Journal of Environmental Analysis and Progress V. 02 N.

03 (2017) 258-265

Journal of Environmental
Analysis and Progress
Journal homepage: www.jeap.ufrpe.br/
ISSN: 2525-815X http://dx.doi.org/10.24221/jeap.2.3.2017.1453.258-265

Rainfall events, high CO2 concentration, and germination of seeds in Caatinga

Fabrício Francisco Santos da Silvaa,b, Gilmara Moreira de Oliveiraa,b, Marcelo do Nascimento
Araújob, Francislene Angelottib, Magna Soelma Beserra de Mourab, Bárbara França Dantasb*
a
Universidade Estadual de Santana-UEFS, Departamento de Ciências Biológicas, Feira de Santana, Bahia, Brazil. CEP
44036-900. E-mail: fabriciofrancisco2006@gmail.com; gilmara_5@hotmail.com.
b
Embrapa Tropical Semi-Arid, Brazilian Agricultural Research Corporation, Embrapa Semiárido, Petrolina, Pernambuco,
Brazil CEP 56302-970. E-mail: dr.marcelo_araujo@outlook.com; francislene.angelotti@embrapa.br;
magna.moura@embrapa.br; barbara.dantas@embrapa.br. *Corresponding author.

ARTICLE INFO ABSTRACT

Received 29 Jun 2017 In semi-arid environments, the distribution of rainfall over time is decisive in
Accepted 10 Jul 2017 emergence in a soil seed bank. The objective of this study was to evaluate the effect
Published 31 Jul 2017 of rainfall events on the soil seed bank emergence of Caatinga species under high
CO2 concentration. The experiment was carried out in an experimental area of
Embrapa Semi-arid in a randomized complete block design, with subdivided plots.
Each plot consisted of different environmental conditions (open top greenhouse with
an injection of 550 ppm CO2; open top greenhouse and environmental CO2; natural
environment). The subplots consisted of the depth at which the seeds of Poincianella
pyramidalis and Myracrodruon urundeuva were sown (superficially and buried at
0.02 m and 0.06 m depth). Seedling emergence was monitored daily after the first
rains. During the experiment, weather data showed a rainfall volume of 83 mm, an
average air temperature of 28.7oC, average soil temperature of 35.4 and 34.9oC, at
depths of 0.02 and 0.06 m, respectively. Seedling emergence started 56 days after
sowing and four days after the first rains. After 154 days of the onset of the
experiment, drip irrigation was performed. The greenhouse environment, regardless
of the addition of CO2 or not, allowed higher emergence percentage of P.
pyramidalis. The seeds of this species sowed on the soil surface only emerged when
irrigation started. M. urundeuva seeds showed low germination even after irrigation.
Keywords: Emergence, Poincianella pyramidalis, Myracrodruon urundeuva, soil
seed bank.

Introduction catingueira-verdadeira (Poincianella pyramidalis

Caatinga Biome covers an extensive area of (Tul.) L.P. Queiroz).
the Brazilian Northeast, characterized by a Myracrodruon urundeuva is a native species
semiarid climate, with stochastic rainfall events, of Brazil, occurring in Northeast, Southeast and
300-1000 mm/year concentrated in three to five Central-West regions, occurring in Caatinga,
months during the year. Its main vegetation is tree Cerrado, and Atlantic Forest biomes. It flourishes
and shrubs, with specific adaptations to its harsh between June and July, and the fructification is
habitats, such as loss of leaves in the dry period, between August and December. This
small leaves, thorns and xerophytic adaptations Anacardiaceae is considered one of the native
(Queiroz, 2009). In this ecosystem is found a species that presents the most resistant wood
diversity of vegetal species of considerable because its core practically doesn’t rot. This
economic and biological value, as aroeira-do- species can be used in reforestation, forage,
sertão (Myracrodruon urundeuva Allemão) and ornamental and medicinal purposes (Maia, 2012).
Due to this characteristic and its medicinal

Silva, F. F. S. da; Oliveira, G. M. de; Araújo, M. do N.; Angelotti, F.; Moura, M. S. B. de 258
 Journal of Environmental Analysis and Progress V. 02 N. 03 (2017) 258-265
properties, it has been widely explored and and conserving population genetic variation in the
included in the official list of endangered species long term (Bossuyt & Honnay, 2008).
of Brazilian flora in the vulnerable class, according Caatinga soil seed bank presents a high
to the Ministry of the Environment (Brasil, 2008). diversity of species (Ferreira et al., 2014;
Seed germination of M. urundeuva occurs Fabricante et al., 2016; Paz, Silva & Almeida-
over a temperature range of 20 a 35oC (Virgens et Cortez, 2016), remaining in the soil during an
al., 2012; Oliveira et al., 2014). However, 40% of unfavorable period for emergency (Mamede &
freshly harvested seeds germinate at 40oC (Oliveira Araújo, 2008). However, Caatinga soil seeds bank
et al., 2015). Seeds of M. urundeuva are more must complete its life cycle within the short rainy
tolerant to osmotic stress induced by PEG 6000 season to maintain a viable population.
than induced by NaCl with germination thresholds Brazilian northeast is, both environmentally
at -0,9 MPa and -0,56 MPa, respectively (Dantas et and socially, the most vulnerable region to climate
al., 2014; Oliveira et al., 2014b; Oliveira, 2015). change in Brazil (Stocker, 2013). Increasing
Poicianella pyramidalis is an endemic temperatures up to 4oC and decreases in rainfall
Leguminosae from Caatinga, with extensive use in events can lead to desertification, implying not
the Brazilian semi-arid region due to its potential only climatic but phytogeographical, economic and
for reforestation, forage, timber, and medicinal. It social changes (Nóbrega, Santiago & Soares,
is a medium-sized tree, of four to 12 meters height 2016). However, these variations are not limited
with an open and irregular canopy. Its trunk has only to water deficit and temperature increase, but
about 50 cm in diameter, light gray bark, with also to the increase in the atmospheric
rhytidome that is detached in elongated and concentration of carbon dioxide [CO2] (Araújo et
irregular blades. Its flowers are yellow and al., 2015).
arranged in racemes. Flowering is from October to In recent years several studies have analyzed
February and fructification from December to the potential effects of increased concentrations of
June. It adapts to the most varied types of soil, greenhouse gases in the atmosphere on seeds,
mainly stony ones (Siqueira-Filho et al., 2009; seedlings and adult plants of various native and
Maia, 2012). cultivated species (Dantas et al., 2017; Costa et al.,
This species presents a wide range of 2017). However, an almost unexplored but
environmental tolerance, reaching 10m height important feature of global climate change is
when water is available during its development and changing patterns of rainfall, with likely increases
a shrub size (2 m) when the water supply is in rainfall variability (Stocker, 2013).
restricted (Antunes, 2012).These shrubby-trees are This study aims to evaluate if the [CO2] in
also adapted to different types of soils, including the atmosphere can influence the seedling
the poorest and stony soils (Maia, 2012), and emergence and recruitment of soil seed banks; how
presents high-density populations throughout small rainfall events onset seeds germination and
Caatinga. Adult plants and seeds of P. pyramidalis emergence of seedlings of two native Caatinga
have very high tolerance to water deficit (Antunes species sowed at different depths of the soil, by
et al., 2011) and saline and sodic environments simulating different conditions of a soil seed bank
(Matias et al., 2013). of these species at Caatinga.
Soil seed banks represent a stock of
regeneration potential and an important component Material and Methods
of resilience in plant communities. Soil seed bank Seeds of P. pyramidalis were harvested from
also plays a key role in vegetation dynamics in 17 trees, in the district of Massaroca, at Juazeiro-
degraded ecosystems and some stressful BA (9°52’09”S, 40°16’42”W, 469 m of altitude),
environments (González-Alday et al., 2009; in July 2016. Myracrodruon urundeuva seeds were
Bossuyt & Honnay, 2008). Knowledge harvested from seven trees in the district of Jutaí,
mechanisms that maintain natural community at Lagoa Grande, PE (08°33’33”S, 40°12’10”W,
dynamics may result in an understanding of soil 409 m altitude), in September 2016.
seed banks and its relation to vegetation (Ooi, The original vegetation of the areas is
2015), which in turn can contribute to improving characterized as hyper xerophilic Caatinga
land management practices (Hopfensperger, 2007). (Amaral & Fernandes, 2007), with shrubs and
The ability of plant species to produce seeds scattered trees. The soil is yellow dystrophic Oxisol
remaining viable in the soil allows them to (IBGE, 2006), with hot and dry climate, classified
overcome temporally unsuitable stressful as BSh, according to Köppen, which is, semi-arid
environmental conditions for germination and with average temperatures above 18°C (SEI, 1998;
establishment, spreading germination risk in time Alvares et al., 2013).

Silva, F. F. S. da; Oliveira, G. M. de; Araújo, M. do N.; Angelotti, F.; Moura, M. S. B. de 259
 Journal of Environmental Analysis and Progress V. 02 N. 03 (2017) 258-265
After harvested, P. pyramidalis fruits were (chalice). Then, seeds were submitted to a seed
dried under a plastic canvas in a shadowed blower to separate the impurities, based on size
protected environment. Afterward, fruits were differences, weight and density (Matias, Oliveira &
beaten and seeds collected. In the laboratory, by Dantas, 2014).
visual evaluation, damaged seeds and other For evaluation of the effect of rainfall
impurities (fruit debris, branches, seedlings and events and environmental [CO2] on the soil seed
other species, among others) were discarded to bank, this trial was carried out in the rainy season,
homogenize and purify the lot (Matias, Oliveira & between October 25, 2016, and April 20, 2017, at
Dantas, 2014). the Experimental Field of Caatinga, Embrapa
Processing of M. urundeuva seed was Semi-Arid, Petrolina-PE.
carried out by removal of the branches and wings

Figure 1. Experimental field of Caatinga, Embrapa Semi-arid, in Petrolina-PE. A. Overview of the

experimental area; B. Aluminium circular structure with PVC rigid film, CO2 injection hoses placed laterally;
C. Seeds sowed on the soil surface and different depths, with different infrared gas analyzer- IRGA sensor at
center; D1. Dead; D2. Live Poincianella pyramidalis seedlings after 45 days with no rain. Photos: Angelotti,
A.; F.; B-C. Silva, F. F. S.; D. Dantas, B. F. (2017).

The experimental design was in randomized Two CO2 injection hoses were placed along
blocks, in split-plots. The main plots were the the structure circumference at 0.6 and 0.8 m
environments in which the seeds were sown, and (Figure 1). Infrared gas analyzers (IRGA),
subplots were sowing depths of each species positioned at 0.5 m from the soil, monitored [CO2]
evaluated. Two blocks with three main plots and and information obtained was sent to a controller
three subplots were evaluated, totaling nine which regulated the opening of valves for CO2
treatments for each species. injection in the greenhouses. Pure CO2 was injected
One hundred seeds of P. pyramidalis and against a blower to ensure a proper mixing and to
200 of M. urundeuva were sown superficially and achieve the required concentration when necessary.
buried at depths of 0.02 m and 0.06 m in each Daily monitoring of emergence and seedling
evaluated environment. The three environments of survival was conducted as of the first rainfall event
soil seed banks were characterized as an open-top after the trial setup (November 13, 2016).
greenhouse and 550 ppm CO2 injection; open-top The evaluation of the condition and viability
greenhouse and environmental CO2; natural of non-germinated seeds after rainfall events were
Caatinga environment (no greenhouse and no CO2 made using plastic trays (0.2 x 0.3 x 0.1m) filled
injection). Open top greenhouses were small free with the same soil from the trial site, in which the
air CO2 enhancement (Mini-FACE) structures with seeds of the two species were sown at the same
modifications. The circular aluminum structure of depths (0.02 m and 0.06 m) and kept at the natural
2.0 m diameter and 1.20 m height was sided by 0.40 environment.
mm PVC rigid film.

Silva, F. F. S. da; Oliveira, G. M. de; Araújo, M. do N.; Angelotti, F.; Moura, M. S. B. de 260
 Journal of Environmental Analysis and Progress V. 02 N. 03 (2017) 258-265
After 154 days from the beginning of the rainfall events there was no emergence of P.
trial, drip irrigation was performed to verify the pyramidalis seedlings (Figure 2).
viability of the seeds, of each species, which had The emergence of P. pyramidalis seedlings
not germinated from the soil seed bank. started four days after a second rain event, at 56
Soil temperature measurements were days after sowing. On this occasion, a 9.5 mm
performed with sensors connected to the CR1000 rainfall event occurred, which resulted in an
data acquisition system (Campbell Scientific Inc., emergence of 5% of P. pyramidalis seedlings of
Logan, UT, USA), programmed to perform this species in greenhouses with CO2 injection
measurements at every 60 s with averages at each (Figure 2BC) and 2.5% of seedlings in greenhouses
30 min. The sensors CS107 (Campbell Scientific without CO2 (Figure 2EF) in the treatments in
Inc., Logan, UT, EUA) were inserted into the soil which the seeds were buried.
at 0.02 and 0.06 m depth. Rainfall data (mm) was After this small rainfall event, few
acquired from climatological stations at a 2000m seedlings, which were approximately 0.05 m high,
distance from the experimental area. survived without additional rainfall for 24 days in
greenhouses with CO2 injection and 40 days in
Results greenhouses without CO2 injection. In this last
During six months in which the soil seed condition, a single P. pyramidalis seedlings
bank was evaluated, there was a pluviometric survived until the third rainfall event, 45 days with
volume of 83 mm, average air temperature of no rain. (Figure 2F).
28.7oC, average soil temperature of 35.4 and In early February, a 17 mm rainfall event
34.9oC, at depths of 0.02 and 0.06 m, respectively occurred, inducing the emergence of additional
(Figure 2). 6.5% and 4.5% of P. pyramidalis seedlings in
Twenty days after initiating the trial there greenhouse environments with and without CO2
were four days of small rainfall events, which injection, respectively (Figures 2BCE).
accumulated 10.3 mm. During or after these

Figure 2. Emergence of Poincianella pyramidalis seeds (columns) at soil surface (ADG); buried at 0.02 m
(BEH) and buried at 0.06 m (CFI) in open top greenhouse with side plastic and injection of 550 ppm CO2 (A-
C); open top greenhouse and environmental CO2 (D-F); and natural environment. Dotted lines refer to daily
average temperature (oC) at seed depth, and full lines refer to rainfall (mm) from 10/25/2016 to 04/25/2017.
Dashed vertical lines indicate irrigation onset. Embrapa Semiárido, Petrolina-PE.

Silva, F. F. S. da; Oliveira, G. M. de; Araújo, M. do N.; Angelotti, F.; Moura, M. S. B. de 261
 Journal of Environmental Analysis and Progress V. 02 N. 03 (2017) 258-265
Greenhouses favored the emergence of P. total rainfall during the year (Sala & Laurenroth,
pyramidalis seedlings, probably by blocking dry 1982). At a time when rains become scarce, due to
winds and maintaining soil moisture. Seeds of this climate changes, little that rains in rare events
species when placed on the soil surface only manage to promote germination onset, but the
emerged once drip irrigation started, after 154 days recruitment of seedlings is compromised (Ooi,
of trail setup (Figure 2ADG). 2015).
For all evaluated conditions, only 0.5% of Poincianella pyramidalis is endemic from
M. urundeuva seeds emerged (data not shown), Caatinga biome and is classified as a pioneer,
even after irrigation. However, when in plastic emerging within four days after the first rains
trays maintained in a natural environment where (Figure 2). However, seeds of P. pyramidalis did
irrigated at the end of the field trial, we found 5% not emerge when sown on the soil surface (Figure
emergence M. urundeuva of seedlings. 2ADG). Direct sowing, with subsequent burial,
protects the seed both by the risks of desiccation,
Discussion as well as predation (Sovo, Tigabu & Odén, 2010).
Establishment of species in the field can be Also, water is not retained in Caatinga soil surface.
facilitated by adult individuals close to the Thus, it is required cover on P. pyramidalis seeds,
seedlings, can be hindered by competition with since at the depths of 0.02 and 0.06 m seedlings
nearby plants (Miranda, Padilla & Pugnaire, 2004). emerged after the rains. In Caatinga, dispersion of
In addition to water availability, interspecific P. pyramidalis fruits occurs in the dry season, a
competition seems to be one of the most influential period of greatest deposition of litter (Souza et al.,
factors in the recruitment of M. urundeuva 2014, 2016).
seedlings, since seedlings of this species that The diversity of species in the Caatinga is
emerged during the trial were not established. maintained through processes that allow
Constant rains are usually crucial to perpetuation, such as seed dispersal and its
establishment of species of fast germination (Vieira accumulation in the soil, forming a soil seed bank
et al., 2008), However, this does not always occur that remains viable for years (Baskin & Baskin,
in semi-arid regions where environmental 2014). Soil seed bank is formed from a balance,
stochasticity, which is random variation in where seeds enter through various dispersion
environmental conditions (Kreyling, Jentsch & mechanisms and exit through recruitment (seed
Beierkuhnlein, 2011), is much more pronounced germination and the establishment of young
than in other environments (Sala & Laurenroth, plants), seed predation and mortality (Gasparino et
1982). al., 2006).
Caatinga biome presents a remarkable In dry forest areas of southern Spain
seasonality, with rainy period concentrated (Caballero et al., 2003, 2005) and desert areas in
between 3-5 months (Maia, 2012). Additionally, China (Wang et al., 2005). The reduction in the
within this gap, rainfall is sparse and limited, number of seeds in the soil is related to death and
favorable for rapid germination, but crucial to the emergence period of seedlings. The risk of
success of the seedling establishment. It tends to mortality becomes more evident when these seeds
aggravate in drought years (from 2010 to 2017) and are exposed to rainfall in early summer, providing
with scenarios predicted for future climates moisture only to germinate and having interrupted
(Marengo, 2014; Marengo, Torres & Alves, 2016). water supply in the initial growth phase (Mesgaran
Highest historical averages of precipitation et al., 2017). However, some species that occur in
in the Caatinga region, in Petrolina-PE, are Caatinga have post-germinative tolerance to
concentrated between November and April. Within desiccation and can resume its initial growth even
this range, the smallest records occurred between after a short period without water as Handroanthus
the years of 2011-2012, with accumulated rainfall impetiginosus (Mart. Ex DC.) Mattos (Vieira et al.,
of 114.5 mm and in the period evaluated in this 2010).
study. In the last 40 years, the average annual Studies aiming to determine the minimum
precipitation is around 510 mm (Embrapa water requirement for germination of seeds that
Semiárido, 2017), approximately 85% more than it present high tolerance to water stress, such as P.
rained during the present study. pyramidalis (Antunes et al., 2011; Matias et al.,
Low soil moisture retention capacity, high 2015), are important tools to evaluate the
evaporation rates, and evapotranspiration are ecological behaviour of these species, such as
aggravating the problem of water deficit (Galvíncio aspects of pre- and post-germination desiccation
& Moura, 2005). This, in turn, can reach up to 70% tolerance, and recruitment and establishment of
per year (Marengo et al., 2011). Thus, events of seedlings in response to current and future
low rainfall (<10mm) can account for up to 80% of environmental conditions. Therefore, a greater

Silva, F. F. S. da; Oliveira, G. M. de; Araújo, M. do N.; Angelotti, F.; Moura, M. S. B. de 262
 Journal of Environmental Analysis and Progress V. 02 N. 03 (2017) 258-265
amount of research should be developed evaluating inicial e nas trocas gasosas do meloeiro amarelo.
the germination ecology of Caatinga seeds. Revista Brasileira de Geografia Física, v. 8,
número especial IV SMUD, p. 439-453.
Conclusion
Environment [CO2] did not influence the BASKIN, C. C.; BASKIN, J. M. 2014. Seeds:
germination of P. pyramidalis and M. urundeuva. Ecology, Biogeography, and Evolution of
Sowing depth influences seeds of these species in Dormancy and Germination. Academic Press, San
the soil seed bank. Diego.
Recruitment of M. urundeuva seedlings is
restrained by both low precipitation and BOSSUYT, B.; HONNAY, O. 2008. Can the seed
interspecific competition. bank be used for ecological restoration? An
Small and stochastic rainfall events, as overview of seed bank characteristics in European
predicted in future climate scenarios and drought communities. Journal of Vegetation Science, v. 19,
years, does not completely inhibit P. pyramidalis n. 6, p. 875-884.
seed germination. However, it compromises the
seedlings survival. BRASIL. MINISTÉRIO DO MEIO AMBIENTE.
2008. Instrução Normativa n. 6, de 23 de setembro
Acknowledgements de 2008. Espécies da flora brasileira ameaçadas de
The authors would like to acknowledge the extinção e com deficiência de dados, Diário Oficial
funding for this research by Embrapa (MP3 [da] República Federativa do Brasil, Poder
03.12.12.004.00) and CAPES/Embrapa (Capes- Executivo, Brasília, DF, 24 set. 2008. Seção 1, pp.
Embrapa – Edital 15/2014 Proposta 171). Graduate 75-83.
scholarships were provided by Coordination for the
Improvement of Higher Education Personnel CABALLERO, I.; OLANO, J. M.; ESCUDERO,
(CAPES) and National Counsel of Technological A.; LOIDI, J. 2008. Seed bank spatial structure in
and Scientific Development (CNPq). semi-arid environments: beyond the patch-bare
area dichotomy. Plant Ecology, v. 195, n. 2, p. 215-
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