An Acad Bras Cienc (2024) 96(2): e20230007 DOI 10.

1590/0001-3765202420230007
Anais da Academia Brasileira de Ciências | Annals of the Brazilian Academy of Sciences
Printed ISSN 0001-3765 I Online ISSN 1678-2690
[Link]/aabc | [Link]/aabcjournal

ECOSYSTEMS

Distribution pattern in the rupiculous

genus Orthophytum (Bromelioideae/
Bromeliaceae) reveals high microendemicity
in different types of rocky outcrops

SWAMI L. COSTA, RAFAEL B. LOUZADA, SILMARA CECÍLIA NEPOMUCENO, JOILSON

V. ALVES & MARIA TERESA BURIL

Abstract: This study aimed to recognize the biogeographic patterns, richness, and
diversity levels of the Brazilian endemic genus Orthophytum and identify their biotic
components through a parsimony analysis of endemicity (PAE), to better understand
the evolutionary history of this group and develop strategies for the conservation of its
species. We prepared a database for the 54 currently known species of Orthophytum,
including their geographical locations as obtained from digital databases of the
principal herbaria of Brazil, Europe, and the USA. A parsimony analysis of endemicity
(PAE) was used to delimit the areas of endemism based on two grids’ sizes (1º x 1º and
2º × 2º). The majority rule consensus tree resulting from the PAE indicated three areas
of endemism with high bootstrap, diversity, and richness indices: the northern portion
of the Espinhaço Range, the southern portion of the Espinhaço Range, and the central
portion of the Atlantic Forest. The recognition of those distribution patterns reveals a
high number of microendemic species, which is discussed here.
Key words: Bromelioideae, Areas of Endemism, Rocky outcrops, Microendemism, PAE,
Poales.

INTRODUCTION of plants observed today (Wiens & Donoghue

2004) – with environmental factors affecting
The patterns and processes of biotic distribution,
their spatial distributions and consequently
also known as biogeographic patterns (Morrone
their richness and species compositions. It
2009), play important roles in understanding the
is therefore necessary to analyze multiple
evolution of biodiversity, and represent essential
variables in biogeographic studies, such as water
elements for the reconstruction of the Earth’s
availability, elevation, temperature, and edaphic
geological history. Those patterns are related
features (Pausas & Austin 2001, Austin 2013), as
to the organization and distribution of groups
those climatic or physical features can function
of organisms in geographic space, which can
as barriers that prevent certain organisms from
be approached in many different ways as well
colonizing certain habitats (Wiens & Donoghue
as at different scales. Important historical and
2004, Wiens 2011).
biogeographic factors are believed to influence
One of the tools used in historical
those patterns, such as dispersal, vicariance, and
biogeography to investigate the natural
extinction (Morrone 2009). Additionally, diverse
distribution patterns of organisms is Parsimony
ecological processes may also play relevant
Analysis of Endemicity (PAE) (e.g., Silva & Oren
roles in shaping the geographic distributions

An Acad Bras Cienc (2024) 96(2)

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

1996, Posadas & Miranda-Esquivel 1999, Garcia- and terrestrial life forms that occupy a wide
Barros et al. 2002, Manrique et al. 2003, Rovito et variety of habitats from sea level up to 4000
al. 2004, Sigrist & Carvalho 2008). PAE basically meters above sea level (masl) (Benzing 2000,
consists of classifying areas of endemism Butcher & Gouda, Last Access May 2023).
and constructing cladograms based on the One of the first divergent lineages of
parsimonious cladistic analysis of presence- the Bromelioideae (the second largest clade
absence matrices of species. An area of of the family) to experience considerable
endemism is defined when the distributions of diversification in (what is today) Brazil was the
different endemic species converge to the same genus Orthophytum Beer, which is saxicolous
region (Morrone 1994, Szumik et al. 2002). The (rarely terrestrial) and endemic to the eastern
identification of such areas provide important region of that country. Orthophytum comprises
information for the field of biogeography and 54 species (Louzada 2020) that inhabit different
for studies of biodiversity conservation, and can types of rocky outcrops: granitic inselbergs in
contribute to effective conservation planning. the Atlantic Forest and Caatinga domain and
Congruent occurrence patterns can easily be rocky quartzite outcrops in Campos Rupestres
recognized where endemic species (those found (rocky altitudinal fields) along the Espinhaço
only in an exclusive region) are frequent, and can Range (Louzada & Wanderley 2010). Martinelli et
be delimited to propose areas of endemism and al. (2008) indicated a possible center diversity
then analyze their inter-relationships (Szumik et of the genus in the Atlantic Forest, where its
al. 2002, Morrone 1994). species usually inhabit inselbergs and rock
The Neotropical biogeographical region outcrops and are often restricted to small
(which stretches from Mexico to southern South geographic areas (with many microendemic
America) comprises tremendous taxonomic and taxa). Those environments, which are spatially
habitat diversity as well as a complex geological and ecologically isolated, exhibit barriers to
history (Morrone 2009). Most studies of dispersal and migration, and evidence high
distributional patterns and areas of endemism levels of species diversity and endemism
in this region have been based on relatively well (Echternacht et al. 2011).
studied animal groups (see Cracraft 1985, Silva Studies of bromeliad species adapted
1995, Amorim & Pires 1996, Ron 2000, Costa et to neotropical rock outcrops have already
al. 2000, Silva & Oren 1996) or on tree species improved the understanding of speciation
(see de Lima et al. 2020, Françoso et al. 2016, and the processes of endemicity in naturally
ter Steege et al. 2013, 2015). Despite the crucial isolated environments (Barbará et al. 2009,
importance of non-tree plants for understanding Palma-Silva et al. 2011). However, despite the
biodiversity and ecosystem functioning, there immense importance of non-arboreal plants
are still large gaps in our knowledge of their to understanding biodiversity, ecosystem
distributions in the Neotropics. One of the most functioning, and macroevolutionary and
species-rich and ecologically important non-tree macroecological processes, there are still
monocotyledonous families in the Neotropics – large gaps in the knowledge of how they are
Bromeliaceae Juss. (Smith & Downs 1974). The distributed in the Neotropics (Engemann et al.
family comprises 83 genera and approximately 2015).
3.744 species (Butcher & Gouda 2023, Last Access In order to better understand the processes
May 2023), and includes epiphytic, saxicolous, that led to the considerable diversification

An Acad Bras Cienc (2024) 96(2) e20230007 2 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

of Orthophytum, it will be necessary to use which include specimen records from the
biogeographic reconstruction analysis and principal herbaria of Brazil, Europe, and the
formulate hypotheses concerning the ancestral USA. Only specimens identified by specialists
area of the genus, when it emerged, how its in the genus were used. The species included
dispersal occurred, and how some species in the analysis followed the taxonomic revision
became fixed in restricted regions. Biogeography of Louzada & Wanderley (2010) and the Species
is used to document and understand the spatial List of Brazilian Flora (Louzada 2020); the latter
patterns of biodiversity, species interactions includes a new species described after the last
and their organization, as well as spatial revision. Records without original geographic
processes (Brown & Lomolino 1998, Troppmair coordinates were assigned to the location or
2002). Therefore, in addition to supporting an municipality recorded on the specimen label.
understanding of the evolution of different The geographic coordinates of the sample
plant groups, biogeography can help reduce location or municipality were obtained from
biodiversity losses by indicating potential areas the GeoNames website ([Link]
for species preservation (Goldani 2012). com) or by using the geoLoc tool (available
We tested here the hypothesis that the from the Environmental Information Reference
Espinhaço Mountain Range and the Atlantic Center website [SpeciesLink - [Link]
Forest in Brazil are areas of endemism of [Link]/geoloc]). Records that could not be
Orthophytum, and that microendemism is the georeferenced to at least the municipality level,
predominant pattern in the genus. To test that vouchers with dubious identifications or without
hypothesis, we sought to identify distribution collector number, as well as duplicates, were
patterns, richness, diversity, and use parsimony excluded from the dataset. After data cleaning,
analysis of endemicity (PAE) to identify areas of the dataset comprised 695 records of 54 species.
endemism and their biotic components. The nomenclature of the species follows The
The identification of areas of endemism and International Plant Names Index ([Link]
testing them as evolutionary geographical units [Link]/), Tropicos ([Link]
are the first steps towards the development home), and New Bromeliad Taxon List digital
of a cladistic biogeography. At the same time, databases (Butcher & Gouda continuously
biogeography can improve our knowledge updated, last accessed in Dec 2022).
of the evolutionary history of this group and
Grid size
help future researchers develop conservation
strategies for its species. The choice of grid cells size is decisive and
important for biogeographic analysis. Several
authors discussed how distinct grid sizes can
MATERIALS AND METHODS affect the identification of the areas of endemism
Taxa and Data collection (see Morrone & Escalante 2002, Morrone 1994,
The geographic coordinates of Orthophytum Szumik et al. 2002, Casagranda et al. 2009, Navarro
specimens were obtained from the Global et al. 2009). Usually, the use of small grid cells
Biodiversity Information Facility ([Link] results in discontinuous distributions, in a more
[Link]/), CRIA-speciesLink ([Link] detailed resolution (where only small areas
[Link]/) and Reflora Virtual Herbarium (http:// of endemism are identified) and can produce
[Link]/) digital databases, poorly resolved area cladograms. Meanwhile,

An Acad Bras Cienc (2024) 96(2) e20230007 3 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

large grid cells generally detected larger areas Endemicity analyses

of endemism, which increases the congruence Morrone (1994) proposed PAE as a tool to
of species (many species appearing as endemic) identify endemic areas as primary biogeographic
and could camouflage disjunctions (Morrone & homology, using grid cells as operational units
Escalante 2002). Therefore, employing different (without inferring relationships or hierarchies
grid sizes is the most suitable way to establish between areas) based on their common species,
the results to recognize endemic areas and to defining an endemism area as a group of grid
be able to identify different patterns if some cells of at least two taxa endemic to the region.
taxonomic groups display congruence at We draw the grid-cells on a map of Brazil,
different scales (Aagesen et al. 2009, Casagranda including grids only where at least one locality
et al. 2009, Navarro et al. 2009). of one species was recorded. Visualizations
For all the analyses presented here (richness, of the species were also performed using the
diversity and PAE), we choose two sizes of grid QGIS software (QGIS Development Team 2021).
cells, 1° x 1° (ca. 100 km x 100 km) and 2° x 2° (ca. Based on the occurrence of the species, a
200 km x 200 km). The grid cells were produced presence/absence matrix (Tables III and IV)
using DIVA-GIS software (Hijmans 2001). was constructed using the software Mesquite,
version 3.16. (Maddison & Maddison 2021), where
Richness and diversity analysis and distribu-
tion patterns the presence of a given taxon in each sampled
area was coded as “1”, and the absence of that
Richness and Diversity (Shannon Index;
taxon was coded as “0”. A hypothetical area
Magurran 1988) analyses were performed using
coded as “0” was added to the matrix as an
DIVA-GIS software. The visualization of the
outgroup to root the tree.
species distribution, the maps construction
The PAE was conducted using PAUP* 4.0
and the visualization of the main conservation
software (Swofford 2003) to infer the centers
reserves was performed using QGIS software
of endemism and to reconstruct the most
(QGIS Development Team 2021).
parsimonious tree, using a heuristic algorithm
Distribution patterns were assessed based
with 1000 replications from random additions
on the plotted dots on the maps, following
and TBR (tree-bisection-reconnection) branch-
Menini-Neto & Forzza (2013), and were coded as
swapping, saving two trees per replication;
follows: (1) widespread distribution (WD), when
the same commands were used for bootstrap
encountered in more than five grid squares; (2)
analysis.
intermediate distribution (ID), when found in two
Here, for more robust results, we define
to four grid squares; (3) restricted distribution
an endemism area when the clades in the
(RD), when found in only one grid square, but
consensus area cladograms received bootstrap
with more than five known sites of occurrence;
support ≥ 50% and shared three or more species.
and (4) microendemic distribution (MED),
The data matrix is available upon request to the
when restricted and exclusive to a vegetation
corresponding author.
physiognomy and found in only one grid square
and only one to four known sites of occurrence.

An Acad Bras Cienc (2024) 96(2) e20230007 4 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

RESULTS coincides with the areas in the larger grid size

Patterns of geographic distribution, richness with the A20 and A26 areas with the greatest
and diversity richness (comprising together 28 species, with
We obtained a database with 692 records 14 exclusives). These areas encompassing the
distributed among 54 species of Orthophytum central portion of the Espinhaço Range (in
using the parameters and methods described the north of Minas Gerais, grid A52 in smaller
above. The distribution maps for all grid size and A20 in the larger one), and the
Orthophytum species are presented in Figures inselbergs of Espírito Santo and its boundary
4 – 10. Its species are endemic to eastern Brazil with Minas Gerais (grid A67 in smaller grid size
with the following limits: north, the municipality and A26 in the larger one). Even the grids with
of Alcântaras, Ceara State (40°30’W, 03°34’S), low to medium richness still included exclusive
where O. cearense Leme & [Link] occurs; species, such as grids A17 (with one exclusive
south, the municipality of Domingos Martins, species), A31 (1 sp.), A38 (2 spp.), A43 (2 spp.),
Espírito Santo State (40°36’W, 20°21’S), where O. A54 (2 spp.), A58 (1 sp.), A63 (1 sp.), A66 (4 spp.)
foliosum [Link]. occurs; east, the municipality and A69 (2 spp.) in the smaller grid size and A10
of Jaqueira, Pernambuco State (35°30’W, (2 spp.), A13 (1 sp.), A17 (2 spp.), A18 (2 spp.), A21
08°25’S), where O. disjunctum [Link]. occurs; (2 spp.), A23 (1 sp.) and A24 (3 spp.) in the larger
and west, the municipality of Água Quente, grid size. Our results therefore indicated two
Bahia State (44°18’W, 13°10’S), where O. harleyi main centers of Orthophytum species richness:
Leme & [Link] has been recorded. The the central portion of Espinhaço Range (CPER)
species with the greatest numbers of records and the central portion of the Atlantic Forest
were O. disjunctum, O. maracasense [Link]., (CPAF).
and O. foliosum with 175, 86 and 70 records The diversity analysis (Fig. 2) as the richness
respectively. Four categories were established analysis also produced 70 grids in 1° x 1° size
to represent the distribution patterns of these and 26 grids in 2° x 2° size. The regions with
taxa: widespread, intermediate, restricted, the greatest diversity are represented by grid
and microendemic. Seven species evidenced squares A52, A58, A59 and A67 (with Shannon
widespread distribution patterns, 19 species index H’ between 1.58 − 2.0) in the smaller grid
have intermediated distributions, four species size, encompassing the north and northeast
have restricted distributions, and 24 species of Minas Gerais and the inselbergs of Espirito
evidenced a microendemic distribution pattern. Santo. The larger grid size has the A17, A18,
The distribution patterns, number of grids, A20, A21 and A26 areas with high diversity (H’
and the phytogeographical domain in which between 1.77 − 2.19), encompassing the central
they occurred are presented in Table I for each portion of Espinhaço Range (in the north and
species. northeast of Minas Gerais and south of Bahia)
The richness analysis found 70 grids in and the inselbergs of Espirito Santo and its
1° x 1° size and 26 grids in 2° x 2° size (Fig. boundaries with Minas Gerais. In summary,
1). The regions with the greatest richness are the main difference between the two analyses
represented by the grid squares A52 and A67 is found in areas A17 and A18 (in larger grid
in the smaller grid size (comprising together 18 size) – the central portion of Espinhaço Range
species, with 7 exclusives to those areas), which in Bahia – which presents a relatively reduced

An Acad Bras Cienc (2024) 96(2) e20230007 5 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

Table I. Distribution patterns of Orthophytum species with the associated vegetation and number of grids (in
2-degree grid size) that each species are present. CA: Caatinga; CE: Cerrado; AF: Atlantic Forest; MED: Microendemic
Distribution; ID: Intermediated Distribution; RD: Restricted Distribution; WD: Widely Distribution.

Number of Phytogeographical Distribution

Species
grids Domains Pattern
Orthophytum alagoanum Leme & [Link] 1 CA MED
Orthophytum alvimii W. Weber 2 CA, AF ID
Orthophytum arcanum Leme 1 AF MED
Orthophytum argentum Louzada & Wand. 1 CA MED
Orthophytum atalaiense [Link] & Leme 1 AF MED
Orthophytum boudetianum Leme & [Link] 1 AF RD
Orthophytum braunii Leme 3 CA ID
Orthophytum buranhense Leme & [Link] 1 AF MED
Orthophytum cearense Leme & [Link] 3 CA, AF ID
Orthophytum conquistense Leme & [Link] 2 AF ID
Orthophytum cristaliense Leme 1 CE MED
Orthophytum diamantinense Leme 2 CE ID
Orthophytum disjunctum [Link]. 9 CA, AF WD
Orthophytum duartei [Link]. 2 AF ID
Orthophytum eddie-estevesii Leme 1 CA MED
Orthophytum elegans Leme 2 AF ID
Orthophytum erigens Leme 1 CA MED
Orthophytum estevesii (Rauh) Leme 1 AF MED
Orthophytum falconii Leme 4 CA, AF ID
Orthophytum foliosum [Link]. 5 CE, AF WD
Orthophytum fosterianum [Link]. 1 AF MED
Orthophytum glabrum (Mez) Mez 2 CE, AF ID
Orthophytum graomogolense Leme & [Link] 2 CE, AF ID
Orthophytum grossiorum Leme & [Link] 1 AF MED
Orthophytum guaratingense Leme & [Link] 1 AF MED
Orthophytum gurkenii Hutchison 2 AF ID
Orthophytum harleyi Leme & Machado 3 CA ID
Orthophytum horridum Leme 5 CA, AF WD
Orthophytum jabrense [Link] & [Link] 2 CA, AF ID
Orthophytum jacaraciense Leme 1 CA MED
Orthophytum lanuginosum Leme & [Link] 1 AF MED
Orthophytum lemei [Link] & [Link] 8 CA, AF WD
Orthophytum leprosum (Mez) Mez 5 CA, CE, AF WD
Orthophytum macroflorum Leme & [Link] 1 CA MED
Orthophytum magalhaesii [Link]. 3 AF ID
Orthophytum maracasense [Link]. 7 CA, AF WD
Orthophytum mello-barretoi [Link]. 3 CE, AF ID
Orthophytum minimum Leme & [Link] 1 CE MED
Orthophytum piranianum Leme & [Link] 2 CE ID
Orthophytum pseudostoloniferum Leme & [Link] 1 AF MED
Orthophytum pseudovagans Leme & [Link] 1 AF RD
Orthophytum riocontense Leme 3 CA, AF ID
Orthophytum roseolilacinum Leme 1 AF MED
Orthophytum santaritense Leme, [Link] & Zizka 1 AF MED
Orthophytum santosianum Leme 2 CA, CE ID
Orthophytum saxicola Ule 7 CA, AF WD
Orthophytum schulzianum Leme & [Link] 1 CE, AF RD
Orthophytum sp. nov. 1 CA MED
Orthophytum striatifolium Leme & [Link] 1 AF MED
Orthophytum sucrei [Link] 1 AF MED
Orthophytum toscanoi Leme 3 CA, CE ID
Orthophytum triunfense [Link] & Leme 1 CA RD
Orthophytum vasconcelosianum Leme 1 AF MED
Orthophytum zanonii Leme 1 AF MED

An Acad Bras Cienc (2024) 96(2) e20230007 6 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

Figure 1. Richness map

showing the analyses in
1-degree (a) and 2-degree
(b) cells. PI: Piauí; CE:
Ceará; PB: Paraíba; PE:
Pernambuco; AL: Alagoas;
SE: Sergipe; ES: Espírito
Santo; BA: Bahia; MG:
Minas Gerais.

richness (nine and ten species respectively) but according to PAE quadrat size, 70 grid-cells in 1°
high diversity (H’ = 1.77 – 2.19). Therefore, our x 1° and 26 in 2° x 2°, which both represent the
results indicate two main centers of diversity of entire distribution of the genus in Brazil.
Orthophytum: the Central Portion of Espinhaço For the 1-degree quadrat matrix, PAE
Range (CPER) and the Central Portion of Atlantic produced 5000 equally parsimonious trees, with
Forest (CPAF). CI (consistency index) = 0.79 and RI (retention
index) = 0.62, based on 28 parsimony-informative
Parsimony analysis of endemicity
characters. The strict consensus tree (Fig. 3)
As the same as the above analyzes, the total revealed a basal polytomy for most areas, but
number of occupied areas (grid-cells) varied

An Acad Bras Cienc (2024) 96(2) e20230007 7 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

Figure 2. Diversity map

showing the analyses in
1-degree (a) and 2-degree
(b) cells. PI: Piauí; CE:
Ceará; PB: Paraíba; PE:
Pernambuco; AL: Alagoas;
SE: Sergipe; ES: Espírito
Santo; BA: Bahia; MG:
Minas Gerais.

15 remaining quadrats were grouped into six effectively be observed in this analysis and
minor clades – AFMG (Atlantic Forest of Minas considered areas of endemism.
Gerais), AFES (Atlantic Forest of Espírito Santo), For the 2-degree quadrat matrix, PAE
NBA (North of Bahia), CD (Chapada Diamantina), produced 1416 equally parsimonious trees, with
GM (Grão Mogol) and CT (Caetité) and in three CI = 0.71 and RI = 0.63, based on 24 parsimony-
major ones – CPAF (Central Portion of Atlantic informative characters. The strict consensus
Forest), NER (North of Espinhaço Range) and tree (Fig. 3) also revealed a basal polytomy for
CPER (Central Portion of Espinhaço Range). 16 quadrats, but ten remaining quadrats were
Thus, three major clades with bootstrap support grouped into three major clades – CPAF, NER
≥ 50% and three or more species shared can and SER (South of Espinhaço Range), considered
areas of endemism.

An Acad Bras Cienc (2024) 96(2) e20230007 8 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

Figure 3. The majority rule

consensus tree obtained with
the 1-degree (a) and 2-degree
(b) size cells analyses, showing
the major and minor areas
of endemism determined by
PAE. Numbers under the lines
indicate bootstrap proportions.
AFMG: Atlantic Forest of Minas
Gerais; AFES: Atlantic Forest of
Espírito Santo; NBA: North of
Bahia; CD: Chapada Diamantina;
GM: Grão Mogol; CT: Caetité;
CPAF: Central Portion of Atlantic
Forest; NER: North of Espinhaço
Range; CPER: Central Portion of
Espinhaço Range; SER: South
of Espinhaço Range; PI: Piauí;
CE: Ceará; PB: Paraíba; PE:
Pernambuco; AL: Alagoas; SE:
Sergipe; ES: Espírito Santo; BA:
Bahia; MG: Minas Gerais.

The results in both analyzes were similar, the Range than the CPER that covers the central
1-degree result shows a big basal polytomy, but region of Espinhaço Range, in the boundaries of
recovered small clades that can be considered Bahia and Minas Gerais states.
small endemism areas. The 2-degree result also Therefore, as there were not so many
shows a basal polytomy, but for being bigger significant differences between both analysis
quadrats, result in larger endemism areas. results, here, we will consider for purposes
The areas NER and CPAF in both degrees are of description and discussion, the areas of
equivalent and the mainly difference between endemism recovered from the 2-degree analysis:
results analyses is in the areas CPER (1-degree) NER, SER and CPAF.
and SER (2-degree), southernmost areas of The first area considered is the northern
Espinhaço Range. The SER of 1-degree analysis region of the Espinhaço Range in Bahia State,
covered areas even further south of Espinhaço formed by the grid groups A8, A13, A14, A17 and

An Acad Bras Cienc (2024) 96(2) e20230007 9 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

A18, (with bootstrap = 62.5%) with five exclusive In this study, smaller grid sizes result in a
species. The second area is the South of finer resolution of distributional patterns and
Espinhaço Range in Minas Gerais, formed by the smaller individual areas of endemism but show
grid group A20, A23 and A25 (bootstrap = 52.8%), a big basal polytomy (Fig. 3). Linder (2001) argues
also with five exclusive species. The last area, that the disadvantage of smaller grid sizes is that
with the highest number of exclusive species they would lead to an increase in the number
(13), is the Central Portion of the Atlantic Forest of false absences. Nelson et al. (1990) showed
located in Espirito Santo and on the boundary that the detection of centers of endemic plant
with Minas Gerais, formed by the grid group A24 species in the Brazilian Amazon may be flawed
and A26 (bootstrap = 96.4%). due to collection errors. However, this distorting
effect could be avoided if was used large grids
sizes. In our case, the most of major patterns
DISCUSSION are recovered in both areas’ cladogram, and the
As previously mentioned, many authors define difference between both analysis results was
an area of endemism when the distribution of not so significant (see in the Results section),
two or more species converge to the same region so we choose to use the 2-degree results for
and do not occur anywhere else (Morrone 1994, purposes of description and discussion, and
Garcia-Barros et al. 2002, Platnick 1991). Such we believe that appears to be an adequate
a region will evidence a higher-than-expected resolution to be explored in future research of
endemism when compared to adjacent regions secondary biogeographical homologies (Costa
(Crisp et al. 2001, Laffan & Crisp 2003). Due to S.L. et al., unpublished data).
the high number of microendemic species and Thus, the present study identified and
species known for only one location, we decided defined three large areas of endemism of
to define here an endemism area when the clades Orthophytum: NER, SER, and CPAF (Fig. 3), with
resulted in the consensus area cladograms from existing legally constituted conservation areas
PAE analysis received bootstrap support ≥ 50% within them; the main conservation reserves in
and shared three or more species. each area are listed in Table II.
The identification of areas of endemism The NER and SER together cover almost the
is important for reconstructing the historic entire Espinhaço Range, except for the region of
and ecological biogeography of taxa, as well Oliveira dos Brejinhos (corresponding to areas
as for biodiversity conservation (Szumik 2002, A12 in central Bahia – which had good support
Echternacht et al. 2011). This is because the in together with A16, however have only one
delimitation of areas of endemism allows the record and therefore were not identified as
identification of priority areas for conservation areas of endemism for Orthophytum). The NER
that harbor unique concentrations of is located in the Bahia portion of the Espinhaço
biodiversity (Myers et al. 2000, Williams et al. Range, while the SER is in Minas Gerais portion
2002) and represent independent geographic of the Espinhaço Range. Both areas evidence
units. The analysis of the relationships between high richness (36 and 22 species respectively),
those areas will generate information about medium endemism (five exclusive species each),
the processes responsible for their formation and medium to high diversity (NER: 0.89 − 2.19;
(Anderson 1994, Morrone 1994, Laffan & Crisp SER: 0.88 − 2.19).
2003).

An Acad Bras Cienc (2024) 96(2) e20230007 10 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

Table II. Main Conservation Units created within the areas of endemism identified by PAE of 2-degrees grid size
and the grid/area in which each one occurs. Numbers in parentheses indicate the total number of conservation
units within the grid/area. Bold indicates that there are Orthophytum species within these conservation units.
RPPN: Reserva Particular do Patrimônio Natural; APA: Área de Proteção Ambiental; RVS: Refúgio de Vida Silvestre;
PN: Parque Nacional; PE: Parque Estadual; RDS: Reserva de Desenvolvimento Sustentável; FN: Floresta Nacional;
ARIE: Área de Relevante Interesse Ecológico; MN: Monumento Natural; RB: Reserva Biológica; ESEC: Estação
Ecológica.

Endemism
Area Main Conservation Units
Areas
Sustainable Use Units Integral Protection Units

RPPN Maria Maria, APA Lago de

A8 (6) Sobradinho, RPPN Toca dos Ossos, APA RVS Riacho Pontal, PN Boqueirão da onça;
Boqueirão da Onça;

ARIE Serra do Orobó, APA Marimbus, APA PN Chapada Diamantina, PE Morro do

A13 (22) Grutas dos Brejões, RPPN Ametista, RPPN Chapéu, PE Sete Passagens, MN Cachoeira do
Pau d’Arco; Ferro Doido;

APA Baía de Todos os Santos; RPPN

North of A14 (14) Curió, RPPN Olho de Fogo, APA Plataforma MN Cânions do Subaé;
Espinhaço Continental do Litoral Norte;
Range
PE Serra dos Montes Altos, PE Verde Grande,
PE Caminho dos Gerais, RVS Serra dos
A17 (7) APA Lajedão;
Montes Altos, PE Mata Seca, PE Lagoa do
Cajueiro;

ARIE Nascente do Rio de Contas, FN

Contendas do Sincorá, APA Serra do PN Boa Nova, PN Chapada Diamantina, RVS
A18 (17)
Barbado, APA da Serra do Ouro, APA Serra Boa Nova;
do Barbado, RPPN Arco Verde;

RB Jaíba, PE Caminho dos Gerais, PE

RDS Nascentes Geraizeiras, APA Serra do Serra Nova e Talhado, PE Lapa Grande, PE
A20 (10)
Sabonetal; Botumirim, PE Grão Mogol, RB Serra Azul, PE
Montezuma;

PE de Botumirim, PE Serra da Candonga, PE

South of APA Águas Vertentes, APA do Alto do Serra do Intendente, MN Estadual Várzea do
A23 (15)
Espinhaço Mucuri; Lageado, PE Serra Negra, PE Pico do Itambé,
Range PN das Sempre Vivas, ESEC Acauã, PE Biribiri;

APA da Perdição, APA Alto Taboão, APA MN Municipal Morro do Pires, PE do Rio Doce,
Cachoeira das Andorinhas, APA Córrego PE Serra do Sobrado, PE Serra do Brigadeiro,
A25 (60) da Mata, RPPN Mata dos Jaca, APA Carste PE Serra Verde, PE Serra do Intendente, PN
de Lagoa Santa, APA Sul-RMBH, RVS da Serra do Gandarela, PE do Itacolomi, PN
Macaúbas; Serra do Cipó;

APA Alto do Mucuri, RPPN Córrego das

RB Córrego do Veado, RB Sooretama, MN Pico
A24 (8) Traíras, RPPN Prati, RPPN Antônio Lopes
do Ibituruna;
Merson, RPPN Lemke;
Central
Portion of FN Pacotuba, FN Goytacazes, APA
Atlantic MN Pontões Capixabas, PE Mata das Flores,
Corredeiras, RPPN Meu Cantinho, RPPN
Forest A26 PE Sete Salões, PN Caparao, MN Pontões
Bei Cantoni, RPPN Olho d’Água, RPPN Três
(120) Capixabas, MN Pedra do Monjolo, MN o Frade
Pontões, RPPN Boa Fé, RPPN Passos, RPPN
e a Freira;
Freisleben, APA Lagoa Jacuné;

An Acad Bras Cienc (2024) 96(2) e20230007 11 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

Both the NER and SER have a large extension plants. These lowlands were also reported as
of “Campos Rupestres” (rocky altitudinal fields) probable geographic barriers for Harley (1988),
– a landscape composed mainly of grasslands Rando & Pirani (2011), and Alves & Buril (2022)
and quartzite-sandstone rock outcrops above in the dispersal of many mountain species and
900 masl. The Espinhaço Range represents this could help to explain the distribution of the
approximately 1% of the total land area of Brazil, Orthophytum species that occur in SER and NER.
but shelters approximately 10% of the country’s The third area of endemism of our study,
plant diversity (Rapini 2010). Most of those the CPAF (areas A24 and A26), harbors the
species have restricted distributions, and the highest diversity, richness, and endemism, with
floristic compositions of the rocky fields of the 13 exclusive species, H’ = 1.76 - 2.19, and the
Espinhaço Range are marked by rare species and highest support in PAE (96.4%). This area was
high rates of endemism – perhaps the highest previously indicated by Martinelli et al. (2008)
among Brazilian plant formations (Giulietti et al. as a possible center of diversity and endemism.
1987, 1997, Giulietti & Pirani 1988, Rapini 2010). The CPAF is located in northeastern Espírito
The geological origin of the constituent Santo State and Minas Gerais State, which
blocks of the Espinhaço Range is dated to the are covered by the Atlantic Forest domain
Pre-Cambrian period. The soil is characterized and characterized by an evergreen tropical
by being shallow, sandy, acidic, and nutrient- forest vegetation (Oliveira-Filho et al. 2006).
poor, which contributes to a phytophysiographic In addition to forest physiognomies, the area
mosaic of rare species, due to the segmentation holds mangrove swamps, shrubby Restinga
of the vegetation in different and small (sandy coastal) vegetation, and patches of high-
populations between rock outcrops (providing altitude grasslands and rock outcrops (Safford
specific niches to the species) besides living 1999, 2007, Scarano 2002).
in several locations with difficult to access, The Orthophytum species that occur in the
remaining undersampled or practically CPAF are found on inselbergs and on exposed
unexplored (Rapini 2010, Schaefer et al. 2016, rock outcrops in the Atlantic Forest domain.
Silveira et al. 2016), even being an area intensely Several authors observed that naturally open
researched (see Rapini et al. 2008, Borges et al. formations (such as inselbergs and granite rock
2011, Echternacht et al. 2011, Bitencourt & Rapini outcrops) found in high altitude montane areas
2013, Colli-Silva et al. 2019, Alves & Loeuille 2021, of the Atlantic Forest harbor a highly endemic
Alves & Buril 2022, Assunção-Silva & Assis 2022, flora (over 20% of its species) that have strong
Barros-Souza & Borges 2023). floristic connections with other montane areas,
The Orthophytum species of the NER occur such as those of the Andes and the Espinhaço
on high elevation rock outcrops (more than 900 Range (e.g., Safford 1999, 2007, Giulietti & Pirani
masl), while the species of the SER occur on 1988, Calio et al. 2008, Porembski 2007).
lower elevation rock outcrops (less than 900 m). Not only Orthophytum, but also most of
Echternacht et al. (2011) observed that the species the Bromeliaceae family, demonstrate high
that occur in the northern section of Espinhaço diversity and endemism in the Atlantic Forest
Range in Minas Gerais (corresponding to SER of domain. Zizka et al. (2019) identified three
our study) occur in lowlands with contrasting large centers of endemism and diversity of the
climatic and edaphic conditions that could American Bromeliaceae: in the Atlantic Forest
represent barriers to many endemic mountain of southeastern Brazil, in the Central Andes,

An Acad Bras Cienc (2024) 96(2) e20230007 12 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

Table III. Presence/absence matrix (area × species) used in parsimony analysis of endemicity of 1-degree grid size.

Orthophytum pseudostoloniferum

Orthophytum vasconcelosianum
Orthophytum eddie-estevesii

Orthophytum graomogolense
Orthophytum diamantinense

Orthophytum roseolilacinum
Orthophytum mello-barretoi

Orthophytum pseudovagans
Orthophytum guaratingense
Orthophytum boudetianum

Orthophytum conquistense

Orthophytum maracasense
Orthophytum lanuginosum

Orthophytum santosianum
Orthophytum macroflorum

Orthophytum schulzianum
Orthophytum santaritense
Orthophytum jacaraciense

Orthophytum striatifolium
Orthophytum fosterianum

Orthophytum magalhaesii
Orthophytum buranhense

Orthophytum riocontense
Orthophytum piranianum
Orthophytum cristaliense

Orthophytum disjunctum

Orthophytum grossiorum
Orthophytum alagoanum

Orthophytum caraibense
Orthophytum atalaiense

Orthophytum triunfense
Orthophytum argentum

Orthophytum minimum
Orthophytum leprosum
Orthophytum horridum
Orthophytum cearence

Orthophytum foliosum
Orthophytum arcanum

Orthophytum estevesii

Orthophytum jabrense

Orthophytum toscanoi
Orthophytum glabrum

Orthophytum gurkenii

Orthophytum saxicola
Orthophytum elegans
Orthophytum erigens

Orthophytum falconii

Orthophytum zanonii
Orthophytum duartei
Orthophytum braunii

Orthophytum harleyi

Orthophytum [Link].
Orthophytum alvimii

Orthophytum sucrei
Orthophytum lemei
Out 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A2 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A3 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A4 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A5 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A6 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0
A7 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A8 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A9 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A10 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A11 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A12 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A13 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A14 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A15 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A16 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A17 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A18 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A19 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A20 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A21 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0
A22 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0
A23 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A24 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A25 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A26 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A27 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A28 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A29 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A30 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A31 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A32 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0
A33 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0
A34 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0
A35 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0
A36 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A37 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0
A38 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 0 0 1 0 0 1 0 0 0 0 0 0
A39 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0
A40 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A41 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A42 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A43 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A44 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A45 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0
A46 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0
A47 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A48 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0
A49 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0
A50 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A51 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A52 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 1 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0
A53 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 1 0 0 0 0 0 1 0 0 0 1 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A54 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A55 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A56 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0
A57 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

An Acad Bras Cienc (2024) 96(2) e20230007 13 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

Table III. Continuation.

A58 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 1 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0
A59 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 1 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0
A60 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0
A61 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0
A62 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A63 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0
A64 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A65 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A66 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 1 0
A67 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 1 1 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 1
A68 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A69 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0
A70 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Table IV. Presence/absence matrix (area × species) used in parsimony analysis of endemicity of 2-degree grid size.

Orthophytum pseudostoloniferum

Orthophytum vasconcelosianum
Orthophytum eddie-estevesii

Orthophytum graomogolense
Orthophytum diamantinense

Orthophytum roseolilacinum
Orthophytum mello-barretoi

Orthophytum pseudovagans
Orthophytum guaratingense
Orthophytum boudetianum

Orthophytum conquistense

Orthophytum maracasense
Orthophytum lanuginosum

Orthophytum santosianum
Orthophytum macroflorum

Orthophytum schulzianum
Orthophytum santaritense
Orthophytum jacaraciense

Orthophytum striatifolium
Orthophytum fosterianum

Orthophytum magalhaesii
Orthophytum buranhense

Orthophytum riocontense
Orthophytum piranianum
Orthophytum cristaliense

Orthophytum disjunctum

Orthophytum grossiorum
Orthophytum alagoanum

Orthophytum caraibense
Orthophytum atalaiense

Orthophytum triunfense
Orthophytum argentum

Orthophytum minimum
Orthophytum leprosum
Orthophytum horridum
Orthophytum cearence

Orthophytum foliosum
Orthophytum arcanum

Orthophytum estevesii

Orthophytum jabrense

Orthophytum toscanoi
Orthophytum glabrum

Orthophytum gurkenii

Orthophytum saxicola
Orthophytum elegans
Orthophytum erigens

Orthophytum falconii

Orthophytum zanonii
Orthophytum duartei
Orthophytum braunii

Orthophytum harleyi

Orthophytum [Link].
Orthophytum alvimii

Orthophytum sucrei
Orthophytum lemei
Out 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A2 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A3 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A4 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A5 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0
A6 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A7 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A8 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0
A9 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0
A10 1 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A11 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A12 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A13 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0
A14 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0
A15 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0
A16 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A17 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 1 0 1 1 1 0 1 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0
A18 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 0 0 1 1 0 1 0 0 0 0 0 0
A19 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0
A20 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 1 1 0 0 1 1 0 1 1 0 0 0 0 1 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0
A21 0 0 0 0 0 0 0 1 0 0 0 1 0 1 0 0 0 0 0 1 0 0 1 0 0 1 0 0 1 0 0 0 1 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0
A22 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A23 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0
A24 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 1 0 1 0 1 0 0 1 0 1 0 1 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0
A25 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A26 0 0 1 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 1 1 0 0 0 0 1 1 0 0 1 1

An Acad Bras Cienc (2024) 96(2) e20230007 14 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

and in Central America. Those authors also Distinct from the CPAF area, however, both the
observed that the subfamily Bromelioideae NER and SER regions (and the Espinhaço Range
was most species-rich and evidenced high as a whole) have few established conservation
endemism in the eastern Atlantic Forest of areas, and those are not sufficient to protect the
Brazil. The high occurrence of Bromeliaceae, diversity found in the entire complex, requiring
especially Bromelioideae, in the Atlantic Forest more parks and reserves to be created (Silva et
is consistent with the extensive diversification al. 2008, Versieux & Wendt 2007). Additionally, the
of the family in eastern Brazil (Smith 1934, Smith existing conservation areas in those regions lack
& Downs 1974). government support, and without satisfactory
The Atlantic Forest of southeastern Brazil physical and administrative infrastructures
has been identified in many studies as a center (Echternacht et al. 2011).
of endemism, diversity, and richness for many According to the Red List of the National
groups of plants, and as one of the most Center of Flora Conservation (CNCFlora 2022,
important regions for the conservation of global continuously updated) there are ten threatened
biodiversity (see Zizka et al. 2019, Myers et al. species of Orthophytum. Louzada & Wanderley
2000, Smith 1934, Smith & Downs 1974, Benzing (2010), however, observed that there is
2000, Mori et al. 1981, Prance 1982, Cracraft 1985, insufficient information available concerning
Soderstrom et al. 1988, Costa et al. 2000, Silva et most of its species to be able to accurately
al. 2004, Santos et al. 2007). determine their conservation statuses, mainly
According to Forza et al. (2020), 87% of due to the scarcity of collection material (most
all Bromeliaceae species within the Brazilian are only known from the type collection and
Atlantic Forest are endemic, and Zizka et al. imprecise information in the literature and in
(2019) and Martinelli (2000) observed that herbaria collections – which leads us to believe
probably the most threatened Bromeliaceae that the number of threatened species is
species occur within that domain. The high probably much higher. We observed that 44% (24
level of habitat destruction in that region (only species) of the Orthophytum species evidence
approximately 7.5% of the original vegetation microendemic distribution patterns (Table
remains) has indicated it as a priority site for I, Fig. 4–10), being restricted to, and exclusive
global biodiversity conservation (Myers et al. to, a certain vegetation physiognomy found in
2000, Mittermeier et al. 2005). The creation of only one grid square of the PAE, and with one to
greater numbers of protected areas within the four known sites of occurrence. Many of those
Atlantic Forest has accordingly been suggested to species are only known from the type collection
help assure the preservation of that biodiversity. – evidencing the necessity for more intensive
It is worth mentioning that the A26 area alone floristic and geographic distribution studies
(CPAF - which was identified as the most endemic within the family.
area, with the greatest richness and diversity) The concept of microendemism includes
already has more than 120 legally established species with highly restricted and geographically
conservation areas (Table II), which may have proximate distributions (Townsend et al.
been one of the factors aiding the conservation 2011), as was also adopted by McCauley et al.
of Orthophytum species – and demonstrates the (2010) for Guaiacum unijugum Brandegee
importance of such conservation areas. (Zygophyllaceae), which occurs in less than 5%
of the Baja California in Mexico. Giulietti et al.

An Acad Bras Cienc (2024) 96(2) e20230007 15 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

Figure 4. Distribution maps

of Orthophytum species. a. O.
alagoanum. b. O. alvimii. c. O.
arcanum. d. O. argentum. e. O.
atalaiense. f. O. boudetianum.
g. O. braunii. h. O. buranhense.

(2005) also determined that approximately 96% only very small populations with reduced ranges
of all Brazilian species of the Eriocaulaceae in rocky fields, or are known only from their type
family are microendemic. Benzing (2000) and collections.
Martinelli et al. (2013, 2008) observed similar Cox & Moore (2011) noted that there are two
situations with other Bromeliaceae genera. Many main factors influencing the degree of endemism
Aechmea, Hohenbergia, and Neoregelia species in an area: isolation and stability, with isolated
characteristic of the Atlantic Forest have taxa islands and mountains (such as inselbergs and
occurring within just a few forest fragments or, in rock outcrops) always being rich in endemic
some cases, are restricted to only one protected organisms. Inselbergs are isolated rock outcrops
area. Some Cryptanthus species are critically that harbor saxicolous vegetation and are
endangered, principally because they exist in inserted within landscapes of contrasting plant

An Acad Bras Cienc (2024) 96(2) e20230007 16 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

Figure 5. Distribution
maps of Orthophytum
species. a. O. cearense. b. O.
conquistense. c. O. cristaliense.
d. O. diamantinense. e. O.
disjunctum. f. O. duartei. g. O.
eddie-estevesii. h. O. elegans.

communities. Because they are geographically (Noguera-Urbano 2016, Morrone 1994), therefore,
disconnected and show marked ecological as endemic taxa are found exclusively in only a
isolation from their surrounding areas, inselbergs single region, that aspect of their distribution is
are often compared to oceanic islands and tend highly important to their conservation, and the
to maintain their typical attributes regardless identification of neighboring areas of endemism
of their geographic locations, occurring in with low floristic similarities can contribute to
areas with humid forests to dry environments effective conservation planning. Benzing (2000)
(Porembski & Barthlott 2000). hypothesized that the high rates of endemism
Describing and recognizing geographic observed in some Bromeliaceae genera, mainly
distribution patterns is one of the essential saxicolous taxa (e.g., Orthophytum and Dyckia),
stages to delimit areas of endemism could be attributed to their morphologies and

An Acad Bras Cienc (2024) 96(2) e20230007 17 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

Figure 6. Distribution
maps of Orthophytum
species. a. O. erigens.
b. O. estevesii. c. O.
falconii. d. O. foliosum.
e. O. fosterianum.
f. O. glabrum. g. O.
graomogolense. h. O.
grossiorum.

to their lower dispersal capacities. Wanderley Orthophytum is present in a diversity of

(1990) observed that the berry fruits and the spatially and ecologically isolated microhabitats
bird (ornithophily) and insect (entomophily) (inselbergs and rock outcrops) that greatly
dispersal observed in Orthophytum (and in the restrict seed dispersal and migration, thus
whole subfamily Bromelioideae), would limit the favoring the establishment of isolated
geographic distribution of its species in relation populations and promoting speciation over
to the subfamily Tillandsioideae, whose seeds successive generations. Studies of these types of
are widely dispersed by the wind. islands have provided fundamental insights for
understanding the ecological and evolutionary

An Acad Bras Cienc (2024) 96(2) e20230007 18 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

Figure 7. Distribution
maps of Orthophytum
species. a. O.
guaratingense. b. O.
gurkenii. c. O. harleyi.
d. O. horridum. e.
O. jabrense. f. O.
jacaraciense. g. O.
lanuginosum. h. O.
lemei.

processes that affect the biodiversity of step forward to understanding the historical
ecosystems (Porembski & Barthlott 2000), and the processes involved in their current geographic
genus Orthophytum offers an interesting model placement. These results reinforce the
system for examining speciation processes and importance of publishing complete lists of
endemism in neotropical habitats. the materials examined in taxonomic reviews
Studies like this provide a precise overview and making them available to scientists and
of species’ distributions, and represent a herbaria to improve the representativeness of

An Acad Bras Cienc (2024) 96(2) e20230007 19 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

Figure 8. Distribution
maps of Orthophytum
species. a. O. leprosum.
b. O. macroflorum.
c. O. magalhaesii.
d. O. maracasense.
e. O. mello-barretoi.
f. O. minimum. g. O.
piranianum. h. O.
pseudostoloniferum.

An Acad Bras Cienc (2024) 96(2) e20230007 20 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

Figure 9. Distribution
maps of Orthophytum
species. a. O.
pseudovagans. b.
O. riocontense. c.
O. roseolilacinum.
d. O. santaritense.
e. O. santosianum.
f. O. saxicola. g. O.
schulzianum. h. O. sp.
nov.

An Acad Bras Cienc (2024) 96(2) e20230007 21 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

Figure 10. Distribution maps of Orthophytum species. a. O. striatifolium. b. O. sucrei. c. O. toscanoi. d. O. triunfense.
e. O. vasconcelosianum. f. O. zanonii.

those collections. The results presented here Prepuna through the study of Poaceae. Cladistics 25:
and the identification of areas with high levels 295-310.

of endemism can act as a baseline to develop ALVES FVS & LOEUILLE BFP. 2021. Geographic distribution
patterns of species of the subtribe Lychnophorinae
more detailed conservation assessments and
(Asteraceae: Vernonieae). Rodriguésia 72: e02072019.
assist public policies for preserving priority [Link]
conservation areas.
ALVES JV & BURIL MT. 2022. Distribution patterns, endemism,
richness and diversity of Convolvulaceae in the Espinhaço
Acknowledgments Range, Brazil. An Acad Bras Cienc 94: e20211380. https://
The authors are grateful to the Federal Rural University of [Link]/10.1590/0001-3765202220211380.
Pernambuco for institutional support. We also thank the
AMORIM DS & PIRES MRS 1996. Neotropical biogeography
Coordenação de Aperfeiçoamento de Pessoal de Nível
and a method for maximum biodiversity estimation. In
Superior - Brazil (CAPES), for a Doctorate’s scholarships
Bicudo CEM & Menezes WA (Eds), Biodiversity in Brazil, a
to SLC (Process: 88882.436300/2019-01).
first approach. São Paulo: CNPq, 326 p.
ANDERSON S. 1994. Area and endemism. Q Rev Bio 69:

REFERENCES 451-471.
ASSUNÇÃO-SILVA CC & ASSIS S. 2022. Areas of endemism
AAGESEN L, SZUMIK C, ZULOAGA FO & MORRONE O. 2009.
Quantitative biogeography in the South America of Lauraceae: New insights on the biogeographic
highlands - recognizing the Altoandina, Puna and regionalization of the Espinhaço Range, Brazil. Cladistics
38(2): 246-263. [Link]

An Acad Bras Cienc (2024) 96(2) e20230007 22 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

AUSTIN MP . 2013. Vegetation and environment: COX CB & MOORE PD. 2011. Biogeografia: uma abordagem
discontinuities and continuities. In: Van Der Mareel E & ecológica e evolucionária. 7th ed., Rio de Janeiro: LTC,
Franklin J (Eds), Vegetation Ecology, 2nd ed. 398 p.
BARBARÁ T, MARTINELLI G, PALMA-SILVA C, FAY MF, MAYO SJ CRACRAFT J. 1985. Historical biogeography and patterns of
& LEXER C. 2009. Genetic relationships and variation differentiation within the South America avifauna: areas
in reproductive strategies in four closely related of endemism. Ornithol Monogr 36: 49-84.
bromeliads adapted to neotropical ‘inselbergs’: CRISP MD, LAFFAN SW, LINDER HP & MONRO A. 2001. Endemism
Alcantarea glaziouana, A. regina, A. geniculata and A. in the Australian flora. J Biogeogr 28: 183-198.
imperialis (Bromeliaceae). Ann Bot 103: 65-77.
DE LIMA RAF, SOUZA VC, DE SIQUEIRA MF & TER STEEGE H. 2020.
BARROS-SOUZA Y & BORGES LM. 2023. Spatial- and lineage-
Defining endemism levels for biodiversity conservation:
dependent processes underpin floristic assembly in tree species in the Atlantic Forest hotspot. Bio Conserv
the megadiverse Eastern South American mountains. J 252: 108825.
Biogeogr 50(2): 302-315. [Link]
ECHTERNACHT L, TROVÓ M, OLIVEIRA CT & PIRANI JR. 2011. Areas
CNCFLORA - BASE DE DADOS DO CENTRO NACIONAL DE
of endemism in the Espinhaço Range in Minas Gerais,
CONSERVAÇÃO DA FLORA . 2022. Available in: <http://
Brazil. Flora 206: 782-791.
[Link]/portal/>. Access in: Apr 2022.
ENGEMANN K, ENQUIST BJ, SANDEL B, BOYLE B, JORGENSEN PM,
BENZING DH. 2000. Bromeliaceae: profile of an adaptive
MORUETA-HOLME N, PEET RK, VIOLLE C & SVENNING J. 2015.
radiation. Cambridge: University Press, 656 p. Limited sampling hampers “big data” estimation of
BITENCOURT C & RAPINI A. 2013. Centres of endemism in the species richness in a tropical biodiversity hostpot. Ecol
Espinhaço Range: identifying cradles and museums of Evol 5(3): 807-820. [Link]
Asclepiadoideae (Apocynaceae). Syst Biodiv 11: 525-536. FRANÇOSO RD, HAIDAR RF & MACHADO RB . 2016.
BORGES RF, CARNEIRO MA & VIANA P . 2011. Altitudinal Tree species of South America central savanna:
distribution and species richness of herbaceous plants endemism, marginal areas and the relationship with
in campos rupestres of the Southern Espinhaço Range, other biomes. Acta Bot Bras 30: 78-86. [Link]
Minas Gerais, Brazil. Rodriguésia 62: 139-152. org/10.1590/0102-33062015abb0244.
nd
BROWN JH & LOMOLINO MV. 1998. Biogeography. 2 ed, GARCÍA-BARROS E, GURREA P, LUCIANEZ MJ, CANO JM, MUNGUIRA
Ribeirão Preto: Funpec Editora, 691 p. ML, MORENO JC, SAINZ H, SANZ MJ & SIMON JC. 2002. Parsimony
analysis of endemicity and its application to animal and
BUTCHER D & GOUDA E. 2023. A list of accepted Bromeliaceae
plant geographical distributions in the Ibero-Balearic
names. Available in: [Link]
region (western Mediterranean). J Biogeogr 29: 109-124.
Access in: May 2022.
GIULIETTI AM & PIRANI JR. 1988. Patterns of geographical
CALIÓ M F, PIRANI JR & STRUWE L. 2008.
distribution of some plant species from Espinhaço
M o r p h o l o g y - b a s e d p h y l o g e n y a n d re v i s i o n
range, Minas Gerais and Bahia, Brazil. In: Vanzolini PE
of Prepusa and Senaea (Gentianaceae: Helieae) - rare
& Heyer WR (Eds), Proc Neotrop Distrib Patterns. Rio de
endemics from eastern Brazil. Kew Bull 63: 169-191.
Janeiro: Academia Brasileira de Ciências, p. 39-69.
CASAGRANDA MD, ARIAS JS, GOLOBOFF PA, SZUMIK C,
GIULIETTI AM, MENEZES NL, PIRANI JR, MEGURO M & WANDERLEY
TAHER LM, ESCALANTE T & MORRONE JJ. 2009. Proximity,
MGL . 1987. Flora da Serra do Cipó, Minas Gerais:
interpenetration and sympatry: A reply to Dos Santos et
Caracterização e lista de espécies. Bol Bot USP 9: 1-152.
al. Syst Bio 58: 271-276.
GIULIETTI AM, PIRANI JR & HARLEY RM. 1997. Espinhaço
COLLI-SILVA M, VASCONCELOS TNC & PIRANI JR . 2019.
range region. Eastern Brazil. In: Davis SD, Heywood VH,
Outstanding plant endemism levels strongly support
Herrera-Macbryde O, Villa-Lobos J & Hamilton AC (Eds),
the recognition of campo rupestre provinces in
Centres of plant diversity. A guide and strategies for the
mountaintops of eastern South America. J Biogeogr 46:
conservation, Cambridge: The Americas, WWF/IUCN, p.
1723-1733.
397-404.
COSTA LP, LEITE YLR, FONSECA GAB & FONSECA MT. 2000.
GIULIETTI AM, HARLEY RM, QUEIROZ LP, WANDERLEY MGL & VAN
Biogeography of South American forest mammals:
DEN BERG C. 2005. Biodiversity and conservation of plants
endemism and diversity in the Atlantic forest. Biotropica
in Brazil. Conserv Bio 3: 632–639.
32: 872-881.

An Acad Bras Cienc (2024) 96(2) e20230007 23 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

GOLDANI Â. 2012. A importância da Biogeografia Histórica of the rare microendemic species, Guaiacum unijugum
na conservação: exemplos de Análise de Parcimônia de (Zygophyllaceae) in the Cape Region of Baja California,
Endemismo e Panbiogeografia na região Neotropical. Mexico. Rev Mex Biodiv 81: 745-758.
Rev Eletr Bio 5: 119-136.
MENINI-NETO L & FORZZA RC . 2013. Biogeography and
HARLEY RM. 1988. Evolution and distribution of Eriope conservation status assessment of Pseudolaelia
(Labiatae), and its relatives, in Brazil. In: Vanzolini P (Orchidaceae). Bot J Linn Soc 171: 191-200.
& Heyer WR (Eds), Proc Neotr Distrib Patterns. Rio de
MITTERMEIER RA, GIL PR, HOFFMAN M, PILGRIM J, BROOKS
Janeiro: Academia Brasileira de Ciências, p. 71–120.
T, MITTERMEIER CG, LAMOREUX J & FONSECA GAB . 2005.
HIJMANS R, GUARINO L, JARVIS A & O’BRIEN R. 2001. Map anal Hotspots revisited: earth’s biologically richest and
spat data. Available in: [Link] Access most endangered terrestrial ecoregions. Washington:
in: 2022. Conservation International.
LAFFAN SW & CRISP MD . 2003. Assessing endemism at MORI SA, BOOM BM & PRANCE GT. 1981. Distribution patterns
multiple spatial scales, with an example from the and conservation of eastern Brazilian coastal forest tree
Australian vascular flora. J Biogeogr 30: 511-520. species. Brittonia 33: 233-245.
LINDER HP. 2001. Plant diversity and endemism in sub- MORRONE JJ . 1994. On the identification of areas of
Saharan tropical Africa. J Biogeogr 28: 169 -182. endemism. Syst Bio 43(3): 438-441.
LOUZADA RB. 2020. Orthophytum in Flora do Brasil 2020. MORRONE JJ . 2009. Evolutionary biogeography: an
Jardim Botânico do Rio de Janeiro. Available in: http:// integrative approach with case studies. New York:
[Link]/reflora/floradobrasil/FB6274. Columbia University Press.
Access in: May 2022.
MORRONE JJ & ESCALANTE T. 2002. Parsimony analysis of
LOUZADA RB & WANDERLEY MGL . 2010. Revision of endemicity (PAE) of Mexican terrestrial mammals at
Orhophytum (Bromeliaceae): The species with sessile different area units: when size matters. J Biogeogr 29: 1095-
inflorescences. Phytotaxa, 13: 1-26. 1104. [Link]
MADDISON WP & MADDISON DR. 2021. Mesquite: a modular MYERS N, MITTERMEIER RA, MITTERMEIER CG & FONSECA GKJ.
system for evolutionary analysis. Version 3.70. Available 2000. Biodiversity hotspots for conservation priorities.
in: [Link] Access in: 2022. Nature 403: 853-858.
MAGURRAN AE . 1988. Diversity indices and species NAVARRO FR, CUEZZO F, GOLOBOFF PA, SZUMIK C, GROSSO ML &
abundance models. Ecol Div Measur 7-45. QUINTANA G. 2009. Can insect data be used to infer areas

MANRIQUE CE, DURÁN R & ARGÁEZ J. 2003. Phytogeographic

of endemism? An example from the Yungas of Argentina.
analysis of taxa endemic to the Yucatan Peninsula using Rev Chi Hist Nat 82: 507-522.
geographic information systems, the domain heuristic NELSON BW, FERREIRA CAC, DA SILVA MF & KAWASAKI ML. 1990.
method and parsimony analysis of endemicity. Divers Endemism centres, refugia and botanical collection
Distrib 9(4): 313-330. density in Brazilian Amazonia. Nature 345: 714-716.
MARTINELLI G. 2000. The bromeliads of the Atlantic forest. NOGUERA-URBANO EA. 2016. Areas of endemism: travelling
Scient Am: 86-93. through space and the unexplored dimension. Syst
MARTINELLI G, VALENTE A, MAURENZA D, KUTSCHENKO D,
Biodiv 14: 131-139
JUDICE D, SILVA D & PENEDO T. 2013. Avaliações de risco de PALMA-SILVA C, WENDT T, PINHEIRO F, BARBARÁ T, FAY MF,
extinção de espécies da flora brasileira In: Martinelli G COZZOLINO S & LEXER C. 2011. Sympatric bromeliad species
& Moraes M (Eds), Livro vermelho da flora do brasil. Rio (Pitcairnia spp.) facilitate tests of mechanisms involved
de Janeiro: Instituto de Pesquisas Jardim Botânico do in species cohesion and reproductive isolation in
Rio de Janeiro, p. 60-102. Neotropical inselbergs. Mol Ecol, 20: 3185-3201.
MARTINELLI G, VIEIRA CM, GONZALEZ M, LEITMAN P, PIRATININGA PAUSAS JG & AUSTIN MP. 2001. Patterns of plant species
A, DA COSTA AF & FORZZA RC. 2008. Bromeliaceae da Mata richness in relation to different environments: an
Atlântica Brasileira: Lista de éspecies, distribuição e appraisal. J Veg Scienc 12(2): 153-166.
conservação. Rodriguésia 59(1): 209-258.
PLATNICK NI. 1991. On areas of endemism. Austr Syst Bot
MCCAULEY RA, CORTÉS-PALOMEC AC & OYAMA K . 2010. 4: unnumbered.
Distribution, genetic structure, and conservation status

An Acad Bras Cienc (2024) 96(2) e20230007 24 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

POREMBSKI S. 2007. Tropical inselbergs: habitats types, SCHAEFER CEGR ET AL . 2016. The physical environment
adaptative strategies and diversity patterns. Rev Bras of rupestrian grasslands (Campos Rupestres) in
Bot 30: 579-586. Brazil: geological, geomorphological and pedological
POREMBSKI S & BARTHLOTT W . 2000. Inselbergs. Biotic
characteristics, and interplays. In: Fernandes GW (Ed),
diversity of isolated rock outcrops in tropical and Ecology and conservation of mountaintop grasslands
temperate regions. Berlin: Springer-Verlag. in Brazil. New York: Springer International Publishing,
p. 15-53.
POSADAS P & MIRANDA-ESQUIVEL DR. 1999. El PAE (Parsimony
SILVA JMC . 1995. Biogeography analysis of the South
Analysis of Endemicity) como una herramienta en la
evaluación de la biodiversidad. Rev Chill Hist Nat 72: American Cerrado avifauna. Steenstrupia 21: 49-67.
539-546. SILVA JMC & OREN DC. 1996. Application of parsimony

PRANCE GT . 1982. A review of the phytogeographic

analysis of endemicity in Amazonian biogeography: an
evidences for Pleistocene climate changes in the example with primates. Bio J Linn Soc 59: 427-437.
Neotropics. Ann Miss Bot Gard 69: 594-624. SILVA JMC, SOUSA MC & CASTELLETTI CHM. 2004. Areas of

QGIS DEVELOPMENT TEAM . 2021. QGIS Geographic

endemism for passerine birds in the Atlantic forest,
Information System. Available in: [Link] South America. Glob Ecol Biogeogr 13: 85-92.
SILVA JA, MACHADO RB, AZEVEDO AA, DRUMOND GM, FONSECA
RANDO JG & PIRANI JR . 2011. Padrões de distribuição
RL, GOULART MF, MORAES-JÚNIOR EA, MARTINS CS & RAMOS-
geográfica das espécies de Chamaecrista sect.
NETO MB. 2008. Identificação de áreas insubstituíveis
Chamaecrista ser. Coriaceae (Benth.) H. S. Irwin &
Barneby, Leguminosae - Caesalpinioideae. Rev Bras Bot para conservação da Cadeia do Espinhaço nos estado de
34: 499-513. Minas Gerais e Bahia, Brasil. Megadiversidade 4: 272-309.
SILVEIRA FAO ET AL. 2016. Ecology and evolution of plant
RAPINI A . 2010. Revisitando as Asclepiadoideae
(Apocynaceae) da Cadeia do Espinhaço. Bol Bot USP 28: diversity in the endangered campo rupestre: a neglected
97-123. conservation priority. Plant Soil 403: 129-152.
SIGRIST MS & CARVALHO CJBD. 2008. Detection of areas
RAPINI A, RIBEIRO PL, LAMBERT S & PIRANI JR . 2008. A
flora dos campos rupestres da Cadeia do Espinhaço. of endemism on two spatial scales using Parsimony
Megadiversidade 4: 16-24. Analysis of Endemicity (PAE): the Neotropical region and
the Atlantic Forest. Biota Neotrop 8: 33-42.
RON SR. 2000. Biogeographic area relationships of low
SMITH LB . 1934. Geographical evidence on the lines
land Neotropical rainforests based on raw distributions
of vertebrate groups. Biol J Linn Soc 71(3): 379-402. of evolution in the Bromeliaceae. Bot Jahrb Syst,
Pflanzengesch Pflanzengeogr 66: 446-468.
ROVITO SM, ARROYO MT & PLISCOFF P. 2004. Distributional
SMITH LB & DOWNS RJ. 1974. Pitcairnioideae (Bromeliaceae).
modelling and parsimony analysis of endemicity of
Senecio in the Mediterranean-type climate area of Flora Neotropica Monograph No 14 Part 1. New York:
Central Chile. J Biogeogr 31(10): 1623-1636. Hafner Press.
SODERSTROM TR, JUDZIEWICZ EJ & CLARK LG. 1988. Distribution
SAFFORD HD. 1999. Brazilian páramos I. An introduction to
the physical environment and vegetation of the campos patterns of Neotropical bamboos. In: Vanzolini PE
de altitude. J Biogeogr 26: 693-712. & Heyer WR (Eds), Proceedings of a workshop on
Neotropical distribution patterns. Rio de Janeiro: Acad
SAFFORD HD. 2007. Brazilian páramos IV. Phytogeography Bras Cienc, p. 121-157.
of the campos de altitude. J Biogeogr 34: 1701-1722.
SWOFFORD DL . 2003. PAUP* - Phylogenetic Analysis
SANTOS AMM, CAVALCANTI DR, SILVA JMC & TABARELLI M. 2007. Using Parsimony (*and Other Methods). Version 4.
Biogeographical relationships in north-eastern Brazil. J Massachusetts: Sinauer Associates.
Biogeogr 34: 437-446.
SZUMIK CA, CUEZZO F, GOLOBOFF PA & CHALUP AE. 2002. An
SCARANO FR . 2002. Structure, function and floristic optimality criterion to determine areas of endemism.
relationships of plant communities in stressful habitats Syst Bio, 51(5): 806–816.
marginal to the Brazilian Atlantic rainforest. Ann Bot
TER STEEGE H ET AL . 2013. Hyperdominance in the
90(4): 517-524.
Amazonian tree flora. Science 342(6156): 1243092.

An Acad Bras Cienc (2024) 96(2) e20230007 25 | 26

SWAMI L. COSTA et al. DISTRIBUTION PATTERNS IN Orthophytum

TER STEEGE H ET AL. 2015. Estimating the global conservation SWAMI L. COSTA1
status of more than 15,000 Amazonian tree species. Sci [Link]

Adv 1(10): e1500936.

RAFAEL B. LOUZADA2
TOWNSEND TM, LEAVITT DH & REEDER TW . 2011. [Link]
Intercontinental dispersal by a microendemic borrowing
reptile (Dibamidae). Proc Roy Soc: Bio Scienc, p. 1-7. SILMARA CECÍLIA NEPOMUCENO2
[Link]
TROPPMAIR H. 2002. Biogeografia e Meio Ambiente. 5ª ed.,
Rio Claro: Technical Books Editora, 252 p. JOILSON V. ALVES2
[Link]
VERSIEUX LM & WENDT T. 2007. Bromeliaceae diversity and
conservation in Minas Gerais state. Bra Biodiv Conserv
MARIA TERESA BURIL2
1: 2989-3009. [Link]

WANDERLEY MGL . 1990. Diversidade e distribuição 1

Federal Rural University of Pernambuco, Laboratory
geográfica das espécies de Orthophytum (Bromeliaceae).
of Integrative Systematics, Post Graduate Program
Acta Bot Bras 4(1).
in Biodiversity, Av. Manoel de Medeiros, s/n,
WILLIAMS PH, LEES D, ARAÚJO M, HUMPHRIES CJ, VANE-WRIGHT Dois Irmãos, 52171-900 Recife, PE, Brazil
RI & KITCHING IJ. 2002. Biodiversity Worldmap, London: 2
Federal Rural University of Pernambuco, Laboratory
Natural History Museum. of Integrative Systematics, Av. Manoel de Medeiros,
WIENS JJ. 2011. The niche, biogeography and species s/n, Dois Irmãos, 52171-900 Recife, PE, Brazil
interactions. Philos Trans R Soc B: Biol Scien 1576(366):
2336-2350. Correspondence to: Swami Leitão Costa
WIENS JJ & DONOGHUE MJ. 2004. Historical biogeography, E-mail: swamilcosta@[Link]
ecology and species richness. Trends Ecol Evol 19(12):
639-644. Author contributions
ZIZKA A, AZEVEDO J, LEME E, NEVES B, COSTA AF, CACERES D & SLC contributed with data collection, analyses, statistical tests,
ZIZKA G. 2019. Biogeography and conservation status of data interpretation, and manuscript preparation. SCN and JVA
the pineapple family (Bromeliaceae). Div Distrib 00: 1-13. contributed with analyses and statistical tests. RBL and MTB
were responsible for designing the research, supervising the
data analysis stages, and reviewing the manuscript. All authors
How to cite
participated in writing the final version of the manuscript.
COSTA SL, LOUZADA RB, NEPOMUCENO SC, ALVES JV & BURIL MT.
2024. Distribution pattern in the rupiculous genus Orthophytum
(Bromelioideae/Bromeliaceae) reveals high microendemicity in
different types of rocky outcrops. An Acad Bras Cienc 96: e20230007.
DOI 10.1590/0001-3765202420230007.

Manuscript received on January 9, 2023;

accepted for publication on June 18, 2023

An Acad Bras Cienc (2024) 96(2) e20230007 26 | 26