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- Source title: Brazilian bryophytes: a national-scale comprehensive dataset of species richness and geographic distribution
- Author(s): Silva, J. V., Marmo, J., Tavares, K. j., Oliveira, H. C., Silva, M. P.
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# **Brazilian bryophytes: a national-scale comprehensive** **dataset of species richness and geographic distribution**


Jailton Venilson Ferreira da Silva &#91;1&#93;, Jhonyd Jhonata de Oliveira Marmo &#91;1&#93;, Kleberson
Jonata da Cruz Tavares &#91;1&#93;, Hermeson Cassiano de Oliveira², Mércia Patrícia Pereira Silva¹

1 Programa de Pós-Graduação em Biologia Vegetal (PPGBV), Laboratório de Biologia de
Briófitas, Departamento de Botânica, Centro de Biociências, Universidade Federal de
Pernambuco, Av. Prof. Moraes Rego s/n - Cidade Universitária, Recife, PE 50670-901,

Brasil.
2 Laboratório de Botânica, Departamento de Biologia, Universidade Estadual do Piauí,
Campus Heróis do Jenipapo, Av. Santo Antônio s/n - São Luís, Campo Maior, PI 64280
000, Brasil.


**Abstract:** Bryophytes (Anthocerotophyta, Marchantiophyta, and Bryophyta) remain


under-sampled in large-scale biodiversity databases in Brazil, despite their ecological


importance. In Brazil, Flora e Funga do Brasil recognizes 1,618 bryophyte species


distributed across six biomes that are increasingly threatened by anthropogenic pressures


such as deforestation, agricultural expansion, and climate change. Nevertheless,


knowledge of species geographic distributions remains fragmented and based on scattered


records. Flora e Funga do Brasil serves as the official taxonomic reference for the country,


providing the accepted species list and serving as a baseline for biodiversity studies.


However, as it was designed as a nomenclatural and taxonomic checklist, it does not


integrate primary occurrence records with geographic coordinates and collection dates.


Consequently, occurrence data remain dispersed across herbaria, digital repositories, and


the scientific literature, without dynamic updates linked to the official checklist. This


disconnect between the taxonomic framework and primary occurrence records limits


macroecological and biogeographical analyses at national scales, which depend on


standardized georeferenced biodiversity databases. To address this gap, we compiled


&#91;bioRxiv preprint doi: https://doi.org/10.64898/2026.11.01.763487; this version posted November 3, 2026. The copyright holder for this preprint&#93;(https://doi.org/10.64898/2026.11.01.763487)

(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
&#91;available under a CC-BY-NC-ND 4.0 International license.&#93;(http://creativecommons.org/licenses/by-nc-nd/4.0/)


102,528 bryophyte occurrence records from herbarium specimens, using data from GBIF


and the speciesLink network, complemented by field collections from the Caatinga and


records from the Cryptogamic Herbarium of the State University of Piauí. All records were


georeferenced and curated, including verification of inconsistent coordinates, removal of


duplicates, and taxonomic standardization using Bryonames. When available, information


such as collection date and collector was retained. The dataset comprises 1,820 species,


125 families, and 442 genera. Bryophyta (mosses) was the most diverse group (1,056 spp.,


84 families, 310 genera), followed by Marchantiophyta (745 spp., 37 families, 124 genera)


and Anthocerotophyta (19 spp., four families, seven genera). Among mosses, the most


species-rich families were Sphagnaceae (79 spp.), Fissidentaceae (76 spp.), and Pottiaceae


(76 spp.). Among liverworts, Lejeuneaceae dominated with 335 species, followed by


Lepidoziaceae (58 spp.) and Plagiochilaceae (51 spp.). Among hornworts,


Anthocerotaceae, Dendrocerotaceae, and Notothyladaceae each included six species.


Regionally, the Southeast showed the highest richness (1,338 spp.), followed by the South


(974 spp.), Central-West (937 spp.), Northeast (897 spp.), and North (763 spp.). Regarding


biomes, the Atlantic Forest (1,523 spp.) and Amazon (1,054 spp.) were the most species

rich, followed by the Cerrado (810 spp.), Caatinga (671 spp.), Pampa (269 spp.), and


Pantanal (61 spp.). These patterns reflect both ecological gradients and historical sampling


biases. This dataset bridges the gap between the official taxonomic checklist and primary


occurrence data by integrating multiple sources into a standardized, openly accessible


database. It provides a robust tool for studies in macroecology, biogeography, and


conservation, enabling analyses of spatial patterns of species richness, endemism, and


taxonomic, functional, and phylogenetic diversity across environmental gradients. It also


&#91;bioRxiv preprint doi: https://doi.org/10.64898/2026.11.01.763487; this version posted November 3, 2026. The copyright holder for this preprint&#93;(https://doi.org/10.64898/2026.11.01.763487)

(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
&#91;available under a CC-BY-NC-ND 4.0 International license.&#93;(http://creativecommons.org/licenses/by-nc-nd/4.0/)


supports species distribution modeling, climate change impact assessments, biodiversity


hotspot identification, and conservation planning at regional and national scales. Users are


encouraged to cite this article when using the dataset and to report downstream applications


derived from it.


_Key words/phrases:_ biodiversity; Brazilian biomes; data paper; hornworts; liverworts;


mosses.


Open Research: The complete data set is available as Supporting Information at: &#91; _to be_


_completed at proof stage_ &#93;. Associated data is also available at &#91;name of data repository&#93;:


&#91;DOI assigned to deposited material&#93;.


      - Corresponding Author E-mail: jailton.venilson@ufpe.br


&#91;bioRxiv preprint doi: https://doi.org/10.64898/2026.11.01.763487; this version posted November 3, 2026. The copyright holder for this preprint&#93;(https://doi.org/10.64898/2026.11.01.763487)

(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
&#91;available under a CC-BY-NC-ND 4.0 International license.&#93;(http://creativecommons.org/licenses/by-nc-nd/4.0/)


1 Title: Brazilian bryophytes: a national-scale comprehensive dataset of species richness and
2 geographic distribution


3 Authors: Jailton Venilson Ferreira da Silva¹, Jhonyd Jhonata de Oliveira Marmo¹, Kleberson
4 Jonata da Cruz Tavares &#91;1&#93;, Hermeson Cassiano de Oliveira², Mércia Patrícia Pereira Silva¹


5 Author Affiliations:


6 1. Programa de Pós-Graduação em Biologia Vegetal (PPGBV), Laboratório de Biologia de
7 Briófitas, Departamento de Botânica, Centro de Biociências, Universidade Federal de
8 Pernambuco, Av. Prof. Moraes Rego s/n - Cidade Universitária, Recife, PE 50670-901,
9 Brasil.

10 2. Laboratório de Botânica, Departamento de Biologia, Universidade Estadual do Piauí,
11 Campus Heróis do Jenipapo, Av. Santo Antônio s/n - São Luís, Campo Maior, PI 6428012 000, Brasil.


13 Corresponding Author: jailton.venilson@ufpe.br


14 Open Research statement: The data is provided as a private submission for peer review via the
15 following link:


16


17


18


19


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(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
&#91;available under a CC-BY-NC-ND 4.0 International license.&#93;(http://creativecommons.org/licenses/by-nc-nd/4.0/)


20 Introduction


21 Studies on biodiversity in tropical plant communities have predominantly focused on trees


22 (Condit et al., 2002; Eisenlohr et al., 2013; Gentry, 1988; Oliveira &amp; Daly, 1999; Oliveira

23 Filho et al., 2005; Pitman et al., 2001, 2002; Ter Steege et al., 2003), whereas bryophytes


24 (Anthocerotophyta—hornworts, Marchantiophyta—liverworts, and Bryophyta—mosses)


25 have often been overlooked despite their ecological importance. These plants contribute to


26 water retention, facilitate the establishment of vascular plants (Welch, 1948), and provide


27 essential microhabitats for a wide range of organisms (Gradstein et al., 2001; Glime, 2007).


28 In Brazil, 1,618 bryophyte species are currently recognized (18 hornworts, 702 liverworts,


29 and 898 mosses; Flora e Funga do Brasil, 2025), distributed across different biomes that


30 are increasingly threatened by anthropogenic pressures such as deforestation, climate


31 change, agricultural expansion, wildfires, and urbanization (Hirota, 2003; INPE, 2016;


32 Imazon, 2020; Walter &amp; Sevilha, 2019). Despite these threats, the country's unique


33 combination of humid and dry forests still provides suitable conditions for bryophyte


34 development. However, knowledge of their distribution remains fragmented and


35 incomplete (Aires &amp; Bordin, 2024; Carvalho et al., 2025; Lima &amp; Peralta, 2021; Marmo &amp;


36 Silva, 2025; Silva et al., 2024, 2025; Sousa et al., 2024).


37 Despite significant advances in the knowledge of the Brazilian bryoflora over recent


38 decades, information on species occurrences remains scattered across biological


39 collections, digital herbaria, regional inventories, and scientific publications. In this


40 context, Flora e Funga do Brasil (2025) represents the first official taxonomic framework


41 for the country, providing a comprehensive checklist of recognized species (Forzza et al.,


42 2012). However, primary occurrence records including geographic coordinates, collector


&#91;bioRxiv preprint doi: https://doi.org/10.64898/2026.11.01.763487; this version posted November 3, 2026. The copyright holder for this preprint&#93;(https://doi.org/10.64898/2026.11.01.763487)

(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
&#91;available under a CC-BY-NC-ND 4.0 International license.&#93;(http://creativecommons.org/licenses/by-nc-nd/4.0/)


43 information, collection dates, and substrate data remain dispersed among biological


44 collections, digital herbaria, and the specialized literature. Initiatives such as SpeciesLink


45 and GBIF have integrated part of these records (Canhos et al., 2004; GBIF.org, 2025), but


46 they do not replace a consolidated and standardized national-scale database. This limitation


47 stems from the fact that Flora e Funga do Brasil was not designed as a dynamic repository


48 of occurrence records, while taxonomic updates and herbarium data continue to follow


49 independent update cycles.


50 Furthermore, collecting effort has historically been uneven across Brazilian regions and


51 biomes, generating spatial and taxonomic biases that hinder a comprehensive


52 understanding of species distribution and richness patterns (Costa &amp; Peralta, 2015).


53 Consequently, the establishment of large, standardized databases is crucial for advancing


54 studies in macroecology, biogeography, conservation, and environmental change


55 monitoring (Antunes et al., 2022; Costa-Pinto et al., 2022; Costa et al., 2020; Hasui et al.,


56 2017; Ramos et al., 2019; Travers &amp; Dunnell, 2009).


57 In this study, we present a dataset comprising more than 100,000 bryophyte occurrence


58 records from Brazil, encompassing all three bryophyte phyla across a wide range of


59 biomes. The dataset integrates taxonomic, spatial, and temporal information and provides


60 a valuable resource for analyses of species richness, geographic distribution, taxonomic


61 representativeness, and collecting effort over time. Furthermore, it serves as a foundation


62 for future research in biodiversity, conservation, macroecology, biogeography, and


63 ecological modeling.


64


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65 **Class I. Data Set Descriptors**
66 **A.** **Data set identity:**
67 **Title:** Brazilian bryophytes: a national-scale comprehensive dataset of species richness
68 and geographic distribution

69

70 **B.** **Data set identification code.**


71 **Suggested Data Set Identity Codes:**


72 BRYOPHYTES_BRAZIL_Occurrence.csv


73 **C.** **Data set description**


74 **Principal investigators**


75 1. Jailton Venilson Ferreira da Silva, Jhonyd Jhonata de Oliveira Marmo, Kleberson Jonata


76 da Cruz Tavares, Mércia Patrícia Pereira Silva


77 Laboratório de Biologia de Briófitas, Departamento de Botânica, Centro de Biociências,
78 Universidade Federal de Pernambuco, Av. Prof. Moraes Rego s/n - Cidade Universitária,
79 Recife, PE 50670-901, Brasil.


80 2. Hermeson Cassiano de Oliveira


81 Laboratório de Botânica, Departamento de Biologia, Universidade Estadual do Piauí,
82 Campus Heróis do Jenipapo, Av. Santo Antônio s/n - São Luís, Campo Maior, PI 6428083 000, Brasil.


84 1. Abstract: Bryophytes (Anthocerotophyta, Marchantiophyta, and Bryophyta) remain


85 under-sampled in large-scale biodiversity databases in Brazil, despite their ecological


86 importance. In Brazil, Flora e Funga do Brasil recognizes 1,618 bryophyte species


87 distributed across six biomes that are increasingly threatened by anthropogenic


88 pressures such as deforestation, agricultural expansion, and climate change.


89 Nevertheless, knowledge of species geographic distributions remains fragmented and


90 based on scattered records. Flora e Funga do Brasil serves as the official taxonomic


91 reference for the country, providing the accepted species list and serving as a baseline


92 for biodiversity studies. However, as it was designed as a nomenclatural and


&#91;bioRxiv preprint doi: https://doi.org/10.64898/2026.11.01.763487; this version posted November 3, 2026. The copyright holder for this preprint&#93;(https://doi.org/10.64898/2026.11.01.763487)

(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
&#91;available under a CC-BY-NC-ND 4.0 International license.&#93;(http://creativecommons.org/licenses/by-nc-nd/4.0/)


93 taxonomic checklist, it does not integrate primary occurrence records with


94 geographic coordinates and collection dates. Consequently, occurrence data remain


95 dispersed across herbaria, digital repositories, and the scientific literature, without


96 dynamic updates linked to the official checklist. This disconnect between the


97 taxonomic framework and primary occurrence records limits macroecological and


98 biogeographical analyses at national scales, which depend on standardized


99 georeferenced biodiversity databases. To address this gap, we compiled 102,528


100 bryophyte occurrence records from herbarium specimens, using data from GBIF and


101 the speciesLink network, complemented by field collections from the Caatinga and


102 records from the Cryptogamic Herbarium of the State University of Piauí. All records


103 were georeferenced and curated, including verification of inconsistent coordinates,


104 removal of duplicates, and taxonomic standardization using Bryonames. When


105 available, information such as collection date and collector was retained. The dataset


106 comprises 1,821 species, 125 families, and 442 genera. Bryophyta (mosses) was the


107 most diverse group (1,056 spp., 84 families, 310 genera), followed by


108 Marchantiophyta (745 spp., 37 families, 124 genera) and Anthocerotophyta (19 spp.,


109 four families, seven genera). Among mosses, the most species-rich families were


110 Sphagnaceae (79 spp.), Fissidentaceae (76 spp.), and Pottiaceae (76 spp.). Among


111 liverworts, Lejeuneaceae dominated with 335 species, followed by Lepidoziaceae (58


112 spp.) and Plagiochilaceae (51 spp.). Among hornworts, Anthocerotaceae,


113 Dendrocerotaceae, and Notothyladaceae each included six species. Regionally, the


114 Southeast showed the highest richness (1,338 spp.), followed by the South (974 spp.),


115 Central-West (937 spp.), Northeast (897 spp.), and North (763 spp.). Regarding


&#91;bioRxiv preprint doi: https://doi.org/10.64898/2026.11.01.763487; this version posted November 3, 2026. The copyright holder for this preprint&#93;(https://doi.org/10.64898/2026.11.01.763487)

(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
&#91;available under a CC-BY-NC-ND 4.0 International license.&#93;(http://creativecommons.org/licenses/by-nc-nd/4.0/)


116 biomes, the Atlantic Forest (1,523 spp.) and Amazon (1,054 spp.) were the most


117 species-rich, followed by the Cerrado (810 spp.), Caatinga (671 spp.), Pampa (269


118 spp.), and Pantanal (61 spp.). These patterns reflect both ecological gradients and


119 historical sampling biases. This dataset bridges the gap between the official


120 taxonomic checklist and primary occurrence data by integrating multiple sources into


121 a standardized, openly accessible database. It provides a robust tool for studies in


122 macroecology, biogeography, and conservation, enabling analyses of spatial patterns


123 of species richness, endemism, and taxonomic, functional, and phylogenetic diversity


124 across environmental gradients. It also supports species distribution modeling,


125 climate change impact assessments, biodiversity hotspot identification, and


126 conservation planning at regional and national scales. Users are encouraged to cite


127 this article when using the dataset and to report downstream applications derived


128 from it.


129 D. Key words/phrases: biodiversity; Brazilian biomes; data paper; hornworts; liverworts;


130 mosses.


131 **Description**


132 A total of 102,528 bryophyte occurrence records were compiled for Brazil, encompassing


133 representatives of all three phyla: Anthocerotophyta, Bryophyta, and Marchantiophyta.


134 The records show a broad distribution across the Brazilian territory, with the highest


135 concentration of occurrences in the Atlantic Forest and along coastal regions of the


136 Southeast and South (Figure 1). In contrast, lower record densities were observed in the


137 Pantanal and Pampa biomes, as well as in inland areas of the Northern region.


138


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(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
&#91;available under a CC-BY-NC-ND 4.0 International license.&#93;(http://creativecommons.org/licenses/by-nc-nd/4.0/)


139


140 To better understand spatial distribution patterns, occurrence records were


141 subsequently analyzed separately for each bryophyte phylum. Bryophyta accounted


142 for 68,478 records across Brazil (Figure 2), whereas Marchantiophyta comprised


143 33,594 records (Figure 3). Bryophyta and Marchantiophyta exhibited the broadest


144 geographic distributions and the highest numbers of records throughout the


145 country. In contrast, Anthocerotophyta was less well represented, with only 455


146 records and a more restricted spatial distribution (Figure 4).


147 The distribution of records was also analyzed across the different Brazilian biomes


148 for all three bryophyte phyla. The Atlantic Forest exhibited the highest numbers of


149 occurrence records and species richness, followed by the Amazon, Caatinga, and


150 Cerrado. In contrast, the Pantanal and Pampa showed the lowest representation in


151 terms of both record numbers and species richness. Bryophyta and


152 Marchantiophyta were widely distributed across all biomes, particularly in humid


153 forested regions, whereas Anthocerotophyta displayed a more restricted


154 distribution concentrated in specific areas.


155 The complete dataset comprises 1,820 identified species distributed across 125


156 families and 442 genera. Bryophyta was the most diverse division, with 1,056


157 species, 310 genera, and 84 families, followed by Marchantiophyta, with 745


158 species, 124 genera, and 37 families, and Anthocerotophyta, with 19 species, seven


159 genera, and four families. Among the mosses (Bryophyta), the most species-rich


160 families were Sphagnaceae (79 spp.), Fissidentaceae (76 spp.), and Pottiaceae (76


161 spp.). Among the liverworts (Marchantiophyta), Lejeuneaceae was the dominant


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162 family, comprising 335 species, followed by Lepidoziaceae (58 spp.) and


163 Plagiochilaceae (51 spp.). Among the hornworts (Anthocerotophyta), the most


164 species-rich families were Anthocerotaceae, Dendrocerotaceae, and


165 Notothyladaceae, each represented by six species.


166 The Atlantic Forest contained the highest number of bryophyte records, totaling


167 54,237 occurrences, followed by the Amazon, with 18,560 records. The Caatinga


168 and Cerrado were also well represented, accounting for 14,561 and 13,856 records,


169 respectively. The lowest numbers of records were observed in the Pampa, with 954


170 occurrences, and the Pantanal, with only 146 records. Species richness was likewise


171 highest in the Atlantic Forest, where 1,523 species were recorded, followed by the


172 Amazon with 1,054 species. The Cerrado harbored 810 species, whereas the


173 Caatinga contained 671 recorded species. The lowest species richness values were


174 found in the Pampa, with 269 species, and the Pantanal, with only 61 recorded


175 species.


176 The Southeast region contained the highest number of bryophyte records, totaling


177 32,069 occurrences. It was followed by the Northeast, with 30,160 records. The


178 North accounted for 15,288 records, whereas the South and Central-West regions


179 contributed 12,607 and 12,403 records, respectively. Species richness was also


180 highest in the Southeast, with 1,338 recorded species, followed by the South, with


181 974 species. The Central-West harbored 937 species, while the Northeast accounted


182 for 897 species. The lowest species richness was observed in the North, with 763


183 recorded species.


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184 At the state level, Bahia exhibited the highest number of bryophyte records, totaling


185 18,367 occurrences, followed by São Paulo, with 15,292 records. Amazonas


186 accounted for 7,598 records, while Minas Gerais and Rio de Janeiro contributed


187 6,741 and 5,291 records, respectively. Other states with substantial representation


188 included Pará (4,785 records), Espírito Santo (4,745), Mato Grosso (4,728), Rio


189 Grande do Sul (4,376), Paraná (4,305), and Santa Catarina (3,926). Intermediate


190 numbers of records were observed in Goiás (3,433 records), Pernambuco (3,347),


191 the Federal District (3,312), Ceará (2,809), and Piauí (2,596). The lowest numbers


192 of records were found in Rondônia (1,126), Alagoas (1,072), Paraíba (932), Mato


193 Grosso do Sul (930), Roraima (769), Sergipe (671), Acre (558), Amapá (380), Rio


194 Grande do Norte (192), Maranhão (174), and Tocantins (72 records).


195 The highest species richness was recorded in São Paulo, with 999 species, followed


196 by Minas Gerais (826 species), Rio de Janeiro (810), Bahia (770), and Mato Grosso


197 (736). High species richness was also observed in Paraná (667 species), Santa


198 Catarina (618), Rio Grande do Sul (610), Amazonas (591), and Espírito Santo


199 (528). Intermediate richness values were recorded in Pará (410 species),


200 Pernambuco (398), Goiás (382), the Federal District (332), Ceará (285), Roraima


201 (235), and Alagoas (221). The lowest richness values were observed in Paraíba


202 (194 species), Piauí (187), Rondônia (184), Mato Grosso do Sul (180), Acre (145),


203 Sergipe (125), Amapá (90), Maranhão (77), Tocantins (48), and Rio Grande do


204 Norte, with only 41 recorded species.


205 We analyzed bryophyte records across five temporal periods and observed a


206 marked increase in the number of records over time. Between 1800–1850 and


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207 1901–1950, the number of records remained relatively low, indicating limited


208 sampling effort during the early stages of bryological exploration in Brazil. From


209 1951–2000 onward, there was a sharp increase in recorded occurrences, while the


210 2001–2025 period exhibited the highest values, exceeding 45,000 records (Figure


211 5). This pattern likely reflects the expansion of floristic surveys, the growth in the


212 number of bryology specialists, the strengthening of biological collections, and the


213 increasing digitization of Brazilian herbaria.


214 We also analyzed bryophyte species richness over time. The results show a


215 progressive increase in the number of species recorded in Brazil throughout the


216 studied periods. Between 1800–1850, only a few species had been documented,


217 whereas subsequent periods exhibited a continuous increase in known species


218 richness. The most pronounced increase occurred between 1951–2000, when the


219 number of recorded species exceeded 1,400. During the 2001–2025 period, the


220 upward trend in species richness was maintained, with a slight increase of two


221 species compared to the previous interval. This pattern may be related to


222 differences in sampling effort, temporal variation in data availability, or the absence


223 of recently collected records that have not yet been fully digitized (Figure 6).


224 The number of bryophyte records increased markedly over time across all Brazilian


225 biomes. In the earliest periods (1800–1850 and 1851–1900), records were relatively


226 scarce, with the Atlantic Forest already showing the highest number of documented


227 occurrences. From 1951–2000 onward, a pronounced increase in records was


228 observed across all biomes, becoming even more evident in the 2001–2025 period.


229 The Atlantic Forest consistently concentrated the highest number of records across


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(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
&#91;available under a CC-BY-NC-ND 4.0 International license.&#93;(http://creativecommons.org/licenses/by-nc-nd/4.0/)


230 all recent periods, reaching 27,157 records between 2001–2025, followed by the


231 Cerrado (8,332 records), Caatinga (6,642 records), and Amazon (4,695 records).


232 The Pantanal and Pampa exhibited the lowest numbers of records throughout the


233 entire time series, indicating lower sampling intensity or limited data availability in


234 these biomes (Figure 7).


235 Species richness also increased over the analyzed periods, following the growth in


236 the number of records. The Atlantic Forest exhibited the highest richness across all


237 periods, reaching 1,166 species between 2001–2025. It was followed by the


238 Cerrado (557 species), Amazon (522 species), and Caatinga (520 species).


239 Although species richness increased substantially up to the 1951–2000 period,


240 some biomes showed a slight reduction in the number of recorded species between


241 2001–2025, including the Atlantic Forest (from 1,196 to 1,166 species), the


242 Amazon (from 540 to 522 species), and the Cerrado (from 591 to 557 species). This


243 reduction may reflect differences in sampling effort, taxonomic updates, or data


244 standardization procedures. The Pampa and Pantanal consistently exhibited the


245 lowest richness values throughout the time series, reaching 147 and 21 species,


246 respectively, in the most recent period. In contrast, the Caatinga showed one of the


247 largest relative increases in richness, rising from only two recorded species in


248 1800–1850 to 520 species in 2001–2025 (Figure 8).


249 The fifteen most representative moss families and genera in the dataset were


250 selected based on both the number of occurrence records and species richness.


251 Sematophyllaceae had the highest number of records (6,938) and 57 species,


252 followed by Leucobryaceae (6,800 records; 66 species), Calymperaceae (5,354; 52


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253 species), and Fissidentaceae (4,634; 76 species). Other well-represented families


254 included Orthotrichaceae, Pylaisiadelphaceae, and Sphagnaceae (Figure 9). At the


255 genus level, Fissidens was the most representative, with 4,634 records and 76


256 species, followed by Campylopus (4,255 records; 48 species), Syrrhopodon (3,116;


257 33 species), and Sphagnum (3,100 records; 79 species). Other relevant genera


258 included Octoblepharum, Microcalpe, and Calymperes (Figure 10).


259 As with mosses, the fifteen most representative liverwort families and genera in the


260 dataset were selected based on the number of occurrence records and species


261 richness. Among families, Lejeuneaceae stood out prominently, with 17,648


262 records and 335 species, followed by Frullaniaceae (3,520 records; 44 species),


263 Plagiochilaceae (2,787; 51 species), and Lepidoziaceae (2,069; 58 species). Other


264 relevant families included Radulaceae, Metzgeriaceae, and Ricciaceae (Figure 11).


265 At the genus level, Lejeunea had the highest number of records, totaling 3,876


266 occurrences and 68 species, followed by Frullania (3,521 records; 44 species),


267 Cheilolejeunea (3,238; 41 species), and Plagiochila (2,764; 47 species). Additional


268 notable genera included Ceratolejeunea, Radula, Metzgeria, and Cololejeunea


269 (Figure 12).


270 Class II. Research origin descriptors
271 A. Overall project description:


272 1. Identity: A compilation of bryophyte occurrence data from Brazil.


273 2. Originators: The Brazilian Bryophytes Project was coordinated by Jailton Venilson


274 Ferreira da Silva (Federal University of Pernambuco, UFPE). All co-authors


275 contributed to the compilation, validation, and organization of the dataset. The


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276 research team is primarily composed of investigators from UFPE, with additional


277 collaboration from a researcher at the State University of Piauí (UESPI).


278 3. Period of study: The dataset covers the period from 1804 to 2025.


279 4. Objectives: Our objectives were to: (1) compile a comprehensive dataset of


280 Brazilian bryophytes, encompassing the three major groups, hornworts


281 (Anthocerotophyta), liverworts (Marchantiophyta), and mosses (Bryophyta), based


282 on occurrence records derived from field collections and herbarium specimens; and


283 (2) characterize the distribution of Brazilian bryophytes in order to identify


284 priorities for future sampling efforts. To our knowledge, this study represents the


285 first nationwide bryophyte dataset for Brazil integrating occurrence records with


286 information on biomes and geographic regions.


287 5. Abstract: Same as above.


288 6. Sources of funding: The compilation of this dataset was supported by Coordenação


289 de Aperfeiçoamento de Pessoal de Nível Superior (CAPES; Finance Code 001) and


290 Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq;


291 Universal Grant No. 409503/2025-8 and PPBio-RABECA Network Grant No.


292 441209/2023-8).


293 B. Specific subproject description


294 1. Site description


295 Brazil is recognized as a country with extensive forest cover, comprising


296 approximately 500 million hectares of natural and planted forests, which


297 correspond to about 59% of its national territory. This places Brazil as the country


298 with the second-largest forest area in the world, surpassed only by Russia. In


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299 addition, Brazil covers approximately 8.5 million km², extending between latitudes


300 5°16'N and 33°45'S and longitudes 34°47'W and 73°59'W (IBGE, 2025; Serviço


301 Florestal Brasileiro, 2019). Brazil's natural resources are under considerable


302 pressure from large-scale agricultural expansion, mining activities, and timber


303 extraction, creating major challenges for balancing economic development with


304 environmental conservation and the sustainable use of ecosystems (Laurance et al.,


305 2001). The country encompasses six major biomes, Amazon, Cerrado, Caatinga,


306 Atlantic Forest, Pantanal, and Pampa, two of which are recognized as global


307 biodiversity hotspots: the Cerrado and the Atlantic Forest. Together, these biomes


308 harbor approximately 2% of the world's endemic plant and vertebrate species and


309 account for more than two-thirds of the species richness recorded in Brazil (Myers


310 et al., 2000). These biomes are characterized by distinct vegetation formations,


311 climatic conditions, and precipitation regimes. The Atlantic Forest and Amazon are


312 predominantly humid forest ecosystems, whereas the Cerrado and Caatinga form a


313 broad corridor of seasonally dry vegetation. The Pampa, located in southern Brazil,


314 is dominated by grassland formations, while the Pantanal is characterized by


315 extensive floodplain vegetation and seasonal inundation dynamics (Oliveira-Filho


316 &amp; Fontes, 2000; Oliveira-Filho &amp; Ratter, 2002; Oliveira-Filho et al., 2005; Silva et


317 al., 2017; Serviço Florestal Brasileiro, 2019). Brazilian biomes have been subjected


318 to intense anthropogenic pressures, particularly as a result of deforestation, climate


319 change, agricultural expansion, wildfires, and urban growth. These impacts have


320 contributed to biodiversity loss, habitat fragmentation, and alterations in natural


321 ecological processes (Hirota, 2003; Imazon, 2020; INPE, 2016; Walter &amp; Sevilha,


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322 2019). In this context, comprehensive inventories and biodiversity databases are


323 essential for understanding patterns of biodiversity distribution in Brazil,


324 particularly for still under-sampled groups such as bryophytes.


325 2. Experimental or sampling design


326 Data compilation: All data were obtained from records of specimens


327 deposited in herbaria, using the Global Biodiversity Information Facility


328 (GBIF; GBIF.org &#91;2025a, 2025b, 2025c&#93;) as the primary data source due to


329 its broad international coverage. Additionally, the speciesLink network was


330 consulted to incorporate data from Brazilian collections not indexed in


331 GBIF. The dataset was further complemented with field collections


332 conducted in the Caatinga domain, as well as records from the Cryptogam


333 Herbarium of the State University of Piauí (UESPI), which is not currently


334 indexed in any online biodiversity platform.


335 3. Research methods


336 The database of moss, liverwort, and anthocerotophyte occurrences was compiled


337 based on records of specimens deposited in herbaria, using the Global Biodiversity


338 Information Facility (GBIF; GBIF.org &#91;2025a, 2025b, 2025c&#93;) as the primary


339 source, selected for its broad international coverage. In addition, the speciesLink


340 network (splink.org.br) was consulted to integrate data from national collections


341 not indexed in GBIF. The search and data extraction followed standardized criteria.


342 In GBIF, the search was limited to the terms “Bryophyta,” “Marchantiophyta,” and


343 “Anthocerotophyta.” The search was conducted individually for each division,


344 filtering for specimens preserved in scientific collections (Record Bases field) with


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345 available geographic coordinates (Location field). For speciesLink, the filters were


346 set to “Preserved specimen” in the record base. The terms referring to bryophytes


347 in the “Phylum” section were: “Bryophytes,” “Bryophyta,” “Marchantiophyta,”


348 “Anthocerotophyta,” “Moss,” “Hepatics,” “Bryidae,” “Hepatophyta,” and


349 “Marchantiidae.” To avoid pseudo-duplication of data across platforms, a manual


350 review of the records was conducted, excluding from the speciesLink dataset those


351 specimens already obtained via GBIF. After consolidation, a taxonomic quality


352 filter was applied: only samples identified or collected by bryophyte specialists


353 were retained in the dataset. This procedure aims to mitigate identification errors,


354 ensuring the taxonomic robustness essential for studies of distribution patterns and


355 ecological modeling. Subsequently, the geographic coordinates of each record


356 included in the dataset were individually verified using Google Maps to confirm


357 they matched the location provided in the records. The nomenclatural updating of


358 species was based on the BryoNames database (bryonames.org/) using the


359 nameMatch function from the U.Taxonstand package in R Studio v4.5 (R Core


360 Team 2025). When the verified species had multiple synonyms, the Flora and


361 Funga of Brazil database (floradobrasil.jbrj.gov.br) was consulted to determine


362 which of these synonyms were present in Brazil. Species with uncertain


363 nomenclature or questionable records were excluded from the dataset. In addition,


364 the data were supplemented with field collections from the Caatinga region, as well


365 as data from the Cryptogam Herbarium at the State University of Piauí; since this


366 herbarium is not indexed on any online platform, we used a portion of the


367 herbarium’s data. Regarding sample collection, the materials were collected


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368 manually using equipment such as a pocket knife, handheld magnifying glass, paper


369 bags and plastic bags, pens, a field notebook, an altimeter, and a GPS device to


370 record geographic coordinates. Samples were stored in kraft paper bags. For macro

371 and microscopic analysis, we examined the morpho-anatomical characteristics of


372 the gametophyte and sporophyte using a stereoscopic microscope and an optical


373 microscope. For sample handling, we used forceps, needles, scalpels, Petri dishes,


374 slides, and cover slips. We organized all the data into a single database containing


375 information on occurrence, biomes, regions, and Brazilian states where these


376 samples were collected.


377 4. Project personnel: The principal investigators are Jailton Venilson Ferreira da Silva


378 and Mércia Patrícia Pereira Silva. The associate researchers are Jhonyd Jhonata de


379 Oliveira Marmo, Kleberson Jonata da Cruz Tavares, and Hermeson Cassiano de


380 Oliveira.


381 Class III. Data set status and accessibility


382 A. Status


383 1. Latest update: June 26


384 2. Latest archive date: June 26


385 3. Metadata status: Last updated on June 26, submitted version.


386 4. Data verification: Geographic coordinates were verified using Google Earth, and


387 corrections were applied when necessary. Specimens not identified by bryophyte


388 specialists were excluded from the dataset.


389 B. Accessibility


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390 1. Contact persons: Jailton Venilson Ferreira da Silva (jailton.venilson@ufpe.br) and


391 Mércia Patrícia Pereira Silva (mercia.psilva@ufpe.br). Programa de Pós

392 Graduação em Biologia Vegetal (PPGBV), Laboratório de Biologia de Briófitas,


393 Departamento de Botânica, Centro de Biociências, Universidade Federal de


394 Pernambuco, Av. Prof. Moraes Rego s/n - Cidade Universitária, Recife, PE 50670

395 901, Brasil. +55 86 981254170


396 2. Storage location and medium: The original dataset will be available on the


397 Ecology Repository. The updated version and additional information can be


398 accessed on the Zenodo repository via the following link:


399 3. Copyright restrictions: Não


400 4. Proprietary restrictions: Please cite this data paper when using this dataset.


401 Researchers and educators are encouraged to report and share outcomes derived


402 from studies based on these data.


403 5. Costs: None


404 Class IV. Data structural descriptors


405 A. Data set file


406 1. Identity: BRYOPHYTES_BRAZIL_Occurrence.csv


407 2. Size: BRYOPHYTES_BRAZIL_Occurrence.csv, 26 MB


408 3. Format and storage mode: The dataset is provided as a comma-separated values


409 (CSV) file encoded in UTF-8. Text variables are stored as character strings,


410 whereas geographic coordinates are stored as floating-point numeric values and


411 temporal variables (day, month, and year) are stored as integers. Geographic


412 coordinates are reported in decimal degrees using the WGS84 datum. Missing


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413 values are represented as "NA". Data cleaning procedures included the removal of


414 duplicated records, standardization of taxonomic names, verification of geographic


415 coordinates, and validation of metadata fields.


416 4. Header information: See column descriptions in section B.


417 5. Alphanumeric attributes: Mixed


418 6. Data Anomalies: If no information is available for a given record, this is indicated


419 as 'NA'.


420 B. Variable information (Table 1).


421 C. Data anomalies: If no information is available for a given record, this is indicated as 'NA'.


422 Class V. Supplemental descriptors


423 A. Data acquisition


424 1. Data acquisition: Our data were obtained from secondary databases and field


425 collections.


426 2. Location of completed data forms: No physical forms were used for data collection.


427 The dataset was compiled exclusively from field collections and digital records


428 available in biodiversity information systems and biological collections.


429 3. Data entry verification procedures: Data verification involved both automated and


430 manual procedures. Scientific names were standardized according to the adopted


431 taxonomic classification. Geographic coordinates were checked for format


432 consistency and for plausibility of occurrence within Brazilian territory. Duplicate


433 records were identified and removed. Records containing incomplete, inconsistent,


434 or incompatible information relative to the established standards were either


435 corrected when possible or excluded from the final version of the database.


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436 B. Quality assurance/quality control procedures: Several quality control procedures were


437 applied to ensure the taxonomic, geographic, and temporal consistency of the dataset.


438 Scientific names were reviewed and standardized, with synonymy updates and spelling


439 corrections performed when necessary, using widely accepted taxonomic references for


440 bryophytes.


441


442 Records without species-level identification were removed from the database. Geographic


443 coordinates were checked for decimal degree format (WGS84), and records with invalid


444 coordinates were excluded. The final dataset underwent multiple verification steps to


445 ensure consistency among taxonomic, spatial, and temporal information prior to public


446 release.


447 C. Related materials: The records used in this study are associated with specimens


448 deposited in national and international herbaria, with relevant information maintained by


449 the respective institutions. Supplementary data include geographic coordinates, taxonomic


450 information, and collection dates. The dataset does not include photographs, videos, or


451 original field notebooks.


452 D. Archiving


453 1. Archival procedures: The final version of the dataset will be made available in CSV


454 format in a public open-access repository, ensuring long-term digital preservation,


455 traceability, and permanent access to the data used in this study.


456 2. Redundant archival sites: In addition to the version deposited in the primary


457 repository, copies of the dataset are maintained by the authors.


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458 E. Publications and results: The dataset was compiled to support studies on the diversity,


459 geographic distribution, and macroecological patterns of bryophytes in Brazil. The


460 associated results will be presented in this Data Paper and may support future research in


461 biodiversity, biogeography, conservation, and bryophyte ecology. Publications derived


462 from this dataset should appropriately cite this Data Paper and the repository where the


463 data are archived.


464 F. History of data set usage


465 1. Data request history: None


466 2. Data set update history: None


467 3. Review history: None


468 4. Questions and comments from secondary users: None


469 **ACKNOWLEDGMENTS:**


470 We thank Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES;


471 Finance Code 001) and Conselho Nacional de Desenvolvimento Científico e Tecnológico


472 (CNPq; Universal Grant No. 409503/2025-8 and PPBio-RABECA Network Grant No.


473 441209/2023-8).


474 **Literature Citations**


475 Aires, E. T., and J. Bordin. 2024. "Avanços no Conhecimento da Brioflora do Pampa Brasileiro."


476 _Hoehnea_ 51: e762023. https://doi.org/10.1590/2236-8906e762023


477 Antunes, A. C., A. Montanarin, D. M. Gräbin, E. C. S. Monteiro, F. F. de Pinho, G. C. Alvarenga,


478 and J. Ahumada, et al. 2022. "AMAZONIACAMTRAP: A Dataset of Mammal, Bird, and Reptile


479 Species Recorded with Camera Traps in the Amazon Forest." _Ecology_ 103(7): e3738.


480 https://doi.org/10.1002/ecy.3738


&#91;bioRxiv preprint doi: https://doi.org/10.64898/2026.11.01.763487; this version posted November 3, 2026. The copyright holder for this preprint&#93;(https://doi.org/10.64898/2026.11.01.763487)

(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
&#91;available under a CC-BY-NC-ND 4.0 International license.&#93;(http://creativecommons.org/licenses/by-nc-nd/4.0/)


481 Barbosa Sousa, M. E., E. B. Valente, and H. C. Oliveira. 2024. "Brioflora do Parque Nacional Serra


482 das Confusões, Piauí, Brasil." _Scientia_ _Plena_ 20(2): 021201.


483 https://doi.org/10.14808/sci.plena.2024.021201


484 The Bryophyte Nomenclature Database (BryNames). 2026. _BryNames: A Synonymized Checklist_


485 _of Bryophyte Names_ . Available at https://brynames.org (accessed June 16, 2026).


486 Canhos, D. A. L., E. A. B. Almeida, A. L. Assad, M. M. C. Bustamante, V. P. Canhos, A. D.


487 Chapman, and R. De Giovanni, et al. 2022. "speciesLink: Rich Data and Novel Tools for Digital


488 Assessments of Biodiversity." _Biota_ _Neotropica_ 22(spe): e20221394.


489 https://doi.org/10.1590/1676-0611-BN-2022-1394


490 Carvalho, F. V., J. P. Silva, A. L. Ilkiu-Borges, and R. S. Fernandes. 2025. "Bryophytes Diversity


491 in Mirador State Park, Maranhão, and Floristic Similarity among Protected Areas in the Brazilian


492 Cerrado." Biota Neotropica 25(3): e20251794. https://doi.org/10.1590/1676-0611-BN-2025-1794


493 Costa, D. P., and D. F. Peralta. 2015. "Bryophytes Diversity in Brazil." _Rodriguésia_ 66(4): 1063–


494 1071. https://doi.org/10.1590/2175-7860201566411


495 Costa-Pinto, A. L., R. S. Bovendorp, A. Bocchiglieri, A. Caccavo, A. C. Delciellos, A. C. Malhado,


496 and A. K. R. Almeida, et al. 2023. "Small Mammals from the Caatinga: A Dataset for the Brazilian


497 Semiarid Biome." _Ecology_ 104(3): e3879. https://doi.org/10.1002/ecy.3879


498 Costa, L. M. S., M. Goetze, A. V. Rodrigues, G. D. S. Seger, and F. Bered. 2020. "Global


499 Rheophytes Data Set: Angiosperms and Gymnosperms." _Ecology_ 101(8): e03056.


500 https://doi.org/10.1002/ecy.3056


501 Condit, R., N. Pitman, E. G. Leigh, J. Chave, J. Terborgh, R. B. Foster, and P. Núñez, et al. 2002.


502 "Beta-diversity in Tropical Forest Trees." _Science_ 295(5555): 666–669.


503 https://doi.org/10.1126/science.1066854


504 Eisenlohr, P. V., L. F. Alves, L. C. Bernacci, M. C. G. Padgurschi, R. B. Torres, E. M. B. Prata,


505 and F. A. M. dos Santos, et al. 2013. "Disturbances, Elevation, Topography and Spatial Proximity


506 Drive Vegetation Patterns along an Altitudinal Gradient of a Top Biodiversity Hotspot."


507 _Biodiversity and Conservation_ 22: 2767–2783. https://doi.org/10.1007/s10531-013-0553-x


508 Flora do Brasil 2026. Jardim Botânico do Rio de Janeiro. Available


509 in:&lt;http;//floradobrasil.jbrj.gov.br/ &gt;.


&#91;bioRxiv preprint doi: https://doi.org/10.64898/2026.11.01.763487; this version posted November 3, 2026. The copyright holder for this preprint&#93;(https://doi.org/10.64898/2026.11.01.763487)

(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
&#91;available under a CC-BY-NC-ND 4.0 International license.&#93;(http://creativecommons.org/licenses/by-nc-nd/4.0/)


510 Forzza, R. C., J. F. A. Baumgratz, C. E. M. Bicudo, D. A. L. Canhos, A. A. Carvalho Jr., M. A. N.


511 Coelho, and A. F. Costa, et al. 2012. "New Brazilian Floristic List Highlights Conservation


512 Challenges." _BioScience_ 62(1): 39–45. https://doi.org/10.1525/bio.2012.62.1.8


513 GBIF.org. 2026. GBIF Occurrence Download. Copenhagen: Global Biodiversity Information


514 Facility. https://www.gbif.org. Accessed June 16, 2026.


515 GBIF.org, 2025a. GBIF occurrence download &#91;https://doi.org/10.15468/dl.s8csnk&#93;(https://doi.org/10.15468/dl.s8csnk) (accessed 5


516 June 2025).


517 GBIF.org, 2025b. GBIF occurrence download https://doi.org/10.15468/dl.jpvrv5 (accessed 26 June


518 2025).


519 GBIF.org, 2025c. GBIF occurrence download &#91;https://doi.org/10.15468/dl.c36ame&#93;(https://doi.org/10.15468/dl.c36ame) (accessed 9


520 January 2026).


521 Gentry, A. H. 1988. "Tree Species Richness of Upper Amazonian Forests." _Proceedings of the_


522 _National Academy of Sciences_ 85(1): 156–159. https://doi.org/10.1073/pnas.85.1.156


523 Glime, J. M. 2007. _Bryophyte Ecology. Volume 1: Physiological Ecology_ . Houghton, MI: Michigan


524 Technological University and International Association of Bryologists. URL.


525 Gradstein, S. R., S. P. Churchill, and N. Salazar-Allen. 2001. _Guide to the Bryophytes of Tropical_


526 _America_ . Bronx, NY: New York Botanical Garden Press.


527 Hasui, É., J. P. Metzger, R. G. Pimentel, L. F. Silveira, A. A. d. A. Bovo, A. C. Martensen, and A.


528 Uezu, et al. 2018. "ATLANTIC BIRDS: A Data Set of Bird Species from the Brazilian Atlantic


529 Forest." _Ecology_ 99(2): 497. https://doi.org/10.1002/ecy.2119


530 Hirota, M. M. 2003. "Monitoring the Brazilian Atlantic Forest Cover." In _The Atlantic Forest of_


531 _South America: Biodiversity Status, Threats, and Outlook_, edited by C. Galindo-Leal and I. G.


532 Câmara, 60–65. Washington, DC: Center for Applied Biodiversity Science and Island Press.


533 Instituto Brasileiro de Geografia e Estatística (IBGE). 2025. Brasil em Síntese: Dados geográficos.


534 &#91;Instituto Brasileiro de Geografia e Estatística, Rio de Janeiro, Brazil. Available from&#93;(https://brasilemsintese.ibge.gov.br/territorio/dados-geograficos.html?utm_source=chatgpt.com)


535 https://brasilemsintese.ibge.gov.br/territorio/dados-geograficos.html.


536 Instituto do Homem e Meio Ambiente da Amazônia. Desmatamento na Amazônia cresce 30% em


537 2020 e bate recorde dos últimos dez anos. 2021. Available:


&#91;bioRxiv preprint doi: https://doi.org/10.64898/2026.11.01.763487; this version posted November 3, 2026. The copyright holder for this preprint&#93;(https://doi.org/10.64898/2026.11.01.763487)

(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
&#91;available under a CC-BY-NC-ND 4.0 International license.&#93;(http://creativecommons.org/licenses/by-nc-nd/4.0/)


538 https://imazon.org.br/imprensa/desmatamento-na-amazonia-cresce-30-em-um-2020-e-bate

539 recorde-dos-ultimos-dez-anos/


540 Instituto Nacional de Pesquisas Espaciais. Centro do INPE avança no mapeamento da Caatinga.


541 Available in: http://www.inpe.br/noticias/noticia.php?Cod_Noticia=4157.


542 Laurance, W. F., M. A. Cochrane, S. Bergen, P. M. Fearnside, P. Delamônica, C. Barber, S.


543 D'Angelo, and T. Fernandes. 2001. "The Future of the Brazilian Amazon." _Science_ 291(5503):


544 438–439. https://doi.org/10.1126/science.291.5503.438


545 Lima, J. S., and D. F. Peralta. 2021. "Brioflora do Parque Nacional da Serra da Bocaina, de São


546 Paulo, Brasil." _Hoehnea_ 48: e802020. https://doi.org/10.1590/2236-8906-80/2020


547 Marmo, J. J. de O., and M. P. P. Silva. 2025. "Bryophytes of a Brazilian Seasonally Dry Tropical


548 Forest: An Overview of Diversity and Environmental Drivers." _Flora_ 330: 152770.


549 https://doi.org/10.1016/j.flora.2025.152770


550 Myers, N., R. A. Mittermeier, C. G. Mittermeier, G. A. B. da Fonseca, and J. Kent. 2000.


551 "Biodiversity Hotspots for Conservation Priorities." _Nature_ 403(6772): 853–858.


552 https://doi.org/10.1038/35002501


553 Oliveira, A. A. de, and D. C. Daly. 1999. "Geographic Distribution of Tree Species Occurring in


554 the Region of Manaus, Brazil: Implications for Regional Diversity and Conservation." _Biodiversity_


555 _and Conservation_ 8(9): 1245–1259. https://doi.org/10.1023/A:1008972901159


556 Oliveira-Filho, A. T., and M. A. L. Fontes. 2000. "Patterns of Floristic Differentiation among


557 Atlantic Forests in Southeastern Brazil and the Influence of Climate." _Biotropica_ 32(4b): 793–810.


558 https://doi.org/10.1111/j.1744-7429.2000.tb00619.x


559 Oliveira-Filho, A. T., and J. A. Ratter. 2002. "Vegetation Physiognomies and Wood Flora of the


560 Bioma Cerrado." In _The Cerrados of Brazil: Ecology and Natural History of a Neotropical_


561 _Savanna_, edited by P. S. Oliveira and R. J. Marquis, 91–120. New York: Columbia University


562 Press.


563 Oliveira-Filho, A. T., W. A. C. Carvalho, E. Tameirão-Neto, E. Tameirão-Neto, W. A. C. Carvalho,


564 M. Werneck, and M. Werneck, et al. 2005. "Análise Florística de Compartimento Arbóreo de Áreas


565 de Floresta Atlântica Sensu Lato da Região das Bacias do Leste (Bahia, Minas Gerais, Espírito


566 Santo e Rio de Janeiro)." _Rodriguésia_ 56: 185–2 35.


&#91;bioRxiv preprint doi: https://doi.org/10.64898/2026.11.01.763487; this version posted November 3, 2026. The copyright holder for this preprint&#93;(https://doi.org/10.64898/2026.11.01.763487)

(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
&#91;available under a CC-BY-NC-ND 4.0 International license.&#93;(http://creativecommons.org/licenses/by-nc-nd/4.0/)


567 Pitman, N. C. A., J. W. Terborgh, M. R. Silman, P. V. Núñez, D. A. Neill, C. E. Cerón, W. A.


568 Palacios, and M. Aulestia. 2001. "Dominance and Distribution of Tree Species in Upper


569 Amazonian Terra Firme Forests." _Ecology_ 82(8): 2101–2117. https://doi.org/10.1890/0012

570 9658(2001)082&#91;2101:DADOTS&#93;2.0.CO;2


571 Ramos, F. N., S. R. Mortara, N. Monalisa-Francisco, J. P. C. Elias, L. Menini Neto, L. Freitas, and


572 R. Kersten, et al. 2019. "ATLANTIC EPIPHYTES: A Data Set of Vascular and Non-Vascular


573 Epiphyte Plants and Lichens from the Atlantic Forest." _Ecology_ 100(2): e02541.


574 https://doi.org/10.1002/ecy.2541


575 R CORE TEAM. 2025. R: A Language and Environment for Statistical Computing. R Foundation


576 for Statistical Computing, Vienna, Austria. Available at: https://www. Rproject.org/.


577 Serviço Florestal Brasileiro. 2019. _Florestas do Brasil em Resumo 2019_ . Brasília, DF: Ministério


578 da Agricultura, Pecuária e Abastecimento, Serviço Florestal Brasileiro.


579 Silva, J. V. F., A. D. T. Lopes, and H. C. Oliveira. 2024. "Briófitas do Município de Corrente, Piauí,


580 Brasil." _Paubrasilia_ 7: e144. https://doi.org/10.33447/paubrasilia.2024.e0144


581 Silva, J. V. F., H. C. Oliveira, P. M. Villa, and E. B. Valente. 2025. "Brioflora Epíxila em


582 Fragmentos de Floresta Atlântica da Serra da Ibiapaba, Ceará, Brasil." _Scientia Plena_ 21(6):


583 061201. https://doi.org/10.14808/sci.plena.2025.061201


584 Silva, J. M. C. Leal, I. R and Tabarelli, M. 2017. _Caatinga: the largest tropical dry forest region_


585 _in South America._ Miami: Springer.


586 Ter Steege, H., N. Pitman, D. Sabatier, H. Castellanos, P. Van Der Hout, D. C. Daly, and M.


587 Silveira, et al. 2003. "A Spatial Model of Tree Alpha-Diversity and Tree Density for the Amazon."


588 _Biodiversity and Conservation_ 12: 2255–2277


589 Travers, S. E., and K. L. Dunnell. 2009. "First-Flowering Dates of Plants in the Northern Great


590 Plains." _Ecology_ 90(8): 2332. _Ecological Archives_ E090-162. https://doi.org/10.1890/08-2135.1


591 Walter, B. M. T., and A. C. Sevilha. 2019. "A Agonia de um Bioma." _Revista Darcy_ 21: 3–66.


592 Welch, W. H. 1948. "Mosses and Their Uses." _Proceedings of the Indiana Academy of Science_ 58:


593 3–46.


594


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&#91;available under a CC-BY-NC-ND 4.0 International license.&#93;(http://creativecommons.org/licenses/by-nc-nd/4.0/)


595 **Table** **1.** Description of the fields contained in the file


596 Brazilian_Bryophytes_Occurrences.csv. The dataset comprises 102,527


597 occurrence records of bryophytes from Brazil compiled from multiple biodiversity


598 databases, herbarium collections, and published sources.




























|Variable|Description|Levels/Units|Example|
|---|---|---|---|
|Source|Source of occurrence&lt;br&gt;record|4 data sources|GBIF|
|Record_Id|Record identification&lt;br&gt;in each data set.&lt;br&gt;Authors were free to&lt;br&gt;use their own&lt;br&gt;identification|The data set&lt;br&gt;comprises&lt;br&gt;102,527 record&lt;br&gt;IDs.|BRYO_000001|
|Phylum|Bryophyte phylum|3 phyla|Bryophyta|
|Family|Taxonomic family|131 families|Fissidentaceae|
|Species|Accepted scientific&lt;br&gt;name|1,835 species|Fissidens&lt;br&gt;crispus|
|Locality|Locality description&lt;br&gt;provided by the&lt;br&gt;source|23,272 localities|Recife|
|Latitude|Geographic latitude&lt;br&gt;(WGS84)|Decimal degrees|&lt;br&gt;-8.0522|
|Longitude|Geographic longitude&lt;br&gt;(WGS84)|Decimal degrees|&lt;br&gt;-34.9286|
|day|Day of collection|1–31|15|
|month|Month of collection|1–12|7|
|Year|Year of collection|184 unique&lt;br&gt;years|2018|
|catalogNumber|Catalog number&lt;br&gt;assigned by the&lt;br&gt;collection|90,318 catalog&lt;br&gt;numbers|PEUFR12345|
|recordNumber|Collector record|30,151 record|234|


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(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
&#91;available under a CC-BY-NC-ND 4.0 International license.&#93;(http://creativecommons.org/licenses/by-nc-nd/4.0/)


















|Col1|number|numbers|Col4|
|---|---|---|---|
|identifiedBy|Person responsible&lt;br&gt;for taxonomic&lt;br&gt;identification|3,170 identifiers|O. Yano|
|recordedBy|Collector(s) of the&lt;br&gt;specimen|5,867 collectors|J. P. Frahm|
|institutionCode|Institution acronym|178 institutions|PEUFR|
|collectionCode|Collection acronym|154 collections|Bryophytes|
|rightsHolder|Institution or person&lt;br&gt;holding data rights|136 rights&lt;br&gt;holders|Universidade&lt;br&gt;Federal de&lt;br&gt;Pernambuco|
|state|Brazilian state of&lt;br&gt;occurrence|27 states|Pernambuco|
|region|Brazilian geographic&lt;br&gt;region|5 regions|Northeast|
|biome|Brazilian biome|7 biomes|Atlantic Forest|



599


600


601


602


603


604


605


606


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607


608 Figure 1. Distribution of bryophyte occurrence records (Anthocerotophyta, Bryophyta, and


609 Marchantiophyta) across Brazilian biomes.


610


611


612


613


614


615


616


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(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
&#91;available under a CC-BY-NC-ND 4.0 International license.&#93;(http://creativecommons.org/licenses/by-nc-nd/4.0/)


617


618


619 Figure 2. Spatial distribution of moss (Bryophyta) occurrence records across the Brazilian


620 biomes.


621


622


623


624


625


626


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(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
&#91;available under a CC-BY-NC-ND 4.0 International license.&#93;(http://creativecommons.org/licenses/by-nc-nd/4.0/)


627


628 Figure 3. Spatial distribution of liverwort (Marchantiophyta) occurrence records across the


629 Brazilian biomes.


630


631


632


633


634


635


636


637


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(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
&#91;available under a CC-BY-NC-ND 4.0 International license.&#93;(http://creativecommons.org/licenses/by-nc-nd/4.0/)


638


639


640 Figure 4. Spatial distribution of hornwort (Anthocerotophyta) occurrence records across the


641 Brazilian biomes.


642


643


644


645


646


647


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(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
&#91;available under a CC-BY-NC-ND 4.0 International license.&#93;(http://creativecommons.org/licenses/by-nc-nd/4.0/)


648


649


650 Figure 5. Temporal distribution of bryophyte occurrence records in Brazil from 1800 to 2025.


651


652


653 Figure 6. Temporal variation in bryophyte species richness in Brazil from 1800 to 2025.


654


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(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
&#91;available under a CC-BY-NC-ND 4.0 International license.&#93;(http://creativecommons.org/licenses/by-nc-nd/4.0/)


655


656 Figure 7. Temporal distribution of bryophyte occurrence records across Brazilian biomes from


657 1800 to 2025.


658


659 Figure 8. Temporal variation in bryophyte species richness across Brazilian biomes from 1800 to


660 2025.


661


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662


663 Figure 9. Number of occurrence records for the 15 most representative moss families, with


664 species richness indicated in parentheses.


665


666 Figure 10. Number of occurrence records for the 15 most representative moss genera, with


667 species richness indicated in parentheses.


668


669


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670


671 Figure 11. Number of occurrence records for the 15 most representative liverwort families, with


672 species richness indicated in parentheses.


673


674 Figure 12. Number of occurrence records for the 15 most representative liverwort genera, with


675 species richness indicated in parentheses.


676


677


