SciELO - Scientific Electronic Library Online

vol.26 número2Influence of Urbanization on the Dynamics of the Urban Vegetation Coverage Index (VCI) in Erechim (RS)Population Structure and Fruit Productivity Analyses in Support of the Use of Caryocar brasiliense índice de autoresíndice de assuntospesquisa de artigos
Home Pagelista alfabética de periódicos  

Serviços Personalizados




Links relacionados


Floresta e Ambiente

versão impressa ISSN 1415-0980versão On-line ISSN 2179-8087

Floresta Ambient. vol.26 no.2 Seropédica  2019  Epub 04-Abr-2019 

Original Article


Soil Seed Bank at Different Depths and Light Conditions in a Dry Forest in Northern Minas Gerais

Josiane Carvalho Menezes1

Ozorino Caldeira Cruz Neto1

Islaine Franciely Pinheiro Azevedo1

Adriana Oliveira Machado1

Yule Roberta Ferreira Nunes1

1 Laboratório de Ecologia Vegetal, Departamento do Biologia Geral, Universidade Estadual de Montes Claros – UNIMONTES, Montes Claros/MG, Brasil


The soil seed bank is an important natural regeneration strategy for plant communities and can determine floristic composition after disturbances. The aim of the present study was to evaluate the seed bank richness and abundance at different soil depths and under different light conditions in a dry forest. Litter and soil samples were collected at depths of 0-5 and 5-10 cm and submitted to two light conditions (light and shady). In total, 1,725 individuals from 85 species and 19 families emerged. Significant differences in richness between soil depths were observed, being greater at 0-5 cm, while abundance was similar. There were no variations in richness or abundance of germinated seeds between light conditions. Malvaceae and Verbenaceae families were the most representative in this study.

Keywords:  seeds stock; soil layers; light conditions; seasonally; deciduous forest


Dry tropical forests (dry deciduous forests or matas secas in Brazil) are conditioned by strong climatic seasonality and are mainly characterized by tree formations showing pronounced leaf loss during the dry season ( Murphy & Lugo, 1986 ). They have wide occurrence in Brazil, both in Cerrado, Atlantic Rainforest and Caatinga biomes ( Scariot & Sevilha, 2005 ); therefore, vegetation has distinct characteristics depending on the area of occurrence ( Pedralli, 1997 ). Although being quite diverse, they are one of the most threatened ecosystems in the world, with 60% of their territory already modified by anthropogenic actions directed toward agriculture ( Sánchez-Azofeifa et al., 2013 ). Land abandonment after agricultural use is among the main causes for composition, structure and ecological function modification of these forests ( Bullock, 1995 ; Colón & Lugo, 2006 ), which can lead to secondary succession, driven by the germination of seeds present in the soil ( Martins et al., 2008 ).

The soil seed bank consists of a set of viable (non-germinated or dormant) seeds that remain in the soil (on the surface or buried) until adequate conditions for their germination are achieved, and adult plants in an environment are replaced ( Garwood, 1989 ). It is a fundamental part of the regeneration process of the plant community, also determining the vegetation composition in frequently disturbed areas ( Luzuriaga et al., 2005 ). Soil seed banks can also be managed to recover degraded areas ( Reubens et al., 2007 ; Nóbrega et al., 2009 ; Pereira et al., 2010 ). Seeds come from the local community (seed rain) and from neighboring or distant areas brought by dispersing agents ( Hall & Swaine, 1980 ).

After dispersion, some seeds are retained in the litter, while others reach different soil depths due to their morphological characteristics and dispersion structures ( Yu et al., 2007 ). Soil texture also plays a role in this process ( Thompson & Grime, 1979 ; Garwood, 1989 ). Seeds can be buried at several depths, being variable according to layer, amount of seeds, germination capacity and viability period. The highest germination percentage occurs in the most superficial soil layer, influenced by light and temperature conditions ( Garwood, 1989 ; Baider et al., 2001 ), with dormancy breaking decreasing the deeper seeds are buried ( Marthews et al., 2008 ).

Temperature and light are among the major environmental factors that influence seed germination in the soil, according to water and oxygen availability ( Fenner & Thompson, 2005 ). Some seeds germinate after exposure to light (positive photoblastic), which exposure may be short or long. Other seeds germinate only in the absence of light (negative photoblastic) while others are indifferent ( Vàzquez-Yanes & Orozco-Segovia, 1990 ). The number of hours of light required for the germination of photosensitive seeds varies according to the ecological requirements of the species ( Araújo et al., 2003 ).

In regions marked by climatic seasonality, plant communities tend to exhibit adaptations to tolerate drought (xeromorphic characteristics) or to be dominated by annual plants that survive water scarcity, with dormant seeds in the soil ( Howe & Smallwood, 1982 ). Thus, seed banks are one of the main strategies for the long-term survival of plant communities regarding climatic seasonality and irregular rainfall ( Kemp, 1989 ; Baskin & Baskin, 1998 ; Reubens et al., 2007 ).

Therefore, dry tropical forests, being environments marked by climatic seasonality and irregular rainfall, may exhibit regenerative potential associated with the soil seed bank. Thus, knowledge about this vegetation component may assist in the management and restoration of these forests. The aim of the present study was to evaluate richness and abundance in the soil seed bank at different soil depths and light conditions in a dry tropical forest in the “Mata Seca” State Park, northern Minas Gerais. Thus, the following questions were addressed: (i) do the soil seed bank richness and abundance vary among different soil depths and under different light conditions? and (ii) what is the contribution of the seed bank to natural regeneration in the area? Richness and abundance are expected to be higher in superficial soil layers and when submitted to greater luminosity conditions.


2.1. Study area

The study was carried out at “Mata Seca” State Park (MSSP), located in the Middle São Francisco River Valley, northern Minas Gerais. MSSP covers an area of 15,446 ha ( Pezzini et al., 2014 ), located among municipalities of Manga, Itacarambi, São João das Missões and Matias Cardoso (14°48’36”-14°56’59” S and 43°55’12”-44°04’12” W). The predominant climate in the region is As (Köppen classification), with pronounced dry season in winter, average annual temperature of 24° C and average annual rainfall of 818 mm ( Madeira et al., 2009 ; Alvares et al., 2013 ). According to Coelho et al. (2013) , the MSSP geology is associated with Cenozoic coverings (Bambuí and Urucuia Groups and quaternary deposits of the São Francisco River), presenting different soil types. In areas where the dry forest occurs, soils are derived from the carbonate-pelitic rocks and present the following taxonomic suborders: Haplic Cambisols, Red Latosols and Red-Yellow Latosols, Haplic and Argiluvic Chernosols, Haplic Vertisols, Petroferric Plinthosol, Haplic and Melanic Gleisols ( Coelho et al., 2013 ).

Prior to the creation of MSSP, the main use of the park area was for cattle grazing. Therefore, vegetation remnants present variable regeneration degrees ( Madeira et al., 2009 ). The study was developed in a dry forest section where the vegetation survey was carried out (see Madeira et al., 2009 ; Nunes et al., 2013 ), which considered 18 plots of 20 m × 50 m, marked in 2006 in an arbitrary manner. These plots represent three successional stages (initial, secondary and late), with six plots in each ( Madeira et al., 2009 ). These plots had minimum distance from each other of 200 m and maximum distance of 5 km. For the present study, successional stages were not considered.

2.2. Soil seed bank sampling

Soil seed bank sampling was performed in plots described, and soil and litter samples were collected over four periods in 2013 (April, July and October) and 2014 (January). For soil sampling, four points were equidistantly marked in each plot, one at each vertex of the plot and 3 m away from the edges. Superficial litter samples (0.09 m2 each sample) and two soil samples were collected at depths of 0-5 cm and 5-10 cm (0.045 m3 each sample) at each point of the plot using a 30 cm × 30 cm wooden template. In total, 864 samples were collected from 18 plots. The four separate litter and soil samples were mixed to form a composite sample of each depth in each plot, totaling 216 composite samples. Samples composed of litter and soil were separately conditioned in properly identified plastic bags and transported to the Laboratory of Plant Ecology at “Unimontes” campus, Montes Claros/MG.

Each sample composed of litter and soil was divided into two plastic pots with dimensions of 22 cm × 12 cm × 8 cm, and litter samples were placed on sterilized sand, which had been previously autoclaved. Each pair of soil and litter samples was distributed in two greenhouses, with different light conditions (light and shade) to monitor the germination of the species, according to the different light intensity conditions. Greenhouses had dimensions of 11 m × 6 m × 2 m, one simulating direct light conditions (light condition), covered by white polyethylene screen on the top and sides, and plastic only on the top; and the other simulating shade conditions (shade condition), obtained through black polyethylene screen (50% shade), covered with black polyethylene screen on the top and sides, and plastic only on the top. The use of transparent plastic covering the top of greenhouses had the objective of controlling the humidity and avoiding infiltration by local rainfall. The ground in the two greenhouses was covered by gravel and black polyethylene sheeting to avoid the emergence of seedlings native to the experimental site. To avoid infestation by local seed rain, 35 plastic pots containing sterilized sand (autoclaved) were distributed in each greenhouse. Samples from both treatments were watered twice daily (morning and afternoon). The total number of experimental samples was 432 (18 plots × 3 depths × 4 collection periods × 2 light conditions).

For the seed bank test, the Brown’s (1992) emergence method was used. Litter and soil samples were weekly monitored throughout the 12-week period. For this, all individuals whose seeds emerged and produced seedlings were counted, being also morphotyped for identification. After individual establishment (post-seedling stage) or when there was reproductive material (flower), the individual was removed for herborization. The botanical material was identified through consultations with specialists and specialized literature. The botanical material was treated according to traditional herborization techniques and deposited in the “Montes Claros” Herbarium (MCMG) at Unimontes. Angiosperm Philogeny Group III system ( APG III, 2009 ) was used to classify species by family.

2.3. Data analysis

To evaluate the soil seed bank richness and abundance according to depth classes (litter, 0-5cm and 5-10 cm) and light availability (greenhouse: light and shade conditions), richness and abundance values were submitted to Analysis of Variance (ANOVA) in a GLM (Generalized Linear Model) ( Nelder & Wedderburn, 1972 ), with “F” test and posterior contrast analysis. For analyses, the number of individuals observed in the soil seed bank amomg the different periods was added, maintaining two seasons (rainy and dry), totaling 108 samples (18 plots × 3 depths × 2 light conditions). Values were presented as mean ± standard error with differences between averages at 5% probability being considered significant. All analyses were performed using the R3.1.3 software ( R Development Core Team, 2015 ).


Considering all depth levels of the soil seed bank sampled at MSSP, there were 1,725 individuals belonging to 85 species, 38 genera and 19 Angiospermae families, and three unidentified pteridophytes ( Table 1 ). Families that presented the greatest richness were Malvaceae, with 13 species, followed by Verbenaceae and Fabaceae, with 12 species each. Malvaceae and Verbenaceae families were also the most abundant (260 and 233 individuals respectively), followed by Poaceae (214 individuals).

Table 1 Seed bank richness (S) and abundance (A) for botanical families of Seasonally Deciduous Forest in “Mata Seca” State Park (Manga, MG) under different soil depths and light conditions.  

Family Depth Light conditions
Litter 0-5 cm 5-10 cm Shade Light
Amaranthaceae 0 0 1 1 1 8 1 4 1 5
Asteraceae 7 30 6 59 8 64 6 43 8 110
Bignoniaceae 0 0 0 0 1 1 0 0 1 1
Boraginaceae 0 0 3 15 2 10 3 10 2 15
Burseraceae 1 186 1 1 0 0 1 112 1 75
Commelinaceae 0 0 1 2 0 0 1 1 1 1
Convolvulaceae 2 10 4 13 2 4 3 15 4 12
Cyperaceae 0 0 1 2 1 3 1 3 1 2
Euphorbiaceae 4 32 4 23 3 71 4 40 4 86
Fabaceae 9 39 7 73 5 27 9 56 8 83
Lamiaceae 1 2 3 68 1 15 2 46 2 39
Malvaceae 3 83 12 125 8 52 8 53 11 207
Oxalidaceae 1 1 1 3 0 0 1 2 1 2
Phyllanthaceae 1 15 2 65 2 51 2 68 2 63
Poaceae 6 13 6 74 6 127 6 177 6 37
Rubiaceae 1 59 1 21 2 4 2 61 1 23
Smilacaceae 1 18 1 7 1 2 1 16 1 11
Solanaceae 2 2 3 4 0 0 3 3 3 3
Verbenaceae 9 75 12 98 9 60 9 151 12 82
Pteridophyta* 1 3 2 3 1 1 0 0 3 7
Total 49 568 71 657 53 500 63 861 73 864

*Group not classified in botanical family.

In relation to depth, 71 species (83.52% of total germinated species) were present at depth of 0-5 cm, 53 (62.35%) at depth of 5-10 cm and 49 (57.65%) in the litter ( Table 1 ). The mean number of germinated species per sample was 8.46 ± 1.15 at the depth of 0-5 cm, 6.42 ± 0.87 at the depth of 5-10 cm and 4.92 ± 0.96 in the litter, being significantly greater at the depth of 0-5 cm in comparison with the other depths ( Figure 1 , Table 2 ). The abundance of seeds at the depth of 0-5 cm was 657 recruited individuals (38.09% of the total), 568 individuals in the litter (32.93%) and 500 individuals at the depth of 5-10 cm (28.99%). Considering the average number of individuals germinated per sample, there were 27.37 ± 5.65 individuals at the depth of 0-5 cm, 23.71 ± 8.89 in the litter and 20.92 ± 4.00 individuals at the depth of 5-10 cm. There were no significant differences in the abundance of seeds germinated among the different soil layers ( Table 2 ). The vertical distribution of the seed bank is derived from the action of different biotic and abiotic mechanisms for incorporation of seeds into the soil ( Garwood, 1989 ). Most studies indicate higher seed abundance in the most superficial soil layers ( Baider et al., 2001 ; Bossuyt et al., 2002 ; Costa & Araújo, 2003 ; Luzuriaga et al., 2005 ; Wang et al., 2009 ; Santos et al., 2010 ; Silva et al., 2013 ; Savadogo et al. 2017 ), which is favored by constant seed replacement, which determines the maintenance of the species in the plant community ( Bossuyt et al., 2002 ). However, few studies have shown variations in richness ( Korb et al. 2005 ; Wang et al., 2009 ; Jia et al., 2017 ), mainly due to the difficulty of species identification. According to Jia et al. (2017) , environmental factors such as soil characteristics, topography, climate and vegetation distribution play an important role in the formation of the soil seed bank.

Figure 1 Soil seed bank richness (average number of species per sample), collected in different depths in a Seasonally Deciduous Forest (Manga, MG). Bars represent the standard error. Different letters indicate significant differences (p < 0.05; n = 108).  

Table 2 Analysis of Variance of the effect of treatments (depth and light) in response variables: soil seed bank richness and abundance of Seasonally Deciduous Forest in “Mata Seca” State Park (Manga, MG).  

Response variable Source of variation DF Deviance Residual DF Residual Deviance F p
Richness SD 2 151.69 69 1657.6 3.194 0.047*
LC 1 51.68 68 1605.9 2.176 0.145
SD × LC 2 38.69 66 1567.2 0.815 0.447
Abundance SD 2 503.58 69 70036 0.239 0.788
LC 1 0.50 68 70036 0.001 0.983
SD × LC 2 623.58 66 69412 0.297 0.744

SD = soil depth; LC = light conditions; DF = Degrees of freedom; F = Fisher's F test.

*Statistically different (p < 0.05).

Between light treatments, the number of species germinated was 73 (85.88% of the total) in the light condition and 63 species (74.12%) in the shade condition ( Table 1 ). The average species per sample in the light condition was 7.44 ± 0.90 and in the shade condition was 5.75 ± 0.76; but these values showed no significant difference ( Table 2 ). Abundance also presented no significant differences between conditions evaluated, with 864 individuals (50.09% of the total, 24.08 ± 5.42 per sample) in the light condition and 861 individuals (49.91% of the total, 23.92 ± 5.16 per sample) in the shade condition. Other studies, such as Braga et al. (2008) and Martins et al. (2008) also showed no differences in the number of germinated seeds between different light intensities. Differences in germination in relation to light treatments are more common when each species is analyzed separately ( Costalonga et al., 2006 ). The influence of light on dry forest seed banks is more likely to be lower, since these environments present marked foliar deciduousness ( Nunes et al., 2012 ), which allows greater exposure to light inside the forest. On the other hand, water availability may be the main triggering factor for seed germination in these environments.

In relation to floristic composition, 14 families were found in the litter, with Fabaceae and Verbenaceae families being the most representative (nine species each), followed by Asteraceae (seven) and Poaceae families (six) ( Table 1 ). The family with the largest number of individuals was Burseraceae, with 186 individuals, with only one representative species, Commiphora leptophloeos (Mart.) J. B. Gillett., which was consequently the most abundant species. Of families with the highest richness and abundance registered in the MSSP seed bank, only Fabaceae family was recorded as one of the richest and densest in floristic composition surveys of the plant community in the same study site ( Madeira et al., 2009 ; Nunes et al. 2013 ). Tropical forest seed banks are largely composed of seeds that are absent or rare in the local vegetation, presenting limited relationship with the plant community ( Baider et al., 2001 ). Moreover, most of the seed bank is represented by seeds from herbaceous species ( Garwood, 1989 ; Baider et al., 2001 ), which were not considered in the floristic studies carried out in MSSP. Additionally, the high abundance of C. leptophloeos, a pioneer tree species ( Carvalho, 2008 ), characteristic of dry forests in the region ( Santos et al., 2007 ; Sales et al., 2009 ; Arruda et al., 2011 ) and of the arboreal component of the studied forest ( Madeira et al., 2009 ; Nunes et al., 2013 ), suggest that this species uses the seed bank as a reproductive strategy.

At the depth of 0-5 cm, 18 families were recorded, with Commelinaceae family being recorded only at this soil layer. Families that presented the greatest species richness were Malvaceae and Verbenaceae (12 species in each), followed by Fabaceae (seven species). Malvaceae was also the most abundant family, with 125 individuals ( Table 1 ). The species of greatest abundance at this soil depth was Sida planicaulis Cav. (Malvaceae), with 100 individuals. This species is a small shrub, 0.5-1.0 m, widely distributed and common in disturbed areas ( Bovini et al., 2001 ). In addition, the Malvaceae family was also listed among the most representative in other studies that evaluated the seed bank in riparian forest ( Tres et al., 2007 ) and Caatinga vegetation ( Silva et al., 2013 ).

At the depth of 5-10 cm, species from15 families were found, with Bignoniaceae being exclusive to this soil depth, although it was represented by only one individual. The most valuable family was Verbenaceae (nine species), followed by Asteraceae and Malvaceae families (eight species each) ( Table 1 ). At this depth, the most abundant family was Poaceae (127 individuals) and the most prevalent species was an unidentified Poaceae, with 73 individuals. Asteraceae and Poaceae are also among the richest and most abundant families in a seed bank study conducted in the dry forests of the Caatinga biome ( Mamede & Araújo, 2007 ).

Among light treatments, there were practically no differences in the number of families. In the light condition, the 19 families registered in the study were represented, while in the shade condition, 18 were recorded ( Table 1 ). Only Bignoniaceae was not recorded in the shade condition. In the light condition, the family with the highest species richness was Verbenaceae (12 species) followed by Malvaceae (11 species). In the shade condition, the families with the greatest richness were Fabaceae and Verbenaceae (nine species each), followed by Malvaceae (eight species). The most abundant families in the light condition were Malvaceae (207 individuals) and Asteraceae (110 individuals), while in the shade, the most abundant family was Poaceae (177 individuals) followed by Verbenaceae (151 individuals) and Burseraceae (112 individuals). The species that germinated with greatest abundance in the light condition was S. planicaulis, with 173 individuals. As in litter, the most abundant species in the shade was C. leptophloeos, with 112 individuals. Although light is considered an important factor for the germination of many species found in the seed bank ( Garwood, 1989 ), shading can also favor germination, since soil retains greater moisture content ( Franco et al., 2012 ). This could explain the great similarity in the amounts of species and individuals germinating under both light and shade conditions found in this study.

The results of this study indicate the potential of the seed bank to regenerate the dry forests in the region. Although most recruited species are herbaceous and generalist, there are tree species that were recruited via soil seed bank. Information on the regenerative potential of tropical dry forests is extremely important, especially in terms of the reproductive strategies of their species, given the great anthropogenic pressures faced by these environments (Calvo Rodriguez et al., 2016). Therefore, understanding the seed bank functioning may be essential to preserve and restore dry forests.


The soil seed bank under study presented greater richness in the superficial soil layer, that is, in the first 5 cm of depth, but its abundance did not vary among layers or between the different light conditions. Malvaceae, Verbenaceae and Asteraceae families were among the most representative in the soil seed bank, considering the different layers and shading conditions. However, although the great majority of germinated seeds belong to herbaceous species, the high prevalence of Commiphora leptophloeos (Burseraceae) in the soil seed bank should be highlighted, as it is an important tree species characteristic of the studied physiognomy.


The authors thank FAPEMIG (Fundação de Amparo à Pesquisa do Estado de Minas Gerais; APQ-02217-12), CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico; 306375/2016-8) and CAPES (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior) for granting funds for this research. We also thank IEF (Instituto Estadual de Florestas) and UNIMONTES (Universidade Estadual de Montes Claros) for the logistical and legal supports; Dr. Rubens Manoel dos Santos (Universidade Federal de Lavras) for helping in botanical identification; to the Laboratory of Plant Ecology students (UNIMONTES) and IEF staff for field assistance.

FINANCIAL SUPPORT Fundação de Amparo à Pesquisa de Minas Gerais (FAPEMIG) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq).


Alvares CA, Stape JL, Sentelhas PC, Gonçalves JLM, Sparovek G. Köppen’sclimateclassificationmap for Brazil. MeteorologischeZeitschrift 2013; 22(6): 711-728. [ Links ]

APG III. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Botanical Journal of the Linnean Society 2009; 161(2): 105-121. [ Links ]

Araújo JC No, Aguiar IB, Ferreira VM. Efeito da temperatura e da luz na germinação de sementes de Acacia polyphylla DC. Revista Brasileira de Botanica. Brazilian Journal of Botany 2003; 26(2): 249-256. [ Links ]

Arruda DM, Brandão DO, Costa FV, Tolentino GS, Brasil RD, D’Ângelo S No et al. Structural aspects and floristic similarity among tropical dry forest fragments with different management histories in Northern Minas Gerais, Brazil. Revista Árvore 2011; 35(1): 133-144. [ Links ]

Baider C, Tabarelli M, Mantovani W. The soil seed bank during Atlantic forest regeneration in Southeast Brazil. Revista Brasileira de Biologia 2001; 61(1): 35-44. PMid:11340460. [ Links ]

Baskin CC, Baskin JM. Seeds: ecology, biogeography, and evolution of dormancy and germination. London: Academic Press; 1998. 666 p. [ Links ]

Bossuyt B, Heyn M, Hermy M. Seed bank and vegetation composition of forest stands of varying age in central Belgium: consequences for regeneration of ancient forest vegetation. Plant Ecology 2002; 162(1): 33-48. [ Links ]

Bovini MG, Carvalho-Okano RM, Vieira MF. Malvaceae A. Juss. no Parque Estadual do Rio Doce, Minas Gerais, Brasil. Rodriguésia 2001; 52(81): 17-47. [ Links ]

Braga AJT, Griffith JJ, Paiva HN, Meira JAA No. Composição do banco de sementes de uma floresta semidecidual secundária considerando o seu potencial de uso para a recuperação ambiental. Revista Árvore 2008; 32(6): 1089-1098. [ Links ]

Brown D. Estimating the composition of a forest seed bank: a comparison of the seed extraction and seedling emergence methods. Canadian Journal of Botany 1992; 70(8): 1603-1612. [ Links ]

Bullock SH. Plant reproduction in Neotropical dry forests. In Bullock S, Mooney HA, Medina E. Seasonally dry tropical forests. Cambridge: Cambridge University Press; 1995. p. 277-303. [ Links ]

Calvo-Rodriguez S, Sanchez-Azofeifa AG, Duran SM, Espírito-Santo MM. Assessing ecosystem services in Neotropical dry forests: a systematic review. Environmental Conservation 2016; 44(01): 34-43. [ Links ]

Carvalho PER. Espécies arbóreas brasileiras. Brasília: Embrapa Informação Tecnológica; Colombo: Embrapa Florestas; 2008. v. 3. 593 p. [ Links ]

Coelho MR, Dart RO, Vasques GM, Teixeira WG, Oliveira RP, Brefin MLM et al. Levantamento pedológico semi-detalhado (1:30.000) do Parque Estadual da Mata Seca, município de Manga - MG. Rio de Janeiro: Embrapa Solos; 2013. 264 p. (Boletim de Pesquisa e Desenvolvimento, no. 217). [ Links ]

Colón SM, Lugo AE. Recovery of a subtropical dry forest after abandonment of different land uses. Biotropica 2006; 38(3): 354-364. [ Links ]

Costa RC, Araújo FS. Densidade, germinação e flora do banco de sementes do solo no final da estação seca, em uma área de caatinga, Quixadá, CE. Acta Botanica Brasílica 2003; 17(2): 259-264. [ Links ]

Costalonga SR, Reis GG, Reis MGF, Silva AF, Borges EEL, Guimarães FP. Florística do banco de sementes do solo em áreas contíguas de pastagem degrada, plantio de eucalipto em Paula Cândido, MG. Floresta 2006; 36(2): 239-250. [ Links ]

Fenner M, Thompson K. The ecology of seeds. Cambridge: Cambridge University; 2005. 260 p. [ Links ]

Franco BKS, Martins SV, Faria PCL, Ribeiro GA. Densidade e composição florística do banco de sementes de um trecho de floresta estacional semidecidual no campus da Universidade Federal de Viçosa, MG. Revista Árvore 2012; 36(3): 423-432. [ Links ]

Garwood NC. Tropical soil seed banks: a review. In: Leck MA, Parker VT, Simpson RL. Ecology of soil seed banks. San Diego: Academic Press; 1989. p. 149-209. [ Links ]

Hall JB, Swaine MD. Seed stocks in Ghanaian forest soils. Biotropica 1980; 12(4): 256-263. [ Links ]

Howe HF, Smallwood J. Ecology of seed dispersal. Annual Review of Ecology and Systematics 1982; 13(1): 201-228. [ Links ]

Jia F, Tiyip T, Wu N, Tian C, Zhang Y. Characteristics of soil seed banks at different geomorphic positions within the longitudinal sand dunes of the Gurbantunggut Desert, China. Journal of Arid Land 2017; 9(3): 355-367. [ Links ]

Kemp PR. Seed banks and vegetation process in deserts. In: Leck MA, Parker VT, Simpson RL. Ecology of soil seed banks. San Diego: Academic Press; 1989. p. 257-281. [ Links ]

Korb JE, Springer JD, Powers SR, Moore MM. Soil seed banks in Pinus ponderosa forests in Arizona: clues to site history and restoration potencial. Applied Vegetation Science 2005; 8: 103-112. [ Links ]

Luzuriaga AL, Escudero A, Olano JM, Loidi J. Regenerative role of seed banks following an intense soil disturbance. Acta Oecologica 2005; 27(1): 57-66. [ Links ]

Madeira BG, Espírito-Santo MM, D’Ângelo S No, Nunes YRF, Arturo Sánchez Azofeifa G, Wilson Fernandes G et al. Changes in tree and liana communities along a successional gradient in a tropical dry forest in south-eastern Brazil. Plant Ecology 2009; 201(1): 291-491. [ Links ]

Mamede MA, Araújo FS. Effects of slash and burn practices on a soil bank of caatinga vegetation in Northeastern Brazil. Journal of Arid Environments 2007; 72(4): 458-470. [ Links ]

Marthews TR, Mullins CE, Dalling JW, Burslem DFRP. Burial and secondary dispersal of small seeds in a tropical forest. Journal of Tropical Ecology 2008; 24(06): 595-605. [ Links ]

Martins SV, Almeida DP, Fernandes LV, Ribeiro TM. Banco de sementes como indicador de restauração de uma área degradada por mineração de caulim em Brás Pires, MG. Revista Árvore 2008; 32(6): 1081-1088. [ Links ]

Murphy PG, Lugo AE. Ecology of tropical dry forest. Annual Review of Ecology and Systematics 1986; 17(1): 67-88. [ Links ]

Nelder JA, Wedderburn RWM. Generalized linear models. Journal of the Royal Statistical Society. Series A (General) 1972; 135(3): 370-384. [ Links ]

Nóbrega AMF, Valeri SV, Paula RC, Pavani MCMD, Silva SA. Banco de sementes de remanescentes naturais e de áreas reflorestadas em uma várzea do rio Mogi-Guaçu-SP. Revista Árvore 2009; 33(3): 403-411. [ Links ]

Nunes YRF, Luz GR, Braga LL. Phenology of tree species populations in tropical dry forests of Southeastern Brazil. In: Zhang X. Phenology and climate change. Rijeka: In Tech; 2012. p. 125-142. [ Links ]

Nunes YRF, Luz GR, Souza SR, Silva DL, Veloso MDM, Espírito Santo MM et al. Floristic, structural, and functional group variations in tree assemblages in a Brazilian tropical dry forest: effects of sucessional stage and soil properties. In: Sanchez Azofeifa A, Powers JS, Fernandes GW, Quesada M. Tropical dry forests in the Americas: ecology, conservation, and management. Boca Raton: CRC Press; 2013. p. 325-349. [ Links ]

Pedralli G. Florestas secas sobre afloramentos de calcário em Minas Gerais: florística e fisionomia. Bios 1997; 5(5): 81-88. [ Links ]

Pereira IM, Alvarenga AP, Botelho SA. Banco de sementes do solo, como subsídios à recomposição de mata ciliar. Floresta 2010; 40(4): 721-730. [ Links ]

Pezzini FF, Ranieri BD, Brandão DO, Fernandes GW, Quesada M, Espírito Santo MM et al. Changes in tree phenology along natural regeneration in a seasonally dry tropical forest. Plant Biosystems 2014; 148(5): 965-974. [ Links ]

R Development Core Team. R: a language and environment for statistical computing [online]. Vienna: R Foundation for Statistical Computing; 2015 [cited 2015 May 10]. Available from: [ Links ]

Reubens B, Heyn M, Gebrehiwot K, Hermy M, Muys B. Persistent soil seed banks for natural rehabilitation of dry tropical forests in northern Ethiopia. Tropicultura 2007; 25(4): 204-214. [ Links ]

Sales HR, Souza SCA, Luz GR, Costa FM, Amaral VB, Santos RM et al. Flora arbórea de uma Floresta Estacional Decidual na APA Estadual do rio Pandeiros, Januária/MG. Biota 2009; 2(3): 31-41. [ Links ]

Sánchez-Azofeifa A, Calvo Alvarado J, Espírito-Santo MM, Fernandes GW, Powers JS, Quesada M. Tropical dryforests in the Americas: the Tropi-dryendeavor. In: Sanchez-Azofeifa A, Powers JS, Fernandes GW, Quesada M. Tropical dry forests in the Americas: ecology, conservation, and management. Boca Raton: CRC Press; 2013. p. 1-15. [ Links ]

Santos DM, Silva KA, Lopes JMFF, Pimentel CGR, Mendonça RM, Araújo EL. Variação espaço-temporal do banco de sementes em uma área de Floresta Tropical Seca (Caatinga) Pernambuco. Revija za Geografijo 2010; 27(1): 234-253. [ Links ]

Santos RM, Vieira FA, Fagundes M, Nunes YRF, Gusmão E. Riqueza e similaridade florística de oito remanescentes florestais no norte de Minas Gerais, Brasil. Revista Árvore 2007; 31(1): 135-144. [ Links ]

Savadogo P, Sanou L, Dayamba D, Bognounou F, Thiombiano A. Relationship between soil seed banks and above-ground vegetation along a disturbance gradient in the W National Park trans-boundary biophere reserve, West Africa. Journal of Plant Ecology 2017; 10(2): 349-363. [ Links ]

Scariot A, Sevilha AC. Biodiversidade, estrutura e conservação de florestas estacionais deciduais no Cerrado. In: Scariot A, Sousa-Silva JC, Felfili, JM. Cerrado: ecologia, biodiversidade e conservação. Brasília: Ministério do Meio Ambiente; 2005. p. 121-140. [ Links ]

Silva KA, Santos DM, Santos JMFF, Albuquerque UP, Ferraz EMN, Araújo EL. Spatial-temporal variation in a seed bank of a semi-arid region in northeastern Brazil. Acta Oecologica 2013; 46: 25-32. [ Links ]

Thompson K, Grime JP. Seasonal variation in the seed banks of herbaceous species 591 in ten contrasting habitats. JournalofEcology 1979; 67: 893-921. [ Links ]

Tres DR, Sant’Anna CS, Basso S, Langa R, Ribas U Jr, Reis A. Banco e chuva de sementes como indicadores para a restauração ecológica de matas ciliares. Revista Brasileira de Biociências 2007; 5: 309-311. [ Links ]

Vàzquez-Yanes C, Orozco-Segovia A. Ecological significance of light controlled seed germination in two contrasting tropical habitats. Oecologia 1990; 83: 171-175. PMid:22160107. [ Links ]

Wang J, Ren H, Yang L, Li D, Guo Q. Soil seed banks in four 22-year-old plantations in South China: implications for restoration. Forest Ecology and Management 2009; 258(9): 2000-2006. [ Links ]

Yu S, Sternberg M, Kutiel P, Chen H. Seed mass, shape, and persistence in the soil seed bank of Israeli coastal sand dune flora. Evolutionary Ecology Research 2007; 9: 325-340. [ Links ]

Received: March 16, 2017; Accepted: November 23, 2017

Yule Roberta Ferreira NunesDepartamento de Biologia Geral, Laboratório de Ecologia Vegetal, Universidade Estadual de Montes Claros – UNIMONTES, Campus Universitário Prof. Darcy Ribeiro, CEP 39401-089, CP 126, Montes Claros, MG, Brasil e-mail:

Creative Commons License  This is an Open Access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.