Open-access Floristic composition and similarity of a conservation area in the semiarid region of Paraíba State, Northeastern Brazil

Composição florística e similaridade de uma área de conservação do semiárido paraibano, Nordeste brasileiro

Abstract

We present here a floristic survey of the angiosperms found in the Reserva Ecológica Olho d’Água das Onças (REODO) in the semiarid region of Paraíba State, Brazil, to provide a basis for proactive conservation projects there. Fifteen excursions were carried out between August/2022 and April/2024 employing traditional methods of collecting and herborization of botanical material. The collections obtained were incorporated into the Manuel de Arruda Câmara Herbarium (HACAM) at the State University of Paraíba (UEPB), Campus I. We examined the floristic relationships between the study area and 15 other floristic surveys throughout the semiarid region of Paraíba State using a similarity analysis based on the Jaccard index. Two hundred and fifty species distributed among 168 genera and 50 families of angiosperms were identified with the following growth forms: herbaceous (74 spp.), subshrub (60 spp.), shrub (48 spp.), climber (45 spp.), arboreal (17 spp.), epiphytic (3 spp.), and parasitic (3 spp.). The most representative families were Fabaceae (38 spp.), Euphorbiaceae (24 spp.), Convolvulaceae (19 spp.), Malvaceae (18 spp.), Asteraceae (11 spp.), and Poaceae (11 spp.). The most representative genera were Ipomoea L. (8 spp.), Croton L. (7 spp.), Sida L. (7 spp.), Chamaecrista (L.) Moench (6 spp.), and Mimosa L. (5 spp.). Heteropterys arcuata C. Pessoa & Amorim (Malpighiaceae) and Solanum fernandesii V.S. Samp. & R. Moura (Solanaceae) were recorded for the first time in Paraíba State; Urvillea stipitata Radlk (Sapindaceae) represents a new record for the Caatinga. Similarity analysis revealed that the flora of REODO is more closely related to that of protected areas throughout the semiarid region of Paraíba State, underscoring its conservation priority. The study area was therefore found to contain significant floristic richness, especially within the context of the Caatinga, with 30 species endemics to this domain.

Keywords Angiosperms; biodiversity; Caatinga; conservation; new records

Resumo

O presente estudo compreende o levantamento florístico das angiospermas da Reserva Ecológica Olho d’Água das Onças (REODO), Picuí, semiárido da Paraíba, Brasil, objetivando fornecer a base para projetos conservacionistas na área. Foram realizadas 15 incursões durante o período de agosto de 2022 e abril de 2024, seguindo os métodos usuais de coleta e herborização de material botânico. A coleção obtida foi incorporada ao Herbário Manuel de Arruda Câmara (HACAM), pertencente à Universidade Estadual da Paraíba (UEPB), Campus I. Verificaram-se as conexões da flora da área estudada com outras áreas do semiárido paraibano por meio de uma análise de similaridade com base no índice de Jaccard, considerando outros 15 estudos florísticos. Nós encontramos 250 espécies distribuídas em 168 gêneros e 50 famílias de angiospermas pertencentes aos hábitos: herbáceo (74 spp.), subarbustivo (60 spp.), arbustivo (48 spp.), trepador (45 spp.), arbóreo (17 spp.), epifítico (3 spp.) e parasita (3 spp.). As famílias mais representativas foram: Fabaceae (38 spp.), Euphorbiaceae (24 spp.), Convolvulaceae (19 spp.), Malvaceae (18 spp.), Asteraceae (11 spp.) e Poaceae (11 spp.), e dentre os gêneros, Ipomoea L. (8 spp.), Croton L. (7 spp.), Sida L. (7 spp.), Chamaecrista (L.) Moench (6 spp.) e Mimosa L. (5 spp.) foram os mais numerosos. Heteropterys arcuata C. Pessoa & Amorim (Malpighiaceae) e Solanum fernandesii V.S. Samp. & R. Moura (Solanaceae) são registradas pela primeira vez para o Estado da Paraíba, e Urvillea stipitata Radlk (Sapindaceae) é um novo registro para a Caatinga. A análise de similaridade evidenciou que a flora da REODO é mais similar às das áreas protegidas no Estado da Paraíba, demonstrando a importância do seu reconhecimento como área prioritária para a conservação. Em suma, evidenciamos que a área de estudo apresenta uma expressiva riqueza florística especialmente no contexto da Caatinga, contando com 30 espécies endêmicas deste domínio.

Palavras-chave Angiospermas; biodiversidade; Caatinga; conservação; novos registros

Introduction

Caatinga (2°45’ S, 17°21’ S) is the predominant phytogeographic domain in Northeastern Brazil, covering 912,529 km2, which is equivalent to 10.7% of the area of that nation (Silva et al. 2017). According to Silva et al. (2017), the Caatinga domain is classified as a seasonally dry tropical forest and woodland (SDTFW), with at least 13 different phytophysiognomies, ranging from rock outcrops (or inselbergs) in the driest areas to semi-deciduous forests where the soils are richer, and being found in the Brazilian states of Maranhão, Piauí, Ceará, Rio Grande do Norte, Paraíba, Pernambuco, Alagoas, Sergipe, Bahia, and Minas Gerais (Silva et al. 2017).

The Caatinga was long considered the least known phytogeographical domain in Brazil (Giulietti et al. 2004) – which was reinforced by the idea that this domain had been greatly modified by human action and held only low plant biodiversity. That latter view is especially misleading and should not be disseminated, as the plant diversity of the Caatinga is now known to be quite high, with many rare and endemic species (Giulietti et al. 2004). The Caatinga domain is known to contain 180 angiosperm families comprising 1249 genera and 5060 species (Flora e Funga do Brazil, continually updated). Fernandes and Queiroz (2018) observed that 29 genera and approximately 23% of the plant species known to the Caatinga are endemic and restricted to only a few locations. Moro et al. (2014) cited the families with the greatest species richness in the Caatinga phytogeographic domain, with Fabaceae, Euphorbiaceae, Malvaceae, Asteraceae, and Convolvulaceae being the five richest.

The predominant Caatinga vegetation type is xerophytic and deciduous, being adapted to low annual rainfall (varying between 250 and 900 mm), with a rainy period generally lasting from three to five months (Alves 2007). Despite this general definition, there are great taxonomic disparities between different areas due to physiognomic, floristic, and seasonal climatic variations (Alves 2007). Considering the particularities of the flora, which is adapted to long water deficit periods, high solar irradiance, and high temperatures, Silva et al. (2021) highlighted the importance of the Caatinga vegetation in maintaining its regional fauna. Additionally, income generated by wood from arboreal plant communities, combined with the edible fruits, the medicinal, useful, and forage plants of the Caatinga domain have long supported local human communities, and can continue to do so as long as those resources are used in sustainable manners (Silva et al. 2021).

European/African colonization of the Caatinga region dates back approximately 500 years. However, the indiscriminate use of its resources, in activities such as grazing, deforestation, and mineral extraction has resulted in serious damage to the Caatinga and has resulted in impoverished water resources, soil erosion, and decreased biodiversity (Alves 2007, Alves et al. 2008). The urgency of conserving areas of Caatinga has become quite evident, and floristic studies have emerged as an essential tool in this process. It is impossible to consider conserving an area that is only poorly known. Understanding which species occur in a given area and which plant groups are most diverse form the basis for any ecological study seeking to preserve and/or maintain a biome and rationally manage its natural resources (Chaves et al. 2013).

The number of floristic surveys undertaken in Caatinga environments has grown in the last 20 years (personal observation). Among the numerous floristic inventories developed in this region we highlight in the Seridó region: Amorim et al. (2005), who focused on the shrub/arboreal flora of an area of Caatinga within the Seridó Ecological Station (ESEC) in Rio Grande do Norte State, and a floristic survey and species association undertaken by Santana et al. (2009) at the same location. Two studies addressing the flora of specific taxonomic groups were undertaken in the western Curimataú microregion: one dealing with the Asteraceae family (Soares et al. 2021) and the other encompassing the order Boraginales (Sousa et al. 2021). Additionally, other studies have been published in the last two decades focusing on the flora of the Paraíba Caatinga: in the Cariri microregion (Barbosa et al. 2007, Lacerda et al. 2007, Araújo et al. 2010, Santos and Melo 2010, Trovão et al. 2010, Pereira-Júnior et al. 2012, Lima and Barbosa 2014, Farias et al. 2017, Marques et al. 2020); in the Agreste mesorregion (Porto et al. 2008), and in the Sertão mesoregion (Silva et al. 2012, Lucena et al. 2015, Silva et al. 2015, Sabino et al. 2016, Medeiros et al. 2021, Sales et al. 2021, Fernando et al. 2022).

Considering the scarcity of floristic surveys carried out in areas of Caatinga in the Curimataú and Seridó regions of Paraíba State (PB), and recognizing the importance of a detailed knowledge of the floras of these areas to be able to elaborate recovery and conservation plans, a floristic survey and a similarity analysis of the angiosperms in the Reserva Ecológica Olho d’Água das Onças (REODO), a conservation area located in the municipality of Picuí in the semiarid region of Paraíba State, Northeastern Brazil, was undertaken, aiming to understand the diversity and floristic relationships within the study area and in comparison with other floristic surveys conducted in the semiarid region of Paraíba State.

Material and Methods

1. Study area

The municipality of Picuí (6°28’– 6°69’S, 36°21’– 36°46’W) is located in Paraíba State in Northeastern Brazil (Francisco et al. 2011). The municipality covers 667 km2 (IBGE 2022) and is situated in the intermediate region of Campina Grande, more specifically in the immediate region of Cuité - Nova Floresta (IBGE 2017). The Reserva Ecológica Olho d’Água das Onças (6°27’S, 36°18’W) is located 11 km distant from the administrative center of Picuí (Figure 1). The Ecological Reserve comprises an area of 20.73 ha, of which 18.26 ha were allocated to the conservation of the Caatinga phytogeographical domain in 2005 (Silva 2021). Elevations in the reserve range between 590 m and 659 m.a.s.l. (pers. comm.).

Figure 1
Location map of the study area, Reserva Ecológica Olho d’Água das Onças, municipality of Picuí, Paraíba State, Brazil. Produced by E. M. Rodrigues (2022).

The regional climate is tropical hot and dry, with prolonged droughts (type BSh according to the Köppen classification, adapted by Francisco et al. 2015). The average annual regional temperatures are between approximately 23° and 24° C (Francisco and Santos 2017). Two representative soil types were identified by Silva et al. (2022): Yellow Oxisoil in the Chapada area and Litholic Neosoil at the edge of the Chapada. In the study area, the present vegetation range from open sketches to shrubby and shrubby-arboreal phytophysiognomies. The expected precipitation in the municipality of Picuí is 339.1mm/year, but in the years during which this study was undertaken (2022 and 2023), the precipitation rates were 453.1 and 481.5mm/year respectively (AESA, continually updated).

2. Fieldwork

Fieldwork was conducted between August/2022 and April/2024 (except May and June/2023) employing random walks throughout the entire length of the reserve to obtain fertile plant specimens (with flowers and/or fruits). A total of 15 incursions were carried out, covering both the dry and rainy periods (Figure 2). The collections and herborization processes were based on the recommendations of Walters (2011) for succulents and Peixoto and Maia (2013) for other groups. The material obtained was incorporated into the Manuel de Arruda Câmara Herbarium (HACAM) at the State University of Paraíba (UEPB), Campus I. Geographic coordinates were obtained using a handheld GPS; photographic images were captured using a cell phone.

Figure 2
Physiognomies of the Reserva Ecológica Olho d’Água das Onças, Picuí, Paraíba State, Brazil. a-d. Rainy season. e-g. Dry season. Photos: B. F. Rocha.

3. Data analysis

The specialized literature was consulted to identify the species, including florulas, regional floras, unpublished taxonomic texts (TCC, master’s dissertations, and doctoral theses), and, principally, floristic and/or floristic-taxonomic articles published in Botany journals in Brazil and abroad. Additionally, comparisons were made based on collections (exsiccates) deposited in herbaria in Paraíba and surrounding states and made available on online platforms (GBIF, Reflora, SpeciesLink). The Flora e Funga do Brasil platform (continuously updated) was used to verify endemism. The checklist of the flora of the Reserva Ecológica Olho d’Água das Onças is based on the Angiosperm Phylogeny Group (APG) IV (2016) and the Flora e Funga do Brazil (continuously updated) and includes the names of the families, genera, and species, accompanied by their respective habit types and testimonial material. The spelling of the authors’ names were based on the Flora e Funga do Brasil (continuously updated) and Tropicos (continuously updated) platforms.

4. Similarity

The similarity analysis was based on floristic surveys undertaken in Caatinga areas throughout Paraíba State on the crystalline basement. These studies were identified through the Google Scholar platform using the descriptors “floristics”, “Caatinga” and “Paraíba” as well as studies obtained through the authors themselves (Costa et al. 2015, Lima and Melo 2015, Melo et al. 2016, Cordeiro and Souza 2024, Machado-Filho et al. 2024). The selected studies employed similar methodologies to ours, and were published between 2009 and 2024, totaling 15 studies. Based on these floristic surveys, a binary matrix of presence/absence was produced, following the recommendations of Hubálek (1984).

Species names were thoroughly verified, and identifications at the family or genus levels, as well as names marked as (cf.) and morphospecies were excluded from the binary matrix. Synonyms were checked using the Flora e Funga do Brasil (continuously updated) and Tropicos (continuously updated) platforms, to avoid erroneous results. The analysis was performed using the UPGMA (Unweighted Pair Group Method with Arithmetic mean) method, based on Jaccard’s similarity index, with the PAST 4.17 software (Hammer et al. 2001), resulting a similarity dendrogram.

Results

We collected and classified 250 species in the Reserva Ecológica Olho d’Água das Onças belonging to 168 genera and 50 angiosperm families (Figures 310). Of those species, 248 were identified to the specific level, one species could only be determined to the generic level, and one remains undetermined (Table 1). Among the families recorded, five are monocotyledons (10%) and 45 are eudicots (90%). The most representative families in terms of the number of species were: Fabaceae (38 spp.), Euphorbiaceae (24 spp.), Convolvulaceae (19 spp.), Malvaceae (18 spp.), Asteraceae (11 spp.), and Poaceae (11 spp.), representing approximately 48.4% of the total number of species found (Figure 11). The genera with the highest species richness were: Ipomoea L. (8 spp.), Croton L. (7 spp.), Sida L. (7 spp.), Chamaecrista (L.) Moench (6 spp.), and Mimosa L. (5 spp.). With regard to monocotyledons, the following family richness were recorded: Poaceae (11 spp.), Bromeliaceae (6 spp.), Commelinaceae (5 spp.), Cyperaceae (4 spp.) and Amaryllidaceae (1 spp.).

Figure 3
a. Harpochilus paraibanus (Acanthaceae). b. Ruellia asperula (Acanthaceae). c. Gomphrena vaga (Amaranthaceae). d. Zephyranthes cearensis (Amaryllidaceae). e. Spondias tuberosa (Anacardiaceae). f. Ibatia nigra (Apocynaceae). g. Aspilia bonplandiana (Asteraceae). h. Centratherum punctatum (Asteraceae). i. Tilesia baccata (Asteraceae). Photos: a-b, h. M. G. M. Gonçalves; c-g, i. B. F. Rocha.
Figure 4
a. Mansoa onohualcoides (Bignoniaceae). b. Mansoa paganuccii (Bignoniaceae). c. Bromelia laciniosa (Bromeliaceae). d. Neoglaziovia variegata (Bromeliaceae). e. Commiphora leptophloeos (Burseraceae). f. Tacinga subcylindrica (Cactaceae). g. Xiquexique gounellei (Cactaceae). h. Cynophalla flexuosa (Capparaceae). i. Physostemon guianense (Cleomaceae). Photos: a-b, d-i. B. F. Rocha; c. M. G. M. Gonçalves.
Figure 5
a. Combretum leprosum (Combretaceae). b. Commelina obliqua (Commelinaceae). c. Evolvulus glomeratus (Convolvulaceae). d. Ipomoea marcellia (Convolvulaceae). e. Ipomoea rosea (Convolvulaceae). f. Jacquemontia pentanthos (Convolvulaceae). g. Varronia leucomalloides (Cordiaceae). h. Cucumis anguria (Cucurbitaceae). i. Cyperus uncinulatus (Cyperaceae). Photos: a-c, f-i. B. F. Rocha; c-d. M. G. M. Gonçalves.
Figure 6
a. Erythroxylum caatingae (Erythroxylaceae). b. Acalypha multicaulis (Euphorbiaceae). c. Croton blanchetianus (Euphorbiaceae). d. Euphorbia comosa (Euphorbiaceae). e. Microstachys corniculata (Euphorbiaceae). f. Ancistrotropis peduncularis (Fabaceae). g. Cenostigma nordestinum (Fabaceae). h. Chamaecrista repens (Fabaceae). i. Macropsychanthus grandiflorus (Fabaceae). Photos: a-f, i. B. F. Rocha; g-h. M. G. M. Gonçalves.
Figure 7
a. Mimosa invisa (Fabaceae). b. Peltogyne pauciflora (Fabaceae). c. Heliotropium angiospermum (Heliotropiaceae). d. Marsypianthes chamaedrys (Lamiaceae). e. Mentzelia aspera (Loasaceae). f. Pusillanthus pubescens (Loranthaceae). g. Cuphea impatientifolia (Lythraceae). h. Diplopterys lutea (Malpighiaceae). i. Helicteres eichleri (Malvaceae). Photos: a-e, g. B. F. Rocha; f, h. M. G. M. Gonçalves; i. L. G. Sá.
Figure 8
a. Sida anomala (Malvaceae). b. Sida galheirensis (Malvaceae). c. Wissadula contracta (Malvaceae). d. Mollugo verticillata (Molluginaceae). e. Boerhavia coccinea (Nyctaginaceae). f. Oxalis divaricata (Oxalidaceae). g. Passiflora foetida (Passifloraceae). h. Angelonia pubescens (Plantaginaceae). i. Plumbago scandens (Plumbaginaceae). Photos: a, c-h. B. F. Rocha; b, i. M. G. M. Gonçalves.
Figure 9
a. Cenchrus echinatus (Poaceae). b. Chloris barbata (Poaceae). c. Setaria parviflora (Poaceae). d. Asemeia violacea (Polygalaceae). e. Portulaca elatior (Portulacaceae). f. Sarcomphalus joazeiro (Rhamnaceae). g. Cordiera rigida (Rubiaceae). h. Urvillea stipitata (Sapindaceae). i. Sideroxylon obtusifolium (Sapotaceae). Photos: a, h. M. G. M. Gonçalves; b-g, i. B. F. Rocha.
Figure 10
a. Capsicum parvifolium (Solanaceae). b. Physalis pubescens (Solanaceae). c. Talinum fruticosum (Talinaceae). d. Piriqueta viscosa (Turneraceae). e. Laportea aestuans (Urticaceae). f. Lantana achyranthifolia (Verbenaceae). g. Pombalia arenaria (Violaceae). h. Cissus erosa (Vitaceae). i. Tribulus terrestris (Zygophyllaceae). Photos: a, c-d, f-g, i. B. F. Rocha; b, h. M. M. Rodrigues; e. M. F. B. Sousa.
Table 1
List of Angiosperm families and species from Reserva Ecológica Olho d’Água das Onças, Picuí, Paraíba State, Brazil. * Endemic to Brazil; ** Endemic to the Caatinga.
Figure 11
Number of species of the richest families in comparison with the others families in the Reserva Ecológica Olho d’Água das Onças, Picuí, Paraíba State, Brazil.

Two species constitute new records for the flora of Paraíba State: Heteropterys arcuata C.Pessoa & Amorim (Malpighiaceae) and Solanum fernandesii V.S. Samp. & R. Moura (Solanaceae). Urvillea stipitata Radlk (Sapindaceae) is recorded here for the Caatinga for the first time. These findings reinforce the importance of undertaking floristic surveys in areas of Caatinga.

In terms of growth form, an herbaceous phytophysiognomy predominated, with 74 species (29.6%) (Figure 12), this stratum being essentially restricted to the rainy season, followed by subshrub (60 spp.; 24%), shrub (48 spp.; 19.2%), climbing plants (45 spp.; 18%), and tree (17 spp.; 6.8%) components. Three epiphytic species (1.2%) were also found attached to shrubs and trees, all belonging to the genus Tillandsia L. (Bromeliaceae). Three parasitic plant species (1.2%) were also identified: two holoparasites belonging to the genus Cuscuta L. (Convolvulaceae), characterized by being leafless and achlorophyllous, and one hemiparasitic plant belonging to the family Loranthaceae, Pusillanthus pubescens (Rizzini) Caires, the latter having leaves and chlorophyll; both had modified roots (haustoria) to obtain nutrients from the host plant.

Figure 12
Distribution of Angiosperm growth form from Reserva Ecológica Olho d’Água das Onças, Picuí, Paraíba State, Brazil.

Among the total species found, 79 are endemic to Brazil (31.6%); the other species have extra-Brazilian distributions (68.4%). It is important to highlight that 30 species (12%) are restricted to the Caatinga phytogeographical domain, indicating the RE Olho d’Água das Onças as a priority conservation center for plant diversity in the Paraíba Caatinga.

Based on the Jaccard similarity index, the cophenetic correlation coefficient for the dendrogram is 82.2%, indicating a relatively strong analysis. The flora of the REODO exhibits greater similarity to floristic surveys carried out in protected areas (Figure 13), forming a well-defined group with them (Lima and Barbosa 2014, Fernando et al. 2022). This group shows greater similarity with studies conducted in inselbergs located in the municipality of Patos, in the Sertão mesoregion (Lucena et al. 2015, Lopes-Silva et al. 2019). Another group includes the rocky outcrops situated in the Borborema Plateau, specifically in the Paraíba Cariri and Agreste regions (Tölke et al. 2011, Sales-Rodrigues et al. 2014, Costa et al. 2015, Lima and Melo 2015, Melo et al. 2016, Machado-Filho et al. 2024). Furthermore, a separate group comprises floristic surveys undertaken in the eastern region of the Caatinga phytogeographical domain in Paraíba (Cordeiro et al. 2015, Cordeiro et al. 2018, Cordeiro and Souza 2024).

Figure 13
Dendrogram of floristic similarity including the present study and others floristic surveys conducted in the Caatinga domain throughout Paraíba State, Brazil.

Discussion

Barbosa et al. (2007) identified 396 species for the flora of the entire Cariri region of Paraíba State (29 municipalities, 11,236,391 km2). The number of species found in the present study (250 species) was therefore equivalent to more than 60% of that total, despite the study area being considerably smaller (20.73 ha). Other surveys carried out in Caatinga areas in Paraíba reported greater numbers of species: Fernando et al. (2022) encountered 320 angiosperm species in an area of Caatinga in the municipality of Passagem, Paraiban Sertão mesoregion; Lima and Barbosa (2014) identified 290 angiosperm species in the RPPN Fazenda Almas, an area of Caatinga located in Paraiban Cariri microregion. The greater numbers of species found by these authors probably reflect the longer durations of their field excursions with longer collection times during flowering and/or fruiting periods (Fernando et al. 2022 – four collection years; Lima and Barbosa 2014 – ten collection years) and the larger sizes of their study areas (Fernando et al. 2022 – 350 hectares; Lima and Barbosa 2014 – 3,505 hectares).

Our study evidenced high species richness as compared to other surveys carried out in Caatinga regions in Paraíba, such as inselbergs (Porto et al. 2008, Tölke et al. 2011, Sales-Rodrigues et al. 2014, Lima and Melo 2015, Lucena et al. 2015, Melo et al. 2016), riparian forests (Lacerda et al. 2007, Trovão et al. 2010, Silva et al. 2015, Farias et al. 2017, Marques et al. 2020), and even areas of preserved Caatinga (Pereira-Júnior et al. 2012, Silva et al. 2012, Cordeiro and Souza 2024, Machado-Filho et al. 2024). It is important to note that some of these studies focused only on certain types of plant habits (only the herbaceous or only the shrub/tree layer).

Fabaceae (38 spp.) was the most species-rich family in the present study, corroborating the findings of Moro et al. (2014) who likewise identified it as the richest family in the Caatinga. Several studies carried out in Paraíba State had previously identified Fabaceae as the richest family in this domain (Porto et al. 2008, Santos and Melo 2010, Lima and Barbosa 2014, Sales-Rodrigues et al. 2014, Lima and Melo 2015, Lucena et al. 2015, Silva et al. 2015, Melo et al. 2016, Fernando et al. 2022, Cordeiro and Souza 2024, Machado-Filho et al. 2024). Surveys undertaken in the Caatinga domain employing different methodologies (focusing only on the herbaceous component - Andrade et al. [2009], or only on the woody component [Lacerda et al. 2007, Trovão et al. 2010, Pereira-Júnior et el. 2012, Farias et al. 2017, Marques et al. 2020) have consistently shown Fabaceae to be the richest family, demonstrating the diverse habits of its component species. Even among surveys in which it is not the richest family, it appears among the three most prominent in terms of the number of species (Costa et al. 2009, Tölke et al. 2011, Silva et al. 2012).

Chamaecrista (6 spp.) and Mimosa (5 spp.) were the richest genera of Fabaceae, corroborating Moro et al. (2014), who identified these genera as being among the four richest in the Caatinga, and the two richest genera of Fabaceae in the caatinga phytogeographic domain. Furthermore, Lima et al. (2015) highlighted that most Fabaceae species have associations with nitrogen-fixing bacteria in their roots, which are largely responsible for the entry of nitrogen compounds into terrestrial ecosystems.

Euphorbiaceae (24 spp.) was found to be the second richest family, again in agreement with Moro et al. (2014), who identified it as the second richest family in the Caatinga. Euphorbiaceae appeared as the second richest family in other studies undertaken in Caatinga regions (Lucena et al. 2015, Fernando et al. 2022), and as the richest in the study by Costa et al. (2009), who analyzed the flora of one hectare of Caatinga in the mid São Francisco depression in Pernambuco State. Additionally, Euphorbiaceae stood out among the richest families in terms of the number of species in studies undertaken in areas of the Caatinga sedimentary basement (Gomes et al. 2006, Lima et al. 2009, Sousa et al. 2021).

Additionally, in studies focusing exclusively on woody Caatinga components, Euphorbiaceae is often cited among the most species-rich families (Lacerda et al. 2007, Trovão et al. 2010, Pereira-Júnior et al. 2012, Farias et al. 2017, Marques et al. 2020). In the present study, there were 17 woody Euphorbiaceae species among shrubs and subshrubs, which is equivalent to approximately 70.8% of the total species of the family encountered in the study area, and approximately 15.7% of all shrub and subshrub species there. Five herbaceous species and two climbing plants (Dalechampia scandens L. and Tragia cearensis Pax & K.Hoffm.) were also identified. The most numerous genus in the family in the present study was Croton L. (7 spp.), which, according to Moro et al. (2014), is the richest Caatinga genus. The prominence of Croton was likewise observed in other studies carried out in Caatinga areas (e.g., Farias et al. 2017, Sousa et al. 2021, Cordeiro and Souza 2024).

Convolvulaceae (19 spp.) and Malvaceae (18 spp.) stood out for their floristic importance in this study, being the third and fourth richest families respectively. Convolvulaceae was notable for its high number of climbing species: of the 19 species found in this study, 12 were climbers, beyond hosting the two species of holoparasites plants found, both of the genus Cuscuta. Convolvulaceae represented approximately 26.7% of all of the climbing species identified in the study, emerging as an important floristic component of climbing plants in the study area. Tree species of Malvaceae, belonging to the Bombacoideae subfamily, are frequently found in areas of Caatinga vegetation (Lacerda et al. 2007, Santos and Melo 2010, Guedes et al. 2012, Pereira-Júnior et al. 2012, Lima and Barbosa 2014, Lima and Melo 2015, Sabino et al. 2016, Fernando et al. 2022, Machado-Filho et al. 2024), but no arboreal species of the family was encountered in the study area.

Convolvulaceae and Malvaceae are cited as among the most species-rich families in several floristic studies of Caatinga vegetation (Araújo et al. 2008, Andrade et al. 2009, Santos and Melo 2010, Tölke et al. 2011, Silva et al. 2012, Lima and Barbosa 2014, Lima and Melo 2015, Lucena et al. 2015, Silva et al. 2021, Fernando et al. 2022, Cordeiro and Souza 2024, Machado-Filho et al. 2024). Species of Convolvulaceae and Malvaceae are not generally included among the richest genera in studies that analyze only the shrub/tree component of Caatinga areas (Lacerda et al. 2007, Trovão et al. 2010, Pereira-Júnior et al. 2012, Farias et al. 2017, Marques et al. 2020), probably due to the predominance of climbers (in Convolvulaceae) and subshrubs (in Malvaceae) in these families, as evidenced in the present study. Additionally, Ipomoea L. (Convolvulaceae) (8 spp.) and Sida L. (Malvaceae) (7 spp.) represent the two richest genera of those families in the area, corroborating the results presented by Moro et al. (2014), with Ipomoea appearing as the third richest genus and Sida as the ninth most taxonomically diverse genus in the Caatinga domain.

The Asteraceae (11 spp.) and Poaceae (11 spp.) families ranked fifth among the richest families. Large numbers of species of both families were found in recently anthropized environments (areas invaded by Neltuma juliflora (Sw.) Raf. and close to the administrative center of the reserve), indicating their colonization of habitats in the initial stages of regeneration. Among the 11 Poaceae species found, seven were cited by Ferreira et al. (2009) in their study at the Seridó Ecological Station (ESEC) in Rio Grande do Norte State as pioneer and/or invasive species: Aristida setifolia Kunth, Cenchrus ciliaris L., Chloris barbata Sw., Chloris virgata Sw., Dactyloctenium aegyptium (L.) Willd., Tragus berteronianus Schult., and Urochloa mollis (Sw.) Morrone & Zuloaga, corroborating our results.

Asteraceae was the richest family on an inselberg located in the rural area of the municipality of Puxinanã, PB (Tölke et al. 2011), which exemplifies its ability to colonize this type of environment as pioneers. Poaceae was the richest family in a study carried out at Fazenda Tamanduá, Santa Terezinha, PB, where collections were carried out in both conserved and anthropized environments (Silva et al. 2012). This latter study focused only on the herbaceous component, which would tend to highlight the predominance of Poaceae species (generally herbaceous) (Silva et al. 2012); Poaceae is also the most taxonomically diverse monocot family in the Caatinga (Flora e Funga do Brasil, continuously updated). Asteraceae and Poaceae are cited in several other studies as being among the richest families (Araújo et al. 2008, Porto et al. 2008, Andrade et al. 2009, Costa et al. 2009, Lima and Barbosa 2014, Sales-Rodrigues et al. 2014, Lima and Melo 2015, Lucena et al. 2015, Silva et al. 2015, Melo et al. 2016, Fernando et al. 2022, Cordeiro and Souza 2024).

Several studies have shown that herbaceous plants predominate to the detriment of other growth form in crystalline basement Caatinga areas (Araújo et al. 2008, Porto et al. 2008, Costa et al. 2009, Tölke et al. 2011, Sales-Rodrigues et al. 2014, Lima e Melo 2015, Lucena et al. 2015, Silva et al. 2015, Melo et al. 2016, Fernando et al. 2022, Machado-Filho et al. 2024), differing from areas of sedimentary basement Caatinga (Gomes et al. 2006, Lima et al. 2009, Sousa et al. 2021), where arboreal and shrub habits predominate. The soils of crystalline basement Caatinga areas are shallow and stony, and water does not accumulate there for very long days after the rains (which themselves last only a few months) – which could explain the predominance of an herbaceous layer – with annual plants spending most of the year as seeds, developing and reproducing only when the rainy season begins again (Silva et al. 2017).

Regarding floristic similarity, the dendrogram indicates that the present study is most closely related to the surveys carried out by Fernando et al. (2022) (J = 27.4) and Lima and Barbosa (2014) (J = 23.2), all of which were undertaken in protected areas, despite the relatively considerable distances between the REODO and these areas (107.3 km from Fazenda ABA – municipality of Passagem and 130.86 km from RPPN Fazenda Almas - municipality of São José dos Cordeiros). This finding reinforces the tendency of certain species of establish themselves in conserved areas. For instance, in our analysis, the species Tanaecium parviflorum (Mart. ex DC.) Kaehler & L.G.Lohmann (Bignoniaceae), Evolvulus frankenioides Moric. (Convolvulaceae), Argythamnia malpighiacea Ule, Croton adenocalyx Baill. (Euphorbiaceae), Rhynchosia minima (L.) DC. (Fabaceae) and Stachytarpheta coccinea Schauer (Verbenaceae) were recorded exclusively in these three studies.

Lopes-Silva et al. (2019) and Lucena et al. (2015) are the most closely related studies (J = 36.3), which can be attributed to both surveys had been undertaken in inselbergs in the municipality of Patos, and thus sharing specialized species adapted to this type of environment. In this context, Porembski (2007) notes that inselbergs are prominent rocky environment that create specific climatic and edaphic conditions, such as high temperature, low air humidity, minimal soil accumulation and limited water retention capacity, factors that favor a highly adapted flora. These surveys exhibit greater similarity to the group which the present study belongs, probably due to the proximity to the areas where Fernando et al. (2022) and Lima and Barbosa (2014) carried out their research.

The geographical proximity and microenvironmental characteristics also account for the relationship between the studies of Lima and Melo (2014) and Machado-Filho et al. (2024) (J = 31.4), both studies conducted in rocky outcrops within the Environmental Protection Area (APA) of Cariri, which encompasses the municipalities of Boa Vista and Cabaceiras. This proximity explains the floristic connections between these studies and those undertaken in rocky outcrops in the Agreste mesoregion, specifically in the municipalities of Pocinhos (Costa et al. 2015) and Puxinanã (Tölke et al. 2011, Sales-Rodrigues et al. 2014, Melo et al. 2016), all of them located on the Borborema Plateau. The lower floristic similarity between Sales-Rodrigues et al. (2014) and the other studies located on Borborema Plateau is likely due to the absence of certain species that are relatively common in rocky outcrops, such as Dyckia spectabilis (Mart. ex Schult. & Schult.f.) Baker (Bromeliaceae) and Senna martiana (Benth.) H.S. Irwin & Barneby. (Fabaceae), which are present in the other studies of this group but not record in the study carried out by Sales-Rodrigues et al. (2014). Additionally, the presence of exclusive species, such as Mandevilla dardanoi M.F. Sales, Kin.-Gouv. & A.O. Simões (Apocynaceae), may had contributed to this difference.

Another group formed on the dendrogram includes the surveys carried out in the pre-coastal Agreste region, on the eastern edge of Caatinga in the Paraíba State (Cordeiro et al. 2015, Cordeiro et al. 2018, Cordeiro and Souza 2024). In this region, the climate type is As’, characterized as hot and humid, with precipitation ranging between 800-1000 mm/year (Francisco et al. 2015), and it is bordered by the Borborema Plateau to the west and the coastal plateaus to the east (Costa et al. 2020). These distinctive conditions probably influenced the selection of flora in this area, resulting in a flora that is notably distinct from those in others analyzed regions, thereby explaining the formation of this group.

The study carried out by Silva et al. (2015), conducted along the Piranhas River, Sertão mesoregion, is the second least similar of the analyzed studies. This lower similarity is probably due to it being the only floristic survey focused on a riparian forest and the unique peculiarities of the study area, such as its low altitude of 148 m (Silva et al. 2015), which contributes to a distinct flora. The area studied by Santos and Melo (2010), despite being in Cariri region, where the surveys of Lima and Melo (2015) and Machado-Filho et al. (2024) were conducted, exhibited the lowest floristic similarity with all other studies (Tölke et al. 2011, Lima and Barbosa 2014, Sales-Rodrigues et al. 2014, Cordeiro et al. 2015, Costa et al. 2015, Lima and Melo 2015, Lucena et al. 2015, Silva et al. 2015, Melo et al. 2016, Cordeiro et al. 2018, Lopes-Silva et al. 2019, Fernando et al. 2022, Cordeiro and Souza 2024, Machado-Filho et al. 2024 and this study). This disparity can be attributed primarily to the limited sampling effort, which encompassed only six months of fieldwork, and the presence of recently anthropized patches in the study area.

Pinto et al. (2020) carried out a scientometric analysis of the Caatinga, listing the species considered invasive in this domain. Among the species they cited, five were encountered during our fieldwork: Ricinus communis L. (Euphorbiaceae), Leucaena leucocephala (Lam.) de Wit, Neltuma juliflora (Sw.) Raf. (Fabaceae), Cenchrus ciliaris L. and Megathyrsus maximus (Jacq.) B.K.Simon & S.W.L.Jacobs (Poaceae). Ricinus communis, L. leucocephala, and C. ciliaris were found near the reserve headquarters, a highly anthropized environment due to recreational activities carried out there; M. maximus was found close to an area invaded by N. juliflora, a frequent invasive species in areas of Caatinga (Andrade et al. 2010) that reduces biodiversity when it becomes well established. The occurrence of these species is likely associated with former uses of the area (agriculture and livestock raising) before becoming established as a protected conservation reserve in 2005.

Conclusion

The number of angiosperm species found in the Reserva Ecológica Olho d’Água das Onças was found to be high when compared to other studies carried out in Caatinga conserved sites in Paraíba State, Brazil. Thirty species (12%) were identified as endemic to this phytogeographic domain, highlighting the importance of proactive conservation measures for managing the study area. Moreover, similarity analysis demonstrates a stronger relationship between REODO and other protected areas in the Paraíba semiarid region, reinforcing its priority for conservation. Some species are naturalized but restricted to anthropized locations. Their presence, however, is concerning, as it reflects the anthropization that took place in some locations in the Ecological Reserve before it was declared a conservation area. More studies will be necessary to fully determine the biodiversity of the Reserva Ecológica Olho d’Água das Onças and implement proactive measures to effectively protect the area.

Acknowledgments

The first author would like to thank FAPESq (Fundação de Apoio à Pesquisa do Estado da Paraíba) and CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico) for granting a Bolsa de Longa Duração - Apoio Técnico/Nível Médio (BLD-AT/Nível Médio), and an Iniciação Científica - Cota 2023-2024 (Proc. n. 135844/2023-1) grant through the Programa de Iniciação Científica of the Universidade Estadual da Paraíba (PIBIC/UEPB); the Bolsa de Produtividade em Pesquisa-PQ Nível E (Proc. n. 306658/2022-4) awarded to J.I.M. Melo; FAPESq (Fundação de Apoio à Pesquisa do Estado da Paraíba) for financing this study through Contract 510/2022 (Call: Emenda Impositiva de Apropriação Nº. 484- LOA 2022); Mr. Rubens Germano Costa, the owner of the Reserva Ecológica Olho d’Água das Onças (REODO) for permission to carry out this study and for the use of the Reserve facilities; Marcio Gleisson Medeiros Gonçalves for capturing images of species during the fieldwork; the specialists, for their assistance in identifying or confirming the following groups: Ibatia ganglinosa – Apocynaceae (Hector Keller); Bromeliaceae (Thaynara Silva); Combretaceae (Maria Iracema Loiola); Commelinaceae and Portulacaceae (Igor Albuquerque); Cyperaceae (André Rodolfo Ribeiro); Erythroxylum pyan – Erythroxylaceae (James Costa-Lima); Croton – Euphorbiaceae (Maria de Fátima Araújo); Fabaceae and Schwenckia americana – Solanaceae (Rubens Queiroz); Malpighiaceae (André Amorim); Oxalidaceae (Maria Carolina Abreu); Poaceae (Carlos Santos); Sapindaceae (María Silvia Ferrucci); Solanum fernandesii – Solanaceae (Yuri Gouvêa); Vitaceae (Diego do Nascimento Silva); and the State University of Paraíba (UEPB), for transportation and other facilities provided to facilitate this research. To Anderson Silva Pinto for his help during the dendrogram obtention.

Data Availability

Supporting data are available at <https://doi.org/10.5281/zenodo.11237488>.

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Edited by

  • Associate Editor
    Alexander Vibrans

Publication Dates

  • Publication in this collection
    28 Oct 2024
  • Date of issue
    2024

History

  • Received
    27 May 2024
  • Accepted
    02 Oct 2024
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