Soil seed bank during succession at an abandoned pasture in the upper Parana river-floodplain, Brazil/Banco de sementes do solo durante sucessao em pastagem abandonada na planicie de inundacao do alto rio Parana, Brasil.
A soil seed bank is formed by all viable seeds on or in the soil (SIMPSON et al., 1989), as a dynamic input and output system (FENNER, 1985; GARWOOD, 1989). The bank is a source of genetic variability for plant communities (HARPER, 1977; FENNER, 1985; SIMPSON et al., 1989). It is also the main seed provider for the regeneration of natural gaps (BAIDER et al., 1999; DALLING et al., 1997; GARWOOD, 1989; SAULEI; SWAINE, 1988) and for areas in early stages of secondary succession (BAIDER et al., 1999; FENNER, 1985). Therefore, when an area is disturbed by natural or anthropic events, the structure of its secondary vegetation is determined primarily by the seeds already in the soil (CAMPOS; SOUZA, 2003; GARWOOD, 1989; UHL; CLARK, 1983).
Tropical forests are frequently destroyed for conversion to agriculture and cattle raising (AIDE et al., 1995, 1996, 2000; UHL et al., 1988; ZAHAWI; AUGSPURGER, 1999). After being used for several years, pastures undergo significant soil degradation and frequent fires, usually in highly fragmented landscapes (AIDE et al., 2000). Cattle raising leads to soil compaction (REINERS et al., 1994), thus inhibiting or rendering difficult vegetation recovery and lowering the quantity and quality of seed banks (CAMPOS, 1999a). These conditions inhibit the recovery of native forest and can result in different ecosystems (AIDE et al., 1996).
Knowledge of the early stages of succession can significantly improve the understanding of ecological processes that relate abandoned pastures and forests in advanced stages of succession (DUPUY; CHAZDON, 1998). Several studies on the regeneration process of forests in abandoned pastures have been conducted (AIDE et al., 1995, 1996, 2000; CAMPOS; DICKINSON, 2005; HOLL, 1999; REINERS et al., 1994; UHL et al., 1988; ZAHAWI; AUGSPURGER, 1999; ZIMMERMAN et al., 2000). Few of them, however, have been on changes occurring in the seed bank during pasture succession (CUBINA; AIDE, 2001; QUINTANA-ASCENCIO et al., 1996; ZAHAWI; AUGSPURGER, 1999), particularly in riparian forests associated with floodplains. Furthermore, very few studies have been repeated on the same site (BEKKER et al., 2000).
This is a study of density and composition of species found in the seed bank of a secondary forest on pasture abandoned for 10 years, to observe (1) changes in density, richness, diversity, and composition of species during secondary succession of abandoned pastures, and (2) implications of such changes for the forest recovery process.
[FIGURE 1 OMITTED]
Material and methods
The study was conducted in Porto Rico Island, Porto Rico, Parana State (Figure 1), in the upper Parana river-floodplain, the only remaining free-flowing portion of Parana river in Brazil (AGOSTINHO et al., 2004).
The area vegetation is classified as alluvial semidecidual seasonal forest (CAMPOS; SOUZA, 1997; IBGE, 1992). According to the Koeppen system, the climate is Cfa--tropica-lsubtropical with hot summers (mean annual temperature of 22[degrees]C) with mean annual precipitation of 1500 mm. Some years have a Cwa climate, with dry winters (MAACK, 2002). The altitude is close to 230 m (CAMPOS; COSTA-FILHO, 1994).
Porto Rico Island was originally covered by dense vegetation, but more than half of the island was deforested between 1952 and 1965. In 1996, only three forest fragments remained and covered 5.98% of the territory, the rest was in pasture (CAMPOS, 1999b).
The ecological importance of the region (AGOSTINHO et al., 2004) led to the creation of the Environmental Protection Area of the Islands and Floodplains of Parana River (Area de Protecao Ambiental das Ilhas e Varzeas do Rio Parana, APA- IVRP) in 1997, a federal conservation unit for sustainable use (CAMPOS; DICKINSON, 2005). The creation of APA--IVRP resulted in several actions sponsored by the state and environmental organizations to forbid the use of island soil for crop or livestock production. It also led to the beginning of the secondary succession process in the region.
Forty soil samples, including litter, were collected randomly from a previously delimited area that had been abandoned for 10 years. The samples were collected in September 2007. The sampling device was made of metal and the samples were collected from soil patches measuring 20 x 20 and 5 cm-deep. The total area sampled equaled 1.6 [m.sup.2].
The samples were kept in dark plastic bags and placed in a greenhouse covered with transparent PVC and sides covered with sunscreen, located at the Forest Nursery of the Instituto Ambiental do Parana (IAP), in Mandaguari, Parana, State. The samples were spread on 20 x 40 cm trays with inert vermiculite substrate. Four trays did not receive the soil samples to observe potential contamination (BROWN, 1992). The trays were watered 3 times a day; 5 months later the ground was revolved. The experiment was conducted during 10 months.
Seedlings were counted weekly, using the emergence or germination method (BROWN, 1992). After collection and identification, seedlings were preserved in alcohol solution at 40%. Unidentifiable seedlings were considered control specimens and collected with reproductive material for later identification.
For the comparative analysis between seed banks of active pasture and secondary forest on abandoned pasture for 10 years, we used the pasture seed bank data collected on the same site, by Campos and Souza (2003), in a 1996 study comparing seed banks from pasture and three forest remnants in Porto Rico Island.
The species were classified according to life form (tree, shrub, herb and liana), following Font Quer (1989). Tree species were classified according to the vegetation succession stages, as follows: pioneer, secondary, climax and invasive (BUDOWSKI, 1963, 1965, 1966; KAGEYAMA, 1992). Classification was also based on personal observations on the field.
The absolute and relative density (MUELLER-DOMBOIS; ELLENBERG, 1974), the Shannon-Weaver index of diversity (H') and equitability (J') (KREBS, 1989) were obtained.
The non-parametric Mann-Whitney test (U-Test) (ZAR, 1999) was used for the independent samples to determine the difference in total density of viable seeds. Significant differences in richness, diversity, and equitability were assessed by using the t test for independent samples at 5% significance level (ZAR, 1999). The rarefaction curve (KREBS, 1989) was designed to compare richness without the density effect. The differences in equitability were determined by the dominance/diversity curve or Whittaker plot (MAGURRAN, 2004).
We used the [chi square] test for contingency table (ZAR, 1999) at 5% significance to determine whether the proportions among species of different life forms depended on the vegetation succession stage, by comparing densities and richness. The statistical tests were processed using BioDiversity Pro (MCALEECE et al., 1997), BioEstat v4.0 (AYRES et al., 2005), and STATISTICA for Windows (STASOFT, 2005).
Results and discussion
The total density of viable seeds in the soil decreased from 6312 [+ or -] 1233 seeds [m.sup.-2] (viable seeds per square meter [+ or -] standard error) in 1996, to 3792 [+ or -] 428 in 2007. The decrease that was not statistically different (Z(U) = 1.16; p = 0.25), probably due to data variability (350-36575 seeds [m.sup.-2] in 1996 and 875-13600 seeds [m.sup.-2] in 2007); such variability is widely observed in seed banks due to their high spatial heterogeneity (BIGWOOD; INOUYE, 1988; BUTLER; CHAZDON, 1998; GARWOOD, 1989).
Most research on tropical forests show that seed bank size reaches its biggest size early in the succession process and gradually decreases (ARAUJO et al., 2001; BAIDER et al., 1999, 2001; DALLING; DENSLOW, 1998; DUPUY; CHAZDON, 1998; GARWOOD, 1989; YOUNG, 1985; YOUNG et al., 1987). We counted between 48 and 18900 seeds [m.sup.-2], which are within the expected range for pastures and secondary tropical forests according to the literature review undertaken by Garwood (1989) and Young (1985).
Campos and Souza (2003) found seed bank densities of 1574, 1338, 2513 seeds [m.sup.-2] in forest remnants of Porto Rico Island. Grombone-Guaratini and Rodrigues (2002) found much lower densities (32, 50 and 44 seeds [m.sup.-2]) in a semidecidual seasonal fragment, whereas Grombone-Guaratini et al. (2004) found densities of243 and 499 seeds [m.sup.-2] in a semidecidual seasonal riparian forest.
Difference in densities within a phytophysiognomic dominium can be due to differences in the history of soil and landscape use (ARAUJO et al., 2004; DUPUY; CHAZDON, 1998), or due to different methodologies used during the study.
Richness and diversity
In 1996, 68 taxa were identified (active pasture); in 2007, taxa increased to 81 (secondary forest on abandoned pasture) (Table 1), a significant difference (t = -2.16; p = 0.03). The rarefaction curve also revealed a higher estimated number of species in secondary forest on abandoned pasture (2007) than in active pasture (1996) (Figure 2A).
The index of diversity for active pasture (1996) (H' = 1.198) differed significantly (t = -3.61; p < 0.01) from the index for secondary forest (2007) (H' = 1.3). The index of equitability was also significantly higher (t = -2.63; p = 0.01) in secondary forest (J' = 0.677) than in active pasture (J' = 0.654).
The Wittaker plot (Figure 2B) showed a tendency for high equitability in active pasture and secondary forest.
Richness can be considered high because the number of species exceeds the limit of 8 to 67 species in pasture and secondary forest, as reported by Garwood (1989). There was also a tendency to increase richness and diversity during succession and a decrease in dominance.
[FIGURE 2 OMITTED]
Campos and Souza (2003) registered richness of 64, 47, and 52 in three remnants of the Island. We obtained higher values for richness in pasture and much higher in secondary forest. Araujo et al. (2001) found the highest richness and diversity in the early stage of succession, when a 6-year-old forest had 72 species (H' = 2.23), and a 17-year-old (H' = 1.9) and a 30-year-old (H' = 1, 12) forest had 62 and 59 species, respectively.
Scherer and Jarenkow (2006) found what they considered low diversity for a seasonal forest in Southern Brazil (H' = 1.639 and H' = 1.717). These, however, were higher than the values we found.
Quintana-Ascencio et al. (1996) found that species richness increases in communities open to sites abandoned for 4 to 12 years, and decreases in communities at more advanced succession stages. Data from our study, in addition to those reported by Campos and Souza (2003) and by other cited authors provide support for the pattern found by Quintana-Ascencio et al. (1996).
Dalling and Denslow (1998), and Wills and Read (2007), however, found no significant correlation between species richness and time after disturbance. There seems to be a significant increase in species richness as pasture approaches forest borders (CUBINA; AIDE, 2001; ZIMMERMAN et al., 2000). According to the review by Garwood (1989), the number of species found in seed banks of mature forests and in tropical secondary forests/pastures is similar.
The results above point to a difficulty in establishing a pattern of richness and diversity for seed banks in succession processes of abandoned pastures. This is probably due to the aggregated distribution pattern of the seed bank (BIGWOOD; INOUYE, 1988; BUTLER; CHAZDON, 1998; GARWOOD, 1989), and consequently, to problems in defining a representative sample. Another reason is the lack of a well-defined methodology for the study of seed banks, including criteria for sample size and number of samples.
According to Garwood (1989) review, seedbanks of tropical forests have high dominance. The high equitability obtained in our study in addition to those obtained by Araujo et al. (2001) suggest that equitability follows the richness pattern, thus increasing at the early stages of succession and gradually decreasing thereafter. Araujo et al. (2001) found the highest equitability in a 6-year-old forest (J' = 0.52), and lower values in older sites, a 17-year (J' = 0.46) and a 30-year (J' = 0. 26) forest.
Species composition--Herb species had the highest number of seeds per square meter both in active pasture (1996) and in secondary forest on abandoned pasture (2007), followed by the shrub, tree, and liana species (Table 2). As far as richness, herb species were also first, followed by the shrub, liana, and tree species in active pasture; in secondary forest, shrub species were followed by both liana and tree species at equal levels (Table 3). The [chi square] test shows that the relation between density and life form depends on vegetation succession stage ([chi square] = 1116; p < 0.01), whereas richness does not ([chi square] = 3.92; p = 0.27).
The dominance of herb species in seed banks supports previous studies that show seed banks of tropical forests as made primarily of herb pioneer species and shrub-tree species of short life cycles (ARAUJO et al., 2004; BAIDER et al., 2001; DUPUY; CHAZDON, 1998; GASPARINO et al., 2006; LOPES et al., 2006; MELO et al., 2007; QUINTANA-ASCENCIO et al., 1996; RAMIREZ-MARCIAL et al., 1992; SAULEI; SWAINE, 1988; ZAHAWI; AUGSPURGER, 1999; YOUNG et al., 1987).
In the study by Wills and Read (2007), herbaceous species were the main cause of dissimilarity in succession stages. The high number of pioneer herb species in seed banks is due to their facultative dormancy and efficient dispersion mechanisms (GASPARINO et al., 2006).
The results above show a decrease in density of herb species and an increase in density and richness of shrub-tree species in seed banks during recovery of abandoned pastures. Also in Porto Rico Island, Campos and Souza (2003) found tree dominance in all three forest remnants studied. Grombone-Guaratini and Rodrigues (2002) found similar results in a semidecidual seasonal forest fragment. However, Grombone-Guaratini et al. (2004) observed the predominance of herb species in a semidecidual seasonal riparian forest, probably due to its high level of degradation (ARAUJO et al., 2004; DUPUY; CHAZDON, 1998).
The increase in shrub-tree species in more advanced succession stages, also reported extensively (BAIDER et al., 2001; GASPARINO et al., 2006; QUINTANA-ASCENCIO et al., 1996; SAULEI; SWAINE, 1988), shows that the seed bank seems to recover considering that banks of mature forests contain tree species (GARWOOD, 1989) with pioneer traits (GARWOOD, 1989; GASPARINO et al., 2006; WHITMORE, 1983, 1992).
Campos and Dickinson (2005) assessed forest succession at the same site, after five years of abandonment. They found dominance of pioneer species in the arboreal stratum, and secondary and climax species in the sub-forest. This might have caused the increase in density and richness of trees in the soil seed bank due to the high levels of seed production by pioneer species (WHITMORE, 1983; VAZQUEZ-YANES; OROZCO-SEGOVIA, 1987).
In 2007, during the phytosociological survey conducted at the same pasture land after 10 years of abandonment, 12 tree species were found in the arboreal stratum (unpublished data), only five of them were found in the seed bank (Table 3). The pioneer tree species typical of hydromorphic soils found in seed banks (LORENZI, 1992) were also found in other seed bank studies (ARAUJO et al., 2001; CAMPOS; SOUZA, 2003; GASPARINO et al., 2006; GROMBONE-GUARATINI et al., 2004).
The species Cecropia pachystachya and Croton urucurana were found in all phytosociological studies conducted in the upper Parana river floodplain (CAMPOS et al., 2000; CAMPOS; SOUZA, 2002, 2003; CAMPOS; DICKINSON, 2005; KITA; SOUZA, 2003; SOUZA; MONTEIRO, 2005). Pioneer species in seed banks are important for the regeneration of tropical forests after natural or anthropic disturbances.
The presence of the two main pioneer species and one secondary species in the seed bank, the more than seven-time increase in tree density and over four-time increase in shrub species (Table 1) shows that the seed bank is ready to establish the ecological groups such as the pioneer species group, and the richness of shrub-tree species (BAIDER et al., 1999).
However, the presence of invasive exotic species in the seed bank, mainly Psidium guajava, is worrisome. The invasive species found in our study were also reported by Gasparino et al. (2006) in a riparian forest seed bank under the dominium of a semidecidual seasonal forest. Both results show the potential formation of seed banks with the species Psidium guajava and Tabernaemontana catharinensis.
The P. guajava has been registered as an invasive species of secondary forests on abandoned pasture in the upper Parana river Floodplain (CAMPOS; DICKINSON, 2005). Understories dominated by P. guajava are poorer in species and specimens than areas dominated by native species, showing an invasion process in the recovery area of Porto Rico island (CHAPLA et al., 2008). Plant invasion is a threat to biodiversity and the stability of native ecosystems, and can affect the functioning and processes of ecosystems (EHRENFELD, 2003; GORDON, 1998; MACK et al., 2000; VITOUSEK et al., 1997).
The succession model proposed for the study area (CAMPOS; DICKINSON, 2005) shows that anthropic disturbances can lead to the establishment and dominance of the invasive species Psidium guajava and Tabernaemontana catharinensis, and render difficult or impossible the establishment of pioneer or secondary species in the area, thus modifying the succession process and allowing for the dominance of Psidium guajava.
Psidium guajava has been reported as an invasive species in pastures (SOMARRIBA, 1985; SOMARRIBA; BEER, 1985) and as an important pioneer in the recovery process of abandoned pastures (AIDE et al., 1996, 2000; BERENS et al., 2008; CUBINA; AIDE, 2001; ZAHAWI; AUGSPURGER, 1999), although found only in young secondary forests (AIDE et al., 2000). Psidium guajava success in occupying pastures is due to its re-growth capacity and easy dispersion by birds and cattle (BERENS et al., 2008; SOMARRIBA, 1985; SOMARRIBA; BEER, 1985).
Forest recovery in abandoned pastures is slow compared with recovery due to several kinds of natural disturbances such as land clearing caused by storms or fall of branches, as well as by anthropic disturbances such as slash-and-burn agriculture (UHL, 1987; UHL et al., 1988; AIDE et al., 1995; CUBINA; AIDE, 2001).
Seed banks change drastically under pasture and lose their natural traits. These changes can alter the secondary succession process (QUINTANA-ASCENCIO et al., 1996).
Finally, the replacement of forests by pasture leads to changes in vegetation, including invasion of exotic species and decrease of most tree species (RAMIREZ-MARCIAL et al., 1992; QUINTANA-ASCENCIO et al., 1996).
Therefore, further intervention in Porto Rico Island will be necessary if the goals for creating the Environmental Protection Area (Area de Protecao Ambiental--APA) go beyond the recovery of the island vegetation and increased Carbon sequestration, aiming also at the restoration of species composition.
Density changes in the soil seed bank during succession of abandoned pastures did not follow a pattern. Richness and diversity increased, herb species were dominant and the number of tree species in the seed bank increased during the first 10 years of abandonment of pastures in riparian forests.
Soil use as pasture can result in a different environment at the end of succession due to seed bank deterioration and establishment of invasive exotic species, which can dominate the phytophysiognomy of the new ecosystem.
I am grateful to Nucleo de Pesquisas em Limnologia, Ictiologia e Aquicultura (Nupelia), Programa de Pos-graduacao em Ecologia de Ambientes Aquaticos Continentais (PEA) at Universidade Estadual de Maringa (UEM), Instituto Ambiental do Parana (IAP), Programa Parana Biodiversidade, Conselho Nacional de Desenvolvimento Cientifico (CNPq), and also to biologists Cristina Moreira Pisicchio, Iuli Pessanha Zviejkovski, Rafael Zampar, Nadson Ressye Simoes da Silva and Janielly Carvalho Camargo. I also thank the staff of the Viveiro Florestal of IAP in Mandaguari.
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Received on February 20, 2009.
Accepted on August 3, 2009.
License information: 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.
Tatiani Elisa Chapla (1) and Joao Batista Campos (2) *
(1) Universidade Estadual de Maringa, Av. Colombo, 5790, 87020-900, Maringa, Parana, Brazil. (2) Instituto Ambiental do Parana, Maringa, Parana, Brazil. *Author for correspondence. Email: firstname.lastname@example.org
Table 1. Density (seeds [m.sup.-2] [+ or -] standard error), life form (LF) (T = tree; S = shrub; H = herb; L = liana), and ecological category (C) (P = pioneer; I = invasive; S = secondary) of species collected from the soil seed bank in Porto Rico Island in active pasture (1996) and secondary forest on 10-year abandoned pasture (2007). Specie Family LF C Cecropia pachystachya Urticaceae T P Trecul Croton urucurana Baill. Euphorbiaceae T P Machaonia brasiliensis Rubiaceae T S Cham. & Schltdl. Psidiumguajava L. Myrtaceae T I Tabernaemontana Apocynaceae T I catharinensis A. DC. Asteraceae sp.1 Asteraceae S Conyza canadensis (L.) Asteraceae S Cronquist Diodia brasiliensis Rubiaceae S Spreng. Eupatorium maximiliani Asteraceae S Schrad. ex DC. Lippia alba (Mill.) Verbenaceae S N. E. Br. Malvaceae Malvaceae S Miconia prasina (Sw.) DC. Melastomataceae S Mimosa pudica L. Fabaceae S Pfaffia glomerata Amaranthaceae S (Spreng.) Pedersen Senna obtusifolia Fabaceae S (L.) H. S. Irwin & Barneby Triumfetta bartramia L. Malvaceae S Vernonia chamaedrys Less. Asteraceae S Vernonia incana Less. Asteraceae S Vernonia westiniana Less. Asteraceae S Ageratum conyzoides L. Asteraceae H Amaranthus lividus L. Amaranthaceae H Ambrosia elatior L. Asteraceae H Apium leptophyllum Apiaceae H F. Muell. ex Benth. Asteraceae sp.2 Asteraceae H Asteraceae sp.3 Asteraceae H Caperonia palustris A. Euphorbiaceae H St. -Hil. Chamaesyce hyssopifolia Euphorbiaceae H (L.) Small Conyza bonariensis (L.) Asteraceae H Cronquist Crotonglandulosus L. Euphorbiaceae H Cuphea carthagenensis Lythraceae H (Jacq.) J. F. Macbr. Cuphea sp. Lythraceae H Cyperaceae sp.1 Cyperaceae H Cyperaceae sp.2 Cyperaceae H Cyperus ferax (L.) Rich. Cyperaceae H Cyperus spp. Cyperaceae H Dichondra microcalyx Convolvulaceae H (Hallier f.) Fabris Digitaria ciliaris Poaceae H (Retz.) Koeler Digitaria sp.1 Poaceae H Digitaria sp.2 Poaceae H Digitaria violascens Link Poaceae H Eclipta alba L. ex Asteraceae H B.D.Jacks. Eleusine tristachya Poaceae H (Lam.) Lam. Erechtites hieracifolius Asteraceae H (L.) Raf. ex DC. Erechtites sp. Asteraceae H Euphorbia heterophylla L. Euphorbiaceae H Euphorbiaceae sp.1 Euphorbiaceae H Euphorbiaceae sp.2 Euphorbiaceae H Gnaphalium pensylvanicum Asteraceae H Willd. Gnaphalium spicatum Lam. Asteraceae H Heliotropium indicum L. Boraginaceae H Heliotropium procumbens Boraginaceae H Mill. Hybanthus communis Violaceae H (A.St.-Hil.) Taub. Hyptis brevipes Poit. Lamiaceae H Hyptis sp Lamiaceae H Ipomoea cairica Convolvulaceae H (L.) Sweet Ipomoea grandifolia Convolvulaceae H (Dammer) O'Donell Ipomoea sp. Convolvulaceae H Lamiaceae Lamiaceae H Ludwigia spp. Onagraceae H Ludwigia suffruticosa Onagraceae H Walter Lynaceae Lynaceae H Marsypianthes chamaedrys Lamiaceae H Kuntze Melanthera latifolia Asteraceae H (Gardn.) Cabrera Unidentified 1 Unidentified H Unidentified 2 Unidentified H Unidentified 3 Unidentified H Unidentified 4 Unidentified H Unidentified 5 Unidentified H Unidentified 6 Unidentified H Unidentified 7 Unidentified H Unidentified 8 Unidentified H Unidentified 9 Unidentified H Unidentified 10 Unidentified H Murdannia nudiflora Commelinaceae H (L.) Brenan Nicotiana bonariensis Solanaceae H Lehm. Nicotiana sp. Solanaceae H Panicum sp.1 Poaceae H Panicum sp.2 Poaceae H Panicum sp.3 Poaceae H Paspalum conjugatum Poaceae H PJ.Bergius Paspalum sp. Poaceae H Phytolacca americana L. Phytolaccaceae H Phyllanthus carolinensis Phyllanthaceae H Walther Phyllanthus niruri L. Phyllanthaceae H Phyllanthus tenellus Phyllanthaceae H Roxb. Phyllanthus sp.1 Phyllanthaceae H Phyllanthus sp.2 Phyllanthaceae H Phyllanthus sp.3 Phyllanthaceae H Physalis angulata L. Solanaceae H Plantago tomentosa Lam. Plantaginaceae H Pluchea sagittalis Asteraceae H (Lam.) Cabrera Poaceae sp.1 Poaceae H Poaceae sp.2 Poaceae H Poaceae sp.3 Poaceae H Poaceae sp.4 Poaceae H Poaceae sp.5 Poaceae H Poaceae sp.6 Poaceae H Polygonum punctatum Polygonaceae H Elliott PorophyUum ruderale Asteraceae H (Jacq.) Cass. Richardia brasiliensis Rubiaceae H Gomez Scleria pterota Presl Cyperaceae H Scoparia montevidensis Plantaginaceae H R.E.Fr. Scrophulariaceae Scrophulariaceae H Sida rhombifolia L. Malvaceae H Solanum americanum Mill. Solanaceae H Solanum sisymbrifolium Solanaceae H Lam. Solanum viarum Dunal Solanaceae H Soliva anthemifolia Asteraceae H (A. Juss.) Sweet Spermacoce capitata Rubiaceae H Ruiz & Pav. Stemodia humilis (Sol.) Plantaginaceae H G. Dawson Stemodia trifoliata Rchb. Plantaginaceae H Synedrellopsis Asteraceae H grisebachii Hieron. & Kuntze Triumfetta lappula L. Malvaceae H Verbena bonariensis L. Verbenaceae H Verbena litoralis Kunth Verbenaceae H Verbenaceae sp.1 Verbenaceae H Verbenaceae sp.2 Verbenaceae H Aristolochia sp. Aristolochiaceae L Dalechampia scandens L. Euphorbiaceae L Dalechampia sp. Euphorbiaceae L Fabaceae Fabaceae L Mikania cordfolia Willd. Asteraceae L Passiflora misera Kunth Passifloraceae L Smilax brasiliensis Smilacaceae L Spreng. Total density Total richness Density Specie 1996 2007 Cecropia pachystachya 28 ([+ or -] 7) 94 ([+ or -] 21) Trecul Croton urucurana Baill. 5 ([+ or -] 3) 90 ([+ or -] 13) Machaonia brasiliensis 40 ([+ or -] 19) Cham. & Schltdl. Psidiumguajava L. 35 ([+ or -] 21) Tabernaemontana 1 ([+ or -] 1) catharinensis A. DC. Asteraceae sp.1 1 ([+ or -] 1) Conyza canadensis (L.) 19 ([+ or -] 6) 185 ([+ or -] 88) Cronquist Diodia brasiliensis 76 ([+ or -] 36) 231 ([+ or -] 71) Spreng. Eupatorium maximiliani 63 ([+ or -] 46) Schrad. ex DC. Lippia alba (Mill.) 10 ([+ or -] 4) 3 ([+ or -] 2) N. E. Br. Malvaceae 4 ([+ or -] 2) Miconia prasina (Sw.) DC. 113 ([+ or -] 37) Mimosa pudica L. 3 ([+ or -] 2) Pfaffia glomerata 1 ([+ or -] 1) (Spreng.) Pedersen Senna obtusifolia 1 ([+ or -] 1) (L.) H. S. Irwin & Barneby Triumfetta bartramia L. 1 ([+ or -] 1) Vernonia chamaedrys Less. 4 ([+ or -] 3) Vernonia incana Less. 27 ([+ or -] 10) Vernonia westiniana Less. 7 ([+ or -] 4) Ageratum conyzoides L. 17 ([+ or -] 4) Amaranthus lividus L. 4 ([+ or -] 3) Ambrosia elatior L. 8 ([+ or -] 5) Apium leptophyllum U1 F. Muell. ex Benth. Asteraceae sp.2 45 ([+ or -] 45) Asteraceae sp.3 65 ([+ or -] 41) Caperonia palustris A. 1 ([+ or -] 1) St. -Hil. Chamaesyce hyssopifolia 1 ([+ or -] 1) (L.) Small Conyza bonariensis (L.) 2 ([+ or -] 2) Cronquist Crotonglandulosus L. 49 ([+ or -] 20) 11 ([+ or -] 11) Cuphea carthagenensis 703 ([+ or -] 538) 98 ([+ or -] 21) (Jacq.) J. F. Macbr. Cuphea sp. 1 ([+ or -] 1) Cyperaceae sp.1 8 ([+ or -] 8) Cyperaceae sp.2 14 ([+ or -] 14) Cyperus ferax (L.) Rich. 1179 ([+ or -] 678) Cyperus spp. 395 ([+ or -] 120) Dichondra microcalyx 1 ([+ or -] 1) (Hallier f.) Fabris Digitaria ciliaris 1 ([+ or -] 1) (Retz.) Koeler Digitaria sp.1 128 ([+ or -] 64) Digitaria sp.2 1 ([+ or -] 1) Digitaria violascens Link 371 ([+ or -] 99) Eclipta alba L. ex 3 ([+ or -] 2) 1 ([+ or -] 1) B.D.Jacks. Eleusine tristachya 3 ([+ or -] 2) (Lam.) Lam. Erechtites hieracifolius 3 ([+ or -] 2) (L.) Raf. ex DC. Erechtites sp. 1 ([+ or -] 1) Euphorbia heterophylla L. 1 ([+ or -] 1) Euphorbiaceae sp.1 5 ([+ or -] 3) Euphorbiaceae sp.2 19 ([+ or -] 6) Gnaphalium pensylvanicum 3 ([+ or -] 2) 2 ([+ or -] 2) Willd. Gnaphalium spicatum Lam. 226 ([+ or -] 105) 99 ([+ or -] 28) Heliotropium indicum L. 12 ([+ or -] 12) 5 ([+ or -] 3) Heliotropium procumbens 594 ([+ or -] 299) 387 ([+ or -] 100) Mill. Hybanthus communis 4 ([+ or -] 2) (A.St.-Hil.) Taub. Hyptis brevipes Poit. 4 ([+ or -] 2) Hyptis sp 29 ([+ or -] 19) Ipomoea cairica 4 ([+ or -] 2) (L.) Sweet Ipomoea grandifolia 1 ([+ or -] 1) (Dammer) O'Donell Ipomoea sp. 1 ([+ or -] 1) Lamiaceae 4 ([+ or -] 3) Ludwigia spp. 6 ([+ or -] 4) Ludwigia suffruticosa 1 ([+ or -] 1) Walter Lynaceae 2 ([+ or -] 2) Marsypianthes chamaedrys 21 ([+ or -] 10) 1 ([+ or -] 1) Kuntze Melanthera latifolia 1 ([+ or -] 1) 1 ([+ or -] 1) (Gardn.) Cabrera Unidentified 1 2 ([+ or -] 2) Unidentified 2 1 ([+ or -] 1) Unidentified 3 1 ([+ or -] 1) Unidentified 4 1 ([+ or -] 1) Unidentified 5 21 ([+ or -] 21) Unidentified 6 4 ([+ or -] 3) Unidentified 7 1 ([+ or -] 1) Unidentified 8 1 ([+ or -] 1) Unidentified 9 40 ([+ or -] 35) Unidentified 10 3 ([+ or -] 3) Murdannia nudiflora 1 ([+ or -] 1) (L.) Brenan Nicotiana bonariensis 2 ([+ or -] 1) Lehm. Nicotiana sp. 4 ([+ or -] 2) Panicum sp.1 80 ([+ or -] 21) Panicum sp.2 210 ([+ or -] 72) Panicum sp.3 128 ([+ or -] 24) Paspalum conjugatum 426 ([+ or -] 125) PJ.Bergius Paspalum sp. 151 ([+ or -] 47) Phytolacca americana L. 1 ([+ or -] 1) Phyllanthus carolinensis 31 ([+ or -] 19) Walther Phyllanthus niruri L. 35 ([+ or -] 10) Phyllanthus tenellus 8 ([+ or -] 3) Roxb. Phyllanthus sp.1 74 ([+ or -] 30) Phyllanthus sp.2 1 ([+ or -] 1) Phyllanthus sp.3 3 ([+ or -] 2) Physalis angulata L. 1 ([+ or -] 1) Plantago tomentosa Lam. 1 ([+ or -] 1) Pluchea sagittalis 1 ([+ or -] 1) 9 ([+ or -] 3) (Lam.) Cabrera Poaceae sp.1 146 ([+ or -] 36) Poaceae sp.2 1064 ([+ or -] 192) Poaceae sp.3 3 ([+ or -] 2) Poaceae sp.4 365 ([+ or -] 60) Poaceae sp.5 12 ([+ or -] 10) Poaceae sp.6 61 ([+ or -] 22) Polygonum punctatum 5 ([+ or -] 4) Elliott PorophyUum ruderale 1 ([+ or -] i) 1 ([+ or -] 1) (Jacq.) Cass. Richardia brasiliensis 23 ([+ or -] 15) 2 ([+ or -] 1) Gomez Scleria pterota Presl 12 ([+ or -] 4) 24 ([+ or -] 5) Scoparia montevidensis 2 ([+ or -] 1) R.E.Fr. Scrophulariaceae 227 ([+ or -] 123) Sida rhombifolia L. 54 ([+ or -] 27) 6 ([+ or -] 2) Solanum americanum Mill. 4 ([+ or -] 2) 2 ([+ or -] 1) Solanum sisymbrifolium 53 ([+ or -] 18) 8 ([+ or -] 3) Lam. Solanum viarum Dunal 4 ([+ or -] 2) 2 ([+ or -] 1) Soliva anthemifolia 49 ([+ or -] 23) 2 ([+ or -] 1) (A. Juss.) Sweet Spermacoce capitata 40 ([+ or -] 16) 11 ([+ or -] 4) Ruiz & Pav. Stemodia humilis (Sol.) 174 ([+ or -] 82) G. Dawson Stemodia trifoliata Rchb. 715 ([+ or -] 219) Synedrellopsis 28 ([+ or -] 22) grisebachii Hieron. & Kuntze Triumfetta lappula L. 12 ([+ or -] 7) Verbena bonariensis L. 1 ([+ or -] 1) Verbena litoralis Kunth 1 ([+ or -] 1) Verbenaceae sp.1 148 ([+ or -] 85) Verbenaceae sp.2 1 ([+ or -] 1) Aristolochia sp. 1 ([+ or -] 1) Dalechampia scandens L. 2 ([+ or -] 1) Dalechampia sp. 2 ([+ or -] 1) Fabaceae 1 ([+ or -] 1) Mikania cordfolia Willd. 1 ([+ or -] 1) 1 ([+ or -] 1) Passiflora misera Kunth 1 ([+ or -] 1) Smilax brasiliensis 5 ([+ or -] 3) Spreng. Total density 6312 ([+ or -] 1233) 3792 ([+ or -] 428) Total richness 68 81 Table 2. Absolute density (seeds [m.sup.-2]) and standard error, and relative density (in parenthesis) for each species life form in the seed bank of Porto Rico Island, in active pasture (1996) and in secondary forest on 10-year abandoned pasture (2007). Year Tree Shrub 1996 33 ([+ or -] 7) (0.53) 131 ([+ or -] 37) (2.08) 2007 261 ([+ or -] 39) (6.88) 628 ([+ or -] 146) (16.55) Year Herb Liana 1996 6144 ([+ or -] 1227) (97.33) 3 ([+ or -]) 2 (0.05) 2007 2895 ([+ or -] 398) (76.32) 10 ([+ or -] 4) (0.26) Table 3. Richness and percentage (in parenthesis) for each species life form in the seed bank of Porto Rico Island, in active pasture (1996) and in secondary forest on 10-year abandoned pasture (2007). Year Tree Shrub Herb Liana 1996 2 (2.9) 5 (7.4) 58 (85.3) 3 (4.4) 2007 5 (6.2) 13 (16) 58 (71.6) 5 (6.2)