Aquatic and palustrine plants of restinga in a Ramsar site in the easternmost Amazon, Brazil

Alves, N. P. F.; Pestana, M. C. A.; Pires, S. F.; Penha, J. C. A.; Cardoso-Neto, V. R.; Scatigna, A. V.; Saraiva, R. V. C.

doi:10.1590/1519-6984.288620

Abstract

Aquatic and palustrine plants are a group of plants that have morphological and anatomical adaptations to occupy permanent or temporary aquatic environments. In this study, we carried out the first floristic survey of aquatic and palustrine plants in restingas (restinga swamps and swamp forests) of a Ramsar site in the municipality of Guimarães, western coast of Maranhão State and easternmost Amazon, Brazil. In total, 52 species of 43 genera and 28 families were collected between July 2022 and October 2023, during the dry and rainy seasons. Voyria aphylla (Jacq.) Pers. is a new record for Maranhão. Melastomataceae (9 spp.) was the most representative family, followed by Fabaceae and Plantaginaceae (4 spp. each). Palustrine and emergent plants accounted for 58% and 31% of the species richness, respectively. This study adds important information for local and national policies to ensure biodiversity conservation of the plant biodiversity in Maranhão’s coastal ecosystems.

Keywords:
aquatic vegetation; macrophytes; new record; Reentrâncias Maranhenses

Resumo

As plantas aquáticas e palustres são um grupo de plantas que possuem adaptações morfológicas e anatômicas para ocupar ambientes aquáticos permanentes ou temporários. Neste estudo, realizamos o primeiro levantamento florístico de plantas aquáticas e palustres em restingas (“brejos de restinga” e “florestas paludosas de restinga”) de um sítio Ramsar no município de Guimarães, costa ocidental do Estado do Maranhão e extremo leste da Amazônia, Brasil. No total, 52 espécies de 43 gêneros e 28 famílias foram coletadas entre julho de 2022 e outubro de 2023, durante as estações seca e chuvosa. Voyria aphylla (Jacq.) Pers. é um novo registro para o Maranhão. Melastomataceae (9 spp.) foi a família mais representativa, seguida por Fabaceae e Plantaginaceae (4 spp. cada). As plantas palustres e emergentes foram responsáveis por 58% e 31% da riqueza de espécies, respectivamente. Este estudo acrescenta informações importantes para as políticas locais e nacionais para garantir a conservação da biodiversidade vegetal nos ecossistemas costeiros do Maranhão.

Palavras-chave:
vegetação aquática; macrófitas; novo registro; Reentrâncias Maranhenses

1. Introduction

The restinga vegetation grows on sandy plains (from narrow strips to extensive areas of sand), covering approximately 79% of the Brazilian coast (Veloso et al., 1991; Zickel et al., 2004). Restinga vegetation is characterized by different plant associations organized in mosaics, which favors a rich and heterogeneous flora (Lima et al., 2017). This vegetation is influenced by the sea in soils of Tertiary or Quaternary origin and includes grasslands, shrubs, and trees that can reach up to 20 m in height (Brasil, 2002; Lima et al., 2017). Hence, these plant communities are distinct in terms of their floristic and phytophysiognomic aspects (Araujo and Henriques, 1984; Cerqueira, 2000; Martins, 2012).

Among the types of vegetation found in restinga areas, swamp formations occur in flood-prone regions. Their soil is permanently waterlogged and associated with swamp deposits due to water table emergence (Rodrigues, 2004; Venzke et al., 2015). These formations encompass environments transitioning between coastal wetlands (restinga swamps, locally known as “brejos de restinga”) and swamp forests (locally known as “florestas paludosas de restinga”). Although restinga swamps and swamp forests exhibit distinct floristic and physiognomic compositions, their species can germinate in flooded hydromorphic soils (Torres et al., 1994; Rodrigues, 2004).

In Brazil, studies on swamp formations have focused on understanding their floristic composition, community structure, and ecosystem services (Marques et al., 2003; Teixeira and Assis, 2009; Terror et al., 2011). However, research on swamp formations in coastal areas is primarily concentrated in the Southeast (Sztutman and Rodrigues, 2002; Teixeira and Assis, 2005; Loures et al., 2007; Kurtz et al., 2013) and South regions of Brazil (Dorneles and Waechter, 2004; Santos-Junior et al., 2015). It is also important to mention that these studies generally do not include aquatic and palustrine plants and focus on the arboreal stratum. In Maranhão State, Northeast Brazil, several studies have been conducted in restinga areas (Cabral-Freire and Monteiro, 1993; Serra et al., 2016; Lima and Almeida Júnior, 2018; Correia et al., 2020; Guterres et al., 2020). However, these studies have focused exclusively on dune formations. Consequently, there is a significant gap in the floristic knowledge of restinga swamp formations in the state, particularly regarding aquatic and palustrine plants.

Aquatic and palustrine plants (APs) are defined as vegetation capable of withstanding permanent or periodic/seasonal submergence of their root system, enabling them to occupy humid environments (Amaral et al., 2008). This concept encompasses a diverse and distinct group of plants, such as angiosperms, bryophytes (e.g., thallose liverworts), and seedless vascular plants (Chambers et al., 2008). Regardless of phylogenetic aspects, APs are classified according to their biological forms, reflecting their anatomical and morphological adaptations to survive in truly aquatic environments or waterlogged soils (Alves et al., 2011; Thomaz and Esteves, 2011). The most common life forms include amphibious (or palustrine), emergent, floating (rooted or free), submerged (rooted or free), and epiphyte species (Pott and Pott, 2000; Amaral et al., 2008).

Knowledge about the floristic composition and biological forms of APs in Maranhão is still incipient (Pestana et al., 2024a). No floristic surveys of this plant group were conducted in the state between 2010 and 2017 (Moura-Júnior and Cotarelli, 2019). Moreover, some existing studies in Maranhão were developed in the Baixada Maranhense Environmental Protection Area, mainly focusing on limnological variables (e.g., dissolved oxygen) and ecological aspects (e.g., biomass) (Barbieri and Pinto, 1999; Costa-Neto et al., 2002; Barbieri and Carreiro, 2017).

Floristic surveys provide essential data to characterize a plant community in terms of species composition, contributing to studies related to the biology, ecology, and conservation of plant biodiversity (Chaves et al., 2013; Oliveira and Amaral, 2004; Corrêa et al., 2018). In the Maranhão coast, floristic studies are essential for monitoring APs and managing activities to ensure biodiversity conservation in this area (Araújo et al., 2020). This need stems from the intense degradation observed in the state’s coast, due to the increase in the human occupational process (Almeida Júnior et al., 2018). Therefore, in this work we carried out the first floristic survey of APs in the restinga of Guimarães (restinga swamps and swamp forest). We provide diagnostic descriptions, distribution data, and photographs of the collected species.

2. Material and Methods

2.1. Study area

The Reentrâncias Maranhenses Environmental Protection Area (EPA) is a Brazilian Conservation Unit with an area of approximately 2,680,911.2 hectares located on the western coast of the state of Maranhão (Hazin, 2008). This EPA was designated as a Ramsar site (Convention on Wetlands of International Importance Especially as Waterfowl Habitat) in 1993 (Ramsar, 2024). The coastline is very diverse, comprising bays, islands, estuaries, and extensive areas of mangroves that predominantly have Amazonian vegetation, but with a diverse influence from the surrounding vegetation, especially coastal (Hazin, 2008; Spinelli-Araujo et al., 2016). The productivity and extensive support of this ecosystem is particularly important for aquatic and terrestrial life forms (Silva, 2019).

Among the 18 municipalities that constitute the Reentrâncias Maranhenses EPA, Guimarães (02°03’04”S, 44°31’28”W) covers a total area of secondary forest vegetation of 599 km² (Figure 1). The region’s topography is characterized by coastal plains shaped by marine and fluvial agents and processes, giving rise to beaches, mangroves, swamps, tidal flats, lagoons, and cliffs (Ramos, 2008). According to the Köppen climate classification system, the local climate is Tropical hot and humid (Aw), with a dry summer and a rainy winter (Alvares et al., 2013). The average annual precipitation ranges from 1250 and 2000 mm and the average annual temperature is 26 °C. The rainy season extends from January to June and the dry season from July to December.

Figure 1
Map of the location of the study area and the collection points in Guimarães, Maranhão, Brazil. Map preparation: M. Pestana.

For this study, we selected two flooded areas in each type of restinga swamp formations of the municipality of Guimarães: 1) Swamp forest, located in Abrantes balneary (02°04’14”S, 44°36’04”W; max. elev. 11 m a.s.l.); and Passaginha balneary (02°04’17”S, 44°36’16”W; max. elev. 21 m a.s.l.). 2) Restinga swamps, located in the Guajerutiua village (02°05’11”S, 44°34’49”W; max. elev. 10 m a.s.l.) and Carapirá village (02°03’42”S, 44°33’56”W; max. elev. 6 m a.s.l.) (Figures 1 and 2).

Figure 2
Examples of the aquatic environments where we collected APs plants in Guimarães: (A) Swamp Forest in Abrantes balneary; (B) Swamp Forest in Passaginha balneary; (C) Restinga Swamps in Guajerutiua village; (D) Restinga Swamps Carapirá village. Photos: R.V.C. Saraiva.

2.2. Data collection

The field expeditions were conducted biannually, in July and September 2022, and in May and October 2023, during the dry and rainy seasons. Each expedition lasted 4 days, with collections made during morning and afternoon shifts (from 8:00 AM to 5:00 PM), involving the participation of three researchers. Botanical specimens were collected in the surroundings, banks, and inside the lotic and lentic aquatic environments of the four selected areas, using the walking method (Filgueiras et al., 1994). Samples in the fertile stage (with flowers and/or fruits) were herborized following the recommendations of Haynes (1984).

Species identification was made by morphological analysis of reproductive and vegetative structures using a stereomicroscope and by comparing them with specialized literature (Cook, 1996; Irgang and Gastal Junior, 1996; Pott and Pott, 2000; Amaral et al., 2008; Gonçalves and Lorenzi, 2011). Additionally, we used identification keys and the integrated herbarium network SpeciesLink (CRIA, 2024) and Reflora (JBRJ, 2024a), prioritizing specimens identified by specialists. Specimens were incorporated into the collection of the Rosa Mochel Herbarium (SLUI, acronym following Thiers, 2024), at the State University of Maranhão, in São Luís.

The terminology used for the diagnostic descriptions of the species follows Gonçalves and Lorenzi (2011). Information on the specimen’s habit and habitat was established through field observations. The geographical distribution of the species was determined by consulting Flora e Funga do Brasil (JBRJ, 2024b) and SpeciesLink (CRIA, 2024). Angiosperm and fern families are presented according to the classification system of the Angiosperm Phylogeny Group (APG IV, 2016) and the Pteridophyte Phylogeny Group (PPG I, 2016), respectively. The species names follow the Flora e Funga do Brasil (JBRJ, 2024b). Species were also categorized according to their life forms, as proposed by Pott and Pott (2000) and Amaral et al. (2008) (Figure 3).

Figure 3
Biological forms of APs plants dawning from the categories proposed by Pott and Pott (2000) and Amaral et al. (2008): (1) Palustrine; (2) Emergent; (3) Rooted-floating; (4) Free-floating; (5) Rooted-submerged; (6) Free-submerged; (7) Epiphyte. Dawning by R.S. Soares.

3. Results

In the restinga swamp formations of Guimarães, 52 species distributed in 43 genera and 28 families were recorded (Table 1; Figures 4-6). Of these, only one is a fern (Lygodium volubile Sw.), whereas the other 51 species belong to the angiosperm group. Among these, Voyria aphylla (Jacq.) Pers. is a new record for Maranhão State (JBRJ, 2024b).

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Table 1
List of families, species and biological forms of the APs plants collected in the restinga swamps formations in Guimarães.

Figure 4
Species of APs plants collected in Guimarães: (A) Sagittaria guayanensis; (B) Lepidagathis alopecuroidea; (C) Dieffenbachia aglaonematifolia (globose fruit in detail); (D) Sphagneticola trilobata; (E) Apteria aphylla; (F) Cabomba aquatica; (G) Costus spiralis; (H) Scleria microcarpa; (I) Tonina fluviatilis; (J) Aeschynomene evenia; (K) Desmodium adscendens; (L) Mimosa pudica. Photos: R.V.C. Saraiva.

Figure 6
Species of APs plants collected in Guimarães: (A) Nymphaea rudgeana; (B) Sauvagesia erecta; (C) Ludwigia affinis; (D) Rugoloa pilosa; (E) Pontederia diversifolia; (F) Bacopa aquatica (flower in detail); (G) Bacopa aubletiana; (H) Bacopa salzmannii; (I) Bacopa scoparioides; (J) Oldenlandia lancifolia; (K) Pentodon pentandrus; (L) Xyris jupicai. Photos: R.V.C. Saraiva.

The most representative family in the floristic survey was Melastomataceae (9 species - 17% of the richness), followed by Fabaceae (4 species - 8%), Plantaginaceae (4 species - 8%), Cyperaceae (3 species - 6%), Rubiaceae (3 species - 6%), and Lentibulariaceae (3 species - 6%). Gentianaceae, Onagraceae, Poaceae, and Xyridaceae had two species each and together accounted for 15% of the richness. The other 18 families were represented by only one species. The richest genus was Bacopa Aubl. (4 species), followed by Utricularia L. (3), Ludwigia L. (2), Clidemia D.Don (2), Pterolepis (DC.) Miq. (2), and Xyris Gronov. ex L. (2). The Melastomataceae and Fabaceae not only had the highest species richness but also the greatest genus diversity, with seven and four genera recorded, respectively.

Regarding the biological forms, palustrine plants were represented by 30 species (58%), emergent plants by 16 (31%), rooted-floating plants by four (8%), and rooted-submerged plants by two species (4%). Among the palustrine species, Melastomataceae (9 spp.) and Fabaceae (4 spp.) were the predominant families. The four emergent species belong to the genus Bacopa of the Plantaginaceae. Nymphaea rudgeana G.Mey. (Nymphaeaceae), Nymphoides humboldtiana (Kunth) Kuntze (Menyanthaceae), Pontederia diversifolia (Vahl) M.Pell. & C.N.Horn (Pontederiaceae), and Sagittaria guayanensis Kunth (Alismataceae) were rooted-floating. Meanwhile, submersed species were represented by Cabomba aquatica Aubl. (Cabombaceae) e Tonina fluviatilis Aubl. (Eriocaulaceae).

3.1. Enumerated list of species

1. Alismataceae Vent.

1.1 Sagittaria guayanensis Kunth (Figure 4A).

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 03-VII-2022; fl., fr., Saraiva, R. 346 (SLUI); ibid., Povoado Carapirá, 24-IX-2022, fl., fr., Penha, J. 50 (SLUI).

Species recognition: Rooted-floating herb. Stem rhizomatous, cylindrical. Leaves simple, alternates, in rosette; blades ovate, base sagittate, apex obtuse, margin entire. Inflorescence racemose. Flowers trimerous, petals white with a yellow center; bracts lanceolate, membranaceous. Fruit achene with tuberculate surface.

Distribution and habitat: Occurs in all Brazilian regions and states. In the study area, was collected in Swamp forest and Restinga swamps.

2. Acanthaceae Juss.

2.1 Lepidagathis alopecuroidea (Vahl) R.Br. ex Griseb. (Figure 4B).

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 23-IX-2022; fl., Penha, J. 56 (SLUI).

Species recognition: Palustrine herb, glabrous. Stem decumbent. Leaves simple, opposite; blades elliptic, base acute, apex rounded, margin entire. Inflorescence thyrse, axillary. Flowers bilabiate, petals white with a purple spot at base.

Distribution and habitat: North (AM, PA), Northeast (AL, MA, PE), and Central-West (GO, MT). In the study area, was collected in Swamp forest.

3. Araceae Juss.

3.1 Dieffenbachia aglaonematifolia Engl. (Figure 4C).

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha,12-V-2023; fl., fr., Penha, J. 68 (SLUI).

Species recognition: Palustrine herb, glabrous, dioecious. Stem climbing. Leaves simple, alternate; blades oblong, base cordate, apex rounded, margin entire. Inflorescence spadix. Pistillate flowers white. Fruit globose, smooth, orange-yellow.

Distribution and habitat: North (PA, TO), Northeast (AL, MA), Central-West (GO, MS, MT), Southeast (MG, SP), and South (PR). In the study area, was collected in Swamp forest.

4. Asteraceae Bercht. & J.Presl

4.1 Sphagneticola trilobata (L.) Pruski (Figure 4D)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 12-V-2023; fl., fr., Saraiva, R. 365 (SLUI).

Species recognition: Emergent herb. Stem decumbent. Leaves simple, three-lobed, alternate; blades oval, pubescent, trichomes simple, rigid, base attenuate, apex acuminate, margin serrated. Inflorescence in capitulum, bracts yellow. Flowers 17–20, petals yellow-gold.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Swamp forest.

5. Burmanniaceae Blume

5.1 Apteria aphylla (Nutt.) Barnhart ex Small (Figure 4E)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 12-V-2023; fl., Saraiva, R. 366 (SLUI).

Species recognition: Mycoheterotrophic palustrine herb. Stem erect, unbranched, whitish. Inflorescence cymose, terminal. Flower trimerous, tepals lilac.

Distribution and habitat: North (AC, AM, PA) and Northeast (AL, BA, MA, PB, PE, SE). In the study area, was collected in Swamp forest.

6. Cabombaceae Rich. ex A.Rich.

6.1 Cabomba aquatica Aubl. (Figure 4F)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 23-IX-2022; fl., fr., Penha, J. 48 (SLUI).

Species recognition: Rooted-submerged herb. Stem reddish. Leaves heterophyllous; submerged blades dissected, palmate; floating blades peltate, elliptical to ovate, apex mucronate, base obtuse, margin entire. Inflorescence axillar, bracteoles ovate, reddish. Flowers trimerous, emersed, tepals intensely yellow.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Swamp forest.

7. Costaceae Nakai

7.1 Costus spiralis (Jacq.) Roscoe (Figure 4G)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Povoado Passaginha, 12-V-2023; fl., fr., Penha, J. 78 (SLUI).

Species recognition: Palustrine herb, glabrous. Stem rhizomatous. Leaves simple, sessile, alternate, rosulate; blades elliptic, glabrous, base cordate, apex acuminate, margin entire. Inflorescence spike, terminal, bracts and bracteoles red. Flowers trimerous, petals pink.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Swamp forest.

8. Cyperaceae Juss.

8.1 Cyperus haspan L.

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 12-V-2023; fl., Nunes, M. 70 (SLUI).

Species recognition: Palustrine herb. Stem rhizomatous, trigonous. Leaves simple, sessile, alternate; blades linear, glabrous, margin entire; ligule absent. Inflorescence in spike terminal. Flowers glumes, lanceolate, glabrous; bracts involucre.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Swamp forest.

8.2 Scleria microcarpa Nees ex Kunth (Figure 4H)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Povoado Guajerutiua, 14-V-2023; fl., fr., Nunes, M. 67 (SLUI).

Species recognition: Palustrine herb. Stem rhizomatous. Leaves simple, sessile, alternate; blades linear, glabrous, apex acute, margin scabrous. Inflorescence in panicle, terminal. Flowers glume, brown. Fruit dome, surface smooth.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Restinga swamps.

8.3 Rhynchospora riparia (Nees) Boeckeler

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 1-VII-2022; fl., Saraiva, R. 354 (SLUI).

Species recognition: Emergent herb. Stem rhizomatous. Leaves simple, sessile, alternate; blades basal, linear, glabrous, apex acute, margin entire. Inflorescence spike, terminal; bracts ciliate; spikelet ovoid, arranged in fascicles. Flowers white. Fruit not seen.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Swamp forest.

9. Eriocaulaceae Martinov

9.1 Tonina fluviatilis Aubl. (Figure 4I)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 23-IX-2022; fl., Penha, J. 58 (SLUI); ibid., Povoado Carapirá, 24-IX-2022; fl., Penha, J. 60 (SLUI).

Species recognition: Rooted-submerged herb. Stem long, erect. Leaves simple, sessile, verticillate; blades lanceolate, pubescent, trichomes simple, base amplexicaul, apex acute, margin entire. Inflorescence in capitulum, bracts lanceolate, light brown. Flowers with petals reduced to lobes.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Swamp forest and Restinga swamps.

10. Fabaceae Lindl.

10.1 Aeschynomene evenia C.Wright & Sauvalle (Figure 4J)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 2-V-2023; fl., fr., Saraiva, R. 367 (SLUI).

Species recognition: Emergent herb. Stem cylindrical, glabrous. Leaves compound, bifoliolate, alternate distichous; blades pinnate, glabrous, base obovate, apex rounded, margin entire; interpetiolar stipule present. Inflorescence axillary. Flowers bilabiate, petals yellow. Fruit legume.

Distribution and habitat: North (PA), Northeast (AL, BA, CE, MA, PE, PI, SE), Central-West (GO, DF, MS), and Southeast (ES, RJ, SP). In the study area, was collected in Swamp forest.

10.2 Chamaecrista diphylla (L.) Greene

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Povoado Guajerutiua, 14-X-2023; fl., fr., Alves, N. 1 (SLUI).

Species recognition: Palustrine herb. Stem ascending, glabrous, stipule basifixed. Leaves compound, alternate, glabrous; leaflets with obtuse base, acuminate apex, entire margin. Inflorescence cymose, axillary. Flowers pentamerous, petals yellow. Fruit legume, glabrous.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Restinga swamps.

10.3 Desmodium adscendens (Sw.) DC. (Figure 4K)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 1-VII-2022; fl., fr., Saraiva, R. 343 (SLUI).

Species recognition: Palustrine herb. Stem prostrate, branch cylindrical, stipule lanceolate. Leaves compound, trifoliate, alternate, distichous; blades pinnate; leaflets with obtuse base, rounded apex, margin entire. Inflorescence pseudoracemose. Flowers pentamerous, petals lilac. Fruit loment, pubescent, trichomes uncinate.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Swamp forest.

10.4 Mimosa pudica L. (Figure 4L)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 1-VII-2022; fl., fr., Penha, J. 79 (SLUI).

Species recognition: Palustrine herb. Stem creeping, trichome hirsute, stipule triangular. Leaves compound, bipinnate, alternate; leaflets glabrous; blades oblong, base asymmetrical, apex mucronate, margin entire. Inflorescence glomerular, axillary. Flowers with pink fillets, yellow anthers. Fruit craspid, hirsute.

Distribution and habitat: North (AC, AM, AP, PA, RO, RR, TO), Northeast (AL, BA, CE, MA, PB, PE, PI, SE), Central-West (DF, GO, MT, MS), Southeast (ES, MG, RJ, SP), and South (PR, RS, SC). In the study area, was collected in Swamp forest.

11. Gentianaceae Juss.

11.1 Chelonanthus viridiflorus (Mart.) Gilg (Figure 5A)

Figure 5
Species of APs plants collected in Guimarães: (A) Chelonanthus viridiflorus; (B) Voyria aphylla (new record); (C) Utricularia pusilla (white arrow); (D) Utricularia gibba (yellow arrow); (E) Lindernia rotundifolia; (F) Nymphoides humboldtiana; (G) Aciotis annua; (H) Comolia villosa; (I) Clidemia biserrata; (J) Miconia ciliata; (K) Nepsera aquatica (flower in detail); (L) Pterolepis glomerata. Photos: R.V.C. Saraiva.

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Povoado Carapirá, 24-IX-2022; fl., fr., Penha, J. 62 (SLUI); ibid., Balneário Passaginha, 1-VII-2022; fl., fr., Saraiva, R. 356 (SLUI).

Species recognition: Palustrine herb. Stem erect, cylindrical, glabrous. Leaves simple, sessile, opposite; blades ovate, glabrous, base attenuate, apex acute, margin entire. Flowers pentamerous, petals greenish-white. Fruit capsule, elongated.

Distribution and habitat: North (AM, PA, RR, TO), Northeast (CE, MA, PE), Central-West (GO, MS, MT), Southeast (MG, SP), and South (PR). In the study area, was collected in Swamp forest and Restinga swamps.

11.2 Voyria aphylla (Jacq.) Pers. (Figure 5B)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 12-V-2023; fl., Penha, J. 67 (SLUI).

Species recognition: Palustrine, mycoheterotrophic herb. Leaves simple, sessile, opposite, reduced in the shape of conical scales in the lower half. Flowers pentamerous, petals yellow, triangular lobes. Fruit not seen.

Distribution and habitat: According to Flora e Funga do Brasil (JBRJ, 2024b), its previous distribution was in North (AC, AM, AP, PA, RR), Northeast (AL, BH, CE, PB, PE, RN, SE), Central-West (MT), Southeast (ES, MG, RJ, SP), and South (PR, RS, SC). However, this is its first record for the flora of Maranhão. In the study area, was collected in Swamp forest.

12. Lamiaceae Martinov

12.1 Hyptis atrorubens Poit.

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 12-V-2023; fl., Saraiva, R. 399 (SLUI).

Species recognition: Palustrine herb. Stem decumbent. Leaves simple, opposite; blades ovate, glabrous, base truncate, apex acute, margin crenate. Inflorescence capitulum; bracteoles involucre. Flowers sessile, petals white with a pink spot on the lower lobe.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Swamp forest.

13. Lentibulariaceae Rich.

13.1 Utricularia pusilla Vahl (Figure 5C)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 23-IX-2022; fl., Penha, J. 55 (SLUI).

Species recognition: Palustrine herb. Stolon branched. Leaves dichotomously branched, simple; blades obovate, apex rounded, margin entire; utricles ovoid. Inflorescence racemose. Flowers trilobed, corolla yellow; upper lip narrowly oval; lower lip flat; spur parallel to lower lip, elongated.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Swamp forest.

13.2 Utricularia subulata L.

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 1-VII-2022; fl., fr., Saraiva, R. 341 (SLUI).

Species recognition: Palustrine herb. Leaves dichotomously branched, simple; blades linear, glabrous, apex obtuse, margin entire; utricles ovoid. Inflorescence racemose. Flowers yellow, trilobed; upper lip ovate, lower lip lobed. Fruit capsule, globose.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Swamp forest and Restinga swamps.

13.3 Utricularia gibba L. (Figure 5D)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 12-X-2023; fl., Alves, N. 2 (SLUI).

Species recognition: Emergent herb. Leaves dichotomously branched, simple, glabrous, margin entire; utricles ovoid. Inflorescence simple, scape flexible, slender. Flowers yellow, palate gibbous with brown stripe; upper lip ovate, tri-crenate; lower lip gibbous, spur slightly curved.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Swamp forest.

14. Linderniaceae (Rchb.) Borsch et al.

14.1 Lindernia rotundifolia (L.) Alston (Figure 5E)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 24-IX-2022; fl., Penha, J. 65 (SLUI).

Species recognition: Emergent herb. Leaves simple, sessile, opposite; blades oval, base round, apex obtuse, margin entire. Flowers axillary, corolla white with blue spots. Fruit not seen.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Swamp forest.

15. Lygodiaceae M.Roem

15.1 Lygodium volubile Sw.

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 1-VII-2022; Saraiva, R. 360 (SLUI).

Species recognition: Palustrine fern. Stem rhizomatous, cylindrical. Leaves compound, alternate; blades bipinnate, pilose, trichomes simple; leaflets opposite, pinnules with base truncate, apex acute, margin dentate. Segments fertile, sporangia marginal, from base towards apex of pinnule.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Swamp forest.

16. Malvaceae Juss.

16.1 Sida linifolia Cav.

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Povoado Carapirá, 1-VII-2022; fl., Saraiva, R. 359 (SLUI).

Species recognition: Palustrine herb. Stem erect, cylindrical, pubescent, trichomes simple. Leaves simple, alternate; blades lanceolate, glabrescent, trichomes simple; base subcordate, apex acute, margin entire; stipule linear. Inflorescence corymbose. Flowers pentamerous, petals white with a reddish base. Fruit not seen.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Restinga swamps.

17. Menyanthaceae Dumort.

17.1 Nymphoides humboldtiana (Kunth) Kuntze (Figure 5F)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Povoado Guajerutiua, 14-V-2023; fl., Nunes, M. 66 (SLUI).

Species recognition: Rooted-floating herb. Stem rhizomatous. Leaves simple, alternate, rosulate; blades membranaceous, base cordate, apex rounded, margin sinuate; adaxial surface green, abaxial surface vinaceous. Inflorescence axillary. Flowers pentamerous, petals white, fimbriate.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Restinga swamps.

18. Melastomataceae Juss.

18.1 Aciotis annua (Mart. ex DC.) Triana (Figure 5G)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Povoado Carapirá, 3-VII-2022; fl., Saraiva, R. 357 (SLUI).

Species recognition: Palustrine herb. Stem erect, villous. Leaves simple, opposite; blades ovate, pilose, trichomes non-glandular; base truncate, apex acute, margin entire; 6 basal veins. inflorescence cymose. Flowers tetramerous, petals white.

Distribution and habitat: North (AC, AM, PA, RR), Northeast (BA, CE, MA, PI), and Central-West (GO). In the study area, was collected in Swamp forest and Restinga swamps.

18.2 Comolia villosa (Aubl.) Triana (Figure 5H)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Povoado Carapirá, 23-IX-2022; fl., Penha, J. 54 (SLUI).

Species recognition: Palustrine herb. Stem erect, pilose, trichomes glandular. Leaves simple, opposite; blades oblong, villous, trichomes hirsute, base obtuse, apex acute, margin serrate; 5 basal veins. Inflorescence terminal. Flowers tetramerous, petals purple.

Distribution and habitat: North (AM, AP, PA, RR) and Northeast (BA, CE, MA, PB, PE, PI, RN, SE). In the study area, was collected in Swamp forest and Restinga swamps.

18.3 Clidemia biserrata DC. (Figure 5I)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Povoado Guajerutiua, 12-V-2023; fl., Saraiva, R. 369 (SLUI).

Species recognition: Palustrine subshrub. Stem pilose, trichomes stellate. Leaves simple, opposite; blades elliptic, adaxial surface with filiform trichomes, abaxial surface with stellate trichomes; base cordate, apex acuminate, margin dentate; 5 basal veins. Flowers pentamerous, petals pink.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Restinga swamps.

18.4 Clidemia hirta (L.) D.Don

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 14-X-2023; fl., fr., Saraiva, R. 395 (SLUI).

Species recognition: Palustrine subshrub. Stem erect, pilose, trichomes simple. Leaves simple, opposite; blades oblong or ovate, pilose, trichomes setulose, base cordate, apex acuminate, margin dentate; 5 basal veins. Inflorescence axillary, bracts filiform. Flowers pentamerous, petals white. Fruit berry.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Swamp forest.

18.5 Miconia ciliata (Rich.) DC. (Figure 5J)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Abrantes, 23-IX-2022; fl., Penha, J. 53 (SLUI).

Species recognition: Palustrine subshrub. Stem erect, pilose, trichomes simple. Leaves simple, opposite; blades elliptic, simple trichomes on the adaxial surface; base obtuse, apex acute, margin serrate; 5 basal veins. Inflorescence thyrse. Flowers pentamerous, lilac.

Distribution and habitat: North (AC, AM, AP, RO, RR, TO), Northeast (AL, BA, CE, MA, PB, PE, PI, RN, SE), Central-West (GO, MT), and Southeast (ES, MG). In the study area, was collected in Swamp forest.

18.6 Nepsera aquatica (Aubl.) Naudin (Figure 5K)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Povoado Carapirá, 3-VII-2022; fl., fr., Saraiva, R. 358 (SLUI); ibid., Povoado Guajerutiua, 14-V-2023; fl., fr., Nunes, M. 53 (SLUI).

Species recognition: Emergent herb. Stem slightly pilose, trichomes simple. Leaves simple, opposite; blades ovate, glabrous, base subcordate, apex acuminate, margin serrate; 5–7 basal veins. Inflorescence panicle, axillary. Flowers tetramerous, petals white. Fruit capsule.

Distribution and habitat: North (AM, PA, AP, TO) and Northeast (MA, RN, PB, PE, AL, SE). In the study area, was collected in Swamp forest and Restinga swamps.

18.7 Pterolepis glomerata (Rottb.) Miq. (Figure 5L)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Povoado Carapirá, 24.IX.2022; fl., Penha, J. 63 (SLUI); ibid., Balneário Passaginha, 3.VII.2022; fl., Saraiva, R. 350 (SLUI).

Species recognition: Emergent herb. Stem cylindrical, pubescent, trichomes filiform. Leaves simple, opposite; blades narrow-lanceolate, base acute, apex acute, margin crenate; 3 basal veins. Inflorescence axillary. Flowers tetramerous, petals lilac, hypanthium with long red trichomes.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Restinga swamps.

18.8 Pterolepis trichotoma (Rottb.) Cogn.

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 12-V-2023; fl., Saraiva, R. 370 (SLUI).

Species recognition: Palustrine herb. Stem erect, pubescent, trichomes strigose. Leaves simple, opposite; blades narrowly elliptic, base attenuate, apex acute, margin serrated; 5 basal veins. Inflorescence thyrse. Flowers tetramerous, petals pink. Fruit not seen.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Swamp forest.

18.9 Tococa guianensis Aubl.

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 12-V-2023; fl., fr., Nunes, M. 65 (SLUI).

Species recognition: Palustrine subshrub. Stem pilose, trichome glandular. Leaves simple, opposite; domatium at petiole apex; blades obovate, base rounded, apex attenuate, margin entire; 5 basal veins. Inflorescence panicle, terminal. Flowers pentamerous, petals white to pink. Fruit globose.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Swamp forest.

19. Nymphaeaceae Salisb.

19.1 Nymphaea rudgeana G.Mey. (Figure 6A)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Abrantes, 12-V-2023; fl., Saraiva, R. 371 (SLUI); ibid., Balneário Passaginha, 12-V-2023; fl., Penha, J. 83 (SLUI).

Species recognition: Rooted-floating herb. Stem rhizomatous. Leaves simple, long-petiolate; petiole cylindrical, reddish; blades cordate, base cordate, apex rounded, margin sinuate. Flowers floating, petals white. Fruit berries.

Distribution and habitat: North (AM, AP, PA, RO, RR, TO), Northeast (AL, BA, CE, MA, RN, PB, PE, PI, SE), Southeast (ES, MG, RJ, SP), and South (PR). In the study area, was collected in Swamp forest.

20. Ochnaceae DC.

20.1 Sauvagesia erecta L. (Figure 6B)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 12-V-2023; fl., fr., Saraiva, R. 372 (SLUI).

Species recognition: Palustrine herb. Stem erect, stipules with reddish hairs. Leaves simple, alternate; blades lanceolate, glabrous, the base and the apex acute, margin serrated. Inflorescence axillary. Flowers pentamerous, solitary, petals pink. Fruit capsule, ellipsoid, greenish.

Distribution and habitat: All Brazilian regions and states. In the study area, was in the study area, was collected in Swamp forest and Restinga swamps.

21. Onagraceae Juss.

21.1 Ludwigia affinis (DC.) H. Hara (Figure 6C)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 12-V-2023; fl., fr., Penha, J. 80 (SLUI).

Species recognition: Palustrine herb. Stem cylindrical, pubescent, trichome simple. Leaves simple, alternate; blades oblong, base attenuate, apex acuminate, margin entire. Inflorescence solitary, axillary. Flowers tetramerous, petals yellow. Fruit capsule, vinaceous.

Distribution and habitat: North (AC, AM, AP, PA, RO, TO), Northeast (BA, MA, RN, PB, PE), Central-West (GO, MT, MS), and Southeast (ES, MG, RJ). In the study area, was collected in Swamp forest.

21.2 Ludwigia hyssopifolia (G.Don) Exell

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 3-VII-2022; fl., fr., Saraiva, R. 373 (SLUI).

Species recognition: Emergent aquatic herb. Stem quadrangular, glabrous, vinaceous. Leaves simple, alternate; blades lanceolate, glabrous, base acuminate, apex acute, margin entire. Inflorescence solitary, axillary. Flowers tetramerous, petals yellow. Fruit capsule, elongated, brown.

Distribution and habitat: North (AC, AM, AP, PA, RO, RR), Northeast (AL, BA, MA, PB, PE, PI), Central-West (GO, MT), and Southeast (ES, MG, RJ). In the study area, was collected in Swamp forest.

22. Poaceae Barnhart

22.1 Ichnanthus hoffmannseggii (Roem. & Schult.) Döll

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 1-VII-2022; fl., fr., Saraiva, R. 345 (SLUI); ibid., Balneário Abrantes, 23-IX-2022; fl., fr., Penha, J. 46 (SLUI).

Species recognition: Palustrine aquatic herb. Stem erect. Leaves simple, sessile, alternate; blades lanceolate, glabrous, base rounded, apex acute, margin entire; ligule present. Inflorescence panicle, branches; spikelet lanceolate, densely pilose. Fruit caryopsis.

Distribution and habitat: North (PA, TO), Northeast (MA, PI, BA), and Central-West (GO). In the study area, was collected in Swamp forest.

22.2 Rugoloa pilosa (Sw.) Zuloaga (Figure 6D)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 12-V-2023; fl., fr., Saraiva, R. 374 (SLUI).

Species recognition: Emergent herb. Stem stoloniferous. Leaves simple, alternate; blades lanceolate, base subcordate, apex acute, margin entire; ligule absent. Inflorescence spike; branches unilateral, alternate, axils pilose. Fruit caryopsis.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Swamp forest and Restinga swamps.

23. Pontederiaceae Kunth

23.1 Pontederia diversifolia (Vahl) M.Pell. & C.N.Horn (Figure 6E)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 1-VII-2022; fl., Saraiva, R. 349 (SLUI).

Species recognition: Rooted-floating herb. Stem prostrate, glabrous. Leaves simple, alternate, distichous, floating; blades ovate, base sub-cordate, apex acute, margin entire; petiole cylindrical. Inflorescence spike. Flowers zygomorphic, tepals lilac, with a nectar guide in the middle of the upper lobe. Fruit capsule.

Distribution and habitat: North (AC, AM, AP, PA, RO, RR, TO), Northeast (AL, BA, CE, MA, PB, PE, PI, RN, SE), Central-West (GO, MT, MS), and Southeast (MG). In the study area, was collected in Swamp forest.

24. Plantaginaceae Juss.

24.1 Bacopa aquatica Aubl. (Figure 6F)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 1-VII-2022; fl., fr., Saraiva, R. 355 (SLUI); ibid., Povoado Carapirá, 24-IX-2022; Penha, J. 61 (SLUI).

Species recognition: Emergent herb. Stem ascending, cylindrical. Leaves simple, opposite, sessile; blades lanceolate, glabrous, base attenuate, apex acuminate, margin serrated. Flowers actinomorphic, pentamerous, axillary, solitary, sepals strongly unequal, petals white; bracteoles 2, opposite. Fruit capsule.

Distribution and habitat: North (AM, AP, PA), Northeast (BA, CE, MA, PE, PI, RN), Central-West (MT, MS), and Southeast (ES, RJ). In the study area, was collected in Swamp forest and Restinga swamps.

24.2 Bacopa aubletiana Scatigna (Figure 6G)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 1-VII-2022; fl., fr., Saraiva, R. 348 (SLUI); ibid., Povoado Carapirá, 24-IX-2022; fl., fr., Penha, J. 59 (SLUI).

Species recognition: Emergent herb. Stem erect. Leaves simple, opposite, sessile; blades ovate, margin serrated, base round, apex cuspidate. Flowers zygomorphic, pentamerous, axillary, with yellow tube, sepals subequal, petals blue. Fruit capsule.

Distribution and habitat: North (AP, PA, RR), Northeast (MA, PI, BA), Central-West (GO), and Southeast (MG). In the study area, was collected in Swamp forest and Restinga swamps.

24.3 Bacopa salzmannii (Benth.) Wettst. ex Edwall (Figure 6H)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 23-IX-2022; fl., fr., Penha, J. 52 (SLUI); ibid., Povoado Carapirá, 3-VII-2022; fl., fr., Saraiva, R. 342 (SLUI).

Species recognition: Emergent herb, pubescent, trichomes simple. Stem erect, cylindrical. Leaves simple, opposite, sessile; blades ovate, margin entire, base truncate, apex cuspidate. Flowers slightly zygomorphic, apparently tetramerous, sepals strongly unequal, petals white; bracteoles absent. Fruit capsule.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Restinga swamps.

24.4 Bacopa scoparioides (Cham. & Schltdl.) Scatigna (Figure 6I)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 12-V-2023; fl., fr., Penha, J. 69 (SLUI).

Species recognition: Emergent herb, aromatic, branched. Stem square, with longitudinal fissures. Leaves simple, sessile, opposite; blades elliptic, base and apex attenuated, margin serrate. Flowers zygomorphic, pentamerous, sepals subequal, petals blue. Fruit capsule.

Distribution and habitat: North (AC, AM, PA), Northeast (AL, BA, CE, MA), Central-West (DF, GO, MT), Southeast (ES, MG, SP), and South (PR). In the study area, was collected in Swamp forest.

25. Rubiaceae Juss.

25.1 Borreria remota (Lam.) Bacigalupo & E.L.Cabral

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 12-V-2023; fl., Penha, J. 82 (SLUI).

Species recognition: Palustrine herb. Stem erect, cylindrical. Leaves simple, opposite; blades elliptic, slightly pubescent, trichomes simple, base acute, apex acuminate, margin entire; stipules sheathed. Inflorescence terminal and axillary glomerulus. Flowers tetramerous, corolla white, hypanthium ciliate.

Distribution and habitat: North (AC, AM, PA), Northeast (BA, CE, MA, PE), Central-West (MT), and Southeast (ES, SP). In the study area, was collected in Swamp forest.

25.2 Oldenlandia lancifolia (Schumach.) DC. (Figure 6J)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Povoado Carapirá, 24-IX-2022; fl., fr., Penha, J. 57 (SLUI).

Species recognition: Palustrine herb. Stem erect. Leaves simple, sessile, opposite; blades lanceolate, glabrous, base attenuate, apex acute, margin entire. Inflorescence uniflorous. Flowers tetramerous, axillary, petals white. Fruit capsule.

Distribution and habitat: North (AC, AM, AP, PA, RO, RR, TO), Northeast (BA, MA, PE), Central-West (GO), Southeast (SP), and South (PR). In the study area, was collected in Swamp forest and Restinga swamps.

25.3 Pentodon pentandrus (Schumach. & Thonn.) Vatke (Figure 6K)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Povoado Carapirá, 24-IX-2022; fl., fr., Penha, J. 64 (SLUI).

Species recognition: Emergent herb. Stem prostrate. Leaves simple, sessile, opposite; blades linear, glabrous, base elongated, apex obtuse, margin entire. Inflorescence racemose. Flowers pentamerous, petals white. Fruit capsule.

Distribution and habitat: North (PA), Northeast (BA, MA, PB, PE), Central-West (GO, MT, MS), Southeast (ES, MG, SP, RJ), and South (PR, SC). Collected in Restinga swamps.

26. Taccaceae Dumort.

26.1 Tacca parkeri Seem.

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 12-V-2023; fl., fr., Penha, J. 66 (SLUI); ibid., Balneário Abrantes, 23-IX-2022; fl., fr., Saraiva, R. 396 (SLUI).

Species recognition: Palustrine herb. Stem rhizomatous, cylindrical. Leaves simple, alternate; blades entire, base attenuate, apex acuminate, margin entire. Inflorescence umbellate. Flowers trimerous, petals purple. Fruit ovoid, vinaceous.

Distribution and habitat: North (AM, AP, PA), Northeast (MA), and Central-West (MT). In the study area, was collected in Swamp forest.

27. Verbenaceae J.St.-Hil.

27.1 Stachytarpheta cayennensis (Rich.) Vahl

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 23-IX-2022; fl., Penha, J. 51 (SLUI).

Species recognition: Palustrine herb, pilose, trichomes strigose. Stem erect. Leaves simple, opposite; blades ovate, pilose, base attenuate, apex acute, margin crenate. Flowers solitary, terminal, corolla lilac.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Swamp forest.

28. Xyridaceae C.Agardh

28.1 Xyris jupicai Rich. (Figure 6L)

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Povoado Guajerutiua, 14-V-2023; fl., fr., Nunes, M. 55 (SLUI).

Species recognition: Emergent, cespitose herb, glabrous. Stem rhizomatous. Leaves simple, sessile, alternate distichous; blades linear, base flattened, apex obtuse, margin entire. Inflorescence spike, ovoid; bracts brown. Flowers yellow. Fruit capsule; seeds obovoid to broad-ellipsoid, translucent.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Swamp forest and Restinga swamps.

28.2 Xyris macrocephala Vahl

Specimens examined: BRAZIL. MARANHÃO: APA das Reentrâncias Maranhenses, Guimarães, Balneário Passaginha, 1-VII-2022; fl., fr., Saraiva, R. 344 (SLUI).

Species recognition: Palustrine, cespitose herb, glabrous. Stem erect. Leaves simple, sessile, alternate distichous, glabrous; blades linear, base flattened, apex acute, margin entire. Inflorescence spike, ovoid; bracts oval. Flowers with yellow petals. Fruit capsule, obovoid; seeds ellipsoid to fusiform, dark brown.

Distribution and habitat: All Brazilian regions and states. In the study area, was collected in Swamp forest.

4. Discussion

Our floristic survey revealed a considerable richness of APs in the swamp formations of the municipality of Guimarães, covering wetlands that had not been explored before. In addition, Voyria aphylla is a new species record for the flora of Maranhão according to the Flora e Funga do Brasil (JBRJ, 2024b) and SpeciesLink (CRIA, 2024) databases.

The species Voyria aphylla was found growing in an environment with decomposing leaves and clay soil, in contrast to the study by Gomes et al. (2022) mentioning this species preference for drier environments. This new record highlights the relevance of more comprehensive botanical studies in aquatic environments for better knowledge of the flora of Maranhão (Pestana et al., 2024a). With increased sampling, new species records may be found as indicated in recent studies (Ferreira et al., 2019; Salazar-Ferreira et al., 2020; Pestana et al., 2024b).

In terms of species recorded, this study can be considered significant for surveys that emphasize APs (Matias et al., 2003; Alves et al., 2011; Lima et al., 2011; Araújo et al., 2012). Given the nature of the study area, which covers a range of aquatic environments with similar characteristics, these results also corroborate the species richness and methodology used in other floristic surveys in Maranhão. For instance, Silva and Fontes (2018) recorded 16 species in northeastern Maranhão, Arouche et al. (2021) recorded 66 taxa in an herbarium-based survey, and Pestana et al. (2024a) cataloged 72 species in wetlands in the eastern of the state. However, the difference in species richness can be attributed to varying sampling efforts. Silva and Fontes (2018), sampled a more restricted area with less collection effort compared to the other studies cited above and the present survey, which led to the recording of fewer species. Although these previous studies have also provided extremely valuable data for knowledge of aquatic vegetation in the state of Maranhão, we were the first to include descriptions of all the species cataloged and information on their distribution in Brazil.

Melastomataceae, Fabaceae, Plantaginaceae, Lentibulariaceae, Cyperaceae, and Rubiaceae stood out in terms of species richness, highlighting their importance in the floristic composition and structure of the restinga in the municipality of Guimarães. These results partially align with other surveys APs in restinga areas (Matias et al., 2003; Alves et al., 2011; Valadares et al., 2011), with minor differences in the representative families observed in this study compared to those in the previously mentioned surveys (see Table 2 for further details). Particularly Melastomataceae, the family with the highest number of species recorded, is characterized by colonizing environments with nutrient-poor soils, typical of swamp environments (Albuquerque et al., 2013; Araújo and Lima, 2013). Some of the genera of this family, e.g., Miconia Ruiz & Pav., have an important ecological role as most species provide essential food resources for pollinators and dispersers (Albuquerque et al., 2013).

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Table 2
Comparison of the most representative families in floristic surveys conducted in restinga areas.

Fabaceae was the second most representative family in this study, corroborating surveys conducted in the Northeast Brazil (Castro et al., 2012; Gomes and Guedes, 2014; Santos-Filho et al., 2015) and Southeast (Amaral et al., 2008). Although Cyperaceae contributed to only 6% of the species cataloged in our survey, it is among the most frequent families found in studies on APs (Matias et al., 2003; Silva and Fontes, 2018; Pestana et al., 2024a). According to Matias et al. (2003), this representativeness is related to the complex underground system (rhizomes and stolons) of Cyperaceae species favorable for effective vegetative reproduction.

Rubiaceae is the fourth-largest family of angiosperms and has a cosmopolitan distribution, being most representative in tropical regions (Taylor et al., 2004). It is generally found among the most cited families in restinga areas of the Northeast of Brazil (Cabral-Freire and Monteiro, 1993; Silva et al., 2008; Almeida Júnior et al., 2016; Guterres et al., 2020). The families Plantaginaceae and Lentibulariaceae have adaptations to survive in moist soil conditions and are commonly found in formations of flooded restinga. However, they have low species richness in floristic inventories of restinga in Maranhão and others states of Northeast Brazil (Santos-Filho et al., 2015; Amorim et al., 2016; Rodrigues et al., 2019; Guterres et al., 2019; Araujo et al., 2020), as it only covers the flooded restinga. The species of Lentibulariaceae represented in the surveys, mainly by the genus Utricularia, have adaptations that allow the capture and digestion of small organisms and favor development in habitats with low nutrient content (Taylor, 1989; Guisande et al., 2007), such as restinga soils.

The taxonomically complex genus Bacopa (Plantaginaceae) stood out in terms of species number. Species belonging to this genus perform important ecological functions in the aquatic environment, such as water purification and nutrient cycling (Araújo et al., 2020). Bacopa exhibits greater species diversity in tropical and subtropical areas, showing wide morphological variation (Scatigna et al., 2022; Pestana et al., 2023). Another relevant and common genus in studies involving APs also identified here was Ludwigia (Onagraceae). Most taxa of the Onagraceae family are drought-tolerant, which significantly expands its distribution range to flooded areas (Amaral et al., 2008). Furthermore, Ludwigia encompasses species with considerable phenotypic plasticity, resulting in a high capacity for these species to explore both aquatic and terrestrial environments (Pott and Pott, 2000).

Concerning life forms, the prevalence of palustrine (58%) and emergent species (30%) can be directly linked to habitats prone to seasonal inundations, shallow areas, and the ability of such species to cope with lower water levels (Neves et al., 2006). Studies on APs emphasize palustrine and emergent life forms over others (e.g., Alves et al., 2011; Lima et al., 2011), especially in Northeast Brazil (Moura-Júnior and Cotarelli, 2019).

The presence of submersed species Cabomba aquatica (Cabombaceae) and Tonina fluviatilis (Eriocaulaceae) in this study may be attributed to water transparency and oxygenation, parameters that favor fixed submersed species to grow more rapidly (Henry-Silva and Camargo, 2006; Sobral-Leite et al., 2010). In locations with lower nutrient concentrations and highly transparent water, C. aquatica can establish itself. This contrasts with the findings of Silva and Fontes (2018) reporting that C. aquatica was more prevalent in environments characterized by reduced transparency and low oxygen concentration, such as those observed at Passaginha Balneary. However, climatic conditions also play a significant role in shaping these ecological groups, especially considering that the coast of Maranhão is subjected to specific patterns of winter, temperature, and humidity typical of the equatorial coast (IMESC, 2011). These climatic variations are recognized as factors influencing the floristic and structural differences observed in various tropical ecosystems (Oliveira-Filho and Fontes, 2000).

The distribution and categorization of life forms of aquatic plants contribute to our understanding of the restinga in the municipality of Guimarães. Nevertheless, investigations focusing on temporal, ecological, and morphological aspects are important for comprehending the dynamics of this region and for local and national policies to ensure biodiversity conservation in this ecosystem.

Acknowledgements

We thank the Conselho Nacional de Desenvolvimento Científico e Tecnológico - CNPq (process 402943/2021-0) “Beyond the Flora of Brazil 2020: boosting the flora of Maranhão” and CNPq (process: 404619/2023-1, “Floristic diversity in areas with a lack of botanical information for Maranhão” for financing this study and the Sistema de Autorização e Informação em Biodiversidade (SISBIO) for the license No. 84049-2 granted for the collection and transport of the botanical material. We also thank Maria Nélia Santos Neto for their reception during the collections, and the members of the GPBio (Mayara Nunes, Krislaynne Braga, Mayara Campos, Josélia Dourado, Adelson Viegas, Hely Aroucha and Lorena Araújo) for all their support in the collection and herborization processes. We would like to thank UFMA for the logistics, space, and equipment used for the morphological analysis; and Dr. Francisca Helena Muniz, curator of the SLUI herbarium at UEMA, for the use of the herbarium. Finally, we would also like to thank the reviewers for their corrections and suggestions in the manuscript.

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Publication Dates

Publication in this collection
10 Jan 2025
Date of issue
2024

History

Received
18 July 2024
Accepted
18 Oct 2024

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Families/Species	Vernacular name (in portuguese)	Biological forms	Vouchers (SLUI)
ALISMATACEAE
Sagittaria guayanensis Kunth	lagartixa	RF	8725, 8715
ACANTHACEAE
Lepidagathis alopecuroidea (Vahl) R.Br. ex Griseb.	-	PL	8731
ARACEAE
Dieffenbachia aglaonematifolia Engl.	aninga	EM	8785
ASTERACEAE	-
Sphagneticola trilobata (L.) Pruski	margaridão	EM	8773
BURMANIACEAE
Apteria aphylla (Nutt.) Barnhart ex Small	-	PL	8774
CABOMBACEAE
Cabomba aquatica Aubl.	lodo	RS	8713
COSTACEAE
Costus spiralis (Jacq.) Roscoe	-	PL	8788
CYPERACEAE
Cyperus haspan L.	-	PL	8764
Scleria microcarpa Nees ex Kunth	-	PL	8757
Rhynchospora riparia (Nees) Boeckeler	tiririca	EM	8748
ERIOCAULACEAE
Tonina fluviatilis Aubl.	-	RS	8751, 8758
FABACEAE
Aeschynomene evenia C.Wright & Sauvalle	-	EM	8775
Chamaecrista diphylla (L.) Greene	-	PL	8789
Desmodium adscendens (Sw.) DC.	carrapicho-beiço-de-boi	PL	8718
Mimosa pudica L.	dormideira	PL	8790
GENTIANACEAE
Chelonanthus viridiflorus (Mart.) Gilg	-	PL	8765, 8752
Voyria aphylla (Jacq.) Pers.*	-	PL	8784
LAMIACEAE
Hyptis atrorubens Poit.	hortelã-brava	PL	8776
LENTIBULARIACEAE
Utricularia pusilla Vahl	lodo	PL	8726
Utricularia subulata L.	lodo	PL	8716
Utricularia gibba L.	boca-de-leão-do-banhado	EM	8791
LINDERNIACEAE
Lindernia rotundifolia (L.) Alston	-	EM	8772
LYGODIACEAE	-
Lygodium volubile Sw.	-	PL	8760
MALVACEAE	-
Sida linifolia Cav.	malva	PL	8759
MENYANTHACEAE	-
Nymphoides humboldtiana (Kunth) Kuntze	estrela-branca	RF	8756
MELASTOMATACEAE
Aciotis annua (Mart. ex DC.) Triana	-	PL	8754
Comolia villosa (Aubl.) Triana	-	PL	8724
Clidemia biserrata DC.	-	PL	8777
Clidemia hirta (L.) D.Don	-	PL	8792
Miconia ciliata (Rich.) DC.	-	PL	8723
Nepsera aquatica (Aubl.) Naudin	-	EM	8771, 8732
Pterolepis glomerata (Rottb.) Miq.	-	EM	8766, 8730
Pterolepis trichotoma (Rottb.) Cogn.	-	PL	8778
Tococa guianensis Aubl.	-	PL	8749
NYMPHAEACEAE
Nymphaea rudgeana G.Mey.	ninféia	RF	8779, 8800
OCHNACEAE
Sauvagesia erecta L.	-	PL	8780
ONAGRACEAE
Ludwigia affinis (DC.) H.Hara	cruz-de-malta	PL	8793
Ludwigia hyssopifolia (G.Don) Exell	cruz-de-malta	EM	8781
POACEAE
Ichnanthus hoffmannseggii (Roem. & Schult.) Döll	-	PL	8722, 8711
Rugoloa pilosa (Sw.) Zuloaga	-	EM	8782
PONTEDERIACEAE	-
Pontederia diversifolia (Vahl) M.Pell. & C.N.Horn	camalotinho	RF	8729
PLANTAGINACEAE
Bacopa aquatica Aubl.	-	EM	8750, 8761
Bacopa aubletiana Scatigna	-	EM	8728, 8753
Bacopa salzmannii (Benth.) Wettst. ex Edwall	agriãozinho	EM	8720, 8717
Bacopa scoparioides (Cham. & Schltdl.) Scatigna	pataqueira	EM	8786
RUBIACEAE
Borreria remota (Lam.) Bacigalupo & E.L.Cabral	vassourinha	PL	8712
Oldenlandia lancifolia (Schumach.) DC.	-	PL	8747
Pentodon pentandrus (Schumach. & Thonn.) Vatke	-	EM	8767
TACCACEAE
Tacca parkeri Seem.	-	PL	8783, 8794
VERBENACEAE
Stachytarpheta cayennensis (Rich.) Vahl	gervão-azul	PL	8719
XYRIDACEAE
Xyris jupicai Rich.	botão-de-ouro	EM	8734
Xyris macrocephala Vahl	campim-estrela	PL	8721

Reference	Location	Most representative families
Present study	Reentrâncias Maranhenses EPA (Guimarães, MA)	Melastomataceae (9 spp.), Fabaceae (5 spp.), Plantaginaceae (3 spp.), Lentibulariaceae, Cyperaceae, Rubiaceae (3 spp. each)
Alves et al. (2011)	Entorno Costeiro EPA (Florianópolis, SC)	Cyperaceae (8 spp.), Polygonaceae (5 spp.), Poaceae (3 spp.), Plantaginaceae (3 spp.), Asteraceae (3 spp.), Onagraceae (3 spp.)
Matias et al. (2003)	Lagoa de Jijoca (Jericoacoara, CE)	Cyperaceae (19 spp.), Fabaceae (4 spp.), Poaceae (4 spp.), Lentibulariaceae (2 spp), Malvaceae (2 spp.), Nymphaeaceae (2 spp.)
Valadares et al. (2011)	Restinga de Morada do Sol (Vila Velha, ES)	Asteraceae (13 spp.), Cyperaceae (12 spp.), Fabaceae (9 spp.), Rubiaceae (8 spp.), Poaceae (7 spp.), Melastomataceae (4 spp.)

Aquatic and palustrine plants of restinga in a Ramsar site in the easternmost Amazon, Brazil

Plantas aquáticas e palustres de restinga em um sítio Ramsar no extremo leste da Amazônia, Brasil

Abstract

Resumo

1. Introduction

2. Material and Methods

2.1. Study area

2.2. Data collection

3. Results

3.1. Enumerated list of species

4. Discussion

Acknowledgements

References

Publication Dates

History