A checklist of the bryophytes from Amapá State, Northern Brazil

Uma lista das briófitas do Estado do Amapá, norte do Brasil

ABSTRACT

A checklist is a good base for gathering registers of biodiversity and supporting further studies. The aim of this paper is to present a checklist of the bryophytes from Amapá State, including the distribution data of the species. In total, 174 species (80 mosses and 94 liverworts) in 78 genera and 30 families are registered for Amapá State, northern Brazil, based on examined specimens and on a review of the literature. A table with the species, distribution data for all municipalities of Amapá State, and the first record for all species are presented.

Keywords:
Amapá State flora; bryoflora; compilation of data; floristic study; taxonomy

RESUMO

Uma lista é uma boa base para reunir registros de biodiversidade e apoiar novos estudos. O objetivo deste artigo é apresentar uma lista de verificação das briófitas do Estado do Amapá, incluindo os dados de distribuição das espécies. No total, 174 espécies (80 musgos e 94 hepáticas) em 79 gêneros e 30 famílias são registradas no Estado do Amapá, norte do Brasil, com base em espécimes examinados e em uma revisão da literatura. Uma tabela com as espécies, dados de distribuição para todos os municípios do Estado do Amapá e o primeiro registro para todas as espécies são apresentados.

Palavras-chave:
brioflora; compilação de dados; estudo florístico; flora do Estado do Amapá; taxonomia

Introduction

The Amapá State presents several Amazonian physiognomies of high biological diversity mainly represented by dense Terra Firme forest, Várzea forest, mangrove, transition areas between forests, and savannah vegetation (IBGE 2004, Costa-Neto et al. 2006, ZEE 2008). The bryophyte flora in the State is still poorly known if compared to other States in Brazil (Lisboa et al. 2006, Oliveira-da-Silva & Ilkiu-Borges 2020), especially considering that it is one of the most preserved, with more than 72% of its territory divided into 19 conservation units (12 national, five States, and two municipal) (Drummond et al. 2008).

Herzog (1931) was the first work that made a study with bryophytes from the State of Amapá. He identified mosses and liverworts collected by von Lützelburg in northern Brazil, especially in the State of Amazonas and Pará. However, some collections of Herzog’s study were cited as “Staat Pará: Rio Oyapock”. The Oiapoque River is in the municipality of Oiapoque, in Amapá State, on the border with French Guiana. At the time of the publication of Herzog, the Amapá territory belonged to the Grão-Pará province (currently Pará State). Only in 1943, the Amapá State was separated from Pará and become a separate federal unit.

Oliveira-da-Silva & Ilkiu-Borges (2020) had already listed most studies on bryophytes in Amapá State (Crosby 1969, Grolle 1984, Yano 1981, 1982, 1984b, 1992, Yano et al. 1985, Yano & Lisboa 1988, Churchill 1998, Gradstein & Costa 2003, Lisboa et al. 2006, Gentil & Menezes 2011) discussing its advances in the State. In this study, they investigated the richness and composition of bryophytes of two Parks in Amapá, registering 63 new records for the State and three new records for the North region of Brazil. Also, they pointed out the need for more comprehensive studies on the bryophyte flora in the State.

A checklist is a good base for gathering registers of biodiversity, serving as a data basis for other studies. The bryophyte flora in Brazil is currently represented by 1.521 species, of which 18.1% are endemic, according to the BFG (2018). This number of species was reached through several studies carried out by different researchers, which enable to know the bryophytes that compose the flora of specific localities or geographic regions (e.g., Gradstein & Costa 2003, Costa & Luizi-Ponzo 2010, Costa et al. 2011, Costa & Peralta 2015, BFG 2018), or on checklists (e.g., Yano 1981, 1984, 1989, 1995, 1996, 2006, 2008, Peralta & Yano 2001, Câmara et al. 2005, Costa et al. 2005).

No comprehensive list with distribution data of bryophyte species is available for Amapá State, but the knowledge is spread in a few local studies (e.g., Oliveira-da-Silva & Ilkiu-Borges 2020). This paper aims to fulfill this gap presenting a checklist of the bryophytes in Amapá State, including distribution data.

Material and methods

The checklist was based on the literature of bryophytes as guides, taxonomic revisions, and specific studies for Amapá already published, as follows: Herzog (1931), Crosby (1969), Yano (1981, 1982, 1984b, 1992), Grolle (1984), Yano et al. (1985), Yano & Lisboa (1988), Gradstein (1994), Churchill (1998), Gradstein & Costa (2003), Lisboa et al. (2006), Gentil & Menezes (2011), BFG (2018), Oliveira-da-Silva & Ilkiu-Borges (2020), and Macedo et al. (in press). The voucher cited for each species in table 1 was preferably the one examined by the authors.

Taxonomic classifications adopted are by Goffinet et al. (2009) and Crandall-Stotler et al. (2009) for Bryophyta and Marchantiophyta, respectively. The classification of Sematophyllaceae is following Carvalho-Silva et al. (2017), while for Dibrachiella and Cryptolophocolea we followed Shi et al. (2015) and Söderström et al. (2013), respectively.

Data on municipalities of Amapá are following the Instituto Brasileiro de Geografia e Estatística - IBGE (2019). They were standardized for all specimens in which locality was registered in the publication. Species classified as “common” present records in more than four municipalities in the State of Amapá.

Results

In Amapá State, 174 species in 78 genera and 30 families of bryophytes were registered. Mosses were represented by 80 species, 39 genera, and 21 families, while liverworts were represented by 94 species, 39 genera, and nine families (table 1). None hornwort was recorded.

Lejeuneaceae Cavers and Calymperaceae Kindb. were the most diverse families found in the State of Amapá, with 73 and 23 species, respectively. The two families represented more than 55% of the most diversified families for the State (figure 1). Genera with a higher number of species were Syrrhopodon Schwägr. and Lejeunea Lib., with 12 and 11 species, respectively. The most common species of the mosses (recorded in four or more municipalities) were Callicostella pallida (Hornsch.) Ångström, Calymperes erosum Müll. Hal., Microcalpe subsimplex (Hedw.) W.R. Buck, Octoblepharum albidum Hedw., Syrrhopodon ligulatus Mont., Taxithelium planum (Brid.) Mitt., and Trichosteleum papillosum (Hornsch.) A. Jaeger, and of the liverworts were Ceratolejeunea coarina (Gottsche) Schiffn., C. cornuta (Lindenb.) Steph., Cheilolejeunea oncophylla (Ångstr.) Grolle & M.E. Reiner, and Leptolejeunea elliptica (Lehm. & Lindenb.) Schiffn.

Among 16 municipalities from Amapá, bryophytes were recorded only from seven ones. The municipality of Serra do Navio presented the highest number of species (109 spp.), followed by Macapá (77 spp.), Amapá (60 spp.), Oiapoque (42 spp.), Mazagão (14 spp.), Calçoene (9 spp.), and Laranjal do Jari (2 spp.) (figure 2).

Herzog (1931) was the first to record 13 species for the Amapá State, followed by Crosby (1969) (1 sp.), Yano (1982) (3 spp.), Yano et al. (1985) (1 sp.), Yano & Lisboa (1988) (37 spp.), Gradstein (1994) (1 sp.), Churchill (1998) (5 spp.), Lisboa et al. (2006) (43 spp.), Oliveira-da-Silva & Ilkiu-Borges (2020) (61 spp.), and Macedo et al. (in press) (9 spp.). According to all studies on bryophytes in Amapá, 175 species were registered in the State (figure 3). The studies carried out by Yano & Lisboa (1988), Lisboa et al. (2006), and Oliveira-da-Silva & Ilkiu-Borges (2020) registered 80% of the bryophyte records for the State.

Discussion

With about 70% of its territory included in legally protected areas, the Amapá State is the twentieth State (from 26 States) in the number of bryophyte species (Drummond et al. 2008, Costa & Peralta 2015). Although the bryoflora of Brazilian Amazon is being studied for over 200 years (Lisboa 1991), the bryophyte richness in Amapá is still poorly known despite the diversity of the biome where it is inserted, but it is notably low the number of studies carried out in this State, not including a great area of its territory. Amazonas and Pará States, although with larger territories, have many physiognomies that also occur in Amapá. However, these two States presented a much higher number of bryophyte species (466 and 332 species, respectively) and a great number of studies (e.g., Costa et al. 2011, Costa & Peralta 2015, BFG 2018). Our results showed that Amapá presented almost as the number of species found in Rondônia (166 spp.), but studies in this State was punctual (Lisboa 1993, Cerqueira et al. 2015, Costa & Peralta 2015, Sobreira et al. 2019).

Macrocolura sagittistipula (Spruce) R.M. Schust. is excluded from the list of species of Amapá. Gradstein & Costa (2003) cited the species as occurring in Amazonas and Amapá States based on the study of Grolle & Zhu (2002), but the latter authors recorded this species in Amazonas only.

Yano & Lisboa (1988) registered Mastigolejeunea auriculata (Wilson & Hook.) Schiffn. [= Thysananthus auriculatus (Wilson & Hook.) Sukkharak & Gradst.] in Mazagão municipality, however, the specimen deposited in MG herbarium (MG109797!) proved to be Thysananthus amazonicus (Spruce) Steph.

Cheilolejeunea savannae L.P.C. Macedo et al. is a species from savanna vegetation recently described by Macedo et al. (2020). This species was referred as C. discoidea (Lehm. & Lindenb.) Kachroo & R.M. Schust. by Oliveira-da-Silva & Ilkiu-Borges (2020).

The most abundant families, Lejeuneaceae and Calymperaceae, were expected to be since many studies in the Amazon recorded these families as the richest (e.g., Gradstein et al. 2001, Macedo & Ilkiu-Borges 2014, Cerqueira et al. 2017, Oliveira-da-Silva & Ilkiu-Borges 2018). The two richest genera (Syrrhopodon and Lejeunea) were also expected since they are well distributed in Brazil (Gradstein et al. 2001, Gradstein & Costa 2003, Costa & Peralta 2015). Syrrhopodon is a widely distributed genus in the tropics and subtropics and assembles acrocarpous plants morphologically diverse (Reese 1993), additionally, it is the fourth richest genera of mosses in Brazil (Costa & Peralta 2015). Lejeunea is the more complex and the largest genus of liverworts in Brazil (Reiner-Drehwald 2007, Costa & Peralta 2015) and well distributed in the neotropical region (Gradstein et al. 2001).

The most common species of our list were already expected to be found in Amapá due to their wide distribution in the Amazon and frequent occurrence in various ecosystems from humid rainforests to open environments (Santos & Lisboa 2003, Brito & Ilkiu-Borges 2013, Moura et al. 2013, Tavares-Martins et al. 2014, Pantoja et al. 2015, Cerqueira et al. 2017, BFG 2018, Oliveira-da-Silva & Ilkiu-Borges 2018). Among these species, only Ceratolejeunea coarina was not reported in Marajó island (Brito & Ilkiu-Borges 2013), a large continental island neighbor to Amapá State.

Table 1
Checklist of bryophytes with the distribution in municipalities and refer­ence of the first record of each species for Amapá State. AM: Amapá municipality; MC: Macapá; SN: Serra do Navio; OP: Oiapoque; CÇ: Calçoene; MZ: Mazagão; LJ: Laranjal do Jari; (n.v.): not seen.
Tabela 1
Lista das briófitas com dados de distribuição geográfica nos municípios e referência do primeiro registro de cada espécie para o Estado do Amapá. AM: município de Amapá; MC: Macapá; SN: Serra do Navio; OP: Oiapoque; CÇ: Calçoene; MZ: Mazagão; LJ: Laranjal do Jari; (n.v.): não visto.

Family/Species	Municipalities of Amapá State							First record for Amapá State	Voucher
	AM	MC	SN	OP	CÇ	MZ	LJ
LIVERWORTS
Aneuraceae
Riccardia regnellii (Ǻngstr.) K.G.Hell		X	X					Yano & Lisboa [1988, as Riccardia alata (Steph.) K.G.Hell]	Gentil 513
Riccardia metzgeriformis (Steph.) R.M. Schust.		X						Yano & Lisboa (1988)	Mori & Souza 17602 (n.v.)
Calypogeiaceae
Calypogeia lechlerii (Steph.) Steph.			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 391
Frullaniaceae
Frullania subtilissima (Nees ex Mont.) Lindenb.		X	X			X		Oliveira-da-Silva & Ilkiu-Borges (2020, as Frullania exilis Taylor)	Gentil 433
Frullania gibbosa Nees	X	X						Lisboa et al. (2006)	Macedo 1717
Frullania vitalii Yuzawa & S. Hatt.		X						Macedo et al. (in press)	Macedo 1672
Lejeuneaceae
Archilejeunea fuscescens (Lehm.) Fulford		X	X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 385
Acrolejeunea emergens (Mitt.) Steph.		X						Macedo et al. (in press)	Macedo 1723
Acrolejeunea torulosa (Lehm. & Lindenb.) Schiffn.		X				X		Macedo et al. (in press)	Macedo 2098
Caudalejeunea lehmanniana (Gottsche) A. Evans	X	X						Lisboa et al. (2006)	Gentil 94
Ceratolejeunea coarina (Gottsche) Schiffn.	X	X	X	X				Herzog (1931, as Ceratolejeunea connata Steph.)	Gentil 347
Ceratolejeunea cornuta (Lindenb.) Steph.	X	X	X	X				Herzog [1931, as Ceratolejeunea megalophysa (Spruce) Steph.]	Gentil 353
Ceratolejeunea cubensis (Mont.) Schiffn.		X	X	X				Herzog (1931)	Gentil 395
Ceratolejeunea desciscens (Sande Lac.) Schiffn.			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 467
Ceratolejeunea guianensis (Nees & Mont.) Steph.			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 472
Ceratolejeunea minuta G. Dauphin			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 345
Ceratolejeunea rubiginosa Steph.	X							Lisboa et al. (2006)	Costa Neto 1307 (n.v.)
Cheilolejeunea acutangula (Nees) Grolle			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 363
Cheilolejeunea adnata (Lehm.) Grolle	X							Lisboa et al. (2006)	Gentil 464
Cheilolejeunea clausa (Nees & Mont.) R.M. Schust.	X							Lisboa et al. (2006)	Costa Neto 1491 (n.v.)
Cheilolejeunea savannae L.P.C. Macedo et al.		X	X					Oliveira-da-Silva & Ilkiu-Borges [2020, as Cheilolejeunea discoidea (Lehm. & Lindenb.) Kachroo & R.M.Schust.]	Macedo 1600 (Paratype)
Cheilolejeunea holostipa (Spruce) Grolle & R.L. Zhu			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 497
Cheilolejeunea oncophylla (Ångstr.) Grolle & M.E. Reiner	X	X	X			X		Lisboa et al. (2006)	Gentil 113
C. rigidula (Mont.) R.M. Schust.	X	X	X					Lisboa et al. (2006)	Gentil 114
Cheilolejeunea trifaria (Reinw. et al.) Mizut.	X		X					Lisboa et al. (2006)	Gentil 504
Cololejeunea cardiocarpa (Mont.) A. Evans	X	X						Lisboa et al. (2006)	Gentil 204
Cololejeunea contractiloba A. Evans		X						Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 192
Cololejeunea crenata (A. Evans) Pócs			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 491
Cololejeunea gracilis (Jovet-Ast) Pócs	X							Lisboa et al. (2006, as Aphanolejeunea gracilis Jovet-Ast)	Costa Neto 1472 (n.v.)
Cololejeunea obliqua (Nees & Mont.) Schiffn.	X		X	X				Herzog [1931, as Leptocolea scabriflora (Steph) A.Evans]	Gentil 510
Cololejeunea sicaefolia (Gottsche) Pócs & Bernecker			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 528
Colura greig-smithii Jovet-Ast	X							Lisboa et al. (2006)	Costa Neto 1499 (n.v.)
Cyclolejeunea chitonia (Lehm.) A. Evans			X	X				Herzog (1931)	Gentil 366
Cyclolejeunea convexistipa (Lehm. & Lindenb.) A. Evans			X	X				Herzog (1931)	Gentil 376
Cyclolejeunea foliorum (Nees) Grolle			X	X				Yano & Lisboa (1988)	Gentil 547
Cyclolejeunea luteola (Spruce) Grolle		X	X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 347
Dibrachiella parviflora (Nees) X.Q. Shi et al.	X	X						Lisboa et al. (2006)	Gentil 88
Dibrachiella auberiana (Mont.) X.Q. Shi et al.	X							Lisboa et al. [2006, as Achilejeunea auberiana (Mont.) A.Evans]	Costa Neto 1286 (n.v.)
Diplasiolejeunea brunnea Steph.	X		X					Lisboa et al. (2006)	Gentil 373
Diplasiolejeunea pellucida (Spreng.) Schiffn.			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 388
Drepanolejeunea araucarae Steph.	X							Lisboa et al. (2006)	Costa Neto 1291 (n.v.)
Drepanolejeunea crucianella (Taylor) A. Evans			X	X				Herzog (1931, as Drepanolejeunea trifida Steph.)	Gentil 471
Drepanolejeunea fragilis L. Söderstr.	X	X	X					Lisboa et al. (2006)	Gentil 433
Harpalejeunea oxyphylla (Nees & Mont.) Steph.			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 401
Harpalejeunea tridens (Besch. & Spruce) Steph.			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 415
Lejeunea adpressa Nees		X						Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 101
Lejeunea aphanes Spruce			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 501
Lejeunea asperrima Spruce			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 531
Lejeunea boryana Mont.			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 537
Lejeunea cerina (Lehm. & Lindenb.) Lehm. & Lindenb.			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 532
Lejeunea controversa Gottsche		X	X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Macedo 1899
Lejeunea flaccida Lindenb. & Gottsche	X							Lisboa et al. [2006, as Taxilejeunea lusoria (Lindenb. & Gottsche) Steph.]	Costa Neto 1486 (n.v.)
Lejeunea flava (Sw.) Nees			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 502
Lejeunea glaucecens Gottsche		X	X					Oliveira-da-Silva & Ilkiu-Borges (2020, as Lejeunea caulicalyx (Steph.) M.E.Reiner & Goda)	Gentil 112
Lejeunea immersa Spruce		X	X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 454
Lejeunea laetevirens Nees & Mont.	X	X	X					Lisboa et al. (2006)	Gentil 135
Lepidolejeunea involuta (Gottsche) Grolle	X		X					Lisboa et al. (2006)	Gentil 348
Leptolejeunea elliptica (Lehm. & Lindenb.) Schiffn.	X	X	X	X				Herzog (1931)	Macedo 2091
Lopholejeunea subfusca (Nees) Schiffn.	X	X	X					Lisboa et al. (2006)	Gentil 503
Metalejeunea cucullata (Reinw. et al.) Grolle			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 410
Microlejeunea epiphylla Bischl.	X	X	X					Lisboa et al. (2006)	Gentil 497
Myriocoleopsis minutissima R.L. Zhu et al. ssp. minutissima		X				X		Macedo et al. (in press)	Macedo 1525
Neurolejeunea breutelii (Gottsche) A. Evans			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 358
Odontolejeunea lunulata (F. Weber) Schiffn.	X		X	X				Herzog (1931)	Gentil 371
Odontolejeunea rhomalea (Spruce) Steph.				X				Gradstein (1994)	von Lützelburg 20199 (n.v.)
Otigoniolejeunea huctumalcensis (Lindenb. & Gottsche) Y.M. Wei et al.			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 352
Pictolejeunea picta (Steph.) Grolle			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 540
Prionolejeunea aemula (Gottsche) A.Evans			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 500
Prionolejeunea denticulata (F. Weber) Schiffn.			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 351
Prionolejeunea mucronata (Sande Lac.) Steph.			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 348
Prionolejeunea muricatoserrulata (Spruce) Steph.			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 372
Pycnolejeunea contigua (Nees) Grolle			X			X		Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 433
Rectolejeunea versifolia (Schiffn.) L. Söderstr. & A. Hagborg	X		X					Lisboa et al. (2006)	Gentil 389
Stictolejeunea squamata (F. Weber) Schiffn.			X	X				Herzog (1931)	Gentil 355
Symbiezidium barbiflorum (Lindenb. & Gottsche) A. Evans		X	X	X				Herzog (1931)	Gentil 372
Symbiezidium transversale (Sw.) Trevis.	X	X		X				Yano & Lisboa (1988)	Mori et al. 17147 (n.v.)
Thysananthus amazonicus (Spruce) Steph.						X		Yano & Lisboa [1988, as Mastigolejeunea auriculata (Wilson & Hook.) Schiffn.]	Mori & Cardoso 17481 (n.v.)
Thysananthus innovans (Spruce) Sukkharak & Gradst.			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Macedo 1640
Xylolejeunea crenata (Nees & Mont.) X.-L. He & Grolle	X	X	X					Lisboa et al. (2006)	Gentil 434
Lepidoziaceae
Protocephalozia sp.		X						Macedo et al. (in press)	Macedo 1639
Zoopsidella serra (Spruce) R.M. Schust.			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 378
Lophocoleaceae
Lophocolea liebmanniana Gottsche			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 447
Cryptolophocolea martiana (Nees) L. Söderstr. et al.			X	X				Yano & Lisboa (1988, as Lophocolea martiana Nees)	Gentil 540
Cryptolophocolea perissodonta (Spruce) L. Söderstr.		X						Yano & Lisboa [1988, as Lophocolea perissodonta (Spruce) Steph.]	Mori et al. 17259 (n.v.)
Metzgeriaceae
Metzgeria aurantica Steph.			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 384
Plagiochilaceae
Plagiochila aerea Taylor			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 537
Plagiochila disticha (Lehm. & Lindenb.) Lindenb.		X	X					Yano & Lisboa (1988)	Gentil 360
Plagiochila montagnei Nees		X	X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 205
Plagiochila gymnocalycina (Lehm. & Lindenb.) Mont. & Nees		X						Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 242
Plagiochila raddiana Lindenb.		X	X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 353
Plagiochila rutilans Lindenb.		X						Yano & Lisboa (1988)	Mori & Souza 17602 (n.v.)
Plagiochila subplana Lindenb.			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 346
Radulaceae
Radula flaccida Lindenb. & Gottsche			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 349
Radula javanica Gottsche	X		X	X				Yano & Lisboa [1988, as Radula complanata (L.) Dumort.]	Mori et al. 17221 (n.v.)
MOSSES
Bartramiaceae
Philonotis hastata (Duby) Wijk. & Marg.		X						Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 266
Brachytheciaceae
Squamidium nigricans (Hook.) Broth.				X	X			Yano & Lisboa (1988)	Mori & Cardoso 17293 (n.v.)
Zelometeorium patulum (Hedw.) Manuel		X						Yano & Lisboa (1988)	Gentil 512
Bryaceae
Bryum coronatum Schwägr.		X				X		Macedo et al. (in press)	Macedo 1534
Calymperaceae
Calymperes afzelii Sw.	X							Lisboa et al. (2006)	Costa Neto 1467 (n.v.)
Calymperes erosum Müll. Hal.	X	X	X	X		X		Yano & Lisboa (1988)	Gentil 275
Calymperes lonchophyllum Schwägr.		X	X	X				Yano & Lisboa (1988)	Gentil 350
Calymperes nicaraguense Renauld & Cardot					X			Yano & Lisboa (1988)	Mori 17413
Calymperes palisotii Schwägr.	X	X				X		Churchill (1998)	Gentil 185
Calymperes platyloma Mitt.					X			Churchill (1998)	Mori & Cardoso 17293 (n.v.)
Calymperes rubiginosum (Mitt.) Reese			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 430
Octoblepharum albidum Hedw.		X	X	X		X		Yano (1982)	Gentil 79
Octoblepharum cocuiense Mitt.		X	X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 85
Octoblepharum cylindricum Schimp.				X				Herzog (1931)	von Lützelburg 20312 (n.v.)
Octoblepharum pulvinatum (Dozy & Molk.) Mitt.	X						X	Yano (1982)	Egler & Irwin 46455 (n.v.)
Syrrhopodon cryptocarpus Dozy & Molk.	X	X	X					Lisboa et al. (2006)	Gentil 114
Syrrhopodon cymbifolius Müll. Hal.	X		X					Lisboa et al. (2006)	Gentil 436
Syrrhopodon disciformis Dusén			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 510
Syrrhopodon flexifolius Mitt.			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 518
Syrrhopodon hornschuchii Mart.		X	X	X				Yano & Lisboa (1988)	Gentil 452
Syrrhopodon incompletus Schwägr. var. incompletus		X	X	X				Yano & Lisboa (1988)	Gentil 118
Syrrhopodon incompletus var. berteroanus (Brid.) W.D. Reese		X	X	X				Churchill (1998)	Mori & Cardoso 17124 (n.v.)
Syrrhopodon leprieurii Mont.		X						Yano & Lisboa (1988)	Rosário 212 (n.v.)
Syrrhopodon ligulatus Mont.	X	X	X		X			Yano & Lisboa (1988)	Gentil 142
Syrrhopodon prolifer Schwägr.	X							Lisboa et al. (2006)	Costa Neto 1279 (n.v.)
Syrrhopodon rigidus Hook. & Grev.			X					Churchill (1998)	Gentil 369
Syrrhopodon simmondsii Steere	X		X					Lisboa et al. (2006)	Gentil 379
Dicranaceae
Dicranella hilariana (Mont.) Mitt.		X						Macedo et al. (in press)	Macedo 1736
Fissidentaceae
Fissidens guianensis Mont.	X	X	X					Lisboa et al. (2006)	Gentil 412
Fissidens lagenarius Mitt. var. muriculatus (Mitt.) Pursell			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 438
Fissidens ornatus Herzog		X						Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 326
Fissidens pellucidus Hornsch.		X	X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 439
Fissidens prionodes Mont.		X	X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 328
Fissidens radicans Mont.			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 512
Fissidens submarginatus Bruch		X						Macedo et al. (in press)	Macedo 1732
Fissidens zollingeri Mont.	X							Lisboa et al. (2006)	Costa Neto 1305 (n.v.)
Hookeriaceae
Crossomitrium patrisiae (Brid.) Müll. Hal.	X		X	X				Herzog (1931)	Gentil 401
Hypnaceae
Chrysohypnum diminutivum (Hampe) W.R. Buck		X	X	X				Yano & Lisboa (1988)	Gentil 410
Ectropothecium leptochaeton (Schwägr.) W.R. Buck			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 361
Rhacopilopsis trinitensis (Müll. Hal.) E. Britton & Dixon				X				Yano & Lisboa (1988)	Mori & Cardoso 17108 (n.v.)
Vesicularia vesicularis (Schwägr.) Broth.			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 371
Leucobryaceae
Campylopus surinamensis Müll. Hal.		X				X		Macedo et al. (in press)	Macedo 2174
Leucobryum martianum (Hornsch.) Müll. Hal.		X	X				X	Yano (1982)	Gentil 417
Ochrobryum subulatum Hampe			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 378
Leucomiaceae
Leucomium strumosum (Hornsch.) Mitt.		X		X				Yano & Lisboa (1988)	Macedo 2006
Neckeraceae
Neckeropsis disticha (Hedw.) Kindb.	X							Lisboa et al. (2006)	Costa-Neto 1301 (n.v.)
Neckeropsis undulata (Hedw.) Reichardt			X	X				Yano & Lisboa (1988)	Gentil 556
Orthotrichaceae
Groutiella tomentosa (Hornsch.) Wijk & Margad.	X							Lisboa et al. (2006)	Costa-Neto 1490 (n.v.)
Macromitrium cirrosum (Hedw.) Brid.		X		X				Churchill (1998)	Mori et al. 17541 (n.v.)
Macromitrium punctatum (Hook. & Grev.) Brid.		X		X				Yano & Lisboa (1988)	Rosário 193 (n.v.)
Phyllodrepaniaceae
Phyllodrephanium falcifolium (Schwägr.) Crosby			X	X	X			Yano & Lisboa (1988)	Gentil 369
Pilotrichaceae
Callicostella merkelii (Hornsch.) A. Jaeger	X							Lisboa et al. (2006)	Costa Neto 1486 (n.v.)
Callicostella pallida (Hornsch.) Ångström	X	X	X	X				Yano & Lisboa (1988)	Gentil 390
Lepidopilum affine Müll. Hal.	X							Lisboa et al. (2006)	Costa Neto 1301 (n.v.)
Lepidopilum scabrisetum (Schwägr.) Steere			X	X				Yano & Lisboa (1988)	Gentil361
Lepidopilum surinamense Müll. Hal.		X						Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 240
Pilotrichum evanescens (Müll. Hal.) Crosby		X						Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 177
Pilotrichum bipinnatum (Schaegr.) Brid.			X					Crosby (1969)	Gentil 512
Pottiaceae
Splachnobryum obtusum (Brid.) Müll. Hal.	X							Lisboa et al. (2006)	Costa Neto 1277 (n.v.)
Pterobryaceae
Henicodium geniculatum (Mitt.) W.R. Buck	X	X						Yano & Lisboa [1988, as Leucodontopsis geniculata (Mitt.) H.A.Crum & Steere]	Mori & Cardoso 17556 (n.v.)
Orthostichopsis tetragona (Hedw.) Broth.	X			X				Yano & Lisboa (1988)	Mori & Cardoso 17137 (n.v.)
Pylaisiadelphaceae
Isopterygium subbrevisetum (Hampe) Broth.	X							Lisboa et al. (2006)	Costa Neto 1468 (n.v.)
Isopterygium tenerum (Sw.) Mitt.	X	X	X					Lisboa et al. (2006)	Gentil 88
Pterogonidium pulchelium (Hook.) Müll. Hal.	X							Lisboa et al. (2006)	Costa Neto 1305 (n.v.)
Taxithelium concavum (Hook.) J. Florsch.	X							Lisboa et al. (2006)	Costa Neto 1478 (n.v.)
Taxithelium planum (Brid.) Mitt.	X	X	X	X	X			Yano & Lisboa (1988)	Gentil 351
Taxithelium pluripunctatum (Renauld & Cardot) W.R.Buck			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 404
Sphagnaceae
Sphagnum palustre L.				X				Yano et al. (1985)	Egler 1448 (n.v.)
Sphagnum perichaetiale Hampe				X				Yano & Lisboa (1988, as Sphagnum erythrocalyx Hampe)	Froés 26752 (n.v.)
Sematophyllaceae
Acroporium pungens (Hedw.) Broth.				X	X			Yano & Lisboa (1988)	Mori & Cardoso 17292 (n.v.)
Brittonodoxa subpinnata (Brid.) W.R.Buck et al.	X		X					Lisboa et al. (2006)	Gentil 497
Microcalpe subsimplex (Hedw.) W.R.Buck		X	X	X	X	X		Yano & Lisboa (1988)	Gentil 92
Trichosteleum papillosum (Hornsch.) A. Jaeger	X	X	X	X	X	X		Yano & Lisboa (1988)	Gentil 520
Trichosteleum sentosum (Sull.) A. Jaeger			X					Oliveira-da-Silva & Ilkiu-Borges (2020)	Gentil 528
Trichosteleum subdemissum (Besch.) A. Jaeger	X	X	X					Lisboa et al. (2006)	Gentil 359
Vitalia caespitosa (Hedw.) P.E.A.S. Câmara et al.						X		Yano & Lisboa [1988, as Sematophyllum caespitosum (Hedw.) Mitt.]	Mori et al. 17491 (n.v.)
Stereophyllaceae
Pilosium chlorophyllum (Hornsch.) Müll. Hall.		X	X	X				Yano & Lisboa (1988)	Gentil 153
­­
Pelekium involvens (Hedw.) Touw		X						Yano & Lisboa (1988)	Rosário s.n. (MG99129)
Pelekium scabrosulum (Mitt.) Touw	X	X	X					Lisboa et al. (2006)	Gentil 269
Pelekium schistocalyx (Müll. Hal.) Touw	X							Lisboa et al. (2006)	Costa Neto 1298 (n.v.)
Total	60	77	109	42	9	14	2

Figure 1
Species diversity by bryophyte families in the Amapá State.
Figura 1
Diversidade de espécies por famílias de briófitas no Estado do Amapá.

Figure 2
Localization map of Amapá State with number of species record from State municipalities.
Figura 2
Mapa de localização do Estado do Amapá com número de espécies registradas nos municípios do Estado.

Figure 3
Measurement of new records by study of bryophytes in the State of Amapá.
Figura 3
Medida de novos registros por estudo de briófitas no Estado do Amapá.

The municipalities of Serra do Navio and Macapá harbored the largest number of bryophytes (figure 2). These results are due to intensive inventories of bryophytes in specific areas from Amapá, such as the studies conducted by Oliveira-da-Silva & Ilkiu-Borges (2020) in two municipal natural parks of upland Amazon forest in these two municipalities and by Lisboa et al. (2006) in coastal areas of the Amapá municipality with collections in different phytophysiognomy such as várzea, mangrove, transition forest, and other vegetation types (figure 3).

There are still large areas lacking investigation works on bryophytes, such as the municipality of Laranjal do Jari. Alone, this municipality covers 30,783 km² which represents more than 20% of the territory of Amapá (IBGE 2019) and comprises important conservation units such as the National Park of Tumucumaque Mountains (Drummond et al. 2008). However, only Octoblepharum pulvinatum (Dozy & Molk.) Mitt. and Leucobryum martianum (Hornsch.) Müll. Hal. were registered in this municipality.

It is expected that with further expeditions to unexplored and important areas of Amapá, such as Parque Nacional Montanhas do Tumucumaque, Reserva Extrativista do Rio Jari and Floresta Nacional do Amapá, the number of species may considerably increase. The results reinforced the need for more comprehensive studies on the bryoflora from Amapá.

Acknowledgements

The authors are gratefull to the Programa de Pós-Graduação em Ciências Biológicas - Botânica Tropical of Museu Paraense Emílio Goeldi/Universidade Federal Rural da Amazônia; to the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for the Doctoral fellowship grant of the first author (process n° 141523/2020-4) and for the productivity fellowship grant of the third author (process n°302374/2016).

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Associate Editor: Regina Hirai

Publication Dates

• Publication in this collection
  13 Dec 2021
• Date of issue
  2021

History

• Received
  29 June 2020
• Accepted
  26 Oct 2020