Plants in the clouds: vascular epiphytes of Pedra Azul, a mountain top in Espírito Santo, Southeastern Brazil

Abstract

Cloud forests usually occur at high-altitude sites of the Atlantic Forest in eastern Brazil, albeit scattered and fragmented along the mountain tops. In this habitat, the vegetation occurs at low-temperature conditions and is usually provided by additional water sources that arise due to the horizontal precipitation of the frequent fogs. Together with the more considerable air movement and higher luminosity, these factors are conditioning for singular floras at high elevations, mainly the vascular epiphytes, which are macro and microclimate dependent. In the mountains range at the center of the Espírito Santo state, Southeastern Brazil, some mountain tops such as Pedra Azul (PA) hold these environmental features. Here, we aimed to present the first checklist of vascular epiphytes in the Pedra Azul State Park and surroundings based on fieldwork and herbarium specimens. The checklist comprises 152 species, 65 genera, and 17 families, the main families being Orchidaceae, Bromeliaceae, and Polypodiaceae, with the main genera represented by Vriesea, Acianthera, and Peperomia. The holoepiphytes were the main category among the epiphytes, although an unusually high number of facultative epiphytes were recorded. Asplenium theciferum and Octomeria cucullata are recorded in Espírito Santo for the first time, and we confirmed the occurrence of Rhipsalis cereuscula in the state. Overall, the richness recorded in PA is amongst the highest of the Atlantic Forest cloud forests. Six species are threatened at the national level and 32 at the state level. These results support the importance of the protected area for conserving the flora; however, several species - including threatened - were only recorded in the surroundings, demonstrating that the buffer zone of the Pedra Azul State Park must be included in the management plans.

Key words
Atlantic Forest; cloud forest; endangered species; floristic; inselberg; Pedra Azul State Park

Resumo

As florestas nebulares ocorrem em locais de altitude elevada na Floresta Atlântica do leste do Brasil, embora dispersas e fragmentadas no topo das montanhas. Neste habitat, a vegetação ocorre em condições de baixa temperatura e geralmente é suprida por fontes de água adicionais que surgem devido à precipitação horizontal oriundas de nevoeiros, que são frequentes nesses locais. Ademais, os maiores movimentos de ar e a luminosidade, constituem fatores condicionantes a floras singulares que ocorrem em altitudes elevadas, principalmente as plantas epifíticas, que são sensíveis ao macro e microclima. Na região serrana do centro do estado do Espírito Santo, no Sudeste do Brasil, alguns locais mais elevados, como a Pedra Azul (PA), possuem essas características ambientais. Aqui, objetivamos apresentar a primeiro inventário de epífitas vasculares no Parque Estadual de Pedra Azul e arredores com base em trabalhos de campo e espécimes de herbário. Nós registramos 152 espécies, 65 gêneros e 17 famílias, sendo as principais famílias Orchidaceae, Bromeliaceae e Polypodiaceae, com os principais gêneros representados por Vriesea, Acianthera e Peperomia. As holoepífitas constituem a categoria principal entre as epífitas, embora um número incomumente alto de epífitas facultativas tenha sido registrado. Asplenium theciferum e Octomeria cucullata foram registradas pela primeira vez no Espírito Santo, e confirmamos a ocorrência de Rhipsalis cereuscula no estado. No geral, a riqueza registrada em PA está entre as mais altas das da Floresta Atlântica. Seis espécies estão ameaçadas em nível nacional e 32 em nível estadual. Esses resultados corroboram a importância da área protegida para a conservação da flora; entretanto, várias espécies - inclusive ameaçadas - foram registradas apenas no entorno, demonstrando que a zona de amortecimento do Parque Estadual da Pedra Azul deve ser incluída nos planos de manejo.

Palavras-chave
Floresta Atlântica; floresta nebular; espécies ameaçadas; florística; inselberg; Parque Estadual da Pedra Azul

Introduction

The Atlantic Forest, which occurs in eastern Brazil, northern Argentina, and eastern Paraguay, is a phytogeographic domain comprising the second largest forest massif of South America, only smaller than the Amazon (Oliveira-Filho & Fontes 2000; Fiaschi & Pirani 2009). It occurs in the most mountainous region of Brazil, thus having a rugged relief and, consequently, a physiognomy conditioned by several environmental factors (e.g., climatic and edaphic), resulting in different types of vegetation (e.g., Semideciduous and Deciduous Seasonal Forests, Dense and Mixed Rain Forest, and Coastal Plain Forest (Restinga) (Oliveira-Filho & Fontes 2000; Fiaschi & Pirani 2009).

At high altitudes, primarily in southern and southeastern Brazil, some particular vegetation types occur in the Atlantic Forest, such as the cloud forests, Araucaria forests, and high-altitude grasslands (Safford 1999; Costa et al. 2018). The main mountain ranges are the Serra da Mantiqueira and the Serra do Mar, with elevations reaching nearly 3,000 m above sea level (e.g., Pico da Bandeira, at 2,890 m) (Costa et al. 2018). Besides these, in the state of Espírito Santo, southeastern Brazil, there is a mountain range locally called Montanhas Capixabas or Serra do Castelo (Chiron & Bolsanello 2015). Although it does not reach elevations as high as the former ones, with the highest peak reaching about 2,000 m on the highest mountain top of the Serra do Castelo (Chiron & Bolsanello 2015; Garbin et al. 2017), it is one of the least botanically studied of the Atlantic Forest mountain ranges, with few floristic studies available.

The vegetation of these highlands, nowadays restricted and scattered on mountain tops, was likely widespread in the past due to the cold climate during the Pleistocene glaciations, forcing them to occur at lower elevations and latitudes (Luna-Vega et al. 2001; Behling et al. 2002). Thus, this vegetation represents vestiges of the past cold climate physiognomies that are nowadays reduced to fragments on the mountain tops (Koehler et al. 2002; Bertoncello et al. 2011) and harbor important and threatened species with restricted distributions and which may disappear with the increase of global temperatures (Foster 2001; Colwell et al. 2008).

High-altitude environments hold peculiar environmental conditions such as the presence and persistence of clouds and mist, influencing edaphic and microclimatic conditions (Bertoncello et al. 2011) and, consequently, the distribution of the flora and vegetation (Gentry & Dodson 1987; Oliveira-Filho & Fontes 2000; Blum et al. 2011a; Leitman et al. 2015).

At the Serra do Castelo, high-altitude vegetation, like occurs in the Pedra Azul, is associated with the granite-gneiss dome-shaped rock massifs (inselberg) (Garbin et al. 2017; de Paula et al. 2020). In such environments, according with the topography, in the rock slopes occurs scattered mats with herbaceous plants, and where patches of shallow soil are formed, a shrub vegetation occurs; also, in flat areas, deepest soils are formed and a scattered dwarf cloud forests developed (Rizzini 1997; Garbin et al. 2017; de Paula et al. 2020). Furthermore, surrounding the inselbergs, taller montane and high-montane cloud forests may occur (Magnago et al. 2007; Thomaz 2010). Such forests are characterized by low temperatures and abundant fog that substantially increase the humidity and may provide water during drought periods (Stadtmüller 1987; Bruijnzeel et al. 2010; Bertoncello et al. 2011; Arcova et al. 2016). Hence, vascular epiphytes, which are greatly dependent of humidity, are favored by the horizontal precipitation provided by the clouds, resulting in great diversity and biomass of epiphytes in cloud forests (Nadkarni 1984; Clark et al. 1998; Freiberg & Freiberg 2000; Köhler et al. 2007).

In Brazil, few floristic surveys of vascular epiphytes have been carried out in cloud forests (e.g., Bianchi et al. 2012; Bonnet et al. 2013; Alves & Menini Neto 2014; Furtado & Menini Neto 2016, 2018). In Espírito Santo, only two studies of vascular epiphytes were performed in high-altitude sites (Couto et al. 2016; Francisco et al. 2019). Gaps in biodiversity knowledge are one of the biggest problems for conservation (e.g., Almeida & Mamede 2014; Giaretta et al. 2015). Therefore, it is essential to carry out floristic studies in areas that have been little studied and present environmental heterogeneity, aiming to direct the conservation efforts (Kessler 2001), besides the great relevance of understanding the origin, maintenance, distribution patterns, and processes to which biodiversity is subject (Werneck et al. 2011; Santos et al. 2014; Zizka & Antonelli 2018).

Here, we aim to present the first checklist of the composition of vascular epiphytes at a mountain top in the center of the state of Espírito Santo (Serra do Castelo), Southeastern Brazil, specifically in the Pedra Azul State Park and surroundings.

Material and Methods

Study site

The Pedra Azul (PA) is situated in the mountainous region of Espírito Santo, which occupies the central-south portion of the state (IPEMA 2005). These mountains are known locally as Serra do Castelo (Castelo Mountain Chain) (Chiron & Bolsanello 2015) and delimited to the North by the Doce River valley, to the East by the coastal plain, to the South by the Itapemirim River valley, to the Southwest by the Serra do Caparaó, and to the Northwest by the state of Minas Gerais and the Serra da Chibata, with an average altitude of 700 m reaching up to 2,050 m (Chiron & Bolsanello 2015). In the collection sites of the PA, the elevation ranges between 1,100 m and 1,600 m.

According to the Köppen classification, the climate in the PA is Cfb, with an average temperature from 14 °C to 16 °C, average precipitation of 1,200 mm to 1,500 mm, and seasonal drought in the winter (May to October), with six months with less than 100 mm of rain (Alvares et al. 2013; INCAPER 2020). In the PA, the occurrence of fogs is usual (personal observations), with an estimated mean annual cloud cover of 80 % (Wilson & Jetz 2016).

In PA, the vegetation is usually classified as montane to high-montane cloud rainforest (dense ombrophilous forest) (Magnago et al. 2007; Garbin et al. 2017). However, there are six months of drought (Alvares et al. 2013; INCAPER 2020), so it is better fitted as a seasonal forest (IBGE 2012). In the collection sites, the vegetation ranges from tall montane and high-montane cloud forests to dwarf high-montane cloud forests, including riverine formations of both. The first occur in the deep valleys, slopes, and some patches among the inselberg drainages, while the latter in the high-altitude inselbergs, between Pedra das Flores and Pedra Azul.

The study site comprises part of the Pedra Azul State Park (PASP) and a valley in the Buffer Zone (BF) (central coordinate 20°25’05”S and 41°00’25”W) (Fig. 1), situated in the district of Aracê, which belongs to the municipality of Domingos Martins, in the state of Espírito Santo (ES), Southeastern Brazil. The PASP was created in 1960 originally named Pedra Azul Forest Reserve (State Decree No. 312, of October 31st, 1960), comprising the steepest areas usually with little or no interest for crop cultivation or pastures (IDAF 2004). Consequently, the protected area includes the high-altitude inselbergs (Pedra Azul at 1,822 m, Pedra das Flores at 1,909 m) and adjacent mountain tops, comprising about 1,240 hectares, surrounded by private properties. The PASP has a well-developed tourism activity.

Figure 1
a-b. Geographical location of Pedra Azul and the delimitation of the Pedra Azul State Park (PASP) in white, in Domingos Martins municipality, Espírito Santo, Brazil – a. Espírito Santo state, highlighting the location of Pedra Azul; b. delimitation of PASP (the letters are according to the photos). c-d. showing the interior of the PASP – c. Pedra Azul, dwarf cloud forest and Pedra das Flores in the background; d. Pedra Azul and Pedra das Flores covered by the clouds showing the tall montane forest in the slopes. e-g. the buffer zone sampled in this study – e. tall montane cloud forest; f. alluvial cloud montane forest; g. secondary montane cloud forest. (a. prepared by Henrique Lauand Ribeiro; b. adapted from Google Earth Pro; c-g. photos of Gabriel Mendes Marcusso).

Collections were made on the touristic trail from the natural pools up to the foothill of Pedra das Flores and Pedra Azul (ranging from 1,350 m to 1,600 m a.s.l., covering about 12 hectares of dwarf cloud forest), in the secondary forest surrounding the research lodge (1,300 m to 1,400 m, covering about 20 ha), and in an area contiguous to the Park, in the slopes and riverine forests, in the headwaters of the Jucu River, which crosses a private property (1,100 m to 1,230 m, covering about 5 ha) (Fig. 1).

Data collection

The floristic inventory of vascular epiphytes was carried out over seven expeditions lasting about one to tree days each, in January, May, and September of 2018, February, August, and October of 2019, and November of 2020. Fertile specimens were deposited at the Herbarium Rioclarense (HRCB), and duplicates were sent to the herbaria BHCB, CESJ, RB, and UNOP (acronyms follow Index Herbariorum, continuously updated). Sterile specimens collected mainly from fallen trees and branches were kept at a greenhouse in Rio Claro until flowering and subsequently processed. To complement the floristic inventory, we used the SpeciesLink network (CRIA 2020) and Reflora - Virtual Herbarium (2020) to analyze previous collections carried out within the Park and deposited at the herbaria MBML, RB, and VIES (acronyms follow Index Herbariorum, continuously updated).

During the expeditions to the field, we observed the habits of the epiphytes and classified them into ecological categories according to their relationship with the phorophytes, following Benzing (1990): characteristic holoepiphytes, facultative holoepiphytes, primary hemiepiphytes, and secondary hemiepiphytes. It was preferred to use this classification instead of that of Zotz (2013a) to facilitate comparison with other studies since most used the former. Terrestrial species occasionally found growing as epiphytes were not considered in this study, as they are deemed ephemeral epiphytes (Kersten 2010; Zotz 2016).

Taxonomic identification was made using taxonomic literature and herbarium collections at HRCB and MBML, as well as by consulting specialists (mentioned in the acknowledgments). The taxonomic classification is according to APG IV (2016) for angiosperms and PPG I (2016) for monilophytes and lycophytes. Species names, synonymy, authors, and distributions are used according to Flora do Brasil 2020 (BFG 2018). The conservation statuses were obtained at national (Martinelli & Moraes 2013; CNC Flora 2020) and, at the state level (Fraga et al. 2019).

Results

In the PA, we recorded 152 species of vascular epiphytes belonging to 65 genera and 17 families (Figs. 2-3; Tab. S1, available on supplementary material <https://doi.org/10.6084/m9.figshare.17161412.v1>). Ferns and Lycophytes account for 37 of the recorded species, and flowering plants represent the vast majority (115 species). In the PASP 104 species were recorded, in the BF, of the 73 species, 48 were not recorded inside the protected area; 25 of them were recorded in both (Tab. S1, available on supplementary material <https://doi.org/10.6084/m9.figshare.17161412.v1>). Overall, the richest families were Orchidaceae (56 species), Bromeliaceae (29), Polypodiaceae (16), Cactaceae and Piperaceae (9 each), and Araceae (7). The richness genera were Vriesea (11 species), Acianthera and Peperomia (9 each), Gomesa (7), Asplenium (6), and Hymenophyllum, Nidularium, Octomeria, Rhipsalis, and Tillandsia (5 each). The main ecological category is the characteristic holoepiphytes (88 species), followed by the facultative holoepiphytes (58) and the secondary hemiepiphytes (6).

Figure 2
a-o. Vascular epiphytes representatives of Pedra Azul State Park and a neighbor particular area, Domingos Martins, Espírito Santo, Brazil, highlighting new records to Espírito Santo, threatened and data deficient species – a. Anthurium fragae (Araceae); b-f. Bromeliaceae – b. Billbergia zebrina; c. Neoregelia diversifolia; d. Neoregelia farinosa; e. Quesnelia kautskyi; f. Vriesea capixabae; g-i. Cactaceae – g. Rhipsalis cereuscula; h. Rhipsalis hoelleri; i. Schlumbergera kautskyi; j-k. Gesneriaceae – j. Codonanthe cordifolia; k. Sinningia douglasii; l-o. Orchidaceae – l. Acianthera binotii; m. Acianthera heringeri; n. Anathallis radialis; o. Centroglossa macroceras.

Figure 3
a-o. Vascular epiphytes representatives of Pedra Azul State Park and a neighbor particular area, Domingos Martins, Espírito Santo, Brazil, highlighting new records to Espírito Santo, threatened and data deficient species – a-m. Orchidaceae – a. Dichaea cogniauxiana; b. Epidendrum chlorinum; c. Gomesa gomezoides; d. Gomesa imperatoris-maximiliani; e. Gomesa praetexta; f. Gomesa uhlii; g. Hadrolaelia pumila; h. Lankesterella ceracifolia; i. Miltonia spectabilis; j. Octomeria cucullata; k. Octomeria palmyrabellae; l. Pabstia jugosa; m. Pabstiella leucopyramis; n-o. Piperaceae – n. Peperomia catharinae; o. Peperomia pilicaulis.

Six species are reported as threatened at the national level, one critically endangered, two endangered, and three vunerable. At state level 32 are cited in the list, four are critically endangered, six endangered, and 22 vunerable (Tab. S1, available on supplementary material <https://doi.org/10.6084/m9.figshare.17161412.v1>). Furthermore, 10 species are Data Deficient (DD) at the state level.

Discussion

We presented the first checklist of the vascular epiphytes of the Pedra Azul, including part of the Pedra Azul State Park and a river valley in the buffer zone. We found new records for the state and species threatened of extinctions at both state and national level. Overall, the number of species is outstanding for the cloud forests of the AF. The great number of facultative epiphytes is a characteristic feature of forest in rockness matrix, like the mountain’s summits. Furthermore, the peculiar climatic conditions -with the prevalence of cloud cover inputing water in the vegetation and the low temperature- are the main drive for the singular flora found in this high-altitude site.

In Brazil, few floristic studies regarding vascular epiphytes have been carried out in mountainous sites above 1,000 m of elevation (Tab. 1). These studies registered a richness ranging from 50 to 312 species, although they used different sample efforts and areas (Tab. 1). The richness recorded in the PA agrees with the data on vascular epiphytes in mountainous sites of the Atlantic Forest. The PA only stands below studies that carried out more comprehensive sampling, considering several years of collections, and those in areas with a wider altitude range (e.g., Fontoura et al. 1997; Furtado & Menini Neto 2016). Furthermore, considering the area sampled in the present study (about 37 hectares) and the entire area of the State Park (1,240 hectares), we may consider that the PA harbors a rich vascular epiphyte flora. Undoubtedly, additional species are expected to occur in the area since most pristine forest patches of the PA are of difficult access due to the mountainous relief and there being few trails that cross such areas. Some species are not mentioned in the list because they were not found fertile [e.g., Acianthera nemorosa (Barb.Rodr.) F.Barros, Epidendrum paranaense Barb.Rodr. (Orchidaceae), Spirotheca rivieri (Decne.) Ulbr. (Malvaceae), and Nematanthus sp. (Gesneriaceae)].

Table 1
Local floristic surveys exclusively of vascular epiphytes in the high-altitude [above (880–)1,000 m] pluvial to seasonal cloud forests in the Brazilian Atlantic Forest organized in descending order of species richness. Spp = number of species; Gen = number of genera; Fam = number of families; m = meters above sea level; mm = millimeters, ha = hectares; NI = data not informed at the source; * = excluded mistletoes.

Location	State	Spp.	Gen.	Fam.	Altitude (m)	Precipitation (mm)	Area (ha)	Source
Macaé de Cima	RJ	312	87	26	880-1720	2128	7200	Fontoura et al. (1997)*
Ibitipoca	MG	224	81	22	1000-1800	1532	300	Furtado & Menini Neto (2018)
Pedra Azul	ES	152	65	17	1100-1600	1200-1500	37	Present study
Serra do Papagaio	MG	138	66	25	1800-2000	NI	NI	Furtado & Menini Neto (2016)
Serra do Cruz	MG	135	62	16	1100-1500	1200-1500	NI	Alves & Menini Neto (2014)
Marumbi	PR	127	59	21	1000	1384	1	Bianchi et al. (2012)
Serra da Prata	PR	121	64	23	1000-1100	2290	0.2	Blum et al. (2011b)
Ponte Alta	SC	84	47	16	1131	NI	2.01	Bonnet et al. (2013)
Cubatão, Joinville	SC	73	44	16	1168	NI	2.01	Bonnet et al. (2013)
Pedra dos Pontões	ES	66	34	10	1150	NI	1	Francisco et al. (2019)
Pedra Roxa	ES	62	46	15	1114	1391	2	Couto et al. (2016)
Guarapuava	PR	54	32	13	1100-1200	NI	NI	Kersten et al. (2009)
Campos do Jordão	SP	51	31	11	1030-2007	NI	NI	Mania (2013)
Nova Trento	SC	50	34	13	1097	NI	2,01	Bonnet et al. (2013)

Low temperatures and frost are constraining factors that affect vascular epiphyte diversity in high-altitude habitats (Gentry & Dodson 1987; Krömer et al. 2005; Blum et al. 2011b; Hsu et al. 2014). In turn, the occurrence of fogs acts as a source of humidity (Arcova et al. 2012) and is positively associated with the epiphytic assemblage, thus being one of the most important factors conditioning the occurrence of vascular epiphytes (Gentry & Dodson 1987; Laube & Zotz 2003; Ding et al. 2016). This additional humidity has great importance in the driest periods, mitigating the drought due to the “horizontal precipitation” (Vogelmann 1973; Arcova et al. 2012). Unfortunately, quantitative climatic data (e.g., on water input by cloud condensation) are scarce in the Brazilian mountains, mainly in Espírito Santo. In the PA, fog at the mountain is common, especially from nightfall to morning (personal observations).

In the PA, the richest families reported follow the expected for cloud forests of eastern Brazil (Alves & Menini Neto 2014; Furtado & Menini Neto 2016, 2018), for the Atlantic Forest in particular (Kersten 2010; Freitas et al. 2016), and to the global patterns, except for the more considerable importance of the Bromeliaceae family, which is a near endemic Neotropical family (Zotz 2013b, 2016). About 80 % of the vascular epiphytes are concentrated in the greatly diversified Orchidaceae family, resulting in the significant taxonomic importance of this family in this synusia (Kersten 2010; Zotz 2013b, 2016). The low representativity of the Araceae family in high-altitude habitats is expected in the Neotropical Region (Krömer et al. 2005; Ortiz et al. 2019), and the Atlantic Forest (Menini Neto et al. 2009; Rogalski et al. 2016), with a better contribution in lowland forests (Coelho 2010), mainly in the Amazon (Irume et al. 2013; Obermuller et al. 2014).

Few ferns and lycophytes were recorded in the PA (21.7 % of the total). This pattern is not usually observed in higher-altitude forests in the Neotropical Region, which usually is richer at high-altitude sites (Krömer et al. 2005, 2013; Cardelús et al. 2006). The pattern appears to be different in the Atlantic Forest, with lycophytes and ferns decreasing in richness when the altitude increases (Paciencia 2008; Blum et al. 2011b; Nóbrega 2013; Couto et al. 2016).

Regarding the distribution, the vast majority of the identified species (95 spp., 62.5% overall) are endemic of the Atlantic Forest, and seven are restricted to Espírito Santo (Anthurium fragae, Gomesa uhlii, Neoregelia diversifolia, Nidularium cariacicaense, Vriesea capixabae, V. pereirae, and Rhipsalis hoelleri). Octomeria cucullata and Asplenium theciferum are being reported for the first time in Espírito Santo and represent the first collections for Brazil after over 50 years, according to Dutra et al. (2015), CRIA (2020), and REFLORA (2020). We recorded the occurrence of Rhipsalis cereuscula in Espírito Santo; the only collection previously reported of it (Dutra et al. 2015) was in the city of Espírito Santo do Pinhal, in the state of São Paulo. Furthermore, 22 species are being recorded in cloud forests of the Southeastern Brazil for the first time (Furtado 2020).

The predominance of holoepiphyte characteristics is an expected pattern (Freitas et al. 2016; Zotz 2016). On the other hand, the great number of facultative holoepiphytes recorded is outstanding (38.1 % overall) when compared with some sites of the Atlantic Forest (Bonnet et al. 2011; Padilha et al. 2015; Couto et al. 2016; Marcusso & Monteiro 2016; Rolim et al. 2016). This different pattern could be explained by the abundance of rock outcrops in the studied area (Biral & Lombardi 2012; Perleberg et al. 2013; Couto et al. 2016). The rocks show a level of ecological equivalence with the trunks and branches of trees, such as the few nutrients available and the higher water drainage (Zotz 2016). Also, when situated below the forest cover, they may hold light conditions similar to those of the lower trunk and inner tree crown (Zotz 2016). These features have demonstrated a high habitat equivalence, allowing the occurrence of epiphytes, mainly accidental, in the trees (Perleberg et al. 2013; Couto et al. 2019). In our study, few accidental epiphytes’ species, not included in the checklist, were observed in forks and horizontal branch (e.g., Bifrenaria tyrianthina (Lodd. ex Loudon) Rchb.f., Epidendrum secundum Jacq. and the exotic Crassulaceae, Kalanchoe fedtschenkoi Raym.-Hamet & H.Perrier. However, several of the facultative species recorded, grows on the rocks. The presence of trees with horizontal and exposed roots perhaps is a determinant factor for the colonization of a high number of typical rupicolous species, like Pseudobombax (Couto et al. 2019), not observed in the PA.

The hemiepiphytes were represented by few species in the PA, in agreement with the low representativity of families with a great prevalence of this habit, such as the Araceae, corroborating the previous data on vascular epiphytes at high altitudes or in rocky outcrop habitats (Alves & Menini Neto 2014; Couto et al. 2016; Furtado & Menini Neto 2016; 2018; Rogalski et al. 2016).

The records of 31,6 % of the species only outside the protected area argue for the conservation issues in the BZ of the PASP, which must be considered in conservationist policies. Special attention regarding the threatened species recorded only outside the PASP are need: three of them are threatened at national level - Acianthera heringeri (Hoehne) F.Barros being critically endangered - and nine at state level - Schlumbergera kautskyi (Horobin & McMillan) N.P.Taylor and Sinningia douglasii (Lindl.) Chautems being critically endangered. On the other hand, the sampling of the inaccessible areas inside the PASP could locate these threatened taxa inside its area. Therefore, a first step is to reach the inaccessible areas of the PASP by means of trails, to monitor and allow new studies in the areas not visited by tourists. Plus, monitoring the populations inside the PASP and the BF is needed, to avoid the illegal collections, the illegal suppression of the vegetation and the occurrence of the fire. In this way, we encourage more botanical collections and studies within the boundaries of this protected area.

Acknowledgements

This study was financed by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Finance Code 001 (scholarship to the first author); Programa de Apoio a Pós-Graduação (PROAP/CAPES) (revision of the English language); and the Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) (financial support, #2018/04121-0). We thank the Instituto Estadual de Meio Ambiente e Recursos Hídricos (IEMA), for the permission to collect in the Pedra Azul State Park (process No. 77257804/17); the taxonomists Janaine Kunrath Hammes, Lívia Godinho Temponi, Marcus Coelho Nadruz, Rodrigo Theófilo Valadares (Araceae), Elton Leme, Dayvid R. Couto, Rafaela C. Forzza, Rebeca P. Romanini (Bromeliaceae), Alexandre Salino (Ferns and Lycophytes), Alain Chateums (Gesneriaceae), Felipe Gonzatti (Hymenophyllaceae), Benjamin Øllgaard (Lycopodiaceae), Antonio Toscano de Brito, Edlley Pessoa, Tiago Luiz Vieira, Wellington Forster (Orchidaceae), who helped in the identification of the epiphytes species; Henrique Lauand Ribeiro, for the map design; Fabiula Moreno Arantes, Janaine Kunrath Hammes, Renan Borgiani, and Vitor de Andrade Kamimura, for the help in the field work; and two anonymous reviewers, for the valuable contributions in the manuscript.

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Supplementary Material

See supplementary material at <https://doi.org/10.6084/m9.figshare.17161412.v1>

Edited by

Area Editor: Dra. Tatiana Carrijo

Publication Dates

• Publication in this collection
  07 Mar 2022
• Date of issue
  2022

History

• Received
  12 Dec 2020
• Accepted
  12 Apr 2021