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. 2025 Aug 4;14(15):2419. doi: 10.3390/plants14152419

Bryophytes of the Serra dos Órgãos National Park: Endemism and Conservation in the Atlantic Forest

Jéssica Soares de Lima 1,*, Allan Laid Alkimim Faria 2, Mateus Tomás Anselmo Gonçalves 1, Denilson Fernandes Peralta 1
Editor: Marta Puglisi
PMCID: PMC12349472  PMID: 40805769

Abstract

This study presents a comprehensive inventory of bryophytes in Serra dos Órgãos National Park (PARNASO), aiming to evaluate species richness, floristic composition and threatened taxa. Despite the state of Rio de Janeiro being one of the most extensively sampled regions for bryophytes in Brazil, detailed surveys of its conservation units remain scarce. Data were obtained through bibliographic review, herbarium specimen analysis, and new field collections. A total of 504 species were recorded, belonging to 202 genera and 76 families. The park harbors three locally endemic species, eight endemic to Rio de Janeiro, and sixty-nine species endemic to Brazil. Additionally, eleven species were identified as threatened, comprising seven Endangered (EN), two Critically Endangered (CR), and two Vulnerable (VU) according to the IUCN guidelines. PARNASO includes four distinct ecosystems along an altitudinal gradient: sub-montane forest (up to 500 m), montane forest (500–1500 m), upper-montane forest (1500–2000 m), and high-altitude fields (above 2000 m). Montane Forest showed the highest species richness, followed by high-altitude fields, upper-montane forest, and sub-montane forest. The findings highlight PARNASO’s importance in preserving bryophyte diversity in a highly diverse yet understudied region. This work contributes valuable baseline data to support conservation strategies and future ecological studies in Atlantic Forest remnants.

Keywords: endangered species, highlands, liverworts, mosses

1. Introduction

Serra dos Órgãos National Park (PARNASO), established on November 30th, 1939, holds the distinction of being the third national park created in Brazil. This protected area is situated within the Atlantic Forest domain, specifically in the Serra do Mar massifs and called “Órgãos” in Portuguese because its peaks resemble and are named as “pipe organs”. The Serra do Mar is an extensive mountain range approximately 1500 km along the Brazilian coast, from the northern Santa Catarina State to Rio de Janeiro State [1,2].

PARNASO currently represents one of the most important remaining biodiversity hotspots in Rio de Janeiro State, since its ecosystems are one of the largest centers of flower plant endemism in the Atlantic Forest [3,4]. It also plays a vital role in preserving essential water sources that feed two main watersheds—Paraíba do Sul and several streams of the Baía de Guanabara [5,6,7,8].

The Atlantic Forest domain is one of the most species-rich forests in the tropics [4,9,10]. It is also recognized for having high levels of biodiversity and endemism, expressed via its complex structural environments [11,12]. Tropical forests harbor the highest diversity of bryophyte species in the world, not only due to their structural complexity and microhabitat heterogeneity, but also because of favorable climatic conditions such as high humidity, frequent precipitation, and relatively stable temperatures [13,14]. Some authors have reported that this diversity may vary along the altitudinal range, especially concerning structural and floristic composition [15,16,17,18].

The Atlantic Forest within PARNASO comprises well-preserved forests characterized by four distinct vegetation formations based on altitude, climate, and soil conditions, among other factors [6,19,20,21,22]. In the lower parts around Sede Guapimirim, sub-montane forest prevails, while near Sede at Teresópolis, the forest transitions from montane forest to upper-montane forest, culminating above 2000 m.a.s.l. and high-altitude fields along the trail from Teresópolis to Petrópolis [19].

The species list of the vascular plants in Flora Organensis [19] is strongly based on the work of 18th-century botanists and on collections made especially by A.C. Brade (1881–1971) and C.T. Rizzini (1921–1992) between the 1930s and 1950s of the 20th century. These collectors also recorded bryophytes, and the specimens were studied by Bartram (1954) [19], who identified 146 species. Most bryophyte specimens from Serra dos Órgãos are deposited in the Herbarium of Jardim Botânico do Rio de Janeiro (RB).

Bryophytes from PARNASO were not sufficiently explored, and even considering that, though several researchers collected and published new taxa, the study was dispersed as examined material in taxonomical revisionary work. For example, Faria et al. (2021) [23] attempted to re-collect species founded by Glaziou in Serra dos Órgaos and the taxonomic reclassification of an Atlantic Forest endemic, and Peralta et al. (2020) [24] reported species of the Andreaceae family from the region.

There is no other information on bryophytes in PARNASO; thus, there is no detailed data on the floristic structure and composition of bryophytes. A comprehensive species list will contribute to filling a significant knowledge gap for the state of Rio de Janeiro and support the assessment of threatened bryophyte species. In addition, the revision of herbarium specimens and the incorporation of new collections will help stimulate further research on the ecology and biogeography of bryophytes within this important national park. This work aims to provide an inventory of the bryophyte species in PARNASO and discuss the following topics: (i) species richness and floristic composition; (ii) substrate preferences; and (iii) endemism and conservation status.

2. Results

2.1. Species Richness and Floristic Composition

The present study documented 504 species in 202 genera and 76 families (Table 1). The Serra dos Órgaos National Park preserves 57% of the known bryophyte flora of Rio de Janeiro State and 31% of Brazil. A total of 3000 specimens were analyzed, resulting in the identification of 428 species and an additional 79 species (indicated by * in Table 1) that remain known only from the literature.

Table 1.

Richness and diversity of bryophyte species. Symbols on the species voucher: species only known from the literature (*).

Group Family Taxon Voucher
Liverworts Acrobolbaceae Tylimanthus laxus (Lehm. & Lindenb.) Spruce Peralta 27761 (SP)
Aneuraceae Aneura pinguis (L.) Dumort. Peralta 27768 (SP)
Aneuraceae Riccardia chamedryfolia (With.) Grolle Silva, AL 515
Aneuraceae Riccardia digitiloba (Spruce ex Steph.) Pagán Peralta 27496 (SP)
Aneuraceae Riccardia emarginata (Steph.) Hell Peralta 27618 (SP)
Aneuraceae Riccardia fucoidea (Sw.) Schiffn. Peralta 20395 (SP)
Aneuraceae Riccardia glaziou (Spruce) Meenks Gaspar 121 (R)
Aneuraceae Riccardia regnellii (Aongström.) Hell Peralta 27487 (SP)
Arnelliaceae Gongylanthus liebmannianus (Lindenb. & Gott.) Steph Gonçalves 358 (SP)
Arnelliaceae Southbya organensis Herzog * Ph. v. Lützelburg 6003 p.p. (Tipo JE)
Balantiopsidaceae Isotachis aubertii (Schwägr.) Mitt. Gaspar 148 (R)
Balantiopsidaceae Isotachis erythrorhiza (Lehm. & Lindenb.) Steph. Peralta 27572 (SP)
Balantiopsidaceae Isotachis multiceps (Lindenb. & Gottsche) Gottsche Lima 783 (SP)
Balantiopsidaceae Isotachis serrulata (Sw.) Gottsche Gonçalves 396 (SP)
Balantiopsidaceae Neesioscyphus carneus (Nees) Grolle Peralta 20398 (SP)
Balantiopsidaceae Neesioscyphus homophyllus (Nees) Grolle Peralta 27722 (SP)
Calypogeiaceae Calypogeia peruviana Nees & Mont. Peralta 27491 (SP)
Cephaloziaceae Cephaloziella divaricada (Sm.) Schiffn. Gaspar 112 (R)
Cephaloziaceae Cephaloziella granatensis (J.B.Jack) Fulford Peralta 27638 (SP)
Cephaloziaceae Cylindrocolea planifolia (Steph.) R.M.Schust. Peralta 27621 (SP)
Cephaloziaceae Kymatocalyx dominicensis (Spruce) Vána Peralta 27755 (SP)
Cephaloziaceae Odontoschisma denudatum (Nees) Dumort. Peralta 20506 (SP)
Cephaloziaceae Odontoschisma longiflorum (Taylor) Trevis. * Fulford (1976)
Cephaloziellaceae Cephaloziopsis intertexta (Gottsche) R.M. Schust. Peralta 27488 (SP)
Cephaloziellaceae Fuscocephaloziopsis crassifolia (Lindenb. & Gottsche) Váňa & L. Söderstr. Dias 1414 (SP)
Dumortieraceae Dumortiera hirsuta Nees & Mont. Peralta 27710 (SP)
Frullaniaceae Frullania atrata Nees Faria 1560 (SP)
Frullaniaceae Frullania beyrichiana (Lehm. & Lindenb.) Lehm. * & Lindenb. Beyrich s.n. (holótipo W)
Frullaniaceae Frullania brasiliensis Raddi Peralta 27631 (SP)
Frullaniaceae Frullania cuencensis Taylor Peralta 27551 (SP)
Frullaniaceae Frullania ericoides (Nees) Mont. * Nees (1833, as Jungermannia)
Frullaniaceae Frullania kunzei (Lehm. & Lindb.) Lehm. & Lindb. Peralta 20492 (SP)
Frullaniaceae Frullania obcordata (Lehm. & Lindenb.) Lehm. & Lindenb. in Gott. et al. * Winter & Schaefer-Verwimp (2020) [25]
Frullaniaceae Frullania obscura (Sw.) Dumort. Peralta 20453 (SP)
Frullaniaceae Frullania riojaneirensis (Raddi) Spruce * Gottsche et al. (1844) [26]
Frullaniaceae Frullania schaefer-verwimpii Yuzawa & Hatt. Schafer-Verwimp 7375 (SP)
Frullaniaceae Frullania setigera Steph. Peralta 27501 (SP)
Frullaniaceae Frullania vitalii Yuzawa & Hatt. Yano 13575 (SP)
Geocalycaceae Saccogynidium caldense (Ångstr.) Grolle Peralta 27620 (SP)
Herbertaceae Herbertus acanthelius Spruce Peralta 27534 (SP)
Herbertaceae Herbertus juniperoideus (Sw.) Grolle Gissi 339 (SP)
Herbertaceae Herbertus pensilis (T. Taylor) Spruce Peralta 27612 (SP)
Herbertaceae Herbertus sendtneri (Nees) A.Evans * Fulford (1963 as Herbertus grossispinus) [27]
Herbertaceae Triandrophyllum subtrifidum (Hook.f. & Taylor) Fulford & Hatch. * Grolle (1964) [28]
Jamesoniellaceae Syzygiella anomala (Lindenb. & Gottsche) Steph. Lima 789 p.p. (SP)
Jamesoniellaceae Syzygiella colorata (Lehm.) K. Feldberg et al. Peralta 27575 (SP)
Jamesoniellaceae Syzygiella concreta (Gottsche) Spruce Peralta 20397 (SP)
Jamesoniellaceae Syzygiella contigua Steph. Peralta 20509 (SP)
Jamesoniellaceae Syzygiella manca (Mont.) J. B. Jack & Steph. Gonçalves 389 (SP)
Jamesoniellaceae Syzygiella perfoliata (Sw.) Spruce Peralta 20401 (SP)
Jamesoniellaceae Syzygiella sonderi (Gottsche) K. Feldberg, Váňa, Hentschel & Heinrichs Peralta 27502 (SP)
Jamesoniellaceae Syzygiella trigonifolia (Steph.) Herzog Peralta 27613 (SP)
Jamesoniellaceae Syzygiella uleana Steph. Peralta 20428 (SP)
Jungermanniaceae Anastrophyllum piligerum (Nees) Steph. Peralta 20380 (SP)
Jungermanniaceae Anastrophyllum tubulosum (Nees) Grolle Peralta 27565 (SP)
Jungermanniaceae Jungermannia amoena Lindenb. & Gottsche Gaspar 131 (R)
Jungermanniaceae Jungermannia hyalina Lyell Lima 792 (SP)
Jungermanniaceae Jungermannia sphaerocarpa Hook. Gaspar 122 (R)
Lejeuneaceae Acanthocoleus aberrans (Lindenb. & Gottsche) Kruijt Peralta 27465 (SP)
Lejeuneaceae Anoplolejeunea conferta (C.F.W.Meissn. ex Spreng.) A.Evans Peralta 27645 (SP)
Lejeuneaceae Blepharolejeunea incongrua (Lindenb. & Gottsche) van Slageren & Kruijt Peralta 27577 (SP)
Lejeuneaceae Brachiolejeunea phyllorhiza (Nees) Kruijt & Gradst. * Gottsche et al. (1844)
Lejeuneaceae Bryopteris diffusa (Sw.) Nees * Stotler & Crandall (1974) [29]
Lejeuneaceae Bryopteris filicina (Sw.) Nees Peralta 27530 (SP)
Lejeuneaceae Ceratolejeunea rubiginosa Gottsche ex Steph. Peralta 27681 (SP)
Lejeuneaceae Cheilolejeunea acutangula (Nees) Grolle Peralta 27523 (SP)
Lejeuneaceae Cheilolejeunea beyrichii (Lindenb.) E. Reiner Peralta 27492 (SP)
Lejeuneaceae Cheilolejeunea filiformis (Sw.) W. Ye, R.L. Zhu & Gradst. * Gottsche et al. (1844, as Omphalanthus) [26]
Lejeuneaceae Cheilolejeunea grosseoleosa C.J.Bastos & Schäf.-Verw. Peralta 27549 p.p. (SP)
Lejeuneaceae Cheilolejeunea holostipa (Spruce) Grolle & R.-L.Zhu Alkimim-Faria 1486
Lejeuneaceae Cheilolejeunea unciloba (Lindenb.) Malombe Lima 750 (SP)
Lejeuneaceae Cheilolejeunea xanthocarpa (Lehm. & Lindenb.) Malombe Peralta 27504 (SP)
Lejeuneaceae Cololejeunea cardiocarpa (Mont.) A.Evans Peralta 27687 p.p. (SP)
Lejeuneaceae Cololejeunea diaphana A.Evans Peralta 27687 p.p. (SP)
Lejeuneaceae Cololejeunea jamesii M.E.Reiner & Pócs Peralta 20511 p.p. (SP)
Lejeuneaceae Cololejeunea microscopica (Taylor) Schiffn. Peralta 27694 (SP)
Lejeuneaceae Cololejeunea papilliloba (Steph). Steph. Peralta 27695 (SP)
Lejeuneaceae Colura itatyana Steph. Gaspar 138 (R)
Lejeuneaceae Colura tenuicornis (A.Evans) Steph. * Jovet-Ast (1953) [30]
Lejeuneaceae Cyclolejeunea luteola (Spruce) Grolle & R.-L.Zhu Schafer-Verwimp 7416 (SP)
Lejeuneaceae Dibrachiella parviflora (Nees) X.Q. Shi, R.L. Zhu & Gradst. Silva, AL 500
Lejeuneaceae Dicranolejeunea axilaris (Nees & Mont.) Schiffn. Peralta 27735 (SP)
Lejeuneaceae Diplasiolejeunea alata Jovet-Ast Peralta 27593 p.p. (SP)
Lejeuneaceae Diplasiolejeunea unidentata (Lehm. & Lindenb.) Schiffn. * Gradstein & Costa (2003) [31]
Lejeuneaceae Drepanolejeunea anoplanta (Spruce) Steph. Peralta 27659 (SP)
Lejeuneaceae Drepanolejeunea araucariae Steph. Gaspar 135 (R)
Lejeuneaceae Drepanolejeunea bidens (Steph.) A.Evans Gaspar 117 (R)
Lejeuneaceae Drepanolejeunea fragilis Bischl. Peralta 27593 p.p. (SP)
Lejeuneaceae Drepanolejeunea lichenicola (Spruce) Steph. * Bischler (1964) [32]
Lejeuneaceae Drepanolejeunea mosenii (Steph.) Bischl. Peralta 27639 (SP)
Lejeuneaceae Frullanoides tristis van Slageren Peralta 27646 (SP)
Lejeuneaceae Haplolejeunea umbrosa Gradst. & Ilkiu-Borges Peralta 27495 (SP)
Lejeuneaceae Harpalejeunea oxyphylla (Nees & Mont.) Steph. Gaspar 124 (R)
Lejeuneaceae Harpalejeunea schiffneri S.W. Arnell Peralta 27462 (SP)
Lejeuneaceae Harpalejeunea stricta (Lindenb. & Gottsche) Steph. Peralta 27526 (SP)
Lejeuneaceae Harpalejeunea subacuta A.Evans Peralta 20450 p.p. (SP)
Lejeuneaceae Lejeunea acanthogona Spruce Peralta 27463 (SP)
Lejeuneaceae Lejeunea aphanes Spruce Lima 750 (SP)
Lejeuneaceae Lejeunea bermudiana (A.Evans) R.M.Schust. Peralta 27719 (SP)
Lejeuneaceae Lejeunea beyrichiana (Steph) Gradst. & C.J.Bastos * Beyrich s.n. (Tipo NY)
Lejeuneaceae Lejeunea cerina (Lehm. & Lindenb.) Gottsche Schaefer-Verwimp 7369 (SP)
Lejeuneaceae Lejeunea flaccida Lindenb. & Gottsche Peralta 27512 (SP)
Lejeuneaceae Lejeunea flava (Sw.) Nees Peralta 27505 (SP)
Lejeuneaceae Lejeunea glaucescens Gottsche Peralta 27476 (SP)
Lejeuneaceae Lejeunea grossitexta (Steph.) E.Reiner & Goda * Schäfer-Verwimp & Vital (1989) [33]
Lejeuneaceae Lejeunea laetevirens Nees & Mont. Peralta 27464 (SP)
Lejeuneaceae Lejeunea longidentata C.J.Bastos, Gradst., S. Vilas Bôas-Bastos & Schäf.-Verw. Peralta 27543 (SP)
Lejeuneaceae Lejeunea oligoclada Spruce Peralta 27498 p.p. (SP)
Lejeuneaceae Lejeunea pterigonia (Lehm. & Lindenb.) Mont. Peralta 27525 (SP)
Lejeuneaceae Lejeunea raddiana Lindenb. Peratla 20484 (SP)
Lejeuneaceae Lejeunea serpillifolioides (Raddi) Gradst. Peralta 27611 (SP)
Lejeuneaceae Lejeunea subplana (Steph.) C.J.Bastos Gonçalves 306 p.p. (SP)
Lejeuneaceae Marchesinia brachiata (Sw.) Schiffn. Peralta 27508 (SP)
Lejeuneaceae Metalejeunea cuculata (Reinw. et al.) Grolle Yano 13558 (SP)
Lejeuneaceae Microlejeunea cystifera Herzog Peralta 27482 (SP)
Lejeuneaceae Microlejeunea epiphylla Bischl. Gonçalves 306 p.p. (SP)
Lejeuneaceae Microlejeunea squarrosa J. Heinrichs, A. Schäfer-Verwimp, Pócs & S.S. Dong Peralta 27671 p.p. (SP)
Lejeuneaceae Microlejeunea stricta (Lindenb. & Gottsche) Steph. Lima 782 p.p. (SP)
Lejeuneaceae Neurolejeunea breutelii (Gottsche) A.Evans Lima 767 p.p. (SP)
Lejeuneaceae Otigoniolejeunea huctumalcensis (Lindenb. & Gottsche) Y.M. Wei, R.L. Zhu & Gradst. Peralta 27474 (SP)
Lejeuneaceae Prionolejeunea aemula (Gottsche) A.Evans Peralta 27727 (SP)
Lejeuneaceae Schiffneriolejeunea polycarpa (Nees) Gradst. Peralta 20468 (SP)
Lejeuneaceae Vitalianthus bischelerianus (Porto & Grolle) R.M.Schust. & Giancotti * Gottsche et al. (1844, as Phragmicoma juliformis)
Lepidoziaceae Bazzania aurescens Spruce Peralta 20394 (SP)
Lepidoziaceae Bazzania hookeri (Lindenb.) Trevis. Fulford (1963)
Lepidoziaceae Bazzania longistipula (Lindenb.) Trevis. Peralta 27480 (SP)
Lepidoziaceae Bazzania nitida (Weber) Grolle Peralta 27529 (SP)
Lepidoziaceae Bazzania stolonifera (Sw.) Trevis. * Nees (1833, as Jungermannia) [34]
Lepidoziaceae Kurzia capillaris (Sw.) Grolle Lima 780 (SP)
Lepidoziaceae Lepidozia coilophylla Taylor Peralta 27532 (SP)
Lepidoziaceae Lepidozia cupressina (Sw.) Lindenb. Gonçalves 311 (SP)
Lepidoziaceae Lepidozia inaequalis (Lehm. & Lindenb.) Lehm. & Lindenb. Peralta 20402 (SP)
Lepidoziaceae Lepidozia incurvata Lindenb. Lima 784 (SP)
Lepidoziaceae Lepidozia macrocolea Spruce Peralta 27539 (SP)
Lepidoziaceae Micropterygium leiophyllum Spruce * Gottsche et al. (1844, as M. vulgare)
Lepidoziaceae Micropterygium pterygophyllum (Nees) Trevis. Nees (1833, as Jungermannia)
Lepidoziaceae Paracromastigum dusenii R.M.Schust. * Dias 1310 (SP)
Lepidoziaceae Paracromastigum pachyrhizum (Nees) Fulford Schafer-Verwimp 7421 (SP)
Lepidoziaceae Telaranea nematodes (Gottsche ex Austin) M.A.Howe Lima 754 (SP)
Lophocoleaceae Lophocolea muricata (Lehm.) Nees in Gottsche et al. Peralta 27508 (SP)
Lophocoleaceae Clasmatocolea vermicularis (Lehm.) Grolle Peralta 27566 (SP)
Lophocoleaceae Cryptolophocolea martiana (Nees) L.Soderstr. et al. subsp. martiana Lima 768 (SP)
Lophocoleaceae Heteroscyphus combinatus (Nees) Schiffn. Peralta 27556 (SP)
Lophocoleaceae Leptoscyphus amphibolius (Nees) Grolle Peralta 27583 (SP)
Lophocoleaceae Leptoscyphus porphyrius (Nees) Grolle Peralta 27628 (SP)
Lophocoleaceae Leptoscyphus spectabilis (Steph.) Grolle Peralta 27658 (SP)
Lophocoleaceae Lophocolea bidentata (L.) Dumort. Peralta 20420 (SP)
Lophocoleaceae Lophocolea connata (Sw.) Nees * Nees (1833, as Jungermannia)
Lophocoleaceae Lophocolea mandonii Steph. Peralta 20379 (SP)
Marchantiaceae Marchantia chenopoda L. Vital 1197 (SP)
Marchantiaceae Marchantia polymorpha L. Schwacke, C.A.W. s.n. (R)
Marchantiaceae Marchantia paleacea Bert. Peralta 20475 (SP)
Metzgeriaceae Metzgeria acuminata Steph. Peralta 27471 (SP)
Metzgeriaceae Metzgeria albinea Spruce Peralta 27475 (SP)
Metzgeriaceae Metzgeria brasiliensis Schiffn. Dias 1234 (SP)
Metzgeriaceae Metzgeria ciliata Raddi * Raddi (1823) [35]
Metzgeriaceae Metzgeria conjugata Lindb. Silva, AL 517 (UB)
Metzgeriaceae Metzgeria consanguinea Schiffn. Peralta 27528 (SP)
Metzgeriaceae Metzgeria convoluta Steph. * Costa (2008) [36]
Metzgeriaceae Metzgeria cratoneura Schiffn. * Costa (2008) [36]
Metzgeriaceae Metzgeria decipiens (C. Massal.) Schiffn. Eiten 7105 (SP)
Metzgeriaceae Metzgeria fruticola Spruce Peralta 27715 (SP)
Metzgeriaceae Metzgeria furcata (L.) Dumort. * Bartram (1954) Gonçalves 326 (SP)
Metzgeriaceae Metzgeria hegewaldii Kuwah. Peralta 27466 (SP)
Metzgeriaceae Metzgeria lechlerii Steph. * Costa et al. (2005) [37]
Metzgeriaceae Metzgeria myriopoda Lindb. Yano 13561 (SP)
Noterocladaceae Noteroclada confluens (Hook.f. & Taylor) Spruce Peralta 20471 (SP)
Pallaviciniaceae Jensenia spinosa (Lindenb. & Gottsche) Grolle Gonçalves 400 (SP)
Pallaviciniaceae Pallavicinia lyelii (Hook.) S.F.Gray Gonçalves 186 (SP)
Pallaviciniaceae Symphyogyna brasiliensis (Nees) Nees & Mont. Peralta 27750 (SP)
Pallaviciniaceae Symphyogyna podophylla (Thunb.) Mont. & Nees Silva, AL 502
Plagiochilaceae Plagiochila amicta Steph. * Heinrichs (2002) [38]
Plagiochilaceae Plagiochila bicuspidata Gottsche Peralta 10518 (SP)
Plagiochilaceae Plagiochila bifaria (Sw.) Lindenb. Peralta 27516 (SP)
Plagiochilaceae Plagiochila boryana Gottsche ex Steph. Peralta 27749 (SP)
Plagiochilaceae Plagiochila corrugata (Nees) Nees & Mont. Peralta 20376 (SP)
Plagiochilaceae Plagiochila crispabilis Lindenb. Peralta 20511 (SP)
Plagiochilaceae Plagiochila cristata (Sw.) Lindenb. Schafer-Verwimp 7401 (SP)
Plagiochilaceae Plagiochila deflexirama Taylor Peralta 27721 (SP)
Plagiochilaceae Plagiochila exigua (Taylor) Taylor Peralta 20468 (SP)
Plagiochilaceae Plagiochila flabelliflora Steph. Gonçalves 334 (SP)
Plagiochilaceae Plagiochila gymnocalycina (Lehm. & Lindenb.) Lindenb. Dias 1397 (SP)
Plagiochilaceae Plagiochila loriloba Herzog ex Carl Perlata 27519 (SP)
Plagiochilaceae Plagiochila macrotachya Lindenb. Peralta 27700 (SP)
Plagiochilaceae Plagiochila martiana (Nees) Lindenb. Peralta 20385 (SP)
Plagiochilaceae Plagiochila montagnei Nees & Mont. * Heinrichs & Gradstein (2000) [39]
Plagiochilaceae Plagiochila pseudoradicans Herzog * Ph. v. Lützelburg 6029b (type JE)
Plagiochilaceae Plagiochila punctata (Taylor) Taylor Lima 783 p.p. (SP)
Plagiochilaceae Plagiochila rutilans Lindenb. * Heinrichs (2002) [38]
Plagiochilaceae Plagiochila simplex (Sw.) Lindenb. Peralta 27780 (SP)
Plagiochilaceae Plagiochila subbidentata Taylor Peralta 27586 (SP)
Plagiochilaceae Plagiochila superba d’Orbigny Peralta 20457 (SP)
Porellaceae Porella brasiliensis (Raddi) Schiffn. Peralta 27724 (SP)
Porellaceae Porella reflexa (Lehm. & Lindenb.) Trevis. Yano 13580 (SP)
Radulaceae Radula angulata Steph. Peralta 27697 (SP)
Radulaceae Radula javanica Gottsche Peralta 27527 (SP)
Radulaceae Radula ligula Steph. Peralta 20407 (SP)
Radulaceae Radula nudicaulis Steph. Gonçalves 408 (SP)
Radulaceae Radula pallens (Sw.) Nees ex Mont Lima 759 (SP)
Radulaceae Radula recubans Taylor Gaspar 142 (R)
Radulaceae Radula schaefer-verwimpii K.Yamada Peralta 27489 (SP)
Radulaceae Radula stenocalyx Mont. Peralta 27691 (SP)
Radulaceae Radula subinflata Lindenb. & Gottsche Gonçalves 322 (SP)
Radulaceae Radula tectiloba Steph. Peralta 27507 (SP)
Radulaceae Radula voluta Taylor ex Gottsche, Lindenb. & Nees Peralta 20429 (SP)
Scapaniaceae Scapania portoricensis Hampe & Gottsche Peralta 27541 (SP)
Trichocoleaceae Trichocolea brevifissa Steph. Peralta 27503 (SP)
Mosses Adelotheciaceae Adelothecium bogotense (Hampe) Mitt. Peralta 27692 (SP)
Andreaceae Andreaea acutifolia Hook.f. & Wilson * Müller (1845) Gonçalves 401 (SP)
Andreaceae Andreaea rupestris Hedw. Gaspar 137 (R)
Andreaceae Andreaea subulata Harv. * Lützelburg, P. von 7041 (tipo JE of Andreaea hamulata)
Bartramiaceae Bartramia mathewsii Mitt. Peralta 27606 (SP)
Bartramiaceae Breutelia grandis (Hampe) Paris Peralta 27559 (SP)
Bartramiaceae Breutelia tomentosa (Sw. ex Brid.) A.Jaeger Lima 798 (SP)
Bartramiaceae Leiomela bartramioides (Hook.) Paris Peralta 27500 (SP)
Bartramiaceae Leiomela piligera (Hampe) Broth. * Brotherus (1924) [40]
Bartramiaceae Philonotis cernua (Wilson) Griffin & W.R.Buck Peralta 20426 (SP)
Bartramiaceae Philonotis hastata (Duby) Wijk & Margad. Alkimim-Faria, 1513
Bartramiaceae Philonotis uncinata (Schwägr.) Brid. * Brotherus (1894)
Brachytheciaceae Aerolindigia capillacea (Hornsch.) M.Menzel Peralta 20418 (SP)
Brachytheciaceae Meteoridium remotifolium (Müll.Hal.) Manuel Peralta 27763 (SP)
Brachytheciaceae Rhynchostegium scariosum (Taylor) A.Jaeger Peralta 20419 (SP)
Brachytheciaceae Rhynchostegium serrulatum (Hedw.) A.Jaeger Gonçalves 333 p.p. (SP)
Brachytheciaceae Squamidium brasiliense Broth. * Allen & Crosby (1986) [41]
Brachytheciaceae Squamidium leucotrichum (Taylor) Broth. Peralta 27486 (SP)
Brachytheciaceae Zelometeorium ambiguum (Hornsch.) Manuel * Manuel (1977) [42]
Brachytheciaceae Zelometeorium patulum (Hedw.) Manuel Gonçalves 332 (SP)
Bruchiaceae Trematodon longicollis Michx. * Brotherus (1924b) [43]
Bryaceae Anomobryum julaceum (Schrad. ex P.Gaertn. et al.) Schimp. Peralta 20377 (SP)
Bryaceae Brachymenium acuminatum Harv. Dias 1372 (SP)
Bryaceae Brachymenium consimile (Mitt.) A.Jaeger Peralta 27654 (SP)
Bryaceae Brachymenium hornschuchianum Mart. * Brotherus (1894) [44]
Bryaceae Brachymenium radiculosum (Schwägr.) Hampe * Brotherus (1924b) [43]
Bryaceae Brachymenium systilium (Müll.Hal.) A.Jaeger Peralta 27673 (SP)
Bryaceae Bryum argenteum Hedw. Vital 1191 (SP)
Bryaceae Bryum atenense Williams Alkimim-Faria 1246 (UB)
Bryaceae Bryum billarderii Schwaegr. Dias 1306a (SP)
Bryaceae Bryum capillare Hedw. Peralta 20465 (SP)
Bryaceae Bryum densifolium Brid. * Bartram (1954, as B. gracilescens)
Bryaceae Bryum huillense Welw. & Duby Peralta 20415 (SP)
Bryaceae Bryum limbatum Müll.Hal * Gonçalves 384 (SP)
Bryaceae Bryum pabstianum Müll.Hal. * Hampe (1879) [45]
Bryaceae Bryum subapiculatum Hampe Peralta 20462 (SP)
Bryaceae Rhodobryum aubertii (Schwägr.) Thér. Peralta 27762 (SP)
Bryaceae Rhodobryum beyrichianum (Hornsch.) Müll.Hal. Peralta 20514 (SP)
Bryaceae Rhodobryum roseum (Hedw.) Limpr. Peralta 27582 (SP)
Bryaceae Rhodobryum subverticillatum Broth. * Bartram (1952) [46]
Bryaceae Schoenobryum concavifolium (Griff.) Gangulee * Bartram (1954) [19]
Calymperaceae Calymperes nicaraguense Renauld & Cardot Alkimim-Faria 1256
Calymperaceae Syrrhopodon gaudichaudii Mont. * Bartram (1954) [19]
Calymperaceae Syrrhopodon helicophyllus Mitt. Alkimim-Faria, AL 1254
Calymperaceae Syrrhopodon ligulatus Mont. Peralta 27490 (SP)
Calymperaceae Syrrhopodon prolifer Schwägr. Peralta 27473 (SP)
Catagoniaceae Catagonium brevicaudatum Müll.Hal. ex Broth. Peralta 27630 (SP)
Daltoniaceae Calyptrochaeta albescens (Hampe) W.R.Buck Lima 775 (SP)
Daltoniaceae Calyptrochaeta setigera (Mitt.) W.R.Buck Peralta 27702 (SP)
Daltoniaceae Daltonia splachnoides (Sm.) Hook. & Taylor Peralta 27546 (SP)
Daltoniaceae Leskeodon aristatus (Geh. & Hampe) Broth. Peralta 27517 (SP)
Dicranaceae Aongstroemia julacea (Hook.) Mitt. Peralta 27550 (SP)
Dicranaceae Atractylocarpus brasiliensis (Müll.Hal.) R.S.Williams Peralta 20490 (SP)
Dicranaceae Atractylocarpus longisetus (Hook.) E.B.Bartram * Bartram (1954) [19]
Dicranaceae Dicranella harrisii Müll.Hal.) Broth. Schafer-Verwimp 13101 (SP)
Dicranaceae Dicranella hilariana (Mont.) Mitt. * Luetzelburg (1923, as D. trematodontifolia) [47]
Dicranaceae Dicranella subsulcata (Hampe) Hampe Schafer-Verwimp 7422 (SP)
Dicranaceae Dicranodontium pulchroalare Broth. * Lützelburg 1519 (holótipo JE)
Dicranaceae Dicranoloma billardieri (Brid. ex Anon) Paris Peralta 27597 (SP)
Dicranaceae Dicranum frigidum Müll.Hal. Dias 1361 (SP)
Dicranaceae Holomitrium crispulum Mart. Peralta 27599 (SP)
Dicranaceae Leucoloma cruegerianum (Müll.Hal.) A.Jaeger Peralta 27678 (SP)
Dicranaceae Leucoloma serrulatum Brid. Peralta 27701 (SP)
Dicranaceae Leucoloma triforme (Mitt.) A.Jaeger Gonçalves 190 (SP)
Dicranaceae Microcampylopus curvisetus (Hampe) Giese & J.-P.Frahm Schafer-Verwimp 7422 p.p. (SP)
Dicranaceae Paraleucobryum longifolium (Hedw.) Loeske Dias 1325 (SP)
Dicranaceae Pilopogon guadalupensis (Brid.) J.-P.Frahm Peralta 1558 (SP)
Diphysciaceae Diphyscium longifolium Griff. Peralta 27723 (SP)
Ditrichaceae Ceratodon purpureus (Hedw.) Brid. Dias 1315 (SP)
Ditrichaceae Cladastomum robustum Broth. Lima 485 (SP)
Ditrichaceae Cladastomum ulei Müll.Hal. Peralta 27635 (SP)
Ditrichaceae Crumuscus vitalis W.R.Buck & Snider Lima 791 (SP)
Ditrichaceae Ditrichum crinale (Taylor) Kuntze Lima 801 (SP)
Ditrichaceae Ditrichum paulense Geh. ex Hampe Peralta 27656 (SP)
Entodontaceae Entodon hampeanus Müll.Hal. Peralta 27614 (SP)
Entodontaceae Entodon jamesonii (Taylor) Mitt. Peralta 27580 (SP)
Entodontaceae Erythrodontium longisetum (Hook.) Paris * Vattimo-Gil & Vattimo (1980) [48]
Entodontaceae Erythodontium squarrosum (Hampe) Paris * Brotherus (1924)
Ephemeraceae Micromitrium tenerum (Bruch & Schimp.) Crosby Alkimim-Faria, AL 1379 (UB)
Fabroniaceae Dimerodontium pellucidum Schwägr. Schafer-Verwimp 13113 (SP)
Fabroniaceae Fabronia ciliaris (Brid.) Brid. Gonçalves 182 (SP)
Fissidentaceae Fissidens anguste-limbatus Mitt. Peralta 27665 (SP)
Fissidentaceae Fissidens asplenioides Hedw. Peralta 27709 (SP)
Fissidentaceae Fissidens bryoides Hedw. Lima 787 (SP)
Fissidentaceae Fissidens crispus Mont. Gonçalves 180 (SP)
Fissidentaceae Fissidens elegans Brid. Peralta 27479 (SP)
Fissidentaceae Fissidens flabellatus Hornsch. * Beyrich s.n. (lectótipo BM)
Fissidentaceae Fissidens intromarginatus (Hampe) Mitt. * Ule 1225 (lectotype H, tipo de F. hemibryoides)
Fissidentaceae Fissidens lagenarius Mitt. * Bartram (1954)
Fissidentaceae Fissidens oblongifolius Hook.f. & Wilson Peralta 20464 (SP)
Fissidentaceae Fissidens oediloma Müll.Hal. ex Broth. Peralta 27571 (SP)
Fissidentaceae Fissidens pellucidus Hornsch. Peralta 27538 (SP)
Fissidentaceae Fissidens prionodes Mont. Peralta 27740 (SP)
Fissidentaceae Fissidens radicans Mont. Peralta 27737 (SP)
Fissidentaceae Fissidens rigidulus Hook.f. & Wilson Peralta 27756 (SP)
Fissidentaceae Fissidens saprofilus Broth. Lima 754 (SP)
Fissidentaceae Fissidens scariosus Mitt. Lima 756 (SP)
Fissidentaceae Fissidens semicompletus Hedw. Alkimim-Faria 1228
Fissidentaceae Fissidens taxifolius Hedw. Alkimim-Faria 1367
Fissidentaceae Fissidens wallisii Müll.Hal. ex Broth. Peralta 27675 (SP)
Fissidentaceae Fissidens weirii Mitt. Peralta 27484 (SP)
Funariaceae Entosthodon bonplandii (Hook.) Mitt. Schaefer-Verwimp 7405 (SP)
Funariaceae Funaria hygrometrica Hedw. * Müller (1845) [49]
Grimmiaceae Grimmia laevigata (Brid.) Brid. Dias 1298 (SP)
Grimmiaceae Grimmia longirostris Hook.f. & Wilson Gaspar 110 (R)
Grimmiaceae Racomitrium subsecundum (Hook. & Grev. ex Harv.) Mitt. Dias 1405 (SP)
Hedwigiaceae Hedwigidium rhabdocarpum (Hampe) A.Jaeger Faria 1547 (SP)
Hookeriaceae Hookeria acutifolia Hook. & Grev. Peralta 27739 (SP)
Hookeriaceae Philophyllum tenuifolium (Mitt.) Broth. Peralta 27581 (SP)
Hypnaceae Chrysohypnum diminutivum (Mitt.) Broth. Peralta 20478 (SP)
Hypnaceae Ectropothecium leptochaeton (Schwägr.) W.R.Buck Peralta 20461 (SP)
Hypnaceae Mittenothamnium reduncum (Mitt.) Ochyra Peralta 27624 (SP)
Hypnaceae Mittenothamnium reptans (Hedw.) Cardot Eiten 7133 (SP)
Hypnaceae Taxiphyllum taxirameum (Mitt.) M. Fleisch. Gonçalves 194 (SP)
Hypnaceae Vesicularia vesicularis (Schwägr.) Broth. Alkimim-Faria, 1364
Hypopterygiaceae Hypopterygium tamariscinum (Hedw.) Brid. Yano 13560 (SP)
Hypopterygiaceae Lopidium concinnum (Hook.) Wilson Peralta 27712 (SP)
Lembophyllaceae Orthostichella versicolor (Müll.Hal.) B.H.Allen & W.R.Buck Peralta 27520 (SP)
Leucobryaceae Campylopus aemulans (Hampe) A.Jaeger * Frahm (1975) [50]
Leucobryaceae Campylopus arctocarpus (Hornsch.) Mitt. Peralta 27547 (SP)
Leucobryaceae Campylopus cryptopodioides Broth. * Frahm (1991) [15]
Leucobryaceae Campylopus cuspidatus (Hornsch.) Mitt. * Frahm (1975) [50]
Leucobryaceae Campylopus densicoma (Müll.Hal.) Paris Gonçalves 409 (SP)
Leucobryaceae Campylopus dichrostis (Müll.Hal.) Paris Gonçalves 330 (SP)
Leucobryaceae Campylopus filifolius (Müll.Hal.) Paris Lima 789 (SP)
Leucobryaceae Campylopus fragilis (Brid.) Bruch & Schimp. Peralta 27610 (SP)
Leucobryaceae Campylopus gemmatus (Müll.Hal.) Paris Vital 7634 (SP)
Leucobryaceae Campylopus heterostachys (Hampe) A.Jaeger Dias 1299 (SP)
Leucobryaceae Campylopus introflexus (Hedw.) Brid. Alkimim-Faria 1510
Leucobryaceae Campylopus julicaulis Broth. Gaspar 104 (R)
Leucobryaceae Campylopus lamellinervis (Müll.Hal.) Mitt. Müller 1845
Leucobryaceae Campylopus occultus Mitt. Frahm sn (SP147028)
Leucobryaceae Campylopus pilifer Brid. Peralta 27605 (SP)
Leucobryaceae Campylopus pyriformis (Schultz) Brid. Gaspar 114 (R)
Leucobryaceae Campylopus richardii Brid. Dias 1324 (SP)
Leucobryaceae Campylopus savannarum (Müll.Hal.) Mitt. Gaspar 139 (R)
Leucobryaceae Campylopus subcuspidatus (Hampe) A.Jaeger Peralta 27641 (SP)
Leucobryaceae Campylopus thwaitesii (Mitt.) A.Jaeger Gonçalves 183 (SP)
Leucobryaceae Campylopus trachyblepharon (Müll.Hal.) Mitt. Paula sn (SP463831)
Leucobryaceae Campylopus uleanus (Müll.Hal.) Broth. Dias 1384 (SP)
Leucobryaceae Campylopus widgrenii (Müll.Hal.) Mitt. Vital 1189 (SP)
Leucobryaceae Leucobryum albicans (Schwägr.) Lindb. Peralta 27522 (SP)
Leucobryaceae Leucobryum clavatum Hampe * Luetzelburg 1923 [47]
Leucobryaceae Leucobryum crispum Müll.Hal. Dias 1235 (SP)
Leucobryaceae Leucobryum giganteum Müll.Hal. * Herzog (1926) [51]
Leucobryaceae Leucobryum sordidum Ångström * Luetzelburg (1923) [47]
Leucobryaceae Ochrobryum gardneri (Müll.Hal.) Lindb. Alkimim-Faria 1364 (UB)
Leucobryaceae Octoblepharum albidum Hedw. * Bartram (1954) [19]
Leucomiaceae Leucomium strumosum (Hornsch.) Mitt. Peralta 27769 (SP)
Meteoriaceae Meteorium deppei (Hornsch.) Mitt. Peralta 27573 (SP)
Meteoriaceae Meteorium nigrescens (Hedw.) Dozy & Molk. * Brotherus (1924)
Meteoriaceae Meteorium pseudoteres W.R.Buck Peralta 27668 (SP)
Meteoriaceae Pilotrichella flexilis (Hedw.) Ångström Peralta 20491 (SP)
Meteoriaceae Toloxis imponderosa (Taylor) W.R.Buck Peralta 20517 (SP)
Mniaceae Epipterygium immarginatum Mitt. Peralta 20432 (SP)
Mniaceae Plagiomnium rhynchophorum (Hook.) T.J.Kop. Peralta 20411 p.p. (SP)
Mniaceae Pohlia camptotrachela (Renauld & Cardot) Broth. Peralta 20433 (SP)
Mniaceae Pohlia elongata Hedw. Lima 790 (SP)
Mniaceae Pohlia papilosa (Müll.Hal. ex A.Jaeger) Broth. Faria 1554 (SP)
Mniaceae Pohlia tenuifolia (A.Jaeger) Broth. Peralta 27579 (SP)
Mniaceae Schizymenium campylocarpum (Arn. & Hook.) A.J.Shaw Lima 761 (SP)
Mniaceae Schizymenium pusillum (Hook.f. & Wilson) A.J.Shaw Dias 1382 (SP)
Neckeraceae Homalia glabella (Hedw.) Schimp. Gonçalves 310 (SP)
Neckeraceae Homaliodendron flabellatum (Sm.) M.Fleisch. Peralta 27569 (SP)
Neckeraceae Neckera ehrenbergii Müll.Hal. Peralta 27588 (SP)
Neckeraceae Porotrichum korthalsianum (Dozy & Molk.) Mitt. Dias 1420 (SP)
Neckeraceae Porotrichum leucocaulon Müll.Hal. Peralta 27674 (SP)
Neckeraceae Porotrichum longirostre (Hook.) Mitt. Peralta 20387 p.p. (SP)
Neckeraceae Porotrichum superbum (Taylor) Hampe Peralta 27585 (SP)
Neckeraceae Porotrichum thieleanum (Müll.Hal.) Mitt. Peralta 20383 (SP)
Neckeraceae Thamnobryum fasciculatum (Hedw.) I.Sastre Peralta 27703 (SP)
Orthotrichaceae Macrocoma orthotrichoides (Raddi) Wijk & Margad. Schafer-Verwimp 7449 (SP)
Orthotrichaceae Macromitrium catharinense Paris * Li et al. (2019) [52]
Orthotrichaceae Macromitrium cirrosum (Hedw.) Brid. * Bartram (1954)
Orthotrichaceae Macromitrium microstomum (Hook. & Grev.) Schwägr. Gonçalves 395 (SP)
Orthotrichaceae Orthotrichum papillosum (Hampe) Lesquereux & James Alkimim-Faria, 1236
Orthotrichaceae Schlotheimia appressifolia Mitt. Atwood 2007
Orthotrichaceae Schlotheimia capillaris Hampe Peralta 20376 p.p. (SP)
Orthotrichaceae Schlotheimia rugifolia (Hook.) Schwägr. Lima 753 (SP)
Orthotrichaceae Schlotheimia tecta Hook.f. & Wilson Gonzatti 4003 (SP)
Orthotrichaceae Zygodon reiwardtii (Hornsch.) A.Braun Peralta 27629 (SP)
Phylogoniaceae Phyllogonium viride Brid. Gonçalves 320 (SP)
Pilotrichaceae Callicostella martiana (Hornsch.) A.Jaeger Gonçalves 197 (SP)
Pilotrichaceae Callicostella pallida (Hornsch.) Ångström Peralta 27689 (SP)
Pilotrichaceae Cyclodictyon albicans (Hedw.) Kuntze Peralta 27587 (SP)
Pilotrichaceae Cyclodictyon marginatum (Hook. & Wilson) Kuntze Schafer-Verwimp 13124 (SP)
Pilotrichaceae Cyclodictyon varians (Sull.) Kuntze Peralta 27619 (SP)
Pilotrichaceae Lepidopilidium brevisetum (Hampe) Broth. Silva, AL 481A (UB)
Pilotrichaceae Lepidopilidium nitens (Hornsch.) Broth. Peralta 27764 (SP)
Daltoniaceae Lepidopilum affine Müll.Hal. ex Broth. Alkimim-Faria, 1360 (UB)
Pilotrichaceae Lepidopilum muelleri (Hampe) Mitt. Peralta 20409 (SP)
Pilotrichaceae Lepidopilum scabrisetum (Schwägr.) Steere Peralta 27666 (SP)
Pilotrichaceae Lepidopilum subsubulatum Geh. & Hampe Peralta 27521 (SP)
Pilotrichaceae Thamniopsis incurva (Hornsch.) W.R.Buck Yano 13564 (SP)
Pilotrichaceae Thamniopsis langsdorffii (Hook.) W.R.Buck Gonçalves 307 (SP)
Pilotrichaceae Trachyxiphium guadalupense (Brid.) W.R.Buck Peralta 27497 p.p. (SP)
Plagiotheciaceae Plagiothecium novogranatense (Hampe) Mitt. Peralta 27570 (SP)
Polytrichaceae Atrichum androgynum (Müll.Hal.) A.Jaeger Gonçalves 195 (SP)
Polytrichaceae Itatiella ulei (Broth. ex Müll.Hal.) G.L.Sm. Peralta 27634 (SP)
Polytrichaceae Oligotrichum riedelianum Mitt. Peralta 27594 (SP)
Polytrichaceae Pogonatum campylocarpum (Müll.Hal.) Mitt. Peralta 20439 (SP)
Polytrichaceae Pogonatum pensilvanicum (E.B.Bartram ex Hedw.) P.Beauv. Paula, J.C. sn (SP463829)
Polytrichaceae Pogonatum perichaetiale (Mont.) A.Jaeger Alkimim-Faria 1236 (UB)
Polytrichaceae Polytrichadelphus pseudopolytrichum (Raddi) G.L.Sm. Eiten 7156 (SP)
Polytrichaceae Polytrichum angustifolium Mitt. Peralta 20378 (SP)
Polytrichaceae Polytrichum commune L. ex Hedw. Goeldi sn (SP219763)
Polytrichaceae Polytrichum juniperinum Willd. ex Hedw. Faria 1562 (SP)
Pottiaceae Barbula indica (Hook.) Spreng. Peralta 27774 (SP)
Pottiaceae Chionoloma angustatum (Mitt.) M.Menzel Peralta 27677 (SP)
Pottiaceae Chionoloma fractum M.J.Cano, J.A.Jiménez & M.Alonso Peralta 27589 (SP)
Pottiaceae Hyophila involuta (Hook.) A.Jaeger Silva, AL 499 (UB)
Pottiaceae Hyophiladelphus agrarius (Hedw.) R.H.Zander * Bartram (1954)
Pottiaceae Leptodontium araucarieti (Müll.Hal.) Paris Gonçalves 413 (SP)
Pottiaceae Leptodontium filicicola Herzog Peralta 27669 (SP)
Pottiaceae Leptodontium flexifolium (Dicks.) Hampe Dias 1276 (SP)
Pottiaceae Leptodontium pungens (Mitt.) Kindb. Dias 1377 (SP)
Pottiaceae Leptodontium stellatifolium (Hampe) Broth. Peralta 27595 (SP)
Pottiaceae Leptodontium viticulosoides (P.Beauv.) Wijk & Margad. Peralta 27607 (SP)
Pottiaceae Leptodontium wallisii (Müll.Hal.) Kindb. Gonçalves 364 (SP)
Pottiaceae Streptopogon calymperes Müll.Hal. Peralta 27450 (SP)
Pottiaceae Tortella humilis (Hedw.) Jenn. Peralta 20417 (SP)
Pottiaceae Tortella tortuosa (Hedw.) Limpr. Lima 806 (SP)
Priodontaceae Prionodon densus (Hedw.) Müll.Hal. Yano 13579 (SP)
Pterobryaceae Orthostichopsis tortipilis (Müll.Hal.) Broth. Peralta 27515 (SP)
Ptychomitriaceae Ptychomitrium sellowianum (Müll.Hal.) A.Jaeger Yano 12555 (SP)
Pylaisiadelphaceae Isopterygium tenerifolium Mitt. Peralta 20516 (SP)
Pylaisiadelphaceae Isopterygium tenerum (Sw.) Mitt. Eiten 7100
Racopilaceae Racopilum tomentosum (Hedw.) Brid. Alkimim-Faria 1402 (UB)
Rhacocarpaceae Rhacocarpus inermis (Müll.Hal.) Lindb. Vital 1192 (SP)
Rhacocarpaceae Rhacocarpus purpurascens (Brid.) Müll.Hal. Peralta 27643 (SP)
Rhizogoniaceae Pyrrhobryum spiniforme (Hedw.) Mitt. Gonçalves 314 (SP)
Rhizogoniaceae Rhizogonium novae-hollandiae (Brid.) Brid. Schaefer-Verwimp 13122 (SP)
Rigodiaceae Rigodium toxarion (Schwägr.) A.Jaeger. Peralta 27537 (SP)
Seligeriaceae Brachydontium notorogenes W.R.Buck & Schaf.-Verw. Peralta 20441 (SP)
Sematophyllaceae Aptychella proligera (Broth.) Herzog Peralta 27650 (SP)
Sematophyllaceae Aptychopsis pungifolia (Hampe) Broth. Gonçalves 200 (SP)
Sematophyllaceae Aptychopsis pyrrhophylla (Müll.Hal.) Wijk & Margad. Peralta 27485 (SP)
Sematophyllaceae Brittonodoxa subpinnata (Brid.) W.R.Buck, P.E.A.S.Câmara & Carv.-Silva * Müller (1845)
Sematophyllaceae Donnellia commutata (Müll.Hal.) W.R.Buck Alkimim-Faria 1342 (UB)
Sematophyllaceae Donnellia lagenifera (Müll.Hal.) W.R.Buck Gonzatti 4005 (SP)
Sematophyllaceae Microcalpe subsimplex (Hedw.) W.R.Buck * Müller (1845)
Sematophyllaceae Kuerschneria laevigata (Herzog) Ochyra & Bedn.-Ochyra * Ochyra & Bednarek Ochyra (2010) [53]
Sematophyllaceae Sematophyllum beyrichii (Hornsch.) Broth. Gonçalves 331 (SP)
Sematophyllaceae Sematophyllum cuspidiferum Mitt. Peralta 20495 (SP)
Sematophyllaceae Sematophyllum decumbens Mitt. Peralta 27494 (SP)
Sematophyllaceae Sematophyllum galipense (Müll.Hal.) Mitt. Peralta 27748 (SP)
Sematophyllaceae Sematophyllum lithophilum (Hornsch.) Ångström * Buck (1998) [54]
Sematophyllaceae Sematophyllum subdepressum (A.Jaeger) Broth. * Brotherus (1924)
Sematophyllaceae Sematophyllum subpinnatum (Brid.) E.Britton Peralta 27743 (SP)
Sematophyllaceae Trichosteleum glaziovii (Hampe) W.R.Buck Peralta 27514 (SP)
Sematophyllaceae Trichosteleum sentosum (Sull.) A.Jaeger Peralta 27497 p.p. (SP)
Sematophyllaceae Vitalia cuspidifera (Mitt.) P.E.A.S.Câmara, Carv.-Silva & W.R.Buck Sampaio, A.J. 2380
Sematophyllaceae Vitalia galipensis (Müll.Hal.) P.E.A.S.Câmara, Carv.-Silva & W.R.Buck Lima 768 p.p. (SP)
Sphagnaceae Sphagnum aciphyllum Müll.Hal Peralta 27592 (SP)
Sphagnaceae Sphagnum brasiliense Warnst. Santos & Costa 759, 760 (RB)
Sphagnaceae Sphagnum brevirameum Hampe Gonçalves 167 (SP)
Sphagnaceae Sphagnum capillifolium (Ehrh.) Hedw. Gonçalves 170 (SP)
Sphagnaceae Sphagnum costae H.A.Crum & D.P.Costa Peralta 20494 (SP)
Sphagnaceae Sphagnum cuspidatum Ehrh. ex Hoffm. * Luetzelburg (1923, as S. serratum) Gonçalves 369 (SP)
Sphagnaceae Sphagnum divisum H.A.Crum Gonçalves 419 (SP)
Sphagnaceae Sphagnum exquisitum H.A.Crum Lima 822 (SP)
Sphagnaceae Sphagnum geraisense H.A.Crum Gonçalves 421 (SP)
Sphagnaceae Sphagnum gracilescens Hampe ex. Müll.Hal. Peralta 27651 (SP)
Sphagnaceae Sphagnum luetzelburgii H.K.G.Paul ex H.A.Crum Costa 4173 (RB)
Sphagnaceae Sphagnum longicomosum Müll.Hal. ex Warnst. Gonçalves 335 (SP)
Sphagnaceae Sphagnum longistolo Müll.Hal. Peralta 27553 (SP)
Sphagnaceae Sphagnum magellanicum Brid. Lima 824 (SP)
Sphagnaceae Sphagnum palustre L. * Andrews (1941) [55]
Sphagnaceae Sphagnum perforatum Warnst. Costa et al. 5017 (RB)
Sphagnaceae Sphagnum perichaetiale Hampe Peralta 27652 (SP)
Sphagnaceae Sphagnum platyphylloides Warnst. Gonçalves 415 (SP)
Sphagnaceae Sphagnum pseudoramulinum H.A.Crum Lima 817 (SP)
Sphagnaceae Sphagnum ramulinum Warnst. Lima 815 (SP)
Sphagnaceae Sphagnum recurvum P.Beauv. Peralta 27560 (SP)
Sphagnaceae Sphagnum sanguinale Warnst. Gonçalves 168 (SP)
Sphagnaceae Sphagnum sehnemii H.A.Crum Luetzelburg 6162 (F, HBR as Sphagnum densum)
Sphagnaceae Sphagnum subsecundum Nees Gonçalves 353 (SP)
Sphagnaceae Sphagnum sucrei H.A.Crum Peralta 27598 (SP)
Stereophyllaceae Entodontopsis leucostega (Brid.) W.R.Buck & Ireland Alkimim-Faria 1456 (UB)
Stereophyllaceae Entodontopsis nitens (Mitt.) W.R.Buck & Ireland Alkimim-Faria 1263 (UB)
Stereophyllaceae Eulacophyllum cultelliforme (Sull.) W.R.Buck & Ireland * Costa et al. (2005) [37]
Thuidiaceae Thuidium pseudoprotensum (Müll.Hal.) Mitt. Peralta 27616 (SP)
Thuidiaceae Thuidium recognitum (Hedwig) Lindberg Lima 797 (SP)
Thuidiaceae Thuidium tamariscinum (Hedw.) Schimp. Gonçalves 359 (SP)
Thuidiaceae Thuidium tomentosum Schimp. Peralta 20510 (SP)
Thuidiaceae Pelekium involvens (Hedw.) A. Touw Lima 751 (SP)
Thuidiaceae Pelekium minutulum (Hedw.) A.Touw Lima 746 (SP)
Thuidiaceae Pelekium muricatulum (Hampe) A.Touw Alkimim-Faria 1376 (UB)
Thuidiaceae Pelekium scabrosulum (Mitt.) A.Touw Alkimim-Faria 1388 (UB)
Hornworts Anthocerotaceae Anthoceros hispidus Steph. Yano 13588 (SP)
Anthocerotaceae Anthoceros punctatus L. * Hooker & Wilson (1844) [56]
Dendrocerotaceae Dendroceros crispatus (Hook.) Nees Peralta 27459 (SP)
Dendrocerotaceae Dendroceros crispus (Sw.) Nees * Gottsche et al. (1844)
Dendrocerotaceae Nothoceros minarum (Nees) J.C.Villarreal Gonçalves 178 SP)
Dendrocerotaceae Nothoceros vincentianus (Lehm. & Lindenb.) J.C. Villareal Peralta 27705 (SP)
Notothyladaceae Phaeoceros bulbiculosus (Broth.) Prosk. Peralta 20452 (SP)
Nothotyladaceae Phaeoceros laevis (L.) Prosk. Peralta 27542 (SP)
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Three species are locally endemic, being known only from the type specimens collected over 100 years ago: Southbya organensis (Lützelburg 6003 p.p. [holotype JE]), Plagiochila pseudoradicans (Lützelburg 6029b [holotype JE]) and Dicranodontium pulchroalare ssp. brasiliense (Lützelburg 1519 [holotype JE]). Additional field research is urgently needed in order to obtain new records and data of these three endemic species.

The most species-rich families of liverworts are Lejeuneaceae (65 spp.) and Lepidoziaceae (16 spp.), while for moss they are Leucobryaceae (30 spp.) and Sphagnaceae (25 spp.). A total of 8 species of hornworts were found, distributed among the families Dendrocerotaceae (4 spp.), Anthocerotaceae (2 spp.) and Nothotyladaceae (2 spp.). The most species-rich genera were Campylopus (Leucobryaceae, 23 spp.); Plagiochila (Plagiochilaceae, 21 spp.); Fissidens (Fissidentaceae, 20 spp.); Lejeunea (Lejeuneaceae, 16 spp.); and Metzgeria (Metzgeriaceae, 14 spp.). Considering the samples analyzed, the most common species with the largest number of records were Campylopus arctocarpus (7 specimens); Lejeunea acanthogona (7); Lejeunea flava (7); Radula nudicaulis (7); Bryum capillare (6); Fissidens wallisii (6); Lejeunea serpillifolioides (6); Rigodium toxarion (6); Trichocolea brevifissa (6); and Zelometeorium ambiguum (6).

2.2. Substrate Preferences

The bryophyte species predominantly colonized rocks (210 spp.), followed by tree trunks (209 spp.), soil (205 spp.), logs (50 spp.), and leaves (29 spp.) (Figure 1).

Figure 1.

Figure 1

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Substrates colonized by bryophytes in Serra dos Órgãos National Park.

The availability of trees within the forests of PARNASO clearly influences the bryophyte species occurrence. Mosses predominantly colonize tree trunks, with only four species being found on leaves: Paraleucobryum longifolium (Dicranaceae); Philophyllum tenuifolium (Hookeriaceae); Toloxis imponderosa (Meteoriaceae); and Schlotheimia tecta (Orthotrichaceae).

The lowest bryophyte colonization was for leaves (epiphyllous). The leaf-colonizing species most frequent were the leafy liverworts of the family Lejeuneaceae (21 spp.), followed by Radulaceae (2 spp.) and the thallose liverwort family Metzgeriaceae (2 spp.). According to Gradstein et al. (2001) [17] and Ilkiu-Borges & Lisboa (2004) [57], species of Lejeuneaceae can colonize different types of substrates, but their occurrence on leaves is indicative of preserved native areas.

2.3. Endemism and Conservation Status

The Serra dos Órgaos National Park protects three species locally endemic, eight from Rio de Janeiro state, and 69 Brazilian (Table 2). Endemism and species richness are high with an increase in altitude, as was also found in Itatiaia National Park, located in Serra da Mantiqueira [18].

Table 2.

List of endemic bryophyte species found in PARNASO. (EB = endemic to Brazil; ERJ = endemic to Rio de Janeiro State; and CS = conservation status category.

Group Táxon EB ERJ CS
Liverworts Cheilolejeunea grosseoleosa C.J.Bastos & Schäf.-Verw. X
Cololejeunea diaphana A.Evans X
Cololejeunea papilliloba (Steph). Steph. X
Colura itatyana Steph. X X
Frullania schaefer-verwimpii Yuzawa & Hatt. X
Frullania vitalii Yuzawa & Hatt. X
Fuscocephaloziopsis crassifolia (Lindenb. & Gottsche) Váňa & L. Söderstr. X
Gongylanthus liebmannianus (Lindenb. & Gottsche) Steph. EN
Haplolejeunea umbrosa Gradst. & Ilkiu-Borges X
Harpalejeunea schiffneri S.W. Arnell X
Harpalejeunea subacuta A.Evans X
Lejeunea beyrichiana (Steph) Gradst. & C.J.Bastos X X
Lejeunea longidentata C.J.Bastos, Gradst., S. Vilas Bôas-Bastos & Schäf.-Verw. X
Lejeunea oligoclada Spruce X
Leptoscyphus spectabilis (Steph.) Grolle X
Metzgeria brasiliensis Schiffn. X
Metzgeria convoluta Steph. X
Metzgeria cratoneura Schiffn. X
Metzgeria hegewaldii Kuwah. X EN
Microlejeunea squarrosa J. Heinrichs, A. Schäfer-Verwimp, Pócs & S.S. Dong X
Micropterygium pterygophyllum (Nees) X
Neesioscyphus carneus (Nees) Grolle X
Paracromastigum dusenii (Steph.) R.M.Schust. X EN
Plagiochila amicta Steph. X
Plagiochila boryana Gottsche ex Steph. EN
Saccogynidium caldense (Ångstr.) Grolle X
Southbya organensis Herzog X CR
Syzygiella uleana Steph. X
Vitalianthus bischlerianus (Porto & Grolle) R.M.Schust. & Giancotti X
Mosses Atractylocarpus brasiliensis (Müll.Hal.) R.S.Williams X EN
Atractylocarpus longisetus (Hook.) E.B.Bartram EN
Brachydontium notorogenes W.R.Buck & Schaf.-Verw. X X CR
Brachymenium hornschuchianum Mart. X
Breutelia grandis (Hampe) Paris X
Callicostella martiana (Hornsch.) A.Jaeger X
Calyptrochaeta albescens (Hampe) W.R.Buck X
Campylopus densicoma (Müll.Hal.) Paris EN
Campylopus dichrostis (Müll.Hal.) Paris X
Campylopus gemmatus (Müll.Hal.) Paris X
Campylopus julicaulis Broth. X
Campylopus thwaitesii (Mitt.) A.Jaeger X
Campylopus uleanus (Müll.Hal.) Broth. X
Campylopus widgrenii (Müll.Hal.) Mitt. X
Cladastomum robustum Broth. X
Cladastomum ulei Müll.Hal. X
Crumuscus vitalis W.R.Buck X
Cyclodictyon marginatum (Hook. & Wilson) Kuntze X
Daltonia splachnoides (Sm.) Hook. & Taylor X
Dicranodontium pulchroalare Broth. X X
Dimerodontium pellucidum Schwägr. X
Ditrichum paulense Geh. ex Hampe X
Itatiella ulei (Broth. ex Müll.Hal.) G.L.Sm. X
Leiomela piligera (Hampe) Broth. X
Lepidopilidium brevisetum (Hampe) Broth. X
Lepidopilum scabrisetum (Schwägr.) Steere X
Leptodontium stellatifolium (Hampe) Broth. X
Leptodontium wallisii (Müll.Hal.) Kindb. VU
Leskeodon aristatus (Geh. & Hampe) Broth. X
Leucobryum clavatum Hampe X
Leucoloma triforme (Mitt.) A.Jaeger X
Microcalpe subsimplex (Hedw.) W.R.Buck X
Polytrichum angustifolium Mitt. X
Rhacocarpus inermis (Müll.Hal.) Lindb. X
Sphagnum brasiliense Warnst. X
Sphagnum costae H.A.Crum & D.P.Costa X
Sphagnum divisum H.A.Crum X
Sphagnum exquisitum H.A.Crum X
Sphagnum geraisense H.A.Crum X
Sphagnum gracilescens Hampe X
Sphagnum longicomosum Müll.Hal. ex Warnst. X
Sphagnum luetzelburgii H.K.G.Paul ex H.A.Crum X X
Sphagnum perforatum Warnst. X VU
Sphagnum pseudoramulinum H.A.Crum X
Sphagnum ramulinum Warnst. X
Sphagnum sehnemii H.A.Crum X
Sphagnum sucrei H.A.Crum X
Trichosteleum glaziovii (Hampe) W.R.Buck X
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Of the 28 species of Brazilian endemic bryophytes exclusive to high-altitude fields, 12 belong to the family Sphagnaceae (42%). The species Sphagnum luetzelburgii and S. sehnemii are endemic to Brazil, occurring in the states of Paraná, Santa Catarina, Rio Grande do Sul, and Rio de Janeiro. However, in Rio de Janeiro State, these species are restricted to mountains of the Atlantic Forest and have only been reported for PARNASO by Costa (2021) [58] and not by the present work.

The lower number of endemic species recorded in sub-montane forest probably reflects the effects of ecological and biogeographic processes, as well as human interference [18]. Among the 19 endemic species exclusively found in montane forest, 13 are liverworts and 6 are mosses. The predominant liverwort family is Lejeuneaceae, with nine species, followed by Metzgeriaceae with three and Jamesoniellaceae with two.

We have re-collected some species considered rare, including Bryum limbatum. Ochi (1980) [59] cites a collection from Metzgeria furcata, collected by Bartram (1954) [19]; Sphagnum cuspidatum, collected by Luetzelburg in 1923; and Andreaea acutifolia, collected by Mueller in 1845. Andreaea acutifolia is always mistakenly identified as A. rupestris, the two differing mainly by perichaetial leaves [24]. The species S. cuspidatum is widespread in Brazil [60], but has been collected only once in PARNASO.

The Serra dos Orgaos Park also protects eleven of the species indicated as threatened by CNCFlora and Flora e Funga do Brasil (Figure 2 and Table 2) in the Rio de Janeiro state: seven as Endangered (EN), two as Critically Endangered (CR) and two as Vulnerable (VU). Among these threatened species, three are known only from the type material, and were not found in the field trips. Southbya organensis was described by Herzog (1949) [61] and is classified as Critically Endangered (CR) by CNCFlora (2012) [58] due to its distribution being restricted to Serra dos Órgãos, specifically Morro do Açu peak, which has excessive tourism and recurrent arson instead. Plagiochila pseudoradicans and Dicranodontium pulchroalare ssp. brasiliense were described by Herzog (1949) [61], and neither was assessed by CNCFlora (2012) [62] because they were considered as Data Deficient (DD) at that time.

Figure 2.

Figure 2

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Pictures of some species indicated as threatened in Serra dos Orgaos National Park. (A) Gongylanthus liebmannianus (Lindenb. & Gottsche) Steph., (B) Metzgeria hegewaldii Kuwah., (C) Plagiochila boryana Gottsche ex Steph., (D) Atractylocarpus longisetus (Hook.) E.B.Bartram, (E) Leptodontium wallisii (Müll.Hal.) Kindb., (F) Sphagnum perforatum Warnst.

3. Discussion

The Serra dos Órgãos National Park (PARNASO) exhibits remarkable bryophyte diversity. This richness highlights the park as a key conservation area for bryophytes in the Atlantic Forest. These findings suggest that even in areas already considered well-studied, significant knowledge gaps remain, highlighting the dynamic nature of bryophyte diversity and distribution.

One important factor contributing to this diversity is the presence of forested areas with abundant tree trunks, which serve as critical substrates for epiphytic bryophytes. The architectural, morphological, phenological, and chemical traits of different tree species create varied microhabitats that support a wide range of bryophyte taxa [57,63]. Shade-loving epiphytes, in particular, are commonly found in these forest environments but are highly sensitive to environmental disturbances. As such, they are often among the first species to disappear following canopy opening or habitat degradation [64], making them reliable indicators of forest health.

In contrast, mosses exhibit a broader ecological amplitude due to their desiccation tolerance and structurally complex adaptations, such as costa development, hyaline leaf apices, papillae, and specialized conducting tissues like hydroids and leptoids [29,65,66]. These features enable many mosses to colonize exposed and drier substrates, including rocky outcrops. Similar patterns of moss dominance on rock surfaces have been observed in other highland regions of Brazil, such as Chapada Diamantina [67,68], Serra do Caraça [69], and Serra da Canastra [70].

The high-altitude ecosystems of PARNASO further enhance bryophyte diversity by creating isolated and unique habitats. Mountain summits, often compared to ecological islands due to their environmental isolation, support specialized flora adapted to harsh conditions such as strong winds, low temperatures, and nutrient-poor soils [71,72,73]. These rupestrian fields form natural refugia, acting as barriers to typical forest species and favoring the emergence of narrow endemics and rare taxa [74,75,76].

Overall, the findings of this study underscore the ecological importance and conservation value of Serra dos Órgãos National Park. Alongside Itatiaia National Park, PARNASO remains one of the most critical remnants of Atlantic Forest in Rio de Janeiro and Brazil, particularly in terms of bryophyte conservation. The park’s diverse topography, wide range of microhabitats, and climatic heterogeneity—marked by high altitudes, low temperatures, and high light availability—create optimal conditions for bryophyte diversity. Furthermore, the presence of endemic and threatened species reinforces the park’s role as a conservation priority. Such studies are valuable as they reveal the significance of endemism and species richness, crucial factors in determining conservation priorities [77], and emphasize the importance of preserving Serra dos Órgãos National Park.

4. Materials and Methods

4.1. Study Area

Serra dos Órgãos National Park (PARNASO) is located in the highlands region of Rio de Janeiro State, Brazil, ranging the municipalities of Teresópolis, Petrópolis, Guapimirim and Magé (22°23′36.96″–22°34′57.72″ S and 43°10′57.72″–42°58′43.68″ W). It is a protected area with 20,024 hectares of Atlantic Forest in a central position in the Serra do Mar Ecological Corridor. Ranging in altitude from 80 to 2275 m.a.s.l. (Figure 3), it is the highest portion of the entire Serra do Mar [2]. The climate is tropical and super humid (80–90% relative humidity), with the annual average temperature ranging from 13 to 23 °C, although minimums can reach −5 °C during winter above 800 m.a.s.l. [6,8,74]. The average annual rainfall ranges from 1500 mm to 3000 mm, with higher concentrations of rain during summer (December to March) and a dry season during winter (June to August), while heat is well distributed throughout the year [78,79].

Figure 3.

Figure 3

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Location of the Serra dos Órgãos National Park, Rio de Janeiro State, Brazil.

The vegetation classification adopted follows Veloso et al. (1991) [20] and Safford (1999) [80]: Sub-montane Forest (below 600 m.a.s.l., typically evergreen, with a high canopy and rich floristic diversity; dominant tree species include Protium spp., Virola spp., and members of the Myrtaceae and Fabaceae families); montane forest (600–1500, characterized by shorter trees and more frequent presence of cloud cover; common tree species include Clethra spp. and various species of Melastomataceae); upper-montane forest (above 1500, vegetation here is more stunted, often with gnarled trees adapted to frequent mist and lower temperatures). Dominant elements include Weinmannia paulliniifoli and Drimys brasiliensis, and high-altitude fields (above 1800–2000) are open grassland formations interspersed with shrubs, herbs, and occasional low woody elements. Common species include Vellozia spp., Paepalanthus spp., and Eryngium spp. (Figure 4).

Figure 4.

Figure 4

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Collection sites in Serra dos Órgãos National Park. (A) Trail of the “Mirante”, (B) aspect of the montane forest with streams, (C) “Véu da Noiva” waterfall, (D) aspect of the upper-montane forest, (E) aspect of high-altitude fields, with Sphagnum among Chusquea sp.

4.2. Studied Taxa

The species list developed here is based on (i) a bibliographic survey conducted by consulting published volumes of major Brazilian botanical journals, as well as international bibliographic databases (Web of Science, SciELO, and Google Scholar), in search of peer-reviewed articles; the search focused on article titles containing combinations of the keywords “Serra dos Órgãos,” “bryophytes,” “mosses,” and “liverworts,” followed by a careful and detailed reading of each selected publication; (ii) herbarium specimens collected in the study area; and (iii) new collections (Figure 4).

Herbarium specimens were obtained from online sources like (GBIF [81], jabot, JStorTypes and SpeciesLink accessed several times). The specimens are requested on loan and re-determined and carefully examined for small species associations, with one of each species cited as voucher in the Table 1. The authors conducted four field excursions to the PARNASO, each lasting one week, two during the rainy season and two during the dry season, with the aim of locating species not previously recorded in the herbarium. The collection, preservation and herborization methodologies follow Frahm (2003) and employ free walks [82] for covering all available substrates like tree trunks, logs, soil, rocks and leaves.

Field trips in the Park intended to check if species collected in the past are yet present, to collect new specimens and to record new taxa. Collected samples were deposited in the herbarium of the University of Brasília (UB), with duplicates sent to the herbarium of Instituto de Pesquisas Ambientais (SP). The classification system used here follows Crandall-Stotler et al. [83] for Marchantiophyta, Goffinet et al. [65] for Bryophyta and Renzaglia et al. [84] for Anthocerotophyta. Data referring to endemic species of Brazil and of Rio de Janeiro State came from Flora e Funga do Brasil (floradobrasil.jbrj.gov.br), while conservation status followed CNCFlora (cncflora.jbrj.gov.br).

5. Conclusions

The Serra dos Órgãos National Park stands out as a key refuge for bryophyte diversity in the Atlantic Forest, hosting numerous endemic and threatened species. The park’s altitudinal variation, diverse microhabitats, and unique environmental conditions favor high species richness and endemism. These findings reinforce the ecological value of PARNASO and underscore the urgency of conserving this unique mountainous ecosystem.

Acknowledgments

We thank the Fundação de Amparo à Pesquisa do Distrito Federal (FAP-DF) for financial support to this project and the park coordinator PARNASO (Jorge Nascimento). We fully thank the support in field collections of the colleagues Amanda Leal, Anderson Eustáquio, Clapton Moura, Cosme Ponte and Flávio Rick.

Abbreviation

PARNASO Serra dos Órgãos National Park
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Author Contributions

Methodology, M.T.A.G. and D.F.P.; Validation, D.F.P.; Investigation, J.S.d.L., A.L.A.F., M.T.A.G. and D.F.P.; Resources, D.F.P.; Data curation, J.S.d.L., A.L.A.F. and D.F.P.; Writing—original draft, A.L.A.F.; Writing—review & editing, J.S.d.L., M.T.A.G. and D.F.P. All authors have read and agreed to the published version of the manuscript.

Data Availability Statement

The original contributions presented in this study are included in the article; further inquiries can be directed to the corresponding author.

Conflicts of Interest

The authors declare no conflicts of interest.

Funding Statement

This research was funded by the Fundação de Amparo à Pesquisa do Distrito Federal (FAP-DF).

Footnotes

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