ABSTRACT
Secondary forests account for more than half of the planet’s tropical formations. It is necessary to understand these forests in order to allow for efficient recovery and restoration processes of altered areas. In Rondônia state, in the Brazilian Amazon, where much of the natural vegetation cover has been altered over the last several decades, secondary forests have become particularly important. In this study, we selected species for restoration in Rondônia based on published work on secondary forests in the state, complemented with field collections by the authors. For each of the selected species, the successional category, dispersal syndrome and functional group was searched in the literature. Information regarding flowering and fruiting times as well as geographic distribution and habitats occupied in the state were obtained from vouchers deposited in herbaria. We listed 55 species from 49 genera and 23 families, of which 40% can be characterized as pioneers, 52.7% as secondary and 5.4% as climax species. Regarding functional group, 49% are diversity species and 34.5% are cover species. As for the dispersal syndrome, 60% are zoochoric, 16.3% anemochoric and 10.9% barochoric. Almost all of the species included in the list are common throughout the state and most are easy to recognize and identify. The species listed are present in all the main environments of Rondônia, and most are from terra firme forest. The listed species have the potential to play a fundamental role in the establishment and development of forests in altered and/or degraded areas in the state.
KEYWORDS:
Amazon; deforestation; secondary forest; restoration
RESUMO
As florestas secundárias representam mais da metade das formações tropicais do planeta. Compreender essas florestas é necessário para permitir processos eficientes de recuperação e restauração de áreas alteradas. No estado de Rondônia, na Amazônia brasileira, onde grande parte da cobertura vegetal natural foi alterada nas últimas décadas, as florestas secundárias tornaram-se particularmente importantes. Neste estudo, selecionamos espécies para restauração em Rondônia com base em trabalhos publicados sobre florestas secundárias no estado, complementados com estudos de campo realizados pelos autores. Para cada uma das espécies selecionadas, a categoria sucessional, a síndrome de dispersão e o grupo funcional foram pesquisados na literatura. Informações sobre o periodo de floração e frutificação, bem como a distribuição geográfica e os habitats ocupados no estado, foram obtidas de comprovantes depositados em herbários. Listamos 55 espécies de 49 gêneros e 23 famílias, das quais 40% podem ser caracterizadas como pioneiras, 52,7% como secundárias e 5,4% como espécies clímax. Em relação ao grupo funcional, 49% são espécies de diversidade e 34,5% são espécies de cobertura. Quanto à síndrome de dispersão, 60% são zoocóricas, 16,3% anemocóricas e 10,9% barocóricas. Quase todas as espécies incluídas na lista são comuns em todo o estado e a maioria é fácil de reconhecer e identificar. As espécies listadas estão presentes em todos os principais ambientes de Rondônia e a maioria é de floresta de terra firme. As espécies listadas têm o potencial de desempenhar papel fundamental no estabelecimento e desenvolvimento de florestas em áreas alteradas e/ou degradadas no estado.
PALAVRAS-CHAVE:
Amazônia; desmatamento; floresta secundária; restauração
INTRODUCTION
In the Brazilian Amazon, deforestation rates have proven resistant to change, year after year, increasing the percentage of deforested areas, notably in the “Deforestation Arch” along the eastern to southwestern rim of the Brazilian Amazon, where the state of Rondônia is located (PRODES 2021; Terrabrasilis 2023). From 1988 to 2024, the Brazilian Amazon lost approximately 500,000 km² of forests, of which more than 67,000 km² were in Rondônia (PRODES 2024), that has gone through several economic cycles based on rubber, minerals and currently on agribusiness, all of which have fomented large-scale environmental degradation (Fearnside 1989; Piontekowski et al. 2014).
Large-scale changes in natural vegetation cover, through clearing and burning, bring harmful consequences to the environment such as loss of local biodiversity, silting of rivers and changes in the hydrological cycle (Fearnside 2005, 2006; Correia et al. 2007; Watanabe et al. 2018; Souza et al. 2019). Another effect is the intense release of greenhouse gases, such as carbon dioxide, methane and nitrous oxide, which in turn, although they are measured regionally, have their effects extended on a global scale due to their contribution to global warming (Fearnside 2006).
The forest restoration/recovery actions can be efficient strategies for reversing forest loss and degradation through the conservation of local biodiversity and environmental services and in mitigating climate change (Joly 2007; Albuquerque and Silva 2008; Santos 2014; Benini et al. 2016). Accordingly, the UN General Assembly proclaimed the period 2021-2030 as the Decade of Ecosystem Restoration aiming to “support and expand efforts to prevent, deter and reverse the degradation of ecosystems” (UN 2019). This initiative highlights the importance and urgency of actions aimed at restoring and recovering degraded areas. At the UN conference, Brazil established goals to restore degraded areas in Brazilian biomes, committing to restore a total of 12 million hectares (Lima et al. 2022).
The Amazonian flora is characterized by a high species richness, with a confirmed occurrence of more than 14,000 plant species (Cardoso et al. 2017). In this scenario, difficulties arise for the various actors, seed collectors, reforesters, NGOs, that interact in the process of ecological recovery and restoration of degraded areas. Selecting the most suitable forest species for these purposes is one of the greatest demands of the sectors involved in recovery/restoration processes in the Amazon. In this sense, reliably identifying and working in situ with species suitable for the recovery and restoration of degraded/altered areas should be the first step to be achieved in the face of the challenges of these processes, allowing for greater diversity in the safe choice of species and probability of recovery (Barbosa et al. 2017).
In Rondônia, degraded areas occupy an increasing proportion of the landscape among the phytophysiognomies in the state (IBGE 2021). Therefore, our objective in this study was to determine tree and palms species known to occur in Rondônia and that have attributes suitable for restoration and recovery programs.
MATERIAL AND METHODS
Species selection
The composition of the list of tree and palm species indicated for restoration and recovery of degraded areas was based firstly on the species being reported in works published about secondary forests and degraded areas for the state of Rondônia, Brazil (Lisboa 1989; Isernhagen 2015; Benini et al. 2016; Paixão and Silveira 2020; Sccoti et al. 2020; Vieira et al. 2021). The list was complemented by field observations carried out by the authors since 2016 in monitoring work on recovered areas in northern Rondônia.
To select the best species for the list, the following criteria were considered: (a) species with abundance greater than 5%; (b) species with natural distribution in Rondônia based on herbarium data; (c) taxa consistently determined at the species level; and (d) availability of relevant ecological information on the species regarding successional category, functional group and dispersal syndrome. The organization of families was based on the APG IV system (Chase et al. 2016) and the species names were updated based on Flora e Funga do Brasil (2023).
Ecological data
Information regarding successional categories, dispersal syndrome and functional group was obtained by consulting published works (Beltrame and Rodrigues 2007; Andrade et al. 2011; Condé and Tonini, 2013; Klippel et al. 2015; Ribeiro 2016; Barbosa 2017; Paixão and Silveira 2020; Barbosa et al. 2021). We determined “successional category” based on seed production, tolerance/intolerance to light and growth time, classifying species into pioneers, secondary and climax species. The characterization of each group can be found in Rodrigues et al. (2009).
Dispersal syndromes result from morphological, chemical and biological characteristics of the propagules that favor the action of specific dispersing agents (Vieira et al. 2002), based on which the species were classified as anemochoric, hydrochoric, autochoric, barochoric and zoochoric. The characterization of each of the syndromes can be found in Rodrigues et al. (2009).
Functional groups (Barbosa 2017), also defined as planting groups (Rodrigues et al. 2009), refer to the system of classifying plants according to their different growth patterns and canopy density. Under this system, plants are divided into two groups: filling or covering species and diversity species. Filling species are fast-growing species with dense canopies, which allow for rapid soil coverage. Diversity species, on the other hand, lack these characteristics but contribute to the species richness of the restored area. The characterization of each group can be found in Rodrigues et al. (2009) and Barbosa et al. (2017).
Data on reproductive phenology and distribution of species in the state of Rondônia were taken from botanical records on online platforms: JABOT from the Herbarium Rondoniense (http://ron.jbrj.gov.br/v2/login.php), the database of the Emílio Goeldi Paraense Museum (https://floradobrasil.jbrj.gov.br/), the New York Botanical Garden (https://sweetgum.nybg.org/science/ih/), and Specieslink (https://specieslink.net/).
A table was created showing the habitats in which each species occurs, based on information from Flora e Funga do Brasil (2023). Based on collection records in Rondônia on the Specieslink platform (Specieslink 2023), we created a distribution map in the state for the 15 species with the highest number of records.
RESULTS
We selected 55 tree and palm species with potential for use in restoration and recovery of degraded areas (Table 1). The species belong to 49 genera and 23 families. The family with the largest number of species on the list is Fabaceae with 17 species, corresponding to 30.9% of the total. Of the remaining families, 14 had only one species (25.4%) and six families had two to five species (43.7%).
Tree and palm species recommended for use in restoration and recovery of degraded areas in the state of Rondônia (Brazil) and their respective ecological attribute, phenology, and habitats with natural occurences. SC = successional category (P = pioneer, S = secondary, C = climax); PG = planting group (R = covering, D = diversity); DS = dispersal syndrome (Zoo = zoochoric, Ane = anemochoric, Bar = barochoric, Aut = autochoric); ID = insufficient data. Asterisks (*) indicate species threatened with extinction according to the Ministry of the Environment ordinance 443 of December 17, 2014). Two asterisks (**) indicate habitat based on the authors field experience.
Regarding ecological groups, 22 of the species are pioneers (40%), 29 are typical of secondary forests (52.7%) and three are typical climax species (5.4%) (Supplementary Material, Table S1). No information was found about the ecological group of Godmania aesculifolia (Kunth) Standl. Regarding functional groups, 27 species belonged to the diversity group (49%) and 19 species belonged to the covering group (34.5%). We found no information regarding functional characteristics for eight species (14.5%). Regarding dispersal syndrome, 33 (60%) of the species are zoochoric, nine (16.3%) are anemochoric, six (10.9%) are barochoric and three (5.4%) are autochoric.
There were two main groups regarding flowering and fruiting periods. One group includes species in that flowering and fruiting occurs throughout the dry season, between June and August. In the second group, flowering and fruiting occurs in the transition period between the dry and rainy seasons, from September to November. For 11 species we only found information on the flowering period, and for 15 only on the fruiting period.
The 15 selected species occur in one or more of the main physiognomies that exist in Rondônia (Figure 1), including open environments such as campinarana and cerrado, seasonal semideciduous forests and floodplain forests by whitewater rivers (várzea) and blackwater rivers (igapós). Terra firme forest was the physiognomy with the largest number of species, with 46 (83.6%), followed by semideciduous forests (30 and 54%), floodplain forests (29 and 52.7%), and different physiognomies of cerrado (13 and 23.6%) (Table 1). Caryocar brasiliense Cambess. was the only species to occur only in cerrado. The vast majority of species occurred in more than one physiognomy.
Distribution of 15 tree species recommended for recovery/restoration of degraded areas in Rondônia state (Brazil) with the highest number of proven records in the surveyed herbaria.
Distribution of species in the state
More than 450 herbarium specimens were analyzed for the 55 species indicated for restoration and recovery of degraded areas. Dialium guianense Aubl. was the species with the highest number of locations with proven occurrence (18 municipalities), followed by Alchornea discolor Poepp. (17). Half of the species (30) occurred in more than eight municipalities. In contrast, Ochroma pyramidale (Cav. ex Lam.) Urb. and Cassia grandis L.f. were recorded only in the municipality of Porto Velho.
We found occurrence records of listed species for 45 of the 52 municipalities in Rondônia, showing the need for a more thorough sampling of the Rondônia flora. The municipalities that presented the highest number of species with proven occurrence were Porto Velho with 53 registered species, Itapuã do Oeste (38) and Guajará-mirim (27). Four municipalities presented one record (Alto Alegre dos Parecis, Brasilândia d´Oeste, Cujubim and Mirante da Serra).
DISCUSSION
We were able to recomend 55 tree and palm species native to the state of Rondônia for environmental restoration and recovery. They were recorded in fieldwork carried out by the authors or were indicated in published references that examined the flora of the state of Rondônia (Lisboa 1989; França 1991; Sanquetta et al. 2017; Vieira et al. 2021). The selected species are widely distributed in Amazonian forest formations (ter Steege et al. 2013), and 16 species are part of the list of 227 hyperdominant species proposed for the Amazon flora (ter Steege et al. 2016).
The family with the largest number of recommended species is Fabaceae, which is one of the most important in the Amazon flora (ter Steege et al. 2013; Cardoso et al. 2017; Santos et al. 2018; Paixão and Silveira 2020), and in tropical forests in general (ter Steege et al. 2013). Fabaceae plays an important role in the recovery of degraded areas (Sprent and Platzmann 2001), as many of its species fix nitrogen (Siddique et al. 2008), and play a role in soil restoration and the formation of organic matter (Resh et al. 2002; Nardoto et al. 2008), which enables the re-colonization of vegetation and an increase in biodiversity (Siddique et al. 2008).
The Arc of Deforestation, which extends from the eastern to the southwestern rim of the Brazilian Amazon, is subject to intense deforestation and burning. Some tree species, although widely distributed, are restricted to the Arc of Deforestation (Lisboa 1989; Silveira and Paixão 2019). They may therefore, to some extent, fit into one of the extinction risk categories for the Arc of Deforestation region, such as Swietenia macrophylla King, Amburana acreana (Ducke) A.C.Sm. and Godmania aesculifolia (Lisboa 1989; Silveira and Paixão 2019). Developing actions that seek to conserve these species are necessary in the northern region of Rondônia, where G. aesculifolia is among the species used in the recovery of degraded areas (personal observation by the authors.
Astrocaryum tucuma Mart. (tucumã), Vismia brasiliensis Choisy and V. guianensis (Aubl.) Choisy (lacre), Cecropia sp. (embaúba) and Solanum sp. (jurubeba) were not selected in this study because they naturally show better results in degraded areas than the planted species, being generalist species that quickly colonize the environment (Monaco et al. 2003; Mesquita et al. 2001). In this way, excessive settlement by these species occurs in degraded areas (personal observation by the authors) and can become a barrier to the development of other species, thus negatively affecting regenerating plant diversity. Therefore, planting these species is neither necessary nor recommended. The use of these species needs to be defined carefully and studied further.
The low number of collections in Rondônia for most species makes it impossible to understand their real distribution. Although they are common and easily identified species, they are rarely collected and recorded in the state, which does not reflect the high dominance and abundance observed in the field. This is, for example, the case of species such as Cassia grandis, Couratari stellata A.C.Sm., Cordia goeldiana Huber, Theobroma grandiflorum (Willd. ex Spreng.) K.Schum., Guazuma ulmifolia Lam. and Peltogyne paniculata Benth., among others. In our many field activities, we observed that these species are widely distributed throughout Rondônia. Even though they are easily found in a fertile state and are widely dispersed, when searching for information to compose the database that originated the present work, few records were found for Rondônia, which makes more detailed study impossible regarding its distribution in the state. Therefore, studies focused on secondary physiognomies are necessary to close knowledge gaps.
A greater proportion of flowering or fruiting species was observed in the dry period (June through August) and in the transition period from dry to rainy (September through November). Likewise, in the Tapajós region of Pará, a greater number of flowering species was observed in the dry season and transition to the rainy season (August through February) (Leão et al. 2001). The latter authors observed, over 14 years, much variation in periods of flowering, fruiting and dispersion of species, both among individuals of the same species and among species, which complicates interpretation of fertile periods of the species.
During our study, we observed the inconsistent or erroneous identification of many species, which led to many collections being left out of the list presented here. These problems arise from the high plant diversity observed in the Amazon (Procópio and Secco 2008; Cysneiros et al. 2018), exacerbated by the scarcity of taxonomists and parabotanists dedicated to the knowledge and study of the regional flora (Cysneiros et al. 2018). Therefore, even though many species on the list are common in altered environments or yield commercial wood, identification is still precarious and insecure, leaving many species with uncertain identification (Cysneiros et al. 2018).
Field observations by the authors made clear the need for greater care in the process of determining species used in recovery/restoration processes. Without precise identification, the objectives of restoration/recovery work may be compromised, since silvicultural treatments are based on the specific requirements of the species. For example, in the north of Rondônia, a species locally called ipê verde is widely used in recovery/restoration processes of degraded areas, and for this reason it has great importance in one of the most degraded areas in the state. Collected testimony material was sent to a specialist who identified it as G. aesculifolia. Previously it had been treated as Handroanthus sp. There is great similarity among species of the genera Handroanthus, Tabebuia and Godmania (Johanes et al. 2022). The similarity is reflected by the popular denomination ipê verde given to these species, as it relates to shared morphological characteristics. Although it occurs naturally in the state, there are no records of this species in floristic works carried out in Rondônia. This fact may be due to the low accuracy of identification, which can have lead the species to be mistakenly grouped with species of Handroanthus. The species has palmate leaves with 5-9 leaflets similar to those of Handroanthus and Tabebuia (Grose and Olmstead 2007). Godmania has fruits with spiral and striated capsules, and flowers with a campanulate corolla, and gibbous and hairy anthers (Grose and Olmstead 2007). Even with observable differences in the reproductive structures, the species was treated in processes of seed collection, seedling production and planting as Handroanthus and called ipê verde. The species has shown potential since 2010 for recovering degraded areas in Rondônia, mainly due to its rapid growth and biomass gain in these areas (Paixão et al. 2023; unpubl. data).
A second positive bias to be considered when using G. aesculifolia in recovery/restoration processes is conservation actions. Its distribution extends from Mexico to central Brazil, both in savanna and tropical forest (Specieslink 2023; Tropics.org 2023). The species has 50 records in the Brazilian Amazon, Caatinga and Cerrado biomes (Lohmann 2023; Specieslink 2023), in the states of Acre, Rondônia, Mato Grosso, Pará, Goiás, Tocantins, Maranhão and Roraima (Specieslink 2023; SiBBr 2023). The records are in anthropisized areas within urban perimeters by the Madeira River in the municipality of Porto Velho, and in pasture areas in Itapuã do Oeste (Specieslink 2023). The low number of G. aesculifolia records may be an indication that it is a rare species in Rondônia and its use in recovery/restoration processes contributed to its conservation in the region in the face of the advance of deforestation in the regions where the species is recorded. On the other hand, the higher number of occurrence records in terra firme forests is possibly due to the high richness of this environment in the Amazon (Oliveira and Mori 1999).
Thirteen species had proven occurrence in the cerrado habitats in the southern part of Rondônia (Silveira and Paixão 2019), which has been intensely altered, with extensive degraded abandoned areas that bear no more value for agriculture, livestock or forestry (Wandelli and Fearnside 2015; Silveira and Paixão 2019). These areas are challenging for restoration since they require species adapted to the restrictive conditions offered by the soils and seasonal climate of these savannas (Duboc et al. 2004; Silva et al. 2015). Therefore, having information on species adapted to these environments improves recovery efforts in a region where knowledge of native species among restoration professionals is still incipient.
CONCLUSIONS
Lack of herbarium records and inconsistencies in the identification to species and, in some cases, to family level, were major hindrances in the definition of the list of species recommended for environmental restoration in this study. A major issue is that incorrectly identified species compromise the use of information and knowledge produced about them, which can limit the success of large-scale restoration projects in the state of Rondônia. Our study shows that there is a need for greater investment of time and resources in botanical surveys, herbarium records, taxonomic studies, phenological characterization, the formation of human resources in botanical taxonomy in Rondônia, as well as the development of identification keys and manuals and their dissemination in the restoration scene. As is true for the Amazon flora in general, most species selected as potentially suitable for the recovery of degraded areas belonged to Fabaceae, with wide occurrence in the different habitats in Rondônia. It was also evident that more studies are needed of the secondary physiognomies for the knowledge of species that are successful in restored areas, in order to assist the actors involved in the ecological restoration/recovery of these areas, which will enable greater security and assertiveness in the processes of implementation of restoration/recovery actions.
ACKNOWLDEGMENTS
The authors would like to thank the support received from the Rio Terra Study Center, the New York Botanical Garden-NYBG, the João Geraldo Kuhlmann Herbarium of Rondônia and the Postgraduate Program in Conservation and Use of Natural Resources - PPGReN of the Biology Department of the Universidade Federal de Rondônia.
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Data availability
The data that support the findings of this study were published in this article and in its attached supplementary material.
SUPPLEMENTARY MATERIAL
Paixão et al. Plant species indicated for the recovery of degraded areas in Rondônia, Brazil
Ecological characterization of the species indicated for recovery of degraded areas in the state of Rondônia. Successional classification: P = pioneer, S = secondary, C = climax; Planting group: R = covering, D = diversity; Dispersion syndrome: Zoo = zoochoric, Ane = anemochoric, Bar = barochoric, Aut = autochoric. -- = unknown
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Edited by
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ASSOCIATE EDITOR:
Geângelo Calvihttps://orcid.org/0000-0002-8631-2325


