Phytotaxa 346 (3): 203–220 ISSN 1179-3155 (print edition)

http://www.mapress.com/j/pt/
Article PHYTOTAXA
Copyright © 2018 Magnolia Press ISSN 1179-3163 (online edition)

https://doi.org/10.11646/phytotaxa.346.3.1

Bryofloristic affinities betwwen Itatiaia National Park and tropical Andean

countries
DENISE PINHEIRO COSTA1, GABRIEL POLITZER COUTO2, MARINEZ FERREIRA SIQUEIRA1 & STEVEN
P. CHURCHILL3
1
Instituto de Pesquisas Jardim Botânico do Rio de Janeiro – JBRJ, Rua Pacheco Leão 915, 22460-030, Rio de Janeiro, RJ, Brazil,
e-mail: dcosta@jbrj.gov.br
2
Pontifícia Universidade Católica – PUC-RJ, Rua Marques de São Vicente 225, 22430-060, Rio de Janeiro, RJ, Brazil,
e-mail: gabrielpolitzer@gmail.com
3
Missouri Botanical Garden – 2345 Tower Grove Avenue, PO Box 299, 63166-0299, St. Louis, Missouri, USA,
e-mail: steve.churchill@mobot.org
Corresponding author: dcosta@jbrj.gov.br
ORCID: 0000-0001-9495-3029

Abstract

Itatiaia National Park (INP) is an interesting biogeographically location in southeastern Brazil, with a unique bryoflora due
to its varied vegetation formations (montane forest, upper montane forest, and campos de altitude) and subtropical climate.
The bryoflora of the INP consists of 519 species, 213 genera, and 81 families (289 mosses and 230 liverworts), with 84 taxa
considered endemic, representing 34% of the total Brazilian bryoflora and 42% of the Atlantic Rainforest. Mosses and liv-
erworts show very similar geographical distributions, with the most common phytogeographical patterns being Neotropical
(219 taxa) and endemic (84 taxa). Endemism is concentrated in the montane forest for liverworts and in the upper montane
forest for mosses. Comparative analyses between the bryoflora of INP and five tropical Andean countries (Bolivia, Peru,
Ecuador, Colombia, and Venezuela) were used to confirm its affinities with the “páramos” phytophysiognomy. Sørensen
similarity tests and cluster analysis were carried out on the similarity matrices for all families, genera, and species. Support-
ing evidence from published data was used to interpret the results. Comparisons made between the bryoflora of INP with
those of five tropical Andean countries mentioned above showed strong familial, generic, and species similarities, which
are explained in the literature by climatic similarities, speciation, long-distance dispersal, and migration through favorable
habitats. At least 40% of the bryoflora species are shared between INP and the other tropical Andean countries, despite their
vast size differences. We emphasize the need for adequate protection for the bryoflora of INP, specially its endemic and dis-
junct species, due to anthropogenic threats (tourism, incorrect trail maintenance, trampling, fire, litter, and construction) and
global warming, which will accelerate the alteration of mountain environments.

Key words: Brazil, conservation, endemic, liverworts, mosses, South America

Introduction

The Atlantic Coastal Region of Brazil has several mountain ranges with cloud forests, Araucaria forests, and high-
altitude fields (campos de altitude). The mountains there extend from the states of Minas Gerais and Espírito Santo
to São Paulo, and then into southern Brazil; the principal ranges are the Serra da Mantiqueira and the Serra do Mar.
The highest elevations are found in the Serra da Mantiqueira: Pico da Bandeira (2,890 m), Serra Fina (2,795 m), and
Agulhas Negras (2,787 m). Occasional snowfalls occur in the Serra da Mantiqueira. Those high mountains have rich
floras with many endemic species with interesting floristic affinities with the Andes. The campos de altitude of the
Itatiaia National Park (INP) above the tree line (ca. 2,300 m), for example, resembles the treeless “páramos” of the
northern Andes (Costa et al. 2015; Gradstein & Costa 2003; Safford 1999a, b).
Although the INP is 2000 km away from any of the nearest Andean subranges and separated from them by
lowlands, the park supports many taxa of Andean origin (Brade 1956; Brown 1987; Martinelli & Bandeira 1989;
Safford 1999a). Safford (2007) noted that this Andean connection has attracted the interest of biogeographers for
more than 50 years, with Brade (1956) being the first to closely examine the biogeography of the flora of the INP and

Accepted by Lars Söderström: 8 Mar. 2018; published: 6 Apr. 2018 203

delimit five source areas for the mountain flora: (1) subtropical rain forest, (2) central Brazil (xerophytic element),
(3) ‘Antarctic’ (widespread in southern South America), (4) ‘austral-Andean’ (southern taxa with extensions into the
tropical Andes), and (5) ‘Andean’ (taxa shared with the tropical Andes). Rambo (1951, 1953, 1956) documented an
Andean affinity of the dominant species in the southern Brazilian Highlands.
Safford (2007) examined the phytogeography of the campos de altitude vascular flora in the mountains
of southeastern Brazil, including the INP, and concluded that many taxa have their ancestries linked to temperate
latitudes, with tropical components primarily derived from the drier, highland environments of the Brazilian interior;
and temperate components increasing in importance with increasing elevation. Costa et al. (2015) focused on the
elevation gradients on the bryoflora of the INP and concluded that the park harbors unique bryofloras. The bryophyte
diversity there is high (comprising 57% of the bryophyte flora of all of Rio de Janeiro State, and 34% of the total
Brazilian bryoflora), and exhibits differences in both species richness and floristic composition along an elevation
gradient.
We therefore evaluated the bryoflora of the INP and analyzed its phytogeographic patterns, as well as its endemism
and floristic affinities with the tropical Andean countries in order to address the following questions: (1) What are
the phytogeographic patterns of bryophyte species in the INP? (2) How do the phytogeographic patterns of the INP
bryophytes change along its elevation gradient? (3) Are endemic species more common in certain vegetation types in
the INP (montane forest, upper montane forest, and campos de altitude)? (4) Does the bryoflora of the INP show the
same degree of Andean affinities shown by its vascular plants?

Material and Methods

Study area
The Itatiaia National Park (“cliff full of peaks” in the indigenous Tupi language) is located in southeastern Brazil (Fig.
1) and comprises approximately 30,000 hectares (22º16’–22º28’S and 44º34’–44º42’W) between the southwestern
Rio de Janeiro State (the municipalities of Resende and Itatiaia) and southern Minas Gerais State (the towns of Alagoa,
Bocaina of Minas, and Itamonte). Park elevations vary between 600–2,787 m, with the highest summits of the park
being the Pico das Agulhas Negras (2,787 m), Maciço das Prateleiras (2,548 m), and Pedra do Altar (2,530 m). The
climate there is subtropical with distinct hot/dry and then humid/cool seasons. Humidity averages 85%, local annual
rainfall averages 2,100 mm, the annual mean? temperature is 11ºC (ranging from 6ºC to a maximum of 27ºC) (Brade
1956; Hueck 1972; Pádua & Coimbra Filho 1979). The vegetation of the INP is characterized as “Dense Atlantic
Rainforest”, with four principal vegetation types that follow an elevation gradient, with distinct soils and floristic
compositions: montane forest (500–1,499 m); upper montane forest (1,500–2,300 m); and campos de altitude (2,300–
2,790 m) (Veloso et al. 1991). The campos de altitude have a peculiar phytophysiognomy on rocky outcrops (Martinelli
& Bandeira 1989). The geographic localization of INP, between two large urban centers, and the fact that the campos
de altitude are phytogeographical islands in an extensive and varied landscape, characterizes this region as a nucleus
of the Atlantic Forest Biosphere Reserve, one of the largest conservation units in the world (Conservation International
of Brazil 2000).

Database
This paper is based on published data (Benitez & Gradstein 2011; Benitez et al. 2012; Buck 2003; Churchill 1988,
1994, 2009; Churchill et al. 2009 a,b, 2000, 2015; Churchill & Griffin 1999; Churchill & Linares C. 1995; Costa
inedit; Costa & Lima 2005; Costa et al. 2011, 2015; Dauphin & Ilkiu-Borges 2002; Dauphin et al. 2008; Gradstein
1999; Gradstein & Ilkiu-Borges 2009; Gradstein et al. 2003, 2006; León-Yanez et al. 2006; Menzel 1984, 1992;
Morales et al. 2007; Nöske et al. 2003; O’Shea 2010; Robinson 1976; Santos & Costa 2010; Schäfer-Verwimp 1992;
Schäfer-Verwimp & Giancotti 1993; Schäfer-Verwimp & Vital 1989; Schäfer-Verwimp et al. 2006, 2013; Uribe
& Gradstein 1998; Winkler 1976); specimens deposited in the Rio de Janeiro Botanical Garden (RB), The New
York Botanical Garden (NY) herbaria, as well as in the private herbarium Schäfer-Verwimp; and on databases from
Churchill (Mosses from the Andes—tropicos.org). A complete species list is presented in Supplementary Table 1.
Precise elevation data were obtained from specimen labels. The lists of bryophytes from tropical Andean countries
are based on the literature cited above (ca. 2600 taxa) and summarized on the following web site: Churchill and
Collaborators 2009–2017 (Mosses from the Andes—tropicos.org). The classification of the Phylum Marchantiophyta
follows Crandall-Stotler et al. (2009), and that of the Phylum Bryophyta follows Buck and Goffinet (2009). Floristic

204 • Phytotaxa 346 (3) © 2018 Magnolia Press COSTA ET AL.

lists are arranged by family, genus, and species; recent changes in taxonomy and nomenclature were updated in the
final checklist to make them comparable across regions. The final species list comprises 2,418 species (1,487 mosses
and 931 liverworts). The Venezuelan species list considered here is not complete.

FIGURE 1. Map illustrating the location of the Itatiaia National Park, Rio de Janeiro State, Brazil (source: Center for Scientific Computing
and GIS-NCCG/JBRJ).

Phytogeographic patterns
Each taxon was assigned to a phytogeographical category based on its known global distribution as determined from
all available information sources (literature, labels, and databases). We analyzed the phytogeographical patterns along
an elevation gradient within the INP, considering intervals of 200 m. The distribution patterns of the species throughout
the world and in Brazil were determined by using Gradstein & Costa (2003) and the database of the Lista de Espécies
da Flora do Brazil (Costa and Peralta 2015).

Floristic affinities
Qualitative and quantitative comparisons were made between the bryofloras of the INP and five tropical Andean
countries (Venezuela, Colombia, Ecuador, Peru, and Bolivia) according to the floristic data and Churchill and
Collaborators database (2009–2017). Families, genera, and species of each of the six areas were used as input for
cluster analyses. Resemblances between the bryoflora of the six areas were quantified using Sørensen similarity index
for distance measurements and Unweighted Par Group Method with Arithmetic Mean—UPGMA (DRYFLOR et al.
2016; Kreft & Jetz 2010; Zhang & Corlett 2003).
We compared the numbers of shared species and the endemism of each country with that of the INP, and constructed
similarity dendrograms to analyze the hierarchical levels of the species, genera and families of mosses and liverworts.
Bootstrap with 1000 replicates was used to analyze the support of each node.
We used the Sørensen coefficient to calculate the similarity between areas. Smith (1986) considered this index
the best choice when only presence/absence is available. The analyses were performed using PAST v. 2.17 software
(Hammer et al. 2001).

BRYOFLORISTIC AFFINITIES Phytotaxa 346 (3) © 2018 Magnolia Press • 205

Results

The current bryophyte species list from the six tropical America sites includes 115 families (72 mosses and 43 liverworts),
495 genera (343 mosses and 152 liverworts), and 2,418 species (1,487 mosses and 931 liverworts) (Supplementary
Table 1).

The bryoflora of INP

The bryoflora of the INP consists of 519 species (in 213 genera and 81 families), of which 289 are mosses and
230 liverworts. The largest families are Lejeuneaceae (71 taxa), Dicranaceae (40 taxa), Sphagnaceae (21 taxa), and
Orthotrichaceae (19 taxa), Metzgeriaceae, Plagiochilaceae and Radulaceae (16 taxa each) and Pilotrichaceae (12
taxa). The largest genera are Sphagnum Linnaeus (1753: 1106) (21 taxa), Metzgeria Raddi (1818: 34), Plagiochila
(Dumortier) Dumortier (1835: 14) and Radula Dumortier (1822: 112) (16 taxa each). Eighty-four taxa are endemic (62
mosses and 22 liverworts) (Supplementary Table 1).

Phytogeographical patterns
Eleven phytogeographical categories were recognized for INP bryophytes: (1) Worldwide distribution—WIDE (the
third largest pattern of the local bryophyte flora [e.g. Metzgeria leptoneura Spruce (1885: 555) and Bryum argenteum
Hedwig (1801: 181–182]); (2) Pantropical—PAN (widely distributed throughout all tropical regions [e.g. Lejeunea
flava (Swartz 1788: 144) Nees (1838: 277) and Campylopus arctocarpus (Hornschuch 1840: 12) Mitten (1869: 87)]);
(3) Tropical and subtropical America—TSBA (distributed in all tropical regions, extending to the subtropics [e.g.
Drepanolejeunea araucariae Stephani (1896: 80) and Rhodobryum grandifolium (Taylor 1847: 336) Schimper (1898:
1116)]); (4) Neotropical—NEO (the largest pattern of the local bryoflora, and prevalent for both liverworts and mosses
[e.g. Adelanthus carabayensis (Montagne 1856: 348) Grolle (1972: 348) and Callicostella pallida (Hornschuch
1840: 64) Ångstr. (1876: 27)]); (5) Tropical South America—TSA [e.g. Drepanolejeunea araucariae Stephani
(1896: 80) and Breutelia subdisticha (Hampe 1872: 50) Hampe (1875: 94)]); (6) Holarctic—HOLO (with only three
liverworts taxa [Lophozia bicrenata (Schmidel 1797: 247) Dumortier (1835: 17), Solenostoma hyalinum (Lyell 1816:
63) Mitten (1870: 319), and Solenostoma sphaerocarpum (Hooker 1816: 74) Stephani (1901: 499)]); (7) Tropical
amphi-atlantic—AF-AM (according to Gradstein et al. [1983], this is a typical feature of tropical liverworts flora,
and for Santos & Costa [2010], Afro-American disjunction predominates in the liverwort flora of Rio de Janeiro [e.g.
Symphyogyna brasiliensis Nees (1836: 67) and Adelothecium bogotense (Hampe 1866: 303) Mitten (1869: 391)]);
(8) Andean Element—AND (according to Gradstein and Costa [2003], this represents the similarity between the high
plateau of southeastern Brazil and the páramos of the northern Andes, particularly in Itatiaia (e.g. Diplasiolejeunea
replicata Stephani (1916: 926), Drepanolejeunea granatensis (Jack & Stephani 1892: 17) Bischler (1964: 150), and
Eurhynchium clinocarpum (Taylor 1848: 194) Paris (1896: 442)); (9) Disjuncts between the high mountains of Central
America, Andes and southeastern Brazil—CAA (Syzygiella anomala (Lindenberg & Gottsche 1844: 646) Stephani
(1902: 471)); (10) Southern temperate element—STE (widely distributed or sometimes scattered [e.g. Symphyogyna
podophylla (Thunberg 1800: 174) Montagne & Nees (1846: 481) and Andreaea subulata Harvey (1840: 201)]); (11)
Endemic of Brazil—BRA (the second largest category of the local bryoflora [e.g. Kurzia brasiliensis Grolle (1963:
174) and Syzygiella uleana Stephani (1917: 118); Sphagnum longicomosum Müller ex Warnstorf (1900: 257) and
Itatiella ulei (Brotherus ex Müller 1898: 26) Smith (1971: 52)]). All taxa with deficient data or data that could not be
allocate to any other pattern were identified as “Pattern not identified—OTHER” (e.g. Cephaloziella divaricata (Smith
1800: 719) Schiffner (1893: 99) and Ctenidium malacodes Mitten (1969: 509)) (Table 2).
The total number of endemic taxa is high for mosses (62 species) and for liverworts (highlighting disjunct patterns)
(a total of 36 species, being three Holarctic, 17 Tropical amphi-atlantic, nine Andino element, four Central America,
Andes and southeastern Brazil, and three southern hemisphere and south-eastern Brazil).

Phytogeographical patterns along the elevation gradient

The phytogeographical patterns were combined into five categories for this analysis (Table 3): (1) Widespread (55
taxa; 12 liverworts and 43 mosses); (2) Holarctic (3 liverwort taxa); (3) Tropical—including PAN, TSBA, NEO, and
TSA (273 taxa; 128 liverworts and 145 mosses); (4) Endemic to Brazil (84 taxa; 22 liverworts and 62 mosses); (5)
Disjunct—including AF-AM, AND, CAA, and STE (41 taxa; 30 liverworts and 11 mosses). The Tropical element is
the largest of the five, comprising ca. 50% of the total taxa in terms of elevation ranges.

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TABLE 2. Phytogeographical patterns of the bryoflora of the INP, with total number for liverworts and mosses. Percentage
in brackets.
Pattern Liverworts Mosses Total (%)
1. Widespread (found at least in three continents) WIDE 12 (2.3) 43 (8.3) 55 (10.6)
2. Pantropical (wide tropical) PAN 14 (2.7) 24 (4.6) 38 (7.3)
3. Tropical and subtropical America TSBA 1 (0.2) 4 (0.8) 5 (1)
4. Neotropical NEO 104 (20) 115 (22.1) 219 (42.1)
5. Tropical South America TSA 9 (1.7) 2 (0.4) 11 (2.1)
6. Holartic (disjunct between temperate regions of northern HOLO 3 (0.6) --- 3 (0.6)
hemisphere and southeastern Brazil)
7. Tropical amphi-atlantic (disjunct between Africa and America) AF-AM 17 (3.3) 9 (1.7) 26 (5.0)
8. Andino Element (disjunct between northern Andes and Atlantic AND 9 (1.7) 1 (0.2) 10 (1.9)
Forest of southeastern of Brazil)
9. Disjunct between the high mountains of Central America, Andes CAA 4 (0.8) --- 4 (0.8)
and southeastern Brazil
10. Southern temperate Element (disjunct between temperate STE 3 (0.6) 1 (0.2) 4 (0.8)
regions of southern hemisphere and south-eastern Brazil)
11. Endemic of Brazil BRA 22 (4.2) 62 (12) 84 (16.2)
12. Pattern not identified OTHER 32 (6.2) 28 (5.4) 60 (11.6)
TOTAL 230 289 519 (100)

The analyses of the phytogeographical patterns along the elevation gradient (at intervals of 200 m), demonstrated
that the liverworts and mosses exhibit distinct patterns (Fig. 2). For liverworts, we found a low number of widespread
species (<10%), highlighting species with disjunct patterns, including taxa disjunct between the northern Andes and
other tropical America mountains. The proportion of disjunct taxa show a peak at 2,000–2,200 m; holartic species were
only found between 2,000–2,400 m (the transition from upper montane forest and campos de altitude). In terms of
mosses, a higher number of endemic and widespread species were represented, with disjunctions being rare (<10%),
except above 2,600 m (in the campos de altitude) (Fig. 2).

Endemic species
The bryophyte flora of INP consists of 84 endemic species (22 liverworts and 62 mosses) among the 519 recognized
species (belonging to 78 families and 213 genera) (Supplementary Table 1).
In terms of liverworts, of the 22 endemic species, 16 are restricted to the Atlantic Rainforest (13 of those are
widespread and occur from northeastern to southern Brazil, while three are limited to the forests of southeastern and
southern Brazil); two taxa area exclusive to southeastern Brazil (Isotachis inflata Stephani (1903: 113) and Riccardia
emarginata (Stephani 1893: 20) Hell (1969: 100); only one taxon is restricted just to INP (Colura itatyana Stephani
(1916: 932)).
In terms of mosses, from the 62 endemic species, 29 are restrict to the Atlantic Rainforest (nine of those are
widespread and occur from northeastern to southern Brazil while 20 occur in the forests of southeastern and southern
Brazil); 10 taxa are exclusive to southeastern Brazil; 11 taxa are restricted to the INP (Ditrichum itatiaiae (Müller
1898: 41) Paris (1900: 131), Eobruchia bruchioides (Müller 1898: 42) Buck (1979: 470), etc.).
In terms of the distributions of endemic species in the different vegetation types of INP, we found distinct results
for liverworts and mosses: among liverworts, most of the endemic species are concentrated in the montane forest; for
mosses, most endemic species are concentrated in the Upper montane forest (Fig. 2).

Comparisons with the tropical Andean countries

There are more species, genera, and families of mosses than liverworts in all the sites analyzed and more endemic
species of mosses than liverworts (with the exception of Colombia, which has more endemic liverworts) (Table 3).
Colombia has the highest number of endemic species (116; being 42 mosses and 74 liverworts), representing 7.3% of
the total bryoflora. When we analyze the percentages of endemics by sites, however, INP was found to have the highest
endemism (16.2%), even representing 1/2-1/3 of the total number of species of the other countries (even though it
comprises a much smaller area) (Table 3).

BRYOFLORISTIC AFFINITIES Phytotaxa 346 (3) © 2018 Magnolia Press • 207

FIGURE 2. Percentage of the phytogeographical patterns of mosses (A) and liverworts (B) along an elevation gradient of the INP.

208 • Phytotaxa 346 (3) © 2018 Magnolia Press COSTA ET AL.

TABLE 3. Species richness (mosses and liverworts) of the INP and Andean countries (endemism and shared families,
genera, and species).M= mosses; L = liverworts.
Sites INP VENEZUELA COLOMBIA ECUADOR PERU BOLIVIA
Group M L M L M L M L M L M L
Families 49 29 61 31 64 39 64 38 61 35 66 37
Families shared 46 26 47 28 46 28 46 28 47 28
(%) - - (75.4%) (83.9%) (73.4%) (71.8%) (71.9%) (73.7%) (75.4%) (80.0%) (71.2%) (75.7%)
Genera 132 81 229 79 261 132 242 112 242 100 268 104
Genera shared 113 56 117 74 113 69 114 63 113 68
(%) - - (49.3%) (70.9%) (44.8%) (56.1%) (46.7%) (61.6%) (47.1%) (63.0%) (42.2%) (65.4%)
Spp. 284 235 711 193 878 710 811 448 799 357 926 402
Total spp. 519 904 1588 1259 1156 1328
Spp. shared (%) 155 62 150 160 152 135 161 112 159 123
- - (21.8%) (32.1%) (17.1%) (22.5%) (18.7%) (30.1%) (20.2%) (31.4%) (17.2%) (30.6%)
Total spp. 217 310 287 273 282
shared - (24%) (19.5%) (22.8%) (22.8%) (23.6%)
Endemic (%) 62 22 12 10 42 74 39 12 28 2 60 20
(12%) (4.2%) (1.7%) (5.2%) (4.8%) (10.4%) (4.8%) (2.7%) (3.5%) (0.6%) (6.5%) (5.0%)
Total endemic 84 22 116 51 30 80
(%) (16.2%) (2.4%) (7.3%) (4.1%) (2.6%) (6.0%)

The numbers of moss and liverwort families and genera in the tropical Andean countries are very similar: (families:
between 61 and 66 for mosses and 31–39 for liverworts; genera: between 229 and 268 for mosses and 79–132 for
liverworts). In terms of species, the numbers are similar for mosses (711 to 926) but very different for liverworts (193–
710). This result may be related, however, to the reduced number of publications available for liverworts in Venezuela
(193 spp.), which may be underestimating their diversity (Table 3).
Although the INP has lower numbers of moss and liverworts families and genera as compared with the Andean
countries, the percentages of shared taxa are high: above 70% for families for both mosses and liverworts, and genera,
above 42% for mosses, and above 56% for liverworts. The percentages of shared species of mosses and liverworts with
the Andean countries are also similar, ranging between 17–32% (Table 3).
The numbers of shared species between the INP and the tropical Andean countries is considered high, averaging
from 40–60% (385 species [189 mosses and 196 liverworts] shared with at least one country). The values of the shared
species are presented in Fig. 3. The Andean country that shared the most species with the INP was Colombia (59.7% of
the total number of species). Venezuela, on the other hand, has the lowest number of species shared with INP (41.8%).
The other countries shared very similar numbers of species with INP (between 53–55%) (Table 3, Fig. 3).

Bryofloristic affinities between INP and tropical Andean countries (Bolivia, Colombia, Ecuador, Peru, and
Venezuela)
Comparisons were made between the bryofloras of INP and those of five tropical Andean countries. The Sørensen
similarities between the six localities analyzed are represented in Figs. 4–5. The bryofloras most similar to INP are
those of Colombia and Ecuador, with 60% and 55% similarities respectively (Table 3 and Fig. 3). Our results show
that greater numbers families, genera, and species of liverworts are shared than those of mosses (Table 3). Bolivia,
Colombia and Ecuador share the most more moss species with INP, while Bolivia, Ecuador and Peru share the most
liverworts species with INP.
For mosses in terms of family-level comparisons, two high similarity clusters were observed (0.83 and bootstrap
100%): INP and VEN-COL-ECU-PER-BOL. The closest family-level affinity found was COL with VEN, with 60
shared families (similarity of 0.96 and bootstrap of 33%) (Fig. 4, Table 4). All sites had similarities above 0.93,
except for INP. For mosses, in terms of genus-level comparisons, three high similarity clusters were observed (0.6 and
bootstrap 100%): INP, BOL-PER, and ECU-COL-VEN. The closest genus-level affinity found was COL with ECU,
with 223 shared genera (similarity of 0.89 and bootstrap of 71%); COL-ECU-VEN (196 shared genera) and PER-
BOL (218 shared genera) formed two groups with similarities of 0.84 and bootstrap of 100% (Fig. 4, Table 4). For
mosses, in terms of species-level comparisons, three clusters of high similarity were similar to those found for genera,
but the INP site was separated from the tropical Andean countries, with a lower similarity value (ca. 0.28, with 118
shared species); the VEN-COL-ECU-PER-BOL group of Andean countries showed a similarity above 0.7 (394 shared
species) (Fig. 4, Table 4).

BRYOFLORISTIC AFFINITIES Phytotaxa 346 (3) © 2018 Magnolia Press • 209

FIGURE 3. Percentages of species shared by INP and the five other tropical Andean countries considered. The sizes of the circles
represent the total numbers of species by locality; species turnovers between localities (proportional to the number of shared species) are
represented by the widths of the lines (values from Table 3).

FIGURE 4. Cluster analyses of mosses families, genera, and species employing the Sørensen index (A. Families. B. Genera. C.
Species).

210 • Phytotaxa 346 (3) © 2018 Magnolia Press COSTA ET AL.

TABLE 4. Sørensen similarities between locations at the levels of family, genus, and species for mosses and liverworts.
0RVVHV)DPLOLHV  0RVVHV*HQHUD  0RVVHV6SHFLHV
 ,13 9(1 &2/ (&8 3(5   ,13 9(1 &2/ (&8 3(5  ,13 9(1 &2/ (&8 3(5
9(1       9(1       9(1     
&2/       &2/       &2/     
(&8       (&8       (&8     
3(5       3(5       3(5     
%2/       %2/       %2/     
                  
/LYHUZRUWV)DPLOLHV  /LYHUZRUWV*HQHUD  /LYHUZRUWV6SHFLHV
 ,13 9(1 &2/ (&8 3(5   ,13 9(1 &2/ (&8 3(5  ,13 9(1 &2/ (&8 3(5
9(1       9(1       9(1     
&2/       &2/       &2/     
(&8       (&8       (&8     
3(5       3(5       3(5     
%2/       %2/       %2/     

In terms of liverworts, family-level comparisons showed three high similarity clusters (0.85 and bootstrap of
100%): INP; VEN; and COL-PER-ECU-BOL. The closest family-level affinity found was COL-PER with 30 shared
families (similarity above 0.95 and bootstrap of 52%). All sites had similarities above 0.88, except for INP (Fig. 5,
Table 4). In terms of liverworts, three clusters were formed in genera-level comparisons: VEN; INP; and COL-ECU-
BOL-PER. The closest genera-level affinity found was COL-ECU-BOL-PER, with 73 shared genera (similarity above
0.81 and bootstrap of 99%). INP and VEN formed isolated groups (0.71 and 0.69, respectively) (Fig. 5, Table 4).
In terms of liverworts, three clusters were formed in species-level comparisons: INP; VEN; and COL-ECU-BOL-
PER. The closest species-level affinity found was COL and ECU, with 369 shared species (similarity above 0.64 and
bootstrap of 99%). Again, INP and VEN formed isolated groups (0.36 and 0.40, respectively) (Fig. 5, Table 4).

FIGURE 5. Cluster analyses of liverwort families, genera, and species employing the Sørensen index (A. Families. B. Genera. C.
Species).

Discussion

Bryoflora of INP and the phytogeographical patterns

Our results reinforce the importance of the INP as one of the most critical Atlantic Forest remnants of southeastern
Brazil for the conservation of the bryophytes as it holds approximately 34% of the total bryophyte diversity of Brazil
and 21% of that nation’s endemic species in a very small area—with the phytogeographical patterns of the liverworts
and mosses changing along the elevation gradient.

BRYOFLORISTIC AFFINITIES Phytotaxa 346 (3) © 2018 Magnolia Press • 211

 The composition of the bryoflora is consistent within a subtropical climate coupled with the climatic limits of
the tropics. The Neotropical element is the main phytogeographical pattern in the INP, representing ca. 50% of the
total taxa occurring along the elevation gradient, although species with restricted distributions (endemics mosses and
disjunct liverworts) are conspicuous.
Several features of the INP bryoflora are typical of tropical floras, exemplified by the presence of 12 dominant families
including the liverworts Frullaniaceae, Lejeuneaceae, Plagiochilaceae, and Radulaceae (Cornelissen & Gradstein,
1990; Gradstein, 1992) and the moss families Dicranaceae, Orthotrichaceae, Pilotrichaceae, Sematophyllaceae, and
Sphagnaceae. Likewise, genera that have their centres of diversity in tropical America include the liverwort genus
Neesioscyphus Grolle (1964: 19) and the moss genus Sphagnum Linnaeus (1753: 1106) (Gradstein et al. 2001; Shaw
et al. 2003). Bryophytes epiphyllous on living leaves are another characteristic feature of tropical forests (Richards
1984), and they are locally well-developed and very visible in the INP, with at least 17% being liverwort species (40
taxa; e.g., Colura itatyana Stephani (1916: 932)).
All 15 bryophyte families that characterize tropical rain forests (Gradstein & Pócs 1984) occur in the INP, as well
as some typical temperate families, such as, Grimmiaceae, Hylocomiaceae, and Plagiotheciaceae (which largely occur
at mostly at high elevations [> 1,900 m] in INP). Two genera characteristic of the Atlantic Rainforest, Balantiopsis
Mitten (1867: 751) and Saccogynidium Grolle (1960: 43), are encountered in INP, emphasizing the tropical America
aspect of its bryoflora (Gradstein et al. 2001).
Examples of Neotropical taxa found below 1,000 m include Brachymenium consimile Jaeger (1875: 120),
Leptostomopsis systylia (Müller 1848: 320) Spence & Ramsay (2005: 70), Herbertus juniperoideus (Swartz 1788:
144) Grolle (1961: 80) and Lophocolea platensis Massalongo (1906: 12). Andean taxa found in INP are more diverse
at higher altitudes (> 2,000 m) (Herbertus sendtneri (Nees 1838: 575) Evans (1917: 212), Lepidozia inaequalis
(Lehmann & Lindenberg 1833: 1) Lehmann & Lindenberg (1845: 209), and Marsupella microphylla Schuster (1978:
249), and South temperate elements found above 2,100 m include Lophocolea muricata (Lehmann 1829: 363) Nees
(1845: 169) and Andreaea subulata Harvey (1840: 201).

Elevation gradient
Liverworts and mosses shown different phytogeographical patterns along the INP elevation gradient. Endemic mosses
predominate above 1,000 m while liverwort endemism is rare, but well spaced above 600 m along the elevation gradient.
Disjunctions liverworts are more common, occurring throughout the elevation gradient, except in the 1,600–1,800 m
interval. Widespread species are more common among mosses, with more than 10% of the moss species occurring
throughout the elevation range, except above 2,600 m. It is generally acknowledged that bryophyte endemism is rare,
while taxa with transoceanic distributions are common (Schofield 1992). Our results corroborate the findings of Costa
& Lima (2005) and Santos & Costa (2010) that attribute the differences observed in the phytogeographical patterns of
the mosses and liverworts of Rio de Janeiro State as a whole to the bryoflora of INP—with its high number of endemic
mosses in contrast to liverwort disjunctions. Costa et al. (2015) found that high endemism observed among mosses in
the Upper montane forests of INP was due to the life forms and gametophyte morphologies of those species, which
enabled them to colonize open environments (e.g., rock outcrops, exposed soils) that generally increase in availability
with increasing elevation.
Among the disjunct patterns, an Afro-American distribution is a typical feature of tropical liverworts and a likely
result of long-distance spores dispersal by transatlantic air currents (Gradstein et al. 1983). The presence of Andean
elements in the liverwort flora of INP can likewise be explained by long-distance spore dispersal or by vegetative
propagule (Santos & Costa 2010; Gradstein & Costa 2003). Disjunct ranges are clearly related to the excellent dispersal
capacities of many tropical liverwort species. According to Gradstein & Costa (2003), the high peaks of the INP and
the “páramo” characteristic of the Campos de Altitude benefit the establishment of large numbers of Andean species
such as Marsupella microphylla Schuster (1978: 249) (INP and Andes of Venezuela), Aureolejeunea fulva Schuster
(1987: 429), Drepanolejeunea granatensis (Jack & Stephani 1892: 17) Bischler (1964: 150), Herbertus oblongifolius
(Stephani 1909: 11) Gradstein & Cleef (1977: 398) (INP and Andes of Colombia), rare Andean disjuncts species such
as Cephalozia crossii Spruce (1885: 397) and Syzygiella liberata Inoue (1974: 301), as well as temperate taxa such as
Paracromastigum dusenii (Stephani 1900: 48) Schuster (1974: 700) (INP and Chile).

Endemism
The Atlantic Forest of southeastern Brazil is considered one of the main centers of diversity and endemism in the
Americas (Guedes-Bruni & Lima 1997; Myers et al. 2000; Mittermeier et al. 2004). The high indices of biodiversity
and endemism appear to have originated during the Quaternary, when geomorphological events (Batalha-Filho et al.
212 • Phytotaxa 346 (3) © 2018 Magnolia Press COSTA ET AL.
2010) and climatic fluctuations associated with glaciations (Batalha-Filho & Miyanki 2011; Ledru et al. 2015) resulted
in successive expansions (interglacial period) and retractions (maximum glacial period) of the Atlantic Forest from
more restricted forests that had served as refuges for the flora and fauna (Haffer 1969; Vanzolini & Williams 1970;
Rizzini 1997).
Those refuges are currently found in southern Bahia, in the Tabuleiros region of Espírito Santo State, and along
the mountains of Rio de Janeiro and north of São Paulo State (Rambaldi et al. 2003). Integrated within the Atlantic
Forest area, the INP is considered a center of diversity and endemism in tropical America (Brade 1956; Chaves et al.
2014; Costa & Faria 2008, Costa & Santos 2009; Martinelli & Bandeira 1989). According to Safford (1999 a,b), low
temperatures at high altitudes have led to selection of many species endemic to mountain peaks.
The 84 bryophyte species endemic to Brazil found in the INP represent 21% of the total number of endemic
species in the country, and 27% of endemic species of the Atlantic Rainforest (Costa & Peralta 2015; Stehmann et
al. 2009). This strongly emphasizes the importance of the INP to bryophyte conservation in Brazil and the Atlantic
Rainforest—as almost a third of all Brazilian endemic bryophyte species are concentrated in that very small area in
southeastern Brazil.
The largest concentration of endemic liverworts in Brazil is in the Serra da Mantiqueira mountain range within
the Itatiaia and Caparaó National Parks (Gradstein & Costa 2003). Some endemic liverworts species found in the
INP are common and widespread in the Atlantic Rainforest, while others have more restricted distributions. Bazzania
heterostipa (Stephani 1909: 532) Fulford (1959: 410), Cheilolejeunea caducifolia (Gradstein & Schäfer-Verwimp
1993: 64) Ye & Zhu (2010: 280), Herbertus juniperoideus spp. bivittatus (Spruce 1885: 343) Feldberg & Heinrichs
(2006: 328), Metzgeria brasiliensis Schiffner (1964: 22), M. convoluta Stephani (1899: 288), M. cratoneura Schiffner
(1964: 24), Neesioscyphus carneus (Nees 1833: 337) Grolle (1964: 188), Riccardia regnellii Hell (1969: 110), and
Saccogynidium caldense (Ångström 1876: 80) Grolle (1960: 44) are common endemic species known from many
localities, while others are known only from one or two localities, such as Colura itatyana Stephani (1916: 932) (only
INP)—a rare species that occurs on shaded rocks at ca. 2,300–2,400 m in campos de altitude (Gradstein & Costa
2003).
Moss endemism is higher in Brazil than in any other Neotropical country (Gradstein et al. 2001). The endemism
rate is approximately 20% in the Atlantic Rainforest. For the liverworts, these endemisms are concentrated in the
Serra da Mantiqueira mountains, especially in INP (Costa et al. 2011). The INP harbors three of the five moss genera
endemic to Brazil (Cladastomum Müller (1898: 21), Crumuscus Buck & Snider (1992: 39) and Itatiella Smith (1971:
52). Among the widespread species in the Atlantic Rainforest are Brachymenium hornschuchianum Martius (1834: 4),
Lepidopilidium brevisetum (Hampe 1878: 266) Brotherus (1907: 944) and Trichosteleum glaziovii (Hampe 1874: 174)
Buck (1998: 243), which occur from northeastern to southern Brazil.
In terms of the endemic species restricted to INP, Andreaea microphylla Müller (1898: 19) and A. squarroso-
filiformis Müller (1898: 19) are restricted to the campos de altitude surrounding the Agulhas Negras peak, a locality
with intense tourism visitation (mainly climbers)—making that microhabitat extremely threatened by trampling.
Sphagnum subhomophyllum Crum (1997: 156) inhabits marshlands in the montane Forest (ca. 1,400 m) and is only
known from the type specimen (Crum 1997), while S. acutirameum Crum (1992: 423) occurs in the upper montane
forest (2,100–2,200 m) in the transition zone with campos de altitude. The latter is apparently rare and very restricted,
as the type is recorded from only one locality and it has been very rarely sampled in the INP (only two collections
known) (Costa et al. 2015; Crum 1992). Sphagnum triporosum Crum (1995: 173) occurs in exposed areas of campos
de altitude (ca. 2,300 m) near the Agulhas Negras peak (Crum 1995).
Endemic liverworts are concentrated in the montane forest, while endemic mosses are found in the upper montane
forest. Global warming is considered the major threat to high-mountain floras worldwide (Grabherr et al. 2010) and
loss of endemic species would result in greatly reduced biodiversity.

Bryoflora of the INP versus bryoflora of the tropical Andean countries

The term “páramos” originated in Peru, Venezuela, and Colombia to designate the high altitude tropical ecosystems
found between ca. 11°N and 8°S (Lauer 1981; Rundel 1994; Hofstede et al. 2003; etc.). Safford (1999 a,b) used the
expression “Brazilian paìramos” for the Brazilian campos de altitude vegetation types in an attempt to designate its
biogeographic patterns. Several authors have referred to the strong floristic affinity between INP and “páramos”, and
our results show strong species, generic, and family level connections between the bryofloras of INP and the five
tropical Andean countries examined here—and despite their size differences, their percentages of shared species are
quite high, varying between 42% and 60% (Fig. 3).
Bryophytes release large quantities of microscopic spores that in some species can remain viable for long periods
BRYOFLORISTIC AFFINITIES Phytotaxa 346 (3) © 2018 Magnolia Press • 213
of time and be carried long distances by the wind, so that wide geographical separations between suitable habitats often
only represent minor impediments to colonization (Sauer 1988; Barrington 1993). The presence of similar bryofloras
may therefore more strongly reflect environmental similarities than shared biogeographical histories.
The relative similarities between the bryoflora of INP and Andean countries are therefore not surprising and,
according to Safford (1999 a,b), in spite of their very different latitudinal and geographic positions they show
macroclimatic congruities that form the basis for strong Andean–southeast Brazilian biogeographic connections, with
evolutionary and ecological parallelisms in terms of their biota. Frahm (1991) likewise discussed the close connections
between the bryophyte floras of the tropical Andes and southeastern Brazilian mountains, with examples especially in
the genera Atractylocarpus Mitten (1869: 13) and Campylopus Bridel (1819: 71). Those views are corroborated by the
results of the present study as well as several other studies in other neotropical areas (Brade 1956; Troll 1959; Hueck
1972; Schnell 1971; Safford 1999 a, 2007).
According to Safford (1999 b, 2007), climatic and environmental congruencies between the tropical Andean
countries and campos de altitude are underlined by strong genus- and species-level floristic connections between their
floras. Safford (1999 b) used an analysis of macroclimatic patterns to show that campos de altitude (including Itatiaia)
are essentially high-latitude variants of the tropical Andean “páramos”. This view was corroborated by our results, as
at least 42% of bryophyte species of the INP were shared with tropical Andean countries (Fig. 3).
The numbers of the families and genera of mosses and liverworts in tropical Andean countries are surprisingly
similarity despite climatic differences in and longitudinal variations and may reflect the high efficiencies of botanical
inventories. The only exception is in terms of liverworts in Venezuela (193 spp.), which are most likely underestimated
due to the availability of only limited data.

Conclusions

The factors discussed here, and confirmed by the literature, show affinities between the INP and “páramos” bryofloras,
and highlight the following: (1) floristic similarities at the family, genera, and species level are very high between
the INP and tropical Andean countries; (2) between 40–60% of the bryophyte species in INP are shared with tropical
Andean countries; (3) the ecological similarities observed could be a result of environmental similarities and shared
biogeographical histories, as demonstrated by Safford (1999a,b) and Maciel et al. (2017).
Our results also emphasize the need to adequately protect the bryoflora of the INP from anthropogenic disturbances
such as tourism, inadequate trail maintenance, trampling, litter, construction, and fire, as well as from the consequences
of global warming (which will accelerate the degradation of mountain environments), since the bryoflora is rich and
the park is one of the most important Atlantic Forest remnants in the country for the conservation of the bryophyte
species diversity.

Acknowledgements

The authors are grateful to William R. Buck and Alfons Schäfer-Verwimp for their help with the data bases of the
floristic surveys undertaken in INP in regard to the specimens deposited in the NY and G herbaria. This research did
not receive any specific grant from public, commercial, or not-for-profit funding agencies.

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Suplementary

TABLE 1. List of the bryophytes species of six localities (Itatiaia National Park, Venezuela, Colombia, Ecuador, Peru,
and Bolivia). * = Endemic to the INP.

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