Taxonomic synopsis of Plantaginaceae and Scrophulariaceae (Lamiales) in Iguazú and Iguaçu National Parks (Argentina and Brazil)

Abstract

The Iguazú region, where the Iguaçu National Park (Brazil) and the Iguazú National Park (Argentina) are located, encompasses one of the largest continuous remnants of the Atlantic Forest. It is considered a hotspot of biodiversity and an example of a region that is poorly known botanically. Even though non-tree taxa represent most of the Atlantic Forest’s plant diversity, some botanical families that are mainly herbaceous or shrubby, such as Plantaginaceae and Scrophulariaceae, are underrepresented in floristic treatments within this domain. Here, we provide a floristic account for the Plantaginaceae and Scrophulariaceae from the Iguazú and Iguaçu National Parks, including identification keys, illustrations, comments on distribution, habitat, and phenology. A total of 13 species and seven genera were documented. Of these, Stemodia is represented by three species, Bacopa, Buddleja, Mecardonia, and Scoparia are each represented by two species, and two genera, namely Plantago and Veronica, are represented by a single species. Our findings expand the knowledge about the flora of the Southern Cone, contributing important information for the establishment of biodiversity management plans and conservation priorities.

Key words:
Atlantic Forest; floristics; Neotropical Flora; Parana Basin; Selva Paranaense

Resumo

A região de Iguazú, onde se encontram o Parque Nacional do Iguaçu (Brasil) e o Parque Nacional Iguazú (Argentina), abrange um dos maiores remanescentes contínuos da Mata Atlântica. É considerada um ponto de grande biodiversidade e um exemplo de uma região botanicamente pouco conhecida. Mesmo que os taxa não relacionados a árvores representem a maior parte da diversidade de plantas da Mata Atlântica, algumas famílias botânicas que são principalmente herbáceas ou arbustivas, como Plantaginaceae e Scrophulariaceae, estão sub-representadas nos tratamentos florísticos dentro deste domínio. Aqui, fornecemos um relato florístico das Plantaginaceae e Scrophulariaceae dos Parques Nacionais Iguazú e Iguaçu, incluindo chaves de identificação, ilustrações, comentários sobre distribuição, habitat e fenologia. Um total de 13 espécies e sete gêneros foram documentados. Destas, Stemodia é representada por três espécies, Bacopa, Buddleja, Mecardonia e Scoparia são representadas por duas espécies cada, e dois gêneros, ou seja, Plantago e Veronica, são representados por uma única espécie. Nossas descobertas ampliam o conhecimento sobre a flora do Cone Sul, contribuindo com informações importantes para o estabelecimento de planos de gestão da biodiversidade e prioridades de conservação.

Palavras-chave:
Mata Atlântica; florística; Flora Neotropical; Bacia do Paraná; Selva Paranaense

Introduction

The family Plantaginaceae (Lamiales) has a cosmopolitan distribution, including about 90 genera and 1,900 species, of which 45 genera and 370-400 species are native to the Neotropical region (Stevens 2023). In Brazil, this family is represented by 28 genera and 150 species, 68 of which are endemic (Souza et al. 2023). In the state of Paraná, 16 genera and 33 species have been recorded (Kaehler 2014). In Argentina, the family is represented by 26 genera and 126 species, of which 23 occur in the Misiones province (O’Leary 2018a; Zanotti et al. 2020). This family is well known for its large number of ornamental species, e.g., snapdragon - Antirrhinum, brazilian snapdragon - Matourea azureaLinden (1862: 6) Colleta & Souza In Coletta et al. (2020: 1356), and the firecracker plant - Russelia equisetiformis Schlechtendal & Chamisso In Garcke et al. (1831: 377). It is also represented by medicinal species such as those of the genus Plantago, used to treat intestinal pain, urinary infections, and tumors (Samuelsen 2000; Judd et al. 2009).

The Scrophulariaceae family also has a cosmopolitan distribution, including 59 genera and 1,880 species distributed worldwide, but with only seven genera and 22 species in Brazil, of which two genera and ten species occur in Paraná (Stevens 2023; Souza et al. 2020). Twenty-two Scrophulariaceae species are found in Argentina, six of which are in the Misiones province (O’Leary 2018b).

These families share many morphological features, e.g., the usually herbaceous habit, absence of stipules, and usually zygomorphic corollas with fused petals, which also represent most of the order Lamiales, leading to uncertainties in familial boundaries, especially between those families segregated from the traditional Scrophulariaceae (prior to phylogenetic studies), e.g., Linderniaceae, Musaceae, Orobanchaceae, and Plantaginaceae (Olmstead & Reeves 1995; Olmstead et al. 2001; Albach et al. 2005; Souza & Lorenzi 2019).

However, even though floristic accounts on a regional scale are essential to progress biodiversity conservation (Giulietti et al. 2005), some important remnants of the Upper Paraná Atlantic Forest (e.g., the Iguaçu and Iguazú National Parks) are still understudied, especially regarding non-tree groups such as Plantaginaceae and Scrophulariaceae.

With that in view, many surveys focusing on floristic studies for the Iguazú and Iguaçu National Parks are currently being conducted, but with only a few published, e.g., a flora inventory for Acanthaceae (Hammes et al. 2021), a synopsis of Bignoniaceae (Hentz Júnior et al. 2022), Leguminosae (Rauber et al. 2021a) and Rubiaceae (Rauber et al. 2021b) a floristic survey focusing on epiphytes (Cervi & Borgo 2007), and a floristic survey encompassing all angiosperms (Trochez et al. 2017) have been published.

Therefore, this study provides a taxonomic synopsis for Plantaginaceae and Scrophulariaceae species in the Iguazú and Iguaçu National Parks. We present identification keys for the species of each family, illustrations of collected species, and comments on their distribution, habitat, conservation, phenology, and taxonomy notes, aiming to increase the knowledge about herbaceous and shrub species of the Upper Paraná Atlantic Forest (Myers et al. 2000).

Material and Methods

Study area

The Iguaçu National Park (ParNa Iguaçu) is a Conservation Unit located within the state of Paraná, Brazil, between 25º05’-25º41’S and 53º40’-54º38’W. The park has an area of 1,860 km² and a perimeter of 420 km, of which 300 km include pristine vegetation. The ParNa Iguaçu includes Seasonal Semideciduous Forest (SSF) and Mixed Ombrophilous Forest (MOF), representing the largest Atlantic Forest remnant in Brazil (IBAMA 1999).

The Iguazú National Park (ParNa Iguazú) is located in the northern portion of the province of Misiones, Argentina, between 25°31’-25°43’S and 54°08’-54°32’W, with 543 km² (SIB 2020). The area is composed of Seasonal Semideciduous Forest vegetation known as “Selva Paranaense” or Interior Atlantic Forest (Di Bitetti et al. 2003). Misiones is considered the most well-preserved province of Argentina, with 40% of its original forest cover (SIB 2020).

For this study, we focused our sampling on five main areas, according to Hentz Júnior et al. (2022), listed as follows: (i) Céu Azul, a transition area between SSF and MOF areas; (ii) Matelândia/Serranópolis do Iguaçu, where collections were mainly carried out in forest borders and in the “Estrada do Colono” stretch; (iii) Capanema, with alluvial SSF; (iv) Foz do Iguaçu, with submontane SSF; and the (v) Iguazú National Park (Argentina) (Fig. 1).

Collection and identification

Monthly visits to the Iguaçu National Park were carried out between February 2019 and February 2020, and Plantaginaceae and Scrophulariaceae specimens were collected and incorporated into the EVB herbarium, following Bridson & Forman (2004). Also, we analyzed herbarium specimens deposited at the CTES, EVB, HCF, MBM, PUC-PR, UNOP, and UPCB herbaria (Thiers B, continuously updated). Specimen data were also analyzed through virtual herbaria (Virtual Reflora Herbarium (<http://reflora.jbrj.gov.br/reflora/herbarioVirtual/>), Flora e Funga do Brasil (<http://floradobrasil.jbrj.gov.br/>), and SpeciesLink (<https://specieslink.net/search/>)). The following literature was used for precise species identification: Souza & Giulietti (2009), Hefler et al. (2011), Scatigna & Mota (2017), Coelho & Miotto (2018) and Lannoy et al. (2018).

Lastly, to determine the conservation status, we checked if the species were already classified on specific literature, CNCFlora (2023), and official governmental lists of threatened species. Then, we assessed the conservation status on a regional level (country) for the species classified as “Not Evaluated” (NE). We used the GeoCat Tool (Bachman et al. 2011) by plotting the species distribution data in Brazil. The analysis was based on the original coordinates from the SpeciesLink database, evaluating the extent of occurrence (EOO) following category B of the IUCN criteria for regional levels (Gärdenfors et al. 2001; IUCN 2001).

Results

In this study, we documented 13 Plantaginaceae species and two Scrophulariaceae species for the Iguaçu and Iguazú National Parks. For Plantaginaceae, Bacopa Aubl., Stemodia L., Mecardonia Ruiz & Pav, and Scoparia L. had two species each, and Plantago L. and Veronica L. were represented by only one species. Scrophulariaceae was represented only by the genus Buddleja L., with two species (Figs. 2-5).

Figure 1
Areas of the Iguaçu and Iguazu National Parks - Area 1 = Céu Azul; Area 2 = Matelândia/Serranópolis; Area 3 = Capanema; Area 4 = Foz do Iguaçu; Area 5 = Iguazu National Park. Source: Modified from Hammes et al. 2021.

Key to the Plantaginaceae from the Iguaçu and Iguazú National Parks

1. Aerial stem absent; leaves rosulate; flowers not showy, disposed in spikes 3.1. Plantago australis

1’. Aerial stem present; leaves opposite; flowers showy, usually axillary, not disposed in spikes 2

2. Sepals strongly unequal 3

3. Corolla yellow; thecae separated by arms of the connective 4

4. Branches erect; leaves with round base; calyx longer than 1.1 cm 2.1. Mecardonia grandiflora

4’. Branches erect or creeping; leaves with acute base; calyx 0.3-0.8 cm long 2.2. Mecardonia procumbens var. flagellaris

3’. Corolla white to purplish 5

5. Bracteoles absent; pedicel 1.5-3.7 cm long; leaf margin entire; external sepal base cordate 1.1. Bacopa australis

5’. Bracteoles present; pedicel 0.4-0.8 cm long; leaf margin serrate; external sepal base truncate 1.2. Bacopa scabra var. scabra

2’. Sepals equal to subequal 6

6. Corolla rotaceous; thecae not separated by arms of the connective 7

7. Sepals 4; corolla white 4.1. Scoparia dulcis

7’. Sepals 5; corolla yellow 4.2. Scoparia montevidensis

6’. Corolla zygomorphic; thecae separated by arms of the connective 8

8. Herbs prostrate; cauline leaves petiolate 9

9. Stamens 4; bracts petiolate; corolla lilac 5.3. Stemodia verticillata

9’. Stamens 2; bracts sessile; corolla blue 6.1. Veronica arvensis

8’. Herbs erect; cauline leaves sessile 10

10. Leaves subcoriaceous; inflorescences dense, terminal 5.2. Stemodia stricta

10’. Leaves membranaceous; inflorescences lax, axillary 5.1. Stemodia diplohyptoides

1. Bacopa Aubl., Hist. Pl. Guiane 128. 1775.

1.1. Bacopa australis V.C.Souza, Acta Bot. Brasil. 15(1): 58 (57-61; fig. 1). 2001.Fig. 2a-b

Bacopa australis is characterized by entire leaves with a rounded base. The species is morphologically similar to Bacopa repens (Swartz 1788: 14) Wettstein (1895: 76), from which it is distinguished by the stamens 4 (vs. stamens 2 in B. repens). The only specimen collected in the study area was previously identified as Bacopa salzmannii (Bentham 1836: 58) Wettstein ex Edwall (1897: 181), from which it is distinguished by the adpressed trichomes and bifid style (vs. erect trichomes and entire style in B. salzmannii) (Souza & Giulietti 2009).

Examined materials: Capanema, Parque Nacional do Iguaçu, 23.XI.1966, J.C. Lindemann & J.H. de Hass 3358 (K[digital image]!; MBM!; NY[digital image]!; UB[digital image]!; US[digital image]!).

Additional specimen examined: BRAZIL. MATO GROSSO DO SUL: Corumbá, Baía do Búfalo, próximo ao Capão de Ingá area, 3.IV.1990, V.J. Pott 1321(MBM!).

Bacopa australis is native from South America, occurring in the Central-West and South regions of Brazil, and in the Corrientes and Misiones provinces of Argentina, mainly in flooded areas (Souza & Giulietti 2009; Sosa et al. 2018). In the ParNa Iguaçu, it was found in Capanema, in a pool by the old “Estrada do Colono” stretch.

This species was found with flowers in November.

1.2. Bacopa scabra var. scabra (Benth.) Descole & Borsini, Genera et Species Plantarum Argentinarum 5: 137. 1954.

Bacopa scabra is characterized by the leaves slightly serrate (vs. entire in B. australis), and the puberulous calyx, with truncate base (vs. cordate base in B. australis).

Examined materials: ARGENTINA. MISIONES: Iguazú, Denis (LIL 70166).

Additional examined materials: BRAZIL. MATO GROSSO DO SUL: Corumbá, margem direita do Rio Miranda, 26.VI.1983, C.A. Conceição 1421 (RB!). SÃO PAULO: São Paulo, próximo a Mogi-Mirim, 7.IX.1956, A.S. Grotta (RB!).

In Argentina it occurs in the Chaco, Corrientes and Misiones provinces (Flora Argentina 2018). In Brazil it is widely distributed in all regions [Flora e Funga do Brasil (continuously updated)].

Figure 2
a-b. Bacopa australis - a. flowering branch; b. calyx with exerted, bifid stigma. c-d. Buddleja elegans subs. elegans - c. flowering branch; d. inflorescence. e-f. Buddleja stachyoides - e. flowering branch; f. inflorescence. g. Plantago australis - habit. h. Mecardonia grandiflora - flowering branch. i. Mecardonia procumbens subs. flagellaris - fruiting branch.

This species was registered with flowers in April (Sosa et al. 2018).

2. MecardoniaRuiz & Pav., Fl. Peruv. Prodr. 95. 1794.

2.1. Mecardonia grandiflora (Benth.) Pennell, Proc. Acad. Nat. Sci. Philadelphia 98: 87. 1946.Fig. 2h

Mecardonia grandiflora presents erect stems, leaves with serrated margin, and calyx longer than 1.1 cm. Differs from Mecardonia procumbens by being only slightly branched and by the larger leaves (> 30 cm long) and calyx (> 12 cm long).

Examined materials: ARGENTINA. MISIONES: Puerto Iguazú, Parque Nacional Iguazú, Sendero Macuco, 12.XII.2014, R. Vanni 4621 (CTES!); Zuloaga et al. 5228 (CTES). BRAZIL. PARANÁ: Capanema, Parque Nacional do Iguaçu, margem do Rio Iguaçu, 30.X.2015, M.G. Caxambu 7104 (HCF!). Foz do Iguaçu, Cataratas do Iguaçu, 2.V.1957, G. Hatschbach, 3916 (MBM!, UPCB!); trilha da desembocadura do Rio São João no Iguaçu, 22.V.2015, M.G. Caxambu 6405 (HCF!, MBM!).

Mecardonia grandiflora occurs in rocky areas near rivers in the South region of Brazil, and in Corrientes and Misiones provinces of Argentina. Found in Foz do Iguaçu and Capanema on rocky borders of the Iguazú and São João Rivers. Also, this species is listed as endemic to Brazil [Souza VC et al. in Flora e Funga do Brasil (continuously updated)] and endemic to Argentina (Flora Argentina 2018).

This species was found with flowers in May and October.

2.2. Mecardonia procumbens var. flagellaris (Cham. & Schltdl.) V.C.Souza, Acta Bot. Brasil. 11(2): 186. 1997. Figs. 2i; 4a-b

Mecardonia procumbens var. flagellaris presents crawling stems, leaves with margins entire or serrated, and calyx with 0.3-0.8 cm long. Differs from M. grandiflora by being significantly branched, with smaller leaves (< 20 cm long) and calyx (< 10 cm long). The subspecies M. procumbens var. flagellaris can be identified by the leaves longer than wide and with margins entire to slightly dentate.

Examined materials: ARGENTINA. MISIONES: Puerto Iguazú, Parque Nacional Iguazú, 7.VIII.1991, R. Vanni 2728 (CTES!). BRAZIL. PARANÁ: Foz do Iguaçu, Parque Nacional do Iguaçu, próximo ao porto, 7.V.1949, A.P. Duarte 1635 (NY[digital image]!, RB!, U[digital image]!). Macuco, G. Hatschbach 49792 (MBM!); G. Hatschbach 11207 (MBM!); 25°38’54.0”S, 54°27’27.0”W, 2019, E.J. Hentz Júnior 74 (EVB!); casa de hóspedes, 12.XII.1999, A.C. Cervi 6943 (UPCB!); trilha do Rio São João, 25°37’37.8”S, 54°28’52.6”W, 22.V.2015, M.G. Caxambu 6412, (HCF!, MBM!); 15.XI.2018, E.L. Siqueira 2841 (HCF!).

In Brazil, M. procumbens subs. flagellaris occurs in the North, Central-West, Southeast, and South regions (Souza et al. 2020). In Argentina, it occurs in Buenos Aires, Chaco, Chubut, Córdoba, Corrientes, Entre Ríos, Formosa, La Pampa, Misiones, Neuquén, Río Negro, Santa Fe, and San Luis (Flora Argentina 2018). In the study area, it was found in humid and disturbed areas of Foz do Iguaçu and in the Iguazú National Park.

This species was found flowering in November.

3. Plantago L., Sp. Pl. 1: 112. 1753.

3.1. Plantago australis Lam., Tabl. Encycl. i. 339, 1792.Fig. 2g

Plantago australis shows leaves arranged in basal rosettes, spike inflorescences, and it can be distinguished from other Plantago species by the leaves with a scarce, ciliated pubescence, and scape with adpressed antrorse trichomes (Hefler et al. 2011). We further identified the specimens from the Park as Plantago australis subs. australis.

Examined materials: Céu Azul, Parque Nacional do Iguaçu, borda do Parque, 24.V.2018, C.R. Rauber 56 (UNOP!). Foz do Iguaçu, trilha do Rio São João, 9.XI.2000, S.M.Hefler 54 (UPCB!); trilha das cataratas, 9.XI.2011, S.M. Hefler 35 (UPCB!); trilha em frente ao museu, 11.XI.2011, S.M. Hefler 34, (SMBD [digital image]!, UPCB!); borda do Parque, 25°39’24.5”S, 54°26’14.9”W, 4.VI.2015, M.G. Caxambu 6485 (HCF!, MBM!); trilha das Bananeiras, 15.VII.2019, E.J. Hentz Júnior 101 (EVB!).

Plantago australis is widely distributed throughout Latin America (Tolaba et al. 1998). In Brazil, it is concentrated in the South and Southeast regions (Hefler et al. 2011). In Argentina, it occurs in all provinces. In the Park, it was found in Céu Azul and Foz do Iguaçu. Probably occurs in all areas as the most common species, found both in disturbed areas, e.g., the borders of BR-469 highway, and in conserved trails.

This species usually flowers from September to February (Hefler et al. 2011). In the Park, it was found flowering in May.

4. Scoparia L., Sp. Pl. 1: 116. 1753.

4.1. Scoparia dulcis L., Sp. Pl. 1: 116. 1753. Figs. 3a-b; 4c-d

Scoparia dulcis can be easily recognized by the calyx with four sepals and the white corolla (vs. five sepals and yellow corolla in S. montevidensis). It can also present a subshrub habit, with rarely opposite 3-4-verticillate leaves (vs. only herbaceous habit and opposite to 3-verticillate leaves in S. montevidensis).

Examined materials: Foz do Iguaçu, Parque Nacional do Iguaçu, 19.III.1963, G. Hatschbach 9922 (MBM!; UPCB!).

Additional examined materials: BRAZIL. PARANÁ: Maringá, Parque do Ingá, 23°25’46.0”S, 51°55’42.0”W, 14.II.2007, M.C. Souza 1804 (HUEM).

Scoparia dulcis has a Pantropical distribution, occurring in all Brazilian states, usually in disturbed forest areas or in urban centers (Souza & Giulietti 2009). In Argentina, it occurs in Chaco, Corrientes, Entre Ríos, Formosa, Jujuy, Misiones, Salta, Santiago del Estero, Santa Fe, and Tucumán (Flora Argentina 2018). In the Park, it was found in Foz do Iguaçu.

This species was found with flowers in March.

4.2. Scoparia montevidensis R.E.Fr., Ark. Bot. 6, no. 9: 22, pl. 4, 8. 1907. Figs. 2c-d; 4e-d

Scoparia montevidensis can be easily recognized by the calyx with five sepals and the yellow corolla (vs. four sepals and white corolla in S. dulcis). It shows only an herbaceous habit and opposite to 3-verticillate leaves (vs. occasionally subshrub habit, with 3-4-verticilate leaves, rarely opposite in S. dulcis).

Examined materials: Capanema, Parque Nacional do Iguaçu, Rio Iguaçu, Ilha do Sol, 25°36’12.5”S, 53°48’35.4”W, 30.X.2015, M.G. Caxambu 7101 (HCF!). Foz do Iguaçu, trilha para o Rio Iguaçu, 17.XII.1992, A.C. Cervi 3931 (MBM!, NY[digital image]!, UPCB!); trilha da Usina, 12.XII.1999, A.C. Cervi 6957, (UPCB!); trilha do Rio São João, 25°37’38.1”S, 54°28’53.1”W, 22.V.2015, M.G. Caxambu 6422 (HCF!, UNOP!); final da trilha do Macuco, 22.III.2019, 25°39’01.0”S, 54°27’22.2”W, M.G. Caxambu 8895 (HCF!).

Scoparia montevidensis occurs from México to Argentina. In Brazil, it occurs in the North, Central-West, and South regions (Souza & Giulietti 2009). In Argentina, it occurs in Buenos Aires, Catamarca, Chaco, Córdoba, Corrientes, Distrito Federal, Entre Ríos, Formosa, Jujuy, La Pampa, La Rioja, Misiones, Salta, Santiago del Estero, Santa Fe, San Luis, Tucumán (Flora Argentina 2018). Found in Capanema, and Foz do Iguaçu, usually in open areas with high solar incidence.

This species was found with flowers in March, May, October, and November.

5. Stemodia L., Syst. Nat., ed. 10. 2: 1091, 1118. 1759.

5.1. Stemodia diplohyptoides M.M.Sosa & Dematt., Phytotaxa 186(5): 272. 2014. Figs. 3e; Fig. 5a-b

Stemodia diplohyptoides can be recognized by the membranaceous leaves and axillary inflorescences (vs. subcoriaceous leaves and terminal inflorescences in Stemodia stricta).

Examined materials: ARGENTINA. MISIONES: Puerto Iguazú, Parque Nacional Iguazú, Isla San Martin, 14.II.2004, M.M. Sosa 131, 133 (CTES!).

Stemodia diplohyptoides occurs in Argentina, in the north and central regions of the Misiones province (Sosa & Dematteis 2014). Found only in the Iguazú National Park, on sandbars and rocky islands next to the Iguazú River.

Flowering from August to December (Sosa & Dematteis 2014). However, it was found with flowers in February in the Iguazú National Park.

5.2. Stemodia stricta Cham. & Schltdl., Linnaea 3: 10. 1828. Figs. 3g; 5c-d

Stemodia stricta can be distinguished from other Stemodia species found in the Park by the sessile leaves (vs. petiolate in S. verticillata), short trichomes in the calyx and leaves margin with less than 20 teeth per side (vs. more than 20 teeth per side in S. diplohyptoides).

Examined materials: ARGENTINA. MISIONES: Puerto Iguazú, Parque Nacional Iguazú, Isla San Martín, 15.X.1993, S.G. Tressens 4562 (MO[digital image]!); Dep. San Miguel, Corrientes, 19.XII.2002, M.M. Sosa (HUEFS[digital image]!). BRAZIL. PARANÁ: Foz do Iguaçu, Parque Nacional do Iguaçu, 19.V.1949, A.P. Duarte 1853 (UEC!); 18.II.1960, E. Pereira 5355 (NY[digital image]!); 8.III.2018, E.L. Siqueria 2477 (HCF!); trilha da Sede, 25°37’38.0”S, 54°28’52.7”W, 16.XII.2016, M.G. Caxambu 7718 (HCF!, MBM!, UNOP!); trilha do hidrometro, 16.XII.2016, M.G. Caxambu 7715 (HCF!, MBM!).

Stemodia stricta occurs in open humid areas susceptible to floods in Argentina, Brazil (Central-West and South regions), Bolivia, Paraguay, and Uruguay (Souza & Giulietti 2009). Found in Foz do Iguaçu and in the Iguazú National Park, in slightly open areas near the Iguazú and São João Rivers.

This species was found with flowers in March and December.

Figure 3
a-b. Scoparia dulcis - a. flowering branch; b. flower detail. c-d. Scoparia montevidensis - c. flowering branch; d. flower detail. e. Stemodia diplohyptoides - flowering branch. f. Stemodia hyptoides - flowering branch. g. Stemodia stricta - flowering branch. h. Stemodia verticillata - flowering branch.

Figure 4
a-b. Mecardonia procumbens - a. flower detail; b. habit. c-d. Scoparia dulcis - c. inflorescence; d. branch side view. e-f. Scoparia montevidensis - flower and branch upper view; f. habit. Pictures by Elmar Hentz Júnior and AVS

Figure 5
a-b. Stemodia diplohyptoides - a. flower front view; b. leaves detail. c-d. Stemodia stricta - c. flower front view; d. leaves detail. e-f. Stemodia verticillata - e. flower front view; f. habit. Pictures by Elmar Hentz Júnior and AVS

Previously to our survey, four Stemodia species were indicated for the Iguaçu National Park. However, upon analyzing the specimen Caxambu 7715 (HCF), identified as Stemodia hyptoides (Fig. 3f), it was possible to indicate its real identity as S. stricta since it showed lax leaves with less than 20 teeth per side and exclusively terminal inflorescences.

5.3. Stemodia verticillata (Mill.) Hassl., Trab. Mus. Farmacol. no. 21: 110. 1909. Figs. 3h; 5e-f

Stemodia verticillata can be distinguished from other Stemodia species found in the Park by the crawling habit (vs. erect), petiolate leaves (vs. sessile), and absent bracteoles (vs. presence).

Examined materials: ARGENTINA. MISIONES: Puerto Iguazú, Parque Nacional Iguazú, borde de camino de acceso al Parque, 14.II.2004, M.M. Sosa 132 (CTES!). BRAZIL. PARANÁ: Céu Azul, Parque Nacional do Iguaçu, trilha da Jacutinga, 25°14’05.5”S, 53°49’08.9”W, 18.III.2018, E.L. Siqueira 2532 (HCF!). Foz do Iguaçu, 14.X.1962, G. Hatschbach 9366 (MBM!, UPCB!); casa da administração, 12.XII.1999, A.C. Cervi 6942 (HUCP!, UPCB!); trilha de acesso à Fazenda Salinet, 25°35’36.0”S, 54°22’51.0”W, 24.VIII.2019, H.T.P. Vieira 28 (EVB!); trilha da represa, 2019, E.J. Hentz Júnior 80 (EVB!). Lindoeste, cachoeira Rio Gonçalves Dias, 25°21’37.2”S, 53°39’28.1”W, 13.II.2020, M.G. Caxambu 9110 (HCF!).

Stemodia verticillata occurs in the Neotropics from Mexico to Argentina, highly widespread (Souza & Giulietti 2009). In Brazil, this species occurs in all states (Scatigna & Souza 2023). Found in Céu Azul, Foz do Iguaçu, and in the Iguazú National Park. Common in the region, occurring in every type of forest formation.

This species was found with flowers in February, March, and December.

6. Veronica L. Species Plantarum 1: 9. 1753.

6.1. Veronica arvensis L., Sp. Pl. 1: 13.

Veronica arvensis can be recognized by the petiolate leaves and sessile bracts, by the rotaceous blue corolla, and by the androecium composed of two stamens.

Examined materials: ARGENTINA. MISIONES: Puerto Iguazú, Parque Nacional Iguazú, 30.X.1960, A.T. Hunziker 15485 (CORD! [digital image]).

Additional examined materials: BRAZIL. RIO GRANDE DO SUL: Porto Alegre, Vila Manresa, 24.IX.1955, B. Rambo 57300 (RB!). SANTA CATARINA: Nova Teutonia, 11.VIII.1944, F. Plaumann 578 (RB!).

Veronica arvensis is native from Eurasia, naturalized in America. In Argentina it occurs in Buenos Aires, Chubut, Córdoba, Corrientes, Entre Ríos, Jujuy, La Pampa, Mendoza, Misiones, Neuquén, Río Negro, Salta, Santa Fe and in Tierra del Fuego (Flora Argentina 2018). In Brazil, this species occurs in Southeast and South regions (Souza 2023). Registered only in the Iguazú National Park.

This species was found with flowers and fruits from November to March (Pérez-Calix 2023). In the Park it was collected with flowers in October.

Key to the Scrophulariaceae from the Iguaçu and Iguazú National Parks

1. Stem cylindrical; corolla white or cream; capsule tomentose 7.1. Buddleja elegans subsp. elegans

1’. Stem quadrangular; corolla yellow; capsule glabrescent 7.2. Buddleja stachyoides

7. Buddleja L., Sp. Pl. 1: 112. 1753.

7.1. Buddleja elegans Cham. & Schltdl., Linnaea 2(4): 594. 1827.Fig. 2c-d

Buddleja elegans has two subspecies, the one found in the Iguaçu National Park being B. elegans subsp. elegans, which is endemic to Brazil and occurs in the Semidecidual Seasonal Forests of Paraná. This subspecies is characterized by a congested cyme with sessile flowers (vs. lax cymes with pedunculated flowers in Buddleja elegans subsp. angustata Bentham (1846: 443) Norman (1995: 192). Also, it differs from the other Buddleja species found in the Park by the coriaceous leaves (vs. membranaceous or chartaceous in B. stachyoides), the cylindrical stem (vs. quadrangular in B. stachyoides), white flowers (vs. yellow in B. stachyoides), and tomentose capsules (vs. glabrescent in B. stachyoides).

Examined materials: Céu Azul, Parque Nacional do Iguaçu, Lagoa Azul, 25°05’22.5”S, 53°40’09.1”W, 5.X.2017, M.G. Caxambu 7931 (HCF!).

Additional examined materials: BRAZIL. PARANÁ: Guaratuba, Morro dos perdidos, 18.IX.1997, E.P. Santos 339 (UPCB!). Tibagi, Parque Estadual do Guartelá, 2.III.2009, C. Snak 230 (UPCB!); Piraquara, arredores da represa, 12.VIII.2015, L.C. Lannoy 1 (UPCB!).

There is one material of Buddleja elegans subsp. angustata for the Iguazú National Park (H.A. Keller 1218), however the material was not examined to really make sure of its identity. But it confirms the occurrence of B. elegans in both parks.

Buddleja elegans subsp. elegans is found in Brazil (Central-West, Southeast, and South regions). In the Park, it was found in Céu Azul. Common in wetlands and forest borders (Coelho & Miotto 2018).

Flowering primarily from September to December, fruiting from October to January. In the Park, it was found flowering in October.

7.2. Buddleja stachyoides Cham. & Schltdl., Linnaea 2: 597 (598). 1827.Fig. 2e-f

Buddleja stachyoides can be distinguished from B. elegans by the membranaceous or chartaceous leaves (vs. coriaceous in B. elegans), quadrangular stem (vs. cylindrical in B. elegans), yellow flowers (vs. white in B. elegans), and glabrescent capsules (vs. tomentose in B. elegans).

Examined materials: Céu Azul, Parque Nacional do Iguaçu, borda do Parque, 25°06’29.1”S, 53°45’09.3”W, altitude 666 m 25.VIII.2016, M.G. Caxambu 7464 (HCF!). Foz do Iguaçu, trilha da sede do ICMBio, 25°37’36.6”S, 54°28’53.7”W, 14.VIII.2015, M.G. Caxambu 6759 (HCF!); trilha do Hidrômetro, 15.IX.2018, C.R. Rauber 165 (UNOP!).

Buddleja stachyoides is widespread, native to Argentina, Bolivia, Brazil (Northeast, Central-West, Southeast, and South regions), Paraguay, and Uruguay (Coelho & Miotto 2018). In the Park, it was found in Céu Azul and Foz do Iguaçu.

This species flowers from July to September, fruits from August to October (Lannoy et al 2018). In the Park, it was found flowering in August and September.

Discussion

Some difficulties were found while working on the specimen M.G. Caxambu 7931 (HCF!). Even though it was already identified as Buddleja elegans, we decided to further identify it down to the subspecies level. That way, while analyzing the possible subspecies, we realized that most of the B. elegans subsp. elegans collections (L.C. Lannoy 1, E.P. Santos 339, C. Snak 230) were made at higher elevations and B. elegans subsp. angustata is common at elevations lower than 100 m, especially on the borders of Paraguay with the Brazilian states of Paraná and Mato Grosso do Sul (Coelho & Miotto 2018).

However, based on the specimens’ morphology, we decided that the subspecies found in the Iguaçu National Park is, indeed, B. elegans subsp. elegans, which is a rare occurrence in the region - especially when there is a collection of B. elegans subsp. angustata in Argentina, possibly showing this subspecies can occur near to each other.

In relation to the areas’ species richness (Tab. 1), Foz do Iguaçu showed the highest among the areas, with eight species, including one not found in any other area, Scoparia dulcis - observed in the Iguazú National Park (Fabian Gatti, personal communication, September 7, 2023). Four species were found in Céu Azul, andBuddleja eleganssubs. elegans was only found in this area in a collection made near a natural deep lake, which can explain its restricted occurrence since similar areas are not well sampled. For the Iguazú National Park, seven species were documented while visiting the CTES herbarium and by online exsiccate images. Two species were found in Capanema and only one in the Matelândia/Serranópolis area.

As for the conservation status, out of the 11 Plantaginaceae species, only Mecardonia grandiflora and Plantago australis had been previously analyzed and classified as Near Threatened (NT) and Least Concern (LC), respectively (CNCFlora 2023). Bacopa australis, Mecardonia procumbens, Scoparia dulcis, Scoparia montevidensis, Stemodia diplohyptoides, Stemodia stricta, and Stemodia verticillata are herein indicated as LC for Brazil, since their EOO is higher than 20,000 km². Both Scrophulariaceae species (Buddleja elegans and Buddleja stachyoides) had their conservation status assessed by Coelho & Miotto (2018) and are categorized as LC. The conservation status of Bacopa scabra and Veronica arvensis were not accessed.

Although some of these species have already been cited in some studies in the region, knowledge about them is still scarce. For example, a floristic survey carried out at the ParNa Iguaçu (Trochezet al.2017) listed seven Plantaginaceae species, which were displayed in this survey. However, no Scrophulariaceae species were previously cited in the Iguaçu National Park. Also, an old management plan for the ParNa Iguaçu listed no species for the two families, and the most recent one does not present a species list for the flora (ICMBio 2018). Thus, the results confirm the importance of detailed floristic surveys in large forest remnants as they are basic resources for Management Plans, and other studies on ecology, environmental education, physiology, phytochemistry, and genetics, all of which depend on identifying and indicating the location of the species within the park, highlighting the importance to conserve the Iguaçu and Iguazú National Parks.

Table 1
Plantaginaceae and Scrophulariaceae species found in each of the study areas within the Iguaçu and Iguazú National Parks.

Species	Céu Azul	Matelândia /Serranópolis	Capanema	Foz do Iguaçu	Argentina
Buddleja elegans	X
Buddleja stachyoides	X			X
Bacopa australis		X
Bacopa scabra					X
Plantago australis	X			X
Mecardonia grandiflora			X	X	X
Mecardonia procumbens				X	X
Scoparia dulcis				X
Scoparia montevidensis			X	X
Stemodia diplohyptoides					X
Stemodia stricta				X	X
Stemodia verticillata	X			X	X
Veronica arvensis					X

Acknowledgements

We would like to thank Dra. Greta Aline Dettke, for her suggestions on the manuscript; ICMBio, for the collection permit; the entire team at the EVB herbarium and the UNOP herbarium, for their help in the fields; the team at the CTES herbarium in Corrientes; the curator of CORD herbaria, for providing the exsicatte images; and CAPES, for the master’s scholarship provided to the first author. AVS thanks Fundação de Amparo à Pesquisa e ao Desenvolvimento Científico e Tecnológico do Maranhão (FAPEMA) (PDCTR-00123/20) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) (301691/2021-5) for financial support. Livia Temponi, Laura Lima and Marcelo Galleazi also thank Fundação de Amparo à Pesquisa e ao Desenvolvimento Científico e Tecnológico do Maranhão (FAPEMA) (PDCTR-00123/20), Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) (301691/2021-5) and NAPI Taxonline (Fundação Araucária) for financial support.

Data availability statement

In accordance with Open Science communication practices, the authors inform that all data are available within the manuscript.

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Edited by

Area Editor:

Dra. Valquíria Dutra

Publication Dates

• Publication in this collection
  22 July 2024
• Date of issue
  2024

History

• Received
  06 June 2023
• Accepted
  21 Jan 2024