperature ranging from 25 - 27°C and average insolation of 2,800 h/year [¹²]. According to these authors, the relative air humidity is on average 50% and the average evaporation rates are approximately 2,000 mm/year. Hyperxerophytic caatinga vegetation predominates in the region where the soils are mostly well-developed Chromic Luvisols with slightly undulating relief [¹³,¹⁴].

Field research was conducted in the Experimental Space reserved for Studies on Ecology and Dynamics of the Caatinga - Area I of the Laboratory of Ecology and Botany - LAEB/CDSA/UFCG (7°39'38.8'' S and 36°53'42.4'' W; 538 m altitude). This space covers an area of 1.05 ha, where 96 plots of 10 x 10 m were arranged (Figure 1).

Figure 1
Location of the Experimental Space reserved for Studies on Ecology and Dynamics of the Caatinga - Area I (LAEB/CDSA/UFCG) and the distribution of monitoring plots in an area of Caatinga in the Cariri region of Paraíba.

Data collection and analysis

A total of 96 contiguous plots of 10 x 10 m were established in the Experimental Space reserved for Studies on Ecology and Dynamics of the Caatinga - Area I (LAEB/CDSA/UFCG). For monitoring the phenophases of Cenostigma pyramidale (Tul.) E. Gagnon & G.P.Lewis, we sought to identify one specimen per plot; however, considering that the species did not occur in two plots, 94 specimens were selected. The individuals in these sampling units were marked and numbered with aluminum platelets.

Monitoring was performed monthly from March 2019 to February 2020, recording budding, senescence, flower bud, flowering, and fruiting data. The intensity of phenological events was estimated for each tree according to Fournier’s (1974) criteria. The obtained data were recorded in Excel spreadsheets, version 2018^®.

The collected data were analyzed based on the activity index (percentage of individuals), which is characterized as a simple method, considering the presence or absence of phenophase in the individual, without estimating the intensity or quantity. Thus, this method has a character of quantitative expression of the observed population, indicating the percentage of individuals in the population and expressing a certain phenological event. Morellato and coauthors (1990) [¹⁸] point out that through this method it is possible to estimate the synchrony between the individuals of a population, because the greater the number of individuals expressing a phenophase at the same time, the greater the synchrony of the population.

The Fournier intensity percentage, proposed by Fournier (1974), was also used. This method can be applied to estimate the percentage of phenophase intensity in each individual, obtaining the values in the field through a semi-quantitative interval scale of five categories (0 - 4), with a 25% interval between each category. Thus, for each monthly monitoring, the intensity values obtained for all individuals of the species were summed and divided by the maximum possible value (number of individuals multiplied by four). The obtained value, corresponding to a proportion, was converted to a percentage by multiplying by 100.

To obtain the intensity of each plant, their crowns were divided into quadrants and applied to the semi-quantitative categories (0 - 4). After observation in the field, the data were recorded in Excel spreadsheets, version 2018^®.

RESULTS

Considering the behavior of phenophases in Cenostigma pyramidale (Tul.) E. Gagnon & G.P.Lewis based on the Fournier intensity and its relationship with precipitation (Figure 2), the species had the highest intensity in the leaf budding phenophase between December 2019, January 2020, and February 2020, when there was a gradual increase in precipitation.

Figure 2
Behavior of budding and senescence phenophases in C. pyramidale based on the Fournier intensity and precipitation from March 2019 to February 2020 in an area of Caatinga in the municipality of Sumé (Paraíba, Brazil).

The senescence reached the intensity peak in December, indicating higher values in October and November 2019, when the absence or reduced levels of precipitation were observed. In January and February 2020, the specimens had low intensity of senescence, when an increase in precipitation levels was recorded.

The flower bud and flowering phenophases in C. pyramidale showed a synchronism, which had practically the same behavior in all monitoring months (Figure 3). These phenophases had the highest values of intensity in April 2019, which stood out with the highest precipitation. From June 2019 to January 2020, the specimens did not express these phenophases, when the absence or low levels of precipitation were recorded. These phenophases were recorded again in February 2020, due to the rainy season.

Figure 3
Behavior of flower bud, flowering, and fruiting phenophases in C. pyramidale based on the Fournier intensity and precipitation from March 2019 to February 2020 in an area of Caatinga in the municipality of Sumé (Paraíba, Brazil).

Regarding fruiting, the highest intensity occurred in April 2019, which had the highest precipitation rate (Figure 3). The presence of fruits was recorded during the entire monitoring period, evidencing they are quite resistant, as their formation occurred at the beginning of the rainy season and, at the end of the dry season, fruits were still recorded in some specimens.

Activity Index (Percentage of Individuals)

Regarding the budding activity index, at the beginning of the monitoring period (March 2019), this phenophase was not observed in the specimens. Between April 2019 and May 2019, some of them developed new leaves and from June 2019 to September 2019 the plants did not express the referred phenophase (Figure 4). In October 2019, budding occurred in some specimens, in response to precipitation pulses, and in the following month, this process no longer occurred. Only in December 2019, a high percentage of individuals started budding again, due to the return of precipitation. In January and February 2020, the highest percentage of individuals was recorded in this phenophase, i.e., budding was observed in 100% of the plants, and these months had an increase in precipitation levels.

Figure 4
Behavior of budding and senescence phenophases in C. pyramidale based on the percentage of individuals and precipitation from March 2019 to February 2020 in an area of Caatinga in the municipality of Sumé (Paraíba, Brazil).

Senescence was observed in some specimens right at the beginning of the monitoring. This phenophase remained until February 2020, i.e., leaf fall was observed in some specimens even in the rainy season (Figure 4). From June 2019 to October 2019, this phenophase had the highest synchronism. Thus, senescence was recorded in 100% of the specimens at the beginning of the dry period.

The percentage of individuals that triggered leaf fall had a slight decrease between November 2019 and December 2019, with practically the same percentage during these months. This behavior abruptly changed between January 2020 and February 2020 when the percentage of individuals greatly decreased, and an increase in precipitation was observed (Figure 4).

The flower bud and flowering phenophases occurred synchronously, i.e., with practically the same activity index, throughout the monitoring period, and between March 2019 and April 2019, flowering had a slight increase in comparison with flower bud (Figure 5). The activity peak of these phenophases was observed in April 2019, when the highest precipitation rate was recorded. Only from February onwards some specimens of C. pyramidale developed flower buds and flowering again when the rainy season began.

Figure 5
Behavior of flower bud, flowering, and fruiting phenophases in C. pyramidale based on the percentage of individuals and precipitation from March 2019 to February 2020 in an area of Caatinga in the municipality of Sumé (Paraíba, Brazil).

From May 2019 to June 2019, there was a significant decrease in the number of specimens expressing the flower bud and flowering phenophases, and from July 2019 to January 2020 none of them went through these processes, because it was the dry season, considering that there were some precipitation pulses, but of short duration and low volume.

Concerning the fruiting, some specimens started this phenophase at the beginning of the monitoring period, with a higher activity index in April 2019, which had the highest level of precipitation (Figure 5). This process remained practically constant from May 2019 to August 2019. Soon after, there was a decrease in the percentage of individuals, again remaining practically constant from November 2019 to December 2019, with lower activity rates from January 2020 to February 2020.

DISCUSSION

Assessing the phenology of woody species from the Caatinga, Amorim and coauthors (2009) [¹⁹] found that leaf emergence was influenced by water availability, and all plants had the complete formation of leaves, from February to April, at the peak of the rainy season. According to these authors, the species did not have the same budding pattern, which may vary due to the sporadic rains in the dry season. In their research, some species such as Mimosa tenuiflora (Willd.) Poir. (“jurema preta”) and Jatropha mollissima (Pohl) Baill. (“pinhão bravo”) started the budding phenophase after the incidence of rainfall, whereas other species such as Anadenanthera colubrina (Vell.) Brenan var. cebil (Griseb.) Altschu (“angico”) and Handroanthus impetiginosus (Mart. ex DC.). Mattos (“ipê roxo”) did not have the same behavior. Lima and coauthors (2018) [¹¹] highlight the importance of precipitation in phenological responses, emphasizing that when interpulses occur, the water availability in the soil tends to decrease for the vegetation; because of this, C. pyramidale starts the leaf senescence phase.

Souza and coauthors (2014) [²⁰] studying the phenology of six tree species in a conserved area of Caatinga, found that C. pyramidale started the senescence in a concentrated manner at the end of the dry season, whereas in other species, senescence occurred at the end of the rainy season and beginning of the drought period.

Parente and coauthors (2012) [²¹] studying the same species in the Cariri Paraibano, found a greater intensity of leaf fall from the beginning of the dry season, thus evidencing the deciduous habit of the species. Amorim and coauthors (2009) [¹⁹] explain that C. pyramidale is among the species that may not suffer total leaf loss in one year but may lose all leaves in just one to two months in another year, usually from September or October, indicating that this species has no uniformity regarding this process.

The results on flower bud phenophase recorded in our research are similar to those found by Lima and coauthors (2018) [¹¹], who also assessed the phenological behavior of C. pyramidale and observed that in 2010 the production of flower buds began in February and reached a maximum peak in April. Rocha and coauthors (2015) [²] studying the phenology of Copernicia prunifera (Mill.) H.E. Moore. (“carnaúba”) also in an area of Caatinga, found that the reproductive phase of the species began with the emergence of flower buds, reaching greater intensity in May 2010 and June 2011, whereas in 2012 the intensity of flower bud emergence had a decrease compared to the previous years.

Parente and coauthors (2012) [²¹] reported similar results, in which flowering occurred during the rainy season, especially in April and May, when precipitation peaks occurred. According to Souza and coauthors (2014) [²⁰], flowering in C. pyramidale can last six months, varying according to environmental conditions.

Melo and coauthors (2021) [²²] explain that assessing flowering and fruiting events in areas of Caatinga is more complex due to variations in soil, relief, microclimate, and vegetation, with species having different flowering periods. Thus, [²³] emphasize that the distribution of rainfall in each month and environment is of great importance for the maintenance and implementation of some phenophases according to each environment.

Melo and coauthors (2021) [²²] observed that in the ten species investigated, the emergence of new leaves occurred in the rainy season and at the beginning of the dry season, showing that the budding phenophase is directly related to precipitation. Regarding the results on the synchronism of the leaf fall phenophase, the same characteristic is found in deciduous species in areas of Caatinga, i.e., in a certain period of the year their physiognomy changes according to the climatic season [²⁴].

Souza and coauthors (2014) [²⁰] observed that the leaf fall in C. pyramidale begins in a concentrated manner at the end of the dry season, a characteristic of species from dry environments, where plants trigger their phenophases mainly depending on precipitation.

Becerra and coauthors (2015) [²⁵] emphasize that the rainfall regime can cause changes in the different phenological events of the vegetation, such as variations in the beginning, end, amplitude, and extension of the plant growth period.

According to Parente and coauthors (2012) [²¹], fruiting depends on rainfall distribution, since fruit formation consequently depends on flowering, and the fruiting phenophase in C. pyramidale occurs in the rainy season.

According to Leite and coauthors (2009) [²⁶] the regrowth ofC. pyramidaleoccurs at the beginning of the rainy season. Souza and coauthors [²⁷] emphasize that the flowering of C. pyramidale occurs over four months, especially in the rainy season, between the months of january and april.

CONCLUSION

The highest Fournier intensity index for budding phenophase in Cenostigma pyramidale (Tul.) Gagnon & G.P.Lewis occurred when a gradual increase in precipitation was evident. The senescence had the highest intensity peaks in the months with the absence or reduced levels of precipitation and low intensity when precipitation levels increased. Flower bud and flowering had practically the same behavior throughout the monitoring period, and the highest intensity was observed in the month with the highest precipitation rate. In this sense, the highest fruiting intensity also occurred in the high precipitation month. As for the budding activity index, all specimens expressed this phenophase as soon as the precipitation levels increased. At the beginning of the dry season, all individuals showed senescence and even in the rainy season some of them expressed this process. The flower bud and flowering phenophases occurred synchronously, i.e., with practically the same activity index during almost the entire monitoring period; in the dry season, these phenophases did not occur. Fruiting had a higher activity index when there was a higher level of precipitation. Therefore, it was observed that rainfall has a great influence on phenological events, especially the reproductive ones, which are more demanding of water.

Acknowledgments

All authors express their gratitude to the members of the Ecology and Botany Laboratory (LAEB/CDSA/UFCG), to the Ecosystem Conservation and Recovery of Degraded Areas in the Semiarid Research Group (CERDES) and everyone from the National Integration Routes Project of the Ministry of Integration and Regional Development - Restoration of Degraded Riparian Ecosystems in the Brazilian Semiarid (REDESAB).

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• Funding:

  This research received no external funding.

Edited by

Editor-in-Chief:

Alexandre Rasi Aoki

Associate Editor:

Marcos Pileggi

Publication Dates

• Publication in this collection
  31 May 2024
• Date of issue
  2024

History

• Received
  11 July 2023
• Accepted
  12 Sept 2023