Bryophytes of Jaú National Park (Amazonas, Brazil): Estimating species detectability and richness in a lowland Amazonian megareserve

Abstract

In the past decade, floristic studies have rebounded as checklists are fundamental for executing meta-analyses which address ecological, biogeographic and evolutionary questions of broad geographic scope. Despite the importance of checklists as baseline records of local diversity and distributions, few attempts have been made to quantify sampling effort and species detectability within and among study sites. Quantitative floristics, which combines the use of checklists with statistical methods for estimating local richness, is a promising method for characterizing the completeness of checklists especially for cryptic components of biodiversity. For bryophytes, quantifying levels of detectability among substrate types is of central importance, especially in tropical forests where much of their diversity is harbored in difficult to access habitats such as the tree canopy. In light of the need to establish quantifiable protocols of detectability in poorly studied tropical regions, we present a bryophyte checklist for the Jaú National Park (JNP), located in the heart of the Amazon, and estimate local species richness and detectability as it relates to five substrate types (epiphytes, epiphylls, epixylic, epipetric and soil). Identifications from 712 collections made during four excursions over the past decade to JNP revealed 150 species consisting of two new country records and five new state records, along with 20 rarely collected Amazonian endemics. Despite our intensive sampling, which included systematic canopy collections during one of the excursions, Chao richness index estimated that ca. 46 species (nearly one-third of those presently observed) remain undetected from JNP. Furthermore, levels of detectability among substrates varied widely, where observed epiphyte richness, in contrast to the other substrates types, most closely approximated the estimates. Our results illustrate the need for quantitative richness estimates as a means to increase the accuracy of checklist data, particularly when used in meta-analyses addressing global-scale questions. Copyright. © 2018 by The American Bryological and Lichenological Society, Inc.

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Bryophytes of Jaú National Park (Amazonas, Brazil): Estimating species
detectability and richness in a lowland Amazonian megareserve
Authors: Adriel M. Sierra, Alain Vanderpoorten, S. Robbert Gradstein, Marta R. Pereira,
Cid José Passos Bastos, et. al.
Source: The Bryologist, 121(4) : 571-588
Published By: American Bryological and Lichenological Society
URL: https://doi.org/10.1639/0007-2745-121.4.571
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Bryophytes of Jau
´National Park (Amazonas, Brazil): Estimating species
detectability and richness in a lowland Amazonian megareserve
Adriel M. Sierra
1
, Alain Vanderpoorten
2
, S. Robbert Gradstein
3
, Marta R. Pereira
1
,
Cid Jos´
e Passos Bastos
4
and Charles E. Zartman
1,5
1
Instituto Nacional de Pesquisas da Amazˆ
onia, Departamento de Biodiversidade, Av. Andr´
eArau
´jo Aleixo, Caixa Postal
478, 69060001 Manaus, Brasil;
2
Institute of Botany, University of Li`
ege, B22 Sart Tilman, 4000 Li`
ege, Belgium;
3
Institut
de Syst´
ematique, ´
Evolution, Biodiversit´
e (UMR 7205), Mus´
eum National d’Histoire Naturelle - Sorbonne Universit´
es, 57
rue Cuvier, CP 39, 75005 Paris, France;
4
Universidade Federal da Bahia, Instituto de Biologia, Laborat´
orio de Taxonomia
de Bri´
ofitas – BrioFLORA, Campus Universita
´rio de Ondina, 40170-280 Salvador, Bahia, Brasil
ABSTRACT.In the past decade, floristic studies have rebounded as checklists are fundamental for executing
meta-analyses which address ecological, biogeographic and evolutionary questions of broad geographic
scope. Despite the importance of checklists as baseline records of local diversity and distributions, few
attempts have been made to quantify sampling effort and species detectability within and among study
sites. Quantitative floristics, which combines the use of checklists with statistical methods for estimating
local richness, is a promising method for characterizing the completeness of checklists especially for
cryptic components of biodiversity. For bryophytes, quantifying levels of detectability among substrate
types is of central importance, especially in tropical forests where much of their diversity is harbored in
difficult to access habitats such as the tree canopy. In light of the need to establish quantifiable protocols
of detectability in poorly studied tropical regions, we present a bryophyte checklist for the Jau
´National
Park (JNP), located in the heart of the Amazon, and estimate local species richness and detectability as it
relates to five substrate types (epiphytes, epiphylls, epixylic, epipetric and soil). Identifications from 712
collections made during four excursions over the past decade to JNP revealed 150 species consisting of two
new country records and five new state records, along with 20 rarely collected Amazonian endemics.
Despite our intensive sampling, which included systematic canopy collections during one of the
excursions, Chao richness index estimated that ca. 46 species (nearly one-third of those presently
observed) remain undetected from JNP. Furthermore, levels of detectability among substrates varied
widely, where observed epiphyte richness, in contrast to the other substrates types, most closely
approximated the estimates. Our results illustrate the need for quantitative richness estimates as a means
to increase the accuracy of checklist data, particularly when used in meta-analyses addressing global-scale
questions.
KEYWORDS.Amazon, epiphyte, epiphyll, Lejeuneaceae, quantitative floristics, non-parametric richness
estimators, Rio Negro, tropical forest.
^^^
The Amazon is the largest tropical ecosystem on
Earth, and its biodiversity remains relatively unex-
plored across large areas. Despite its vast expanse,
the Amazon is widely threatened by anthropogenic
pressures contributing to continued loss of its
biodiversity and native habitats (Laurance et al.
2001; Mahli et al. 2008). The past two decades have
brought about a surge in the documentation of
Brazil’s bryoflora (Flora do Brasil: Costa & Peralta
2015; Gradstein & Costa 2003; Yano 2006a,b);
however, most of this recent floristic work has been
concentrated in extra-Amazonian biomes. To date,
Richard Spruce’s Hepaticae Amazonicae et Andinae
(Spruce 1884–1885) and Churchill’s (1998) checklist
of Amazonian mosses remain the most comprehen-
5
Corresponding author’s e-mail: chaszartman@gmail.com
DOI: 10.1639/0007-2745-121.4.571
The Bryologist 121(4), pp. 571–588 Published online: December 10 2018 0007-2745/18/$1.95/0
Copyright Ó2018 by The American Bryological and Lichenological Society, Inc.
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sive resources on hepatics and mosses, respectively,
of the Amazon Basin.
Quantitative studies of lowland Amazonian
epiphytic bryophytes (Campos et al. 2015; Mota &
ter Steege 2013) have revealed that many recently
described taxa assumed to be narrow endemics (e.g.,
Dauphin 2003; Ilkiu-Borges & Gradstein 2008;
Zartman & Ackerman 2002) are common and
widespread at the continental scale, suggesting that
bryophytes, particularly those of the forest canopy
and rheophytes remain systemically under collected
throughout their Amazonian ranges (see also
Gradstein 1994; Gradstein & Costa 2003). In support
of this hypothesis as well are results from bryo-
floristic studies compiled from both historical (see
Costa et al. 2017) and recent (Costa 2017; Costa
2000; Sierra et al. in press) expeditions which
repeatedly report large range expansions for numer-
ous taxa. Furthermore, a recent acceleration in the
rates of species descriptions from the Amazon
provides additional evidence that taxonomic knowl-
edge of Amazonian bryophytes is far from complete
(Bastos & Zartman 2017; Bastos et al. 2016; Bastos &
Zartman 2016; Brito & Ilkiu-Borges 2012a; Gradstein
& Costa 2018; Gradstein & Ilkiu-Borges 2018; Ilkiu-
Borges & Gradstein 2008; Ilkiu-Borges 2011; Moura
et al. 2012; P ´
ocs 2002; Zartman & Ackerman 2002).
Checklists are an important source of taxonom-
ic, systematic and distributional data that may be
used to address questions rooted in ecological and
biogeographical frameworks (S¨
oderstr¨
om et al.
2008). Bryophyte-based meta-analyses have taken
advantage of these valuable sources of information
to tackle global-scale topics such as, for example, the
presence of latitudinal diversity gradients (Wang et
al. 2016), spatial patterns in beta diversity (Norhaz-
rina et al. 2016), range sizes and reproductive life
history traits (Laenen et al. 2015), and climate
change impacts on distributions (Pati˜
no et al. 2016).
However, particularly pertinent for tropical bryo-
phytes, such as epiphytes and epiphylls, character-
ized by reduced gametophyte sizes of mixed colonies
and inaccessible habitats, is the question of detect-
ability. That is, how can sampling completeness be
quantified among checklists to account for variation
in detectability? This question is of exceptional
concern when dealing with survey projects in
conservation units, such as national parks and
biological reserves, as they may serve as baseline
sites for distributional and demographic studies
since such areas are interminably protected from
local anthropogenic disturbance.
The Rio Negro is the largest tributary of the
Amazon Basin, and its upper reaches along the
Colombian and Venezuelan borders harbor numer-
ous bryophyte taxa known only from either the
holotype or few other historical collections (Costa et
al. 2017; P ´
ocs 2002; Reese 1993; Schuster 1999, 1991;
Spruce 1884–1885). Indeed, major floristic gaps have
been documented for flowering plants (Cardoso et
al. 2017; Hopkins 2007) and hepatics alike (Grad-
stein & Costa 2003) throughout the Amazon Basin.
To our knowledge, no bryophyte checklist exists
from any National Park or Biological Reserve within
the 5,500,000 km
2
of the Amazon forest. Jau
´
National Park (JNP), of the middle Rio Negro,
covers ca. 2.4 million hectares of lowland Amazonian
rainforest (Fig. 1), is the second largest national park
in Brazil, and the third largest unit of protected
tropical terrestrial ecosystem on Earth. More im-
portantly, it is the only federal conservation unit in
the Amazon Basin that encompasses an entire
watershed of a major tributary. Located 220 km
northwest of Manaus, JNP harbors dense (77%),
open (14%), and transitional (7%) rainforest as well
as white-sands campinarana (2%) forests (FVA
1998).
In light of the importance of quantifying
detectability and estimating species richness of less
conspicuous components of Amazonian biodiversi-
ty, we present a bryophyte checklist for JNP resulting
from multiple collecting expeditions made by several
of the authors over the past decade. Our principal
objectives with the present data are to: 1) present a
list of the bryophyte species from JNP with their
updated geographical distributions; 2) employ non-
Figure 1. Map of Jau
´National Park in Amazonas state, Brazil.
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parametric quantitative methods to estimate local
species richness among Amazonian substrate types;
and 3) quantify variability in species detectability as
it relates to substrate type. A secondary goal of this
study is to establish a protocol for bryophyte
documentation in tropical regions in the hopes that
tropical bryologists may standardize quantitative
floristic surveys, thus allowing for more statistically
confident inter-site comparisons.
MATERIALS AND METHODS
Sampling and checklist.Specimens were col-
lected in October of 2007, February of 2008,
November of 2016 and September of 2017 from
the lower Jau
´and lower to middle Carabinani rivers
(Table 1) resulting in a total of 712 specimens.
Collections were made in terra firme (dense rain-
forest), campinarana (high white-sand forest), cam-
pina (low white-sand forest), and igapˆ
o(seasonally
inundated forest) (Fig. 2). During the September
2017 excursion, collections were made from five
height zones of full size canopy trees in igapˆ
oand
terra firme forests. The tree canopy bryoflora was
collected using traditional Amazonian methods,
which entails the use of a belt (peconha) cinched
tightly around the ankles to allow for the climber to
shimmy up the trunk of mid-sized trees. From the
vantage point of the canopy, collecting pruners with
aluminium pole extenders are employed to cut outer
branches for study once fallen to the ground. From
the inner canopy, collections are made with
machetes and knives by removal of live bark upon
which the epiphyte resides. In the other three
excursions, collections were haphazardly made at
various locations within the study region as cited
above. All specimens are deposited at one of several
herbaria (INPA,LG,PC,UFBA) as indicated for each
sample. Nomenclature follows the Liverwort World
Checklist (S¨
oderstr¨
om et al. 2015) for liverworts,
with updates, and Flora do Brasil for mosses (Costa
& Peralta 2015).
Species richness estimation.To estimate the
total bryophyte species richness for JNP based on the
current sampling, species accumulation curves (100
permutations) were generated from sample based
rarefactions using the Coleman method (Coleman et
al. 1982). Species accumulation curves were also
generated using the same method for five substrate
types (epiphytic, epiphyllic, epixylic, epipetric and
soil) in order to estimate substrate specific differ-
ences in species detectability.
Using the entire data set, species richness was
estimated by major taxonomic groups (liverworts
and mosses) as well as for substrate type by
extrapolating the number of undetected species
based on the current species pool using the non-
parametric estimating method Chao Index with
standard error (Chao 1987). Since the Chao
estimation did not stabilize with the present
sampling number for some substrates (e.g., epixylic,
epipetric and soil), we applied first-order Jackknife
and Bootstrap estimators, which may reduce bias of
small sample sizes (Gotelli & Colwell 2001), with
standard error estimated following Smith and van
Belle (1984). All analyses were carried out using R
software (version 3.2.5) (2017). The species accu-
mulation curves and species estimation analysis were
done using the vegan package (Oksanen et al. 2016).
RESULTS AND DISCUSSION
Bryophyte checklist of the Jau
´National Park.
The bryophyte flora of the Jau
´National Park is
represented by 19 families, 58 genera, and 150
species, (109 liverworts and 41 mosses), seven of
which we were not able to identify to species. The
most diverse families were Lejeuneaceae (81 spp.)
and Lepidoziaceae (11 spp.) for liverworts, and
Calymperaceae (16 spp.) and Sematophyllaceae (9
spp.) for mosses, in total accounting for 78% of the
JNP flora. The richest genera Cheilolejeunea 15 spp.,
Cololejeunea 12 spp. and Ceratolejeunea 6 spp. for
Table 1. Description of localities sites in Jau
´National Park.
Locality Latitude Longitude Forest type
Mouth of the Jau
´River 0185401200 S6182504400 W Campinarana/seasonal flooded forest (igapˆ
o)
Jau
´River, Itaubal trail 0185203900 S6183501600 W Dense rainforest/ campinarana
ICMBIO Base, Seringalzinho trail 0185003000 S6183703500 W Secondary rainforest
Carabinani River 0280203800 S6183302800 W Rocky vegetation/dense rainforest
Carabinani River, upstream of Preto creek 0280300700 S61833034 00 W Dense rainforest
Sierra et al.: Bryophytes of Jau
´National Park, Brazil 573
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liverworts, and Syrrhopodon 12 spp. for mosses
account for 30% of the bryophyte richness.
Two species of epiphyllous liverworts are new
country records for Brazil (Cololejeunea moralesiae and
C. cornutissima). Five species are new state records for
Amazonas (Cheilolejeunea ornata, C. polyantha, C.
intertexta, Cololejeunea panamensis, Fissidens steerei).
The most common liverworts are Cheilolejeunea
aneogyna,Pycnolejeunea contigua,Cheilolejeunea ne-
blinensis,Cololejeunea surinamensis,Archilejeunea fus-
cescens,Acrolejeunea torulosa,Vitalianthus aphanellus,
and the mosses Leucobryum martianum and Syrrho-
podon xanthophyllus. Various species common in JNP
were previously considered rare with a restricted
distribution (Cheilolejeunea neblinensis, C. assurgens,
C. asperiflora and Vitalianthus aphanellus)astheywere
previously recorded from only a few localities (Bastos
2017; Gradstein & Costa 2003).
Diversity and species richness estimation.A
total of 150 species from 1,152 records in 712
samples were collected from the five substrate types
as follows: 104 spp. as epiphytes (trees, shrubs,
branches or twigs), 38 spp. as epiphylls (leaf
surfaces), 31 spp. epixylic (decomposing logs), 16
spp. epipetric (rocks) and 24 spp. on soil (white
sand, or dirt). Considering the entire dataset, 68 spp.
(45.3%) are singletons (sensu Chao 1987), either
recorded once (44 spp.) or twice (24 spp.). Among
substrates our data include a high percentage of
singletons, 53.8% of epiphytes (56 spp.), 47.4% of
epiphylls (18 spp.), 71.0% of epixylic (22 spp.),
68.7% of epipetric (11 spp.), and 62.5% on soil (15
spp.). A singleton within a substrate might occur
more frequently on another substrate as a generalist.
Rarefaction curves show that species richness is close
to saturation when the complete data set is
Figure 2. Forest types in Jau
´National Park. A. Water-fall on dense forest in Itaubal trail. B. Seasonal flooded forest (igap ˆ
o). C. Rocky vegetation along
Carabinani River. D. Campinarana rocky forest along Carabinani River.
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considered (Fig. 3A); however, when subdivided by
substrate only the epiphyte curve approximated
saturation (Fig. 4A compared to Fig. 4B–E).
All three quantitative estimators, Chao index,
Jackknife and Bootstrap, showed that the bryophyte
flora of the JNP is not completely represented, as
many species remain undetected across all substrates
(Table 2). Specifically, estimates suggest that the
local species richness of the region sampled in JNP is
ca. 196 species (Fig. 3B), corresponding to 134 spp.
of liverworts and 73 spp. of mosses (Table 2),
suggesting that nearly one-third (~46 spp.) remain
undetected for the JNP.
Even though the accumulation curve showed
that the current species richness recorded was close
to saturation (Fig. 3A), around ~20–40 species
remained undetected based on two of the quantita-
tive estimators (Fig. 3B). The same estimation was
observed for the richest substrate studied (epiphyte)
with estimates of ~40 species more than observed
(Fig. 5A). In the other substrates, (epiphyll, epixylic,
epipetric and soil), the species accumulation curves
were far from reaching the asymptote, but the
species richness estimations were inconsistent,
pointing to one third to a half of the species more
than observed (Fig. 5B–E). As illustrated by the
disparity among observed and estimated richness for
Figure 3. A. Species accumulation curves (Coleman method) for the complete dataset in the JNP. Gray shades represent the intervals of confidence. B.
Number of species observed compared to non-parametric species richness estimator (Chao Index, Jackknife, Bootstrap) for the complete dataset in the
JNP.
Table 2. Species estimation for the complete dataset, by substrate, and by taxonomic group (mosses and liverworts), with the following non-parametric
methods: Chao Index, First order Jackknife and Bootstrap.
Total Epiphyte Epiphyll Epixylic Epipetric Soil Mosses Liverworts
Species pool 151 104 38 31 16 24 42 110
Chao Index 196.96 141.94 101.38 59.54 64.21 65.22 73.84 134.95
Standard error 18.16 16.54 55.46 19.57 57.47 37.60 23.21 11.94
First order Jackknife 195.94 140.92 53.84 47.79 25.64 36.68 57.92 139.94
Standard error 7.13 6.99 4.20 4.31 3.05 3.52 3.98 5.99
Bootstrap 171.48 120.87 44.35 37.95 19.92 29.21 48.61 124.21
Standard error 3.90 3.77 2.11 2.08 1.38 1.71 2.02 3.38
Sample number (n)712 449 103 80 28 41 200 550
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´National Park, Brazil 575
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the JNP dataset, species richness estimates generated
from datasets well represented by singletons are
predicted to have proportionally more undetected
species than those datasets dominated by repeats
(Gotelli & Cowell 2001).
Epiphylls are rich communities, with species
usually diffusely associated with local habitat or
microclimate (Sonnleitner et al. 2009)—a feature
that decreases their detectability. Continued sam-
pling of different habitats would most likely allow us
to reach the estimates of ~101 epiphylls expected for
JNP. Indeed, the only comprehensive epiphyll survey
conducted at a much smaller scale in the Amazon
forest accounted for ~67 species (Zartman & Ilkiu-
Borges 2007).
Species estimation of epixylic, epipetric and soil
bryophytes tend to surpass more than half of the
observed species richness with Chao Index. On the
Figure 4. Species accumulation curves (Coleman method) by substrate in the JNP. A. Epiphyte. B. Epiphyll. C. Epixylic. D. Epipetric. E. Soil. Gray shades
represent the intervals of confidence.
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contrary, Jackknife and Bootstrap estimated that
~10 species remain undetected. These incongruent
estimates could be a bias due to the unevenness of
the rank abundance distribution, the sampling
intensity or the performance of the estimators.
Moreover, the patchy characteristic and lower
availability of suitable substrates (S¨
oderst¨
orm &
During 2005), may be another reason for the
apparently low detectability of epixylic, epipetric
and soil bryophytes.
When comparing the three quantitative estima-
tors, the Chao index presented the highest estima-
Figure 5. Number of species observed compared to non-parametric species richness estimator (Chao Index, Jackknife, Bootstrap) by substrate in the JNP.
A. Epiphyte. B. Epiphyll. C. Epixylic. D. Epipetric. E. Soil.
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tion and standard error for substrates where the rank
abundance was represented more than half by
singletons. Chao Index uses the number of singletons
(those 2 registers) in the dataset to estimate species
richness. Once species have been recorded more than
twice, the Chao Index assumes that all species have
been detected. Likewise, Jackknife considers the
number of singletons in its richness estimation
algorithm; however, it utilizes a narrow optimum
sample size range making it likely to underestimate
true species richness in small sample sizes and
overestimate it in large ones (Gotelli & Cowell 2001).
According to Brose et al. (2003), the performance of
species estimators depends on the presence of
singletons in the data (evenness of the rank
abundance distribution), the sampling intensity,
and the true species richness of the study system.
A floristic survey of the Upper Kabalebo River
in West Suriname (Florsch¨
utz-deWaard & Bekker
1987), an area comparable in size, altitude and
habitat types to JNP, shows similar results in species
richness. In their inventory, however, the canopy
was not completely explored and epiphylls were
also not included. Following an update in the
nomenclature of the bryophytes of Kabalebo River
and considering the epiphyllous species, it is most
likelythatspeciesrichnessinthisareawillbeshown
to be higher than that of our estimates for JNP. This
putatively higher species number in Kabalebo River
couldbeduetotheinfluenceofitsproximitytothe
Caribbean coast and influence from maritime
winds and higher annual precipitation than com-
pared to JNP.
Biogeography of the Jau
´National Park: Rare
and endemic species.The Jau
´National Park is
represented by 85 species with Neotropical distribu-
tions, 16 spp. pantropical, 14 spp. Afro-American, 28
spp. Amazonian endemics, and seven unidentified
species. Two genera registered in the JNP, Verdoor-
nianthus (2 spp.) and Schusterolejeunea (1 sp.), are
Amazonian endemics (Gradstein & Costa 2003). The
upper reaches of the Amazon (particularly in the Rio
Negro Basin) are considered a center of endemism
for certain genera of Lejeuneaceae such as Cheilole-
jeunea, Cololejeunea and Ceratolejeunea, which also
correspond to the most diverse genera in the JNP
(Costa et al. 2017; Gradstein & Costa 2003).
Moss richness is relatively low with only four
families representing more than 50% of the total
number of moss species in the Amazon forest
(Churchill 1998). This is also reflected in JNP where
two families represent 61% of all collected moss
species. The Calymperaceae are a major component
of the epiphytic flora in lowland Amazonia, and in
the JNP are highly diverse accounting for the
majority of moss species (16 spp.) with various
endemic and rare species, such as Calymperes
mitrafugax, Syrrhopodon annotinus, S. fimbriatus, S.
helicophyllus and S. xanthophyllus (Reese 1993).
Twenty nine species collected during this study
were considered rare due to their current worldwide
distributions; 20 are restricted to the Amazon Basin.
Four of these taxa are endemic to the Brazilian
Amazon (Cheilolejeunea asperiflora, C. papulosa, C.
polystachya and Octoblepharum leucobryoides). Of the
29 rare species, eight are scattered throughout the
Neotropics, representing disjunct distributions: Cen-
tral America-Amazon (Cololejeunea moralesiae, C.
panamensis, C. schusteri, Syrrhopodon flexifolius),
Caribbean-Amazon (Cololejeunea spruceana, C. cor-
nutissima), Amazon-Atlantic forest (Vitalianthus
aphanellus, Syrrhopodon cymbifolius) and Andes-
Amazon-Atlantic forest (Cheilolejeunea ornata).
CONCLUSIONS
Here we present a checklist of bryophytes and
quantify variability in substrate detectability from
one of the largest federally protected reserves in the
Amazon Basin. Despite our intensive long-term
sampling approach, non-parametric richness indices
estimate that more than one-third of the total
observed species remain undetected. Coincidentally,
the area sampled for this study (lower and middle
Carabinani and Jau
´Rivers) is roughly one-third of
the total area of the JNP, provoking the question as
to whether more widely scattered sampling points
would have resulted in greater detectability. None-
theless, bryophytes are inconspicuous plants, partic-
ularly in tropical forests with complex canopy
architecture, thereby necessitating intensive surveys
to ensure checklist accuracy.
Evaluating gaps in floristics surveys by using
non-parametric estimators is an important tool for
quantifying patterns of diversity and distribution.
Clearly sampling error, that is species left undetected
in a given floristic survey, is inevitable. However, the
variation in sampling error among sites, whether due
to differences in sampling intensity or the unique
collecting challenges inherent to every biome, is
quantifiable. Such information would offer an
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understanding as to how detectability levels vary
among checklist sites which would ultimately
contribute to reducing the ‘‘ noise’’ inherent in
meta-analysis based on checklist data.
Nonetheless, checklists remain the baseline
source of information for addressing large-scale
ecological and evolutionary questions. The principal
assumption, however, of meta-analyses conducted
with checklist data is that there is little to no inter-
site variation in species detectability. Here we suggest
that the degree of species detectability of a particular
survey is not only based on sampling intensity, but
rather based on the combination of sampling
intensity, approach and inherent collecting challeng-
es, all of which can be jointly accounted for through
a quantitative floristic approach.
CHECKLIST
Liverwort and moss taxa are classified into
families and listed in alphabetical order by genera.
For each species we include voucher numbers and
information on habitat conditions. When pertinent,
notes are included for species about currently known
geographical distributions as well as novel morpho-
logical features observed. New country records for
Brazil are marked with (**) and new state records for
Amazonas are marked with (*).
MARCHANTIOPHYTA (Liverworts)
Aneuraceae
Riccardia regnellii (A
˚ngstr.) G.K.Hell (¼R. amazon-
ica (Spruce) Gradst. & Hekking), Vanderpoort-
en 221.5 (LG); Zartman 7509, 7487, 7164 (INPA),
flat flooded rocks and logs by creek in white
sand forest, with Symphyogyna brasiliensis,
Pictolejeunea sprucei, and Zoopsidella integrifo-
lia. Note: As shown by Gradstein & Reeb
(2018), the name Riccardia regnellii has been
misapplied in the past and is an older name for
the common and widespread R. amazonica.
Calypogeiaceae
Calypogeia tenax (Spruce) Steph., Sierra 4279 (INPA),
on rocks by creek in white sand forest.
Mnioloma nephrostipum (Spruce) R.M.Schust., Zart-
man 7525,10162,10181,10148 (INPA). Note:
rare species of Amazonia, Guyana Highland and
the Choc ´
o of Colombia.
Mnioloma parallelogrammum (Spruce) R.M.Schust.,
Sierra 4328 (INPA), epiphyte on tree base.
Cephaloziaceae
Odontoschisma variabile (Lindenb. & Gottsche)
Trevis., Vanderpoorten 216.3, 216.5 (LG); Zart-
man 10182,10159 (INPA), on bark of living trees
along creek and logs in white sand forest, with
Drepanolejeunea palmifolia, Micropterygium tra-
chyphyllum, Leucobryum martianum, Cheilole-
jeunea aneogyna.
Frullaniaceae
Frullania caulisequa Gottsche, Zartman 10145 (INPA),
on branches of tree in white sand forest, with
Cheilolejeunea neblinensis, Pycnolejeunea conti-
gua.
Frullania gibbosa Nees, Zartman 10051, 9979, 10014,
10164 (INPA), epiphyte on open burned areas or
in tree trunk in Igap ˆ
o forest, with Acrolejeunea
torulosa, Cheilolejeunea rigidula, Cheilolejeunea
trifaria, Pycnolejeunea macroloba, Lopholejeunea
subfusca.
Frullania kunzei (Lehm. & Lindenb.) Lehm. &
Lindenb., Zartman 7473, 7478 (INPA). Note:
Fertile plants with dentate bracts.
Frullania nodulosa (Reinw. et al.) Nees, Zartman
10013, 10098, 10087 (INPA), epiphyte on open
burned areas or in tree trunk canopy in igapˆ
o
forest, with Acrolejeunea torulosa.
Frullania (subg. Diastaloba)sp.,Zartman 7248
(INPA).
Lejeuneaceae
Acrolejeunea torulosa (Lehm. & Lindenb.) Schiffn.,
Vanderpoorten 171.1, 174.1 (LG); Sierra 4283,
4290; Zartman 7206, 7290, 9973, 9977, 9980,
9985, 9989, 9991, 9993, 9998, 10001, 10013,
10014, 10015, 10016, 10021, 10051, 10044,
10047, 10103, 10104, 10093, 10164, 10143,
10160, 10105, 10107, 10108, 10110, 10115 (INPA),
common epiphyte on bark in open area or
canopy in white sand forest.
Archilejeunea badia (Spruce) Steph., Vanderpoorten
178.5, 173.3 (LG); Sierra 4310, 4278, 4377;
Zartman 7184, 7537, 9999, 10074 (INPA), on
bark in white sand forest with Bazzania pallid-
evirens, Calymperes lonchophyllum, C. aneogyna,
C. neblinensis, C. cubensis.
Archilejeunea crispistipula (Spruce) Steph., Sierra
4309, 4303, 4334, 4270; Zartman 7496,10060,
10142 (INPA), on bark in white sand forest.
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Archilejeunea fuscescens (Lehm.) Fulford, Vander-
poorten 220.4, 227.6, 187.1, 193.2, 192.1, 205.5,
209.1, 201.2, 191.1, 186.1, 172.5, 185.6, 179.2,
210.1, 188.4, 182.6, 222.1, 194.1 (PC); Sierra
4311, 4320, 4305, 4338, 4341; Zartman 7446,
7180, 10015, 10100, 10107, 10131, 10044, 10046,
10047, 10102, 10058, 10092, 10070, 10128 (INPA),
one of the commonest epiphyte on bark in both
terra firme and white sand forest. Note: Plants
with regenerants on dorsal leaf surfaces.
Archilejeunea ludoviciana (Lehm.) P.Geissler &
Gradst. subsp. porelloides (Spruce) Gradst. &
P.Geissler, Sierra 4312, 4298, 4269, 4335, 4292,
4344; Zartman 7176, 7252, 7499, 7539, 7462,
10026, 10167, 10140 (INPA), robust epiphyte
usually on twigs with projecting growth.
Caudalejeunea lehmanniana (Gottsche) A.Evans,
Zartman 7437, 10164 (INPA), epiphyte on twigs
or epiphyll.
Ceratolejeunea coarina (Gottsche) Steph., Vander-
poorten 190.4 (LG); Sierra 4378, 4305, 4341,
4349, 4364; Zartman 7169, 10136 (INPA), dead
logs in white sand forest with C. cubensis,
Octoblepharum pulvinatum, Syrrhopodon sim-
mondsii, S. hornschuchii, S. cryptocarpos, epi-
phyte on liana and epiphyllic.
Ceratolejeunea confusa R.M.Schust., Zartman 7468,
7479 (INPA), on bark.
Ceratolejeunea cornuta (Lindenb.) Schiffn., Sierra
4274, 4280, 4290, 4340, 4366, 4367, 4368, 4372,
4373, 4387; Zartman 7262, 7194, 7193, 7244,
7188, 7505.1, 7506.1, 7500, 7497, 9989, 10058,
10093, 10178, 10140, 10115, 7521 (INPA).
Ceratolejeunea cubensis (Mont.) Schiffn., Vander-
poorten 190.4a (LG); Sierra 4310, 4298, 4335,
4270;Zartman 7516, 7532, 7469, 7260, 7294,
7497.1, 7182, 10011, 10053, 10090, 10065, 10069,
10142, 10114, 10117 (INPA), on dead logs and on
bark in white sand forest.
Ceratolejeunea minuta G.Dauphin, Zartman 7246
(INPA).
*Ceratolejeunea rubiginosa Steph., Zartman 10109
(INPA), epiphyte on twigs in white sand forest.
Cheilolejeunea acutangula (Nees) Grolle, Zartman
10120 (INPA;UFBA), epiphyte.
Cheilolejeunea adnata (Lehm.) Grolle, Zartman 7309,
7287 (INPA), epiphyte on bark.
Cheilolejeunea aneogyna (Spruce) A.Evans, Vander-
poorten 216.8 (LG); Pereira 766;Sierra 4314,
4310, 4275, 4308, 4276, 4257, 4306, 4277, 4301,
4300, 4268, 4285, 4258, 4330, 4286, 4288, 4290,
4259, 4291, 4266, 4359; Zartman 7297, 7160,
7286, 7284, 7507.1, 7471, 7472, 7473, 7476.1,
7477, 7465.1, 7439, 7305, 7311 (INPA), one of the
commonest liverworts growing on bark, leaves
and dead logs in white sand forest with
Odontoschisma variabile, Drepanolejeunea pal-
mifolia, Micropterygium trachyphyllum, Leuco-
bryum martianum.
Cheilolejeunea asperiflora (Spruce) Gradst. & Ilk.-
Borg., Zartman 7487, 7465, 7474.1, 7526, 10035,
10162, 10183, 10144, 10149, 10152, 10154,
10155, 10158 (INPA). Note: This rare Amazonian
species is known from Spruce’s type locality in
Venezuela, along R´
ıo Negro near San Carlos
and more recently in Brazil from the Serra
Araca
´(CEZ pers. obs.). The ecology of this
species from field observations entails that it
usually grow on log or soil in white sand forest.
Cheilolejeunea assurgens (Spruce) Steph., Sierra 4271,
4333, 4295;Zartman 7315, 7294, 7296, 7171,
7186, 7182, 7289, 7493.1, 7490, 7276, 9982, 9983,
9987, 10023, 10024, 10042, 10045, 10101, 10073,
10076, 10168, 10142, 10109, 10116, 10131 (INPA),
common epiphyte same as C. aneogyna. Note:
This species was considered a synonym of C.
aneogyna (Bastos 2012), but is currently ac-
cepted as a distinct species (Bastos 2017). It
differs from C. aneogyna by the distinct trigones
and intermediate thickenings and the Lejeunea-
type subgynoecial branch (true, Radula-type
innovation absent).
*Cheilolejeunea intertexta (Lindenb.) Steph., Zart-
man 7441, 10140 (INPA), epiphyte.
Cheilolejeunea neblinensis Ilkiu-Borges & Gradst.,
Vanderpoorten 220.4a (LG); Sierra 4272, 4317,
4307, 4278, 4284, 4299, 4336, 4339, 4343, 4327;
Zartman 7316, 7192, 7299, 7306, 7302, 7300,
7160, 7161, 7188, 7187, 7185, 7251, 7508.1, 7470,
7477, 7496.1, 7503.1, 7468.1, 7266, 7159, 10019,
10091, 10070, 10176, 10179, 10180, 10142,
10145, 10146, 10149, 10152, 10154, 10155,
10157, 10158 (INPA), one of the commonest
liverworts growing on bark, leaves and flat
flooded rocks by creek in white sand forest,
with Micropterygium trachyphyllum, Leuco-
bryum martianum, Xylolejeunea crenata, Archi-
lejeunea fuscescens. Note: Plants from JNP vary
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considerably in size, lobule length and shape,
and presence of low lens shaped papilla on leaf
cells. Fertile plants differ from typical C.
neblinensis (Ilkiu-Borges & Gradstein 2008) by
being autoicous, with small trigones and
gynoecia on very short branches lacking inno-
vations. Although Costa et al. (2017) claimed
this species was only known from its type
locality and Serra Bela Adormicida of the upper
Rio Negro, it is actually widely distributed
throughout Amazonia (Mota & ter Steege 2013;
Campos et al. 2014; Bastos 2017), and is a
common epiphyte in JNP.
Cheilolejeunea oncophylla (A
˚ngstr.) Grolle &
E.Reiner, Zartman 10119, 10126 (INPA), epi-
phyte.
*Cheilolejeunea ornata C.Bastos, Zartman 10011
(INPA), epiphyte.
Cheilolejeunea papulosa Schiffn., Zartman 7197,
7178, 10092, 10049, 10033;Sierra 4327 (INPA,
UFBA), rare epiphyte. Note: Also found in
collections from Serra do Araca
´(Zartman
9622,INPA) and Roraima (G.T. Prance et al.
19993,INPA). The species seems restricted to the
Brazilian Amazon region, since there are no
references to other localities.
*Cheilolejeunea polyantha A.Evans, Zartman 9999,
10049 (INPA), epiphyte.
Cheilolejeunea polystachya (Spruce) Gradst. & Ilk.-
Borg., Zartman 7469.1, 10122 (INPA), rheophyte
on sand near river. Note: This species is a
rheophyte growing on rocks in fast flowing
streams and rivers of the Rio Negro basin in the
Amazon. Also, found in collections from Sa
˜o
Gabriel, along Rio Cariua from Rio Curicuriari
to Cachoeira Piraiauara (Schuster 79-16-831,F).
Endemic from Brazil. The species seems re-
stricted to the Amazon region, since there are
no references to other localities.
Cheilolejeunea rigidula (Mont.) R.M.Schust., Van-
derpoorten 186.4a (LG); Sierra 4267; Zartman
7312, 7174.1, 7208, 7486, 7209, 7474, 9979,
10011, 10014, 10020, 10021, 10027, 10051,
10036, 10044, 10094, 10098, 10055, 10090,
10064, 10164, 10115, 10117, 10129 (INPA), on
bark and dead logs in white sand forest with
Leucobryum martianum, Octoblepharum pulvi-
natum, Syrrhopodon cryptocarpos.
Cheilolejeunea trifaria (Reinw. et al.) Mizut., Sierra
4260, 4287, 4263; Zartman 7204, 7203, 7481.1,
10011, 10051, 10095, 10090, 10065, 10069,
10079, 10141, 10153, 10117 (INPA), occurring
on bark in exposed area or logs in igapˆ
o forest.
Cheilolejeunea urubuensis (Zartman & I.L.Ackerman)
R.L.Zhu & Y.M.Wei, Zartman 10031, 10091
(INPA), occurs as epiphyte on canopy branches.
Cololejeunea camillii (Lehm.) A.Evans, Sierra 4374,
4351; Zartman 7503, 10111, 10113, 10114, 10134
(INPA), epiphyll.
Cololejeunea cardiocarpa (Mont.) A.Evans, Zartman
7279, 10030, 10172, 10111, 10113 (INPA),
epiphyll.
**Cololejeunea cornutissima (R.M.Schust.) Stotl. &
Crand.-Stotl., Zartman 7490 (PC;INPA), epiphyll.
Note: This species is extinct at its type locality
in Florida and is a rare species in the Neotropics
known only from Cuba, the Dominican Re-
public and the Peruvian Amazon (T. P ´
ocs
pers.comm.).
Cololejeunea diaphana A.Evans, Sierra 4329, 4345,
4346, 4349, 4350, 4353, 4355, 4356, 4358, 4359,
4360, 4361, 4362, 4363, 4364, 4365, 4380, 4384,
4387; Zartman 7267.1, 7261, 7244, 7493 (INPA),
one of the commonest epiphyllic bryophyte.
Cololejeunea gracilis (Jov.-Ast) P ´
ocs, Sierra 4345,
4357, 4360, 4364, 4365; Zartman 10113, 10114,
10134 (INPA), epiphyll.
**Cololejeunea moralesiae (Bernecker) Bernecker &
P´
ocs, Zartman 7542 (INPA), epiphyll. Note: A
rare species known only from Costa Rica. This
represents the first record for Brazil and South
America.
Cololejeunea obliqua (Nees & Mont.) Schiffn, Sierra
4357; Zartman 7506, 7267, 7262, 7261, 7267.1,
7478 (INPA), epiphyll.
*Cololejeunea panamensis G.Dauphin & P ´
ocs, Zart-
man 10054, 10055, 10074 (INPA), epiphyte on
bark. Note: A rare species known from its type
locality in lowland forest in Panama (Dauphin
et al. 2006) and for Brazilian Amazon in the
state of Para
´(Brito & Ilkiu-Borges 2012b)
Cololejeunea schusteri P´
ocs, Zartman 7509 (INPA),
epiphyll. Note: A rare species, known only from
the type locality (P ´
ocs 2002) and one collection
from lowland forest in Panama (Dauphin et al.
2006).
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Cololejeunea spruceana Tixier, Sierra 4359 (PC;INPA),
epiphyll. Note: A rare species collected from a
few localities in the Brazilian and Venezuelan
Amazon (Dauphin et al. 2008; Gradstein &
Costa 2003), and from Trinidad (P ´
ocs et al.
2014).
Cololejeunea surinamensis Tixier, Sierra 4329, 4347,
4352, 4353, 4354, 4356, 4357, 4358, 4362, 4363,
4365, 4375, 4388; Zartman 7489, 7307, 7267,
7264, 7191, 7157, 7245, 7244, 7298, 7170, 7251,
10030, 10172, 10151, 10111, 10113, 10114,
10132, 10134, 10135, 10136 (INPA), one of the
commonest epiphyllic bryophyte.
Cololejeunea winkleri (M.I.Morales & Bernecker)
Bernecker & P ´
ocs, Sierra 4350 (INPA), epiphyll.
Colura greig-smithii Jov.-Ast, Sierra 4329, 4357, 4360,
4370; Zartman 7173, 10113 (INPA), epiphyll.
Colura tortifolia (Nees & Mont.) Trevis., Zartman
7173.1, 7478, 7437 (INPA), epiphyll.
Cyclolejeunea convexistipa (Lehm. & Lindenb.)
A.Evans, Sierra 4371, 4372, 4382; Zartman
7267.1, 7264, 7262, 7254, 7261, 7265, 7246,
7245, 7244, 7253, 7172, 7496, 10114, 10133,
10136 (INPA), one of the commonest epiphyllic
bryophytes.
Cyclolejeunea luteola (Spruce) Grolle, Vanderpoorten
217.2a, P424a (LG); Zartman 7165, 10142,
10125 (INPA), on humus along the creek with
Micropterygium trachyphyllum, Syrrhopodon
simmondsii, Monodactylopsis monodactyla, Oc-
toblepharum pulvinatum, Leucobryum martia-
num.
Cyclolejeunea peruviana (Lehm. & Lindenb.)
A.Evans, Zartman 7264, 7253, 7496 (INPA),
epiphyll.
Diplasiolejeunea brunnea Steph., Sierra 4329, 4346,
4357, 4358, 4364, 4366, 4371, 4375, 4376, 4381,
4384, 4385; Zartman 7264, 7278, 7261, 7247,
7493, 7481, 10172, 10151, 10113 (INPA), one of
the commonest epiphyllic bryophytes.
Drepanolejeunea crucianella (Tayl.) A.Evans, Sierra
4373, 4357; Zartman 7261, 7506 (INPA), epiphyll.
Drepanolejeunea orthophylla (Nees & Mont.) Bischl.,
Zartman 7478 (INPA), epiphyll.
Drepanolejeunea palmifolia (Nees) Steph., Vander-
poorten 216.5b (LG); Sierra 4327, 4336; Zartman
7503, 7502, 7495 (INPA), on soil on base of trees
and on bark of trees along creek in white sand
forest, with Syrrhopodon fimbriatus and Odon-
toschisma variabile.
Drepanolejeunea polyrhiza (Nees) Grolle & R.-
L.Zhu, Sierra 4329, 4348, 4350, 4352, 4361,
4363, 4366, 4367, 4368, 4375, 4378, 4379, 4381,
4385, 4386, 4387; Zartman 7265, 7247, 7250
(INPA), one of the commonest epiphyllic
bryophytes.
Drepanolejeunea (subg. Rhaphidolejeunea) sp., Zart-
man 7510 (INPA),epiphyllic.
Harpalejeunea stricta (Lindenb. & Gottsche) Steph.,
Zartman 7465 (INPA), epiphyte.
Lejeunea boryana Mont., Sierra 4300; Zartman 7310,
7167, 7166, 7283, 7484, 10120 (INPA), on bark in
terra firme forest.
Lejeunea flava (Sw.) Nees, Sierra 4367, 4373, 4378;
Zartman 7262, 7254, 7267 (INPA), epiphyll.
Lejeunea tapajosensis Spruce, Zartman 10021, 10027,
10041 (INPA), epiphyte.
Leptolejeunea jamaicensis Schust., Sierra 4367, 4375,
4376; Zartman 7492, 10172, 10151, 10185 (INPA),
epiphyll.
Leptolejeunea elliptica (Lehm. & Lindenb.) Schiffn.,
Sierra 4329, 4357; Zartman 7278, 7533, 7522,
10043, 10185, 10113 (INPA), epiphyll.
Leptolejeunea moniliata Steph., Zartman 7196, 10151,
10185 (INPA), epiphyll.
Leptolejeunea sp., Zartman 10023, 10026, 10055, 9987
(INPA), epiphyll.
Lopholejeunea nigricans (Lindenb.) Schiffn., Zartman
10161 (INPA), epiphyte.
Lopholejeunea subfusca (Nees) Schiffn., Sierra 4260;
Zartman 7205, 7208, 9984, 9994, 10021, 10027,
10051, 10036, 10041, 10094, 10095, 10096,
10098, 10090, 10074, 10164, 10105, 10108,
10110, 10115 (INPA), epiphyte.
Metalejeunea cucullata (Reinw. et al.) Grolle, Zart-
man 7189, 10000 (INPA), on bark.
Microlejeunea bullata (Tayl.) Steph., Sierra 4267,
4354 (INPA), epiphyll.
Otolejeunea schnellii (Tixier) R.L.Zhu & M.L.So,
Zartman 7259 (INPA), epiphyll on filmy fern
(Hymenophyllaceae). Note: A rare Amazonian
liverwort endemic to Amazonas, Brazil, known
from type collection near Manaus (Tixier 1995),
the Biological Dynamics of Forest Fragments
Project north of Manaus (Zartman & Ilkiu-
Borges 2007), Serra da Bela Adormecida on the
upper Rio Negro (Costa et al. 2017), Choc ´
oof
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Colombia (Benavides & Sastre-De Jesu
´s 2011),
and now reported from JNP along the middle
Rio Negro.
Pictolejeunea sprucei Grolle, Vanderpoorten 221.5b
(LG); Sierra 4265, 4318, 4337; Zartman 7505
(INPA), flat flooded rocks by creek in white sand
forest, with Riccardia regnellii, Symphyogyna
brasiliensis, Zoopsidella integrifolia. Note: This
rare species is endemic to the Amazon.
Pycnolejeunea contigua (Nees) Steph., Sierra 4273,
4290; Zartman 7206, 7290, 7181, cf. 7485, 7483,
7488, 9973, 9974, 9977, 9980, 9985, 9986, 9989,
9991, 9993, 9998, 10001, 10015, 10016, 10022,
10026, 10028, 10031, 10051, 10038, 10044,
10046, 10047, 10100, 10104, 10053, 10055,
10057, 10089, 10093, 10065, 10067, 10070,
10074, 10083, 10085, 10145, 10105, 10107,
10108, 10110, 10128 (INPA), on bark of trees.
Pycnolejeunea macroloba (Nees & Mont.) Schiffn.,
Vanderpoorten 221.8 (LG); Sierra 4316, 4274,
4308, 4332; Zartman 7179, 7195, 7192, 7188,
7255, 10051, 10038, 10070, 10163, 10166, 10176,
10141, 10146, 10147, 10156, 10160, 10117, 10129
(INPA), bark of trees along creek in white sand
forest, with Archilejeunea fuscescens.
Rectolejeunea versifolia (Schiffn.) L. S ¨
oderstr. &
Hagborg, Zartman 7265 (INPA), only collected
as epiphyll.
Rectolejeunea emarginuliflora (Schiffn.) A.Evans,
Zartman 7291 (INPA), occurring as epiphyte
and epiphyll.
Schiffneriolejeunea amazonica Gradst., Zartman 9987,
10094, 10110 (INPA), epiphyte.
Schusterolejeunea inundata (Spruce) Grolle, Sierra
4313; Zartman 7547 (INPA), rheophyte on sand
along Carabinani River. Note: This is a rare
rheophyte belonging to a monotypic genus
endemic to lowland Amazonia. Known only for
localities along the Rio Negro in Brazil and
Venezuela (Costa et al. 2017; Dauphin et al.
2008; Gradstein & Costa 2003).
Symbiezidium barbiflorum (Lindenb. & Gottsche)
A.Evans, Zartman 10098 (INPA).
Symbiezidium transversale (Sw.) Trevis., Zartman
7305, 10051 (INPA), on bark of trees.
Thysananthus amazonicus (Spruce) Schiffn.,Zart-
man 7189, 7538, 10140 (INPA), on tree branch-
es.
Thysananthus auriculatus (Wilson) Sukkharak &
Gradst. (¼Mastigolejeunea auriculata (Wils.)
Schiffn.), Sierra 4262; Zartman 7192, 7203,
10011, 10015, 10021, 10027, 10041, 10095,
10090, 10107, 10108, 10117 (INPA), on bark in
exposed habitat. Note: This species unusually
presents long flagelliform branches at both stem
bases and apices.
Thysananthus innovans (Spruce) Sukkharak &
Gradst. (¼Mastigolejeunea innovans (Spruce)
Steph.), Zartman 10161 (INPA), epiphyte.
Verdoornianthus griffinii Gradst., Zartman 10066
(INPA), canopy epiphyte.
Verdoornianthus masupiifolius (Spruce) Gradst.,
Zartman 10000, 10031 (INPA), canopy epiphyte.
Vitalianthus aphanellus (Spruce) Bechteler et al.,
Sierra 4304, 4300, 4297, 4296, 4359; Zartman
7175, 7497, 9980, 9982, 9983, 9992, 10040,
10052, 10053, 10054, 10057, 10061, 10071,
10073, 10076, 10081, 10082, 10083, 10168,
10118, 10120, 10121, 10123, 10124, 10131 (INPA),
on bark in white sand forest along the
Carabinani River, growing with Pycnolejeunea
species. Note: Originally considered a very rare
species known only from the type locality from
Amazonia, one collection made by Yano &
Zartman (Bechteler et al. 2016), and one from
the Colombian Amazon (Campos et al. 2014).
In JNP it is a common and locally abundant
epiphyte along the Carabinani River.
Xylolejeunea crenata (Nees & Mont.) X.-L.He &
Grolle, Vanderpoorten 220.3 (LG); Sierra 4314,
4342;Zartman 7186, 7501.1, 7466, 10130 (INPA),
on logs and flat flooded rocks by creek in white
sand forest, with Micropterygium trachyphyllum,
Leucobryum martianum, Archilejeunea fusces-
cens, Cheilolejeunea neblinensis.
Lepidoziaceae
Bazzania hookeri (Lindenb.) Trevis., Vanderpoorten
188.1, 200.1, 204.2, 205.3, 215.1, 221.6, P426
(LG), epiphytic in terra firme and white sand
forest with Syrrhopodon cryptocarpos, S. pro-
lifer, Archilejeunea fuscescens,Octoblepharum
pulvinatum, Leucobryum martianum, Micro-
pterygium trachyphyllum, Microcalpe subsim-
plex.
Bazzania longistipula (Lindenb.) Trevis., Zartman
7508 (INPA), epiphyte in white sand forest.
Sierra et al.: Bryophytes of Jau
´National Park, Brazil 583
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Bazzania pallidevirens (Steph.) Fulford, Vanderpoort-
en 214.4, 223.4, 178.3, P431 (LG); Zartman 7181
(INPA), epiphyte in white sand forest with
Micropterygium trachyphyllum, Syrrhopodon
prolifer, Octoblepharum albidum, O. pulvina-
tum, Leucobryum martianum, Calymperes lon-
chophyllum.
Micropterygium leiophyllum Spruce, Sierra 4315,
4264, 4319, 4286, 4331 (INPA), forms large mats
on rock and base of living trees in both terra
firme and white sand forest.
Micropterygium parvistipulum Spruce, Zartman 7162
(INPA), on log.
Micropterygium pterygophyllum (Nees) Trevis., Zart-
man 7500.1, 7470 (INPA), on soil.
Micropterygium trachyphyllum Reimers,Vanderpoort-
en 178.2, 195.2, 199.1, 204.4, 205.3bis, 214.1,
216.6, 217.1, 218.1, 220.1, 227.1, 230.1 (LG); Sierra
4289;Zartman 7163 (INPA), common on dead
logs and bark of living trees in both terra firme
and white sand forest, with Syrrhopodon anno-
tinus, Bazzania hookeri, B. pallidevirens, Calym-
peres lonchophyllum, Archilejeunea badia, A.
fuscescens, Leucobryum martianum, Odontoschis-
ma variabile, Sematophyllum subsimplex.
Monodactylopsis monodactyla (Spruce) R.M.Schust.,
Vanderpoorten 217.2a (LG); Zartman 10170,
10181 (INPA), mixed with Syrrhopodon sim-
mondsii and Cyclolejeunea luteola on humus
along creek in white sand forest.
Pteropsiella metzgeriiformis Steph., Zartman 10173
(INPA), on soil.
Telaranea pecten (Spruce) J.J.Engel & G.L.Merr.,
Zartman 7500.1, 7473 (INPA). Note: This species
is common to the greater Amazon Basin (Brazil,
Colombia, Guyana and French Guiana; Grad-
stein 1997; Campos et al. 2014).
Zoopsidella integrifolia (Spruce) R.M.Schust., Van-
derpoorten 221.5c (LG); Sierra 4314; Zartman
7526, 10173, 10181, 10148 (INPA), flat flooded
rocks by creek in White sand forest with
Riccardia regnellii, Symphyogyna brasiliensis
and Pictolejeunea sprucei.
Pallaviciniaceae
Symphyogyna brasiliensis (Nees) Nees & Mont.,
Vanderpoorten 221.5a (PC); Sierra 4281 (INPA),
on rocks near river fall and flat flooded rocks by
creek in white sand forest with Riccardia
regnellii, Zoopsidella integrifolia and Pictolejeu-
nea sprucei.
Plagiochilaceae
Plagiochila disticha (Lehm. & Lindenb.) Lindenb.,
Sierra 4282 (PC,INPA), on bark in white sand
forest.
Plagiochila montagnei Nees, Vanderpoorten 182.3,
175.1 (LG), on bark of living trees in both terra
firme and white sand forest, with Octoblepharum
albidum, O. pulvinatum, Syrrhopodon graminico-
la, S. cryptocarpos, Archilejeunea fuscescens.
Radulaceae
Radula flaccida Lindenb. & Gottsche, Sierra 4375;
Zartman 7267.1, 7291, 7170 (INPA), one of the
commonest epiphyllous bryophytes.
Radula javanica Gottsche, Zartman 10161 (INPA),
epiphyte.
BRYOPHYTA (Mosses)
Bartramiaceae
Philonotis uncinata (Schw¨
agr.) Brid., Sierra 4261
(INPA), on rock in exposed area near river.
Calymperaceae
Calymperes lonchophyllum Schw¨
agr., Vanderpoorten
178.4, 195.1 (LG), epiphyte.
Calymperes mitrafugax Florsch., Zartman 7469 (IN-
PA), epiphyte.
Calymperes rubiginosum (Mitt.) W.D.Reese, Zartman
7484, 10119, 10126 (INPA); Vanderpoorten 190.3
bis (LG), epiphyte.
Syrrhopodon annotinus W.D.Reese & D.G.Griffin,
Vanderpoorten 172.1, 172.AD, 174.2, 173.4,
178.1, 205.4, 212.1, 217.4 (LG); Pereira 756;
Zartman 7476, 7480, 7488, 10169, 10173, 10175,
10148 (INPA), species growing on soil and logs in
white sand forest.
Syrrhopodon cryptocarpos Dozy & Molk., Vander-
poorten 182.4, 182.7, 186.4, 188.2, 205.2, 206.1,
213.3 (LG); Pereira 755 (INPA), occurs in various
substrate as epiphyte, on log and rocks.
Syrrhopodon cymbifolius M¨
ull.Hal., Sierra 4366;
Zartman 10015, 10020, 10164 (INPA), epiphyte
on twigs and epiphyll.
Syrrhopodon fimbriatus Mitt., Vanderpoorten 210.5,
216.2 (LG); Pereira 778; Zartman 7301 (INPA),
epiphyte.
Syrrhopodon flexifolius Mitt., Zartman 7550 (INPA),
epiphyte.
584 The Bryologist 121(4): 2018
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Syrrhopodon graminicola R.S.Williams, Vanderpoort-
en 182.2 (LG), epiphyte.
Syrrhopodon helicophyllus Mitt., Vanderpoorten 221.2,
224.1 (LG); Zartman 7501 (INPA), epiphyte.
Syrrhopodon hornschuchii Mart., Vanderpoorten
180.1, 190.2, 197.1, 198.1, 211.1, P431.2 (LG);
Pereira 773, 761; Zartman 7259.1, 7267 (INPA),
epiphyte.
Syrrhopodon ligulatus Mont., Zartman 7158 (INPA),
epiphyte.
Syrrhopodon prolifer Schw¨
agr. var. prolifer, Pereira
779 (INPA), epiphyte. var. scaber (Mitt.)
W.D.Reese, Pereira 752 (INPA), epiphyte.
Syrrhopodon rigidus Hook. & Grev., Pereira 751, 771
(INPA), epiphyte.
Syrrhopodon simmondsii Steere, Vanderpoorten 173.2,
178.7, 185.5, 190.1, 203.1, 205.1, 216.1, 217.2,
228.3 (LG); Pereira 770, 772; Zartman 7507,
10039, 10050, 10182 (INPA), occurs in various
substrate as epiphyte, on log, soil and rocks in
white sand forest.
Syrrhopodon xanthophyllus Mitt., Pereira 762, 777,
750; Sierra 4280, 4300; Zartman 10005, 10118,
10120, 10123, 10127 (INPA); Vanderpoorten
172.1, 210.2, 227.2, 227.7, 181.1, 190.3, 190.4,
197.1, 210.4, 213.1, 210.3, 172.1bis2, P420.2 (LG),
occurs in various substrate as epiphyte, on log
and soil in white sand forest.
Fissidentaceae
Fissidens pellucidus Hornsch. var. pellucidus,Pereira
763 (INPA), on soil.
Fissidens prionodes Mont., Pereira 753 (INPA), on soil.
*Fissidens steerei Grout, Zartman 7520 (INPA), on soil
in terra firme.
Leucobryaceae
Leucobryum martianum (Hornsch.) M¨
ull.Hal., Van-
derpoorten 172.2, 177.2, 178.6, 184.2, 186.2, 187.2,
193.1, 195bis.2, 195.3, 197.2, 199.2, 204.3, 210.9,
214.5, 216.7, 217.5, 220.2, 221.7, 221.9, 223.2,
225.2, 227.4, 228.1, 230.2, P423 (LG); Pereira 774,
757, 775, 768; Zartman 10142, 10062, 10150
(INPA), occurs in various substrate as epiphyte,
on log, rock and soil in white sand forest.
Octoblepharaceae
Octoblepharum albidum Hedw., Pereira 776, 758,
769; Zartman 10003, 10012, 10035, 10102 (INPA),
occurs as epiphyte, on rock and soil.
Octoblepharum cocuiense Mitt., Pereira 760; Zartman
7156, 7524, 10118, 10075, 10123 (INPA), occurs
as epiphyte, on logs and rocks.
Octoblepharum leucobryoides O.Yano, Zartman
10169, 10170, 10177, 10148 (INPA), on soil in
white sand forest.
Octoblepharum pulvinatum (Dozy & Molk.) Mitt.,
Zartman 10037, 10039, 10050 (INPA), ussually
occurs as epiphyte and on log.
Octoblepharum rhaphidostegium Broth., Pereira 759
(INPA), epiphyte.
Octoblepharum stramineum Mitt., Vanderpoorten
227.3, 230.3 (LG); Zartman 10035, 10099 (INPA),
epiphyte.
Orthotrichaceae
Macromitrium punctatum (Hook. & Grev.) Brid.,
Zartman 7495.1 (INPA), epiphyte.
Pilotrichaceae
Callicostella pallida (Hornsch.) ˚
Angstr., Zartman
7509.1 (INPA), on log.
Crossomitrium patrisiae (Brid.) M¨
ull.Hal., Zartman
7444 (INPA), epiphyll.
Pottiaceae
Hyophila involuta (Hook.) A.Jaeger, Vanderpoorten
174.2 (LG), on soil.
Pterobryaceae
Henicodium geniculatum (Mitt.) W.R.Buck, Zartman
10015, 10021, 10036, 10096, 10098 (INPA),
epiphyte on mid trunk and canopy.
Sematophyllaceae
Brittonodoxa subpinnata (Brid.) W.R.Buck,
P.E.A.S.Cˆ
amara & Carv.-Silva, Pereira 749;
Zartman 7205, 9978, 9994 (INPA), occurs as
epiphyte and on log.
Vitalia galipensis (M¨
ull. Hal.) P.E.A.S.Cˆ
amara et al.,
Pereira 754, 767 (INPA), on log.
Microcalpe subsimplex (Hedw.) W.R.Buck, Pereira
765; Zartman 7491, 7285, 10032, 10039, 10050,
10080, 10084, 10164, 10107, 10112 (INPA);
Vanderpoorten 215.1 (LG), occurs as epiphyte
and on log.
Taxithelium planum (Brid.) Mitt., Pereira 764 (INPA),
on log in white sand forest.
Trichosteleum papillosum (Hornsch.) A.Jaeger, Van-
derpoorten 221.1 (LG); Zartman 7474, 9988,
10004, 10007, 10009 (INPA), flat flooded rocks
by creek with Riccardia regnellii,Symphyogyna
Sierra et al.: Bryophytes of Jau
´National Park, Brazil 585
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brasiliensis, Pictolejeunea sprucei, Zoopsidella
integrifolia.
Trichosteleum vincentinum (Mitt.) A.Jaeger, Zartman
7280, 7281 (INPA), on log in white sand forest.
Trichosteleum subdemissum (Besch.) A.Jaeger, Zart-
man 7277, 7282, 10174, 7277, 7282, 9981, 10017,
10018, 10034 (INPA), epiphyte on base of tree, on
soil, log, and rocks in inundated forest.
ACKNOWLEDGMENTS
Fieldwork was funded by the Projecto Bilateral Brasil/B´
elgica
processo 49518/2013-3 (CNPq), and by the Conselho Nacional de
Desenvolvimento Cient´
ıfico e Tecnol ´
ogico (CNPq, Grant No.
441590/2016-0) and the Funda¸ca
˜o de Amparo `
a Pesquisa do Estado
do Amazonas (FAPEAM, Grant No. 015/2016) for funding through
the project PELD/MAUA (Ecologia, Monitoramento e Uso
Sustenta
´vel de ´
Areas ´
Umidas Amazˆ
onicas). AMS acknowledges
financial support for a Master grant fellowship from CAPES. MRP
acknowledges financial support for Doctoral grant fellowship from
CNPq. CJPB is grateful to Conselho Nacional de Desenvolvimento
Cient´
ıfico e Tecnol ´
ogico (CNPq) for the grant of the Research
Productivity Fellowship. CEZ acknowledges financial support from
MCT/CNPq N8017/2013: Coopera ¸ca
˜o Internacional – Acordos
bilaterais. The authors thank Kleuto Moraes da Silva for logistical
support and help during fieldwork.
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