General characteristcs of Polybotrya. A-E. Polybotrya cylindrica. A. Holodimorphic fertile leaf (left) and sterile leaves (right). B. Fertile pinna, detail. C. Climbing habit of a fertile individual, note erect fertile leaves and spreading sterile ones. D. Climbing habit of a sterile individual. E. Rhizome in cross section, showing five large meristeles surrounded by a dark sclerenchymatous sheath, and many smaller vascular traces of the leaves. F. High-climbing rhizome of P. goyazensis attached firmly to a phorophyte by means of clasping roots produced on the ventral surface. (All photos by Bianca K. Canestraro.) 

General characteristcs of Polybotrya. A-E. Polybotrya cylindrica. A. Holodimorphic fertile leaf (left) and sterile leaves (right). B. Fertile pinna, detail. C. Climbing habit of a fertile individual, note erect fertile leaves and spreading sterile ones. D. Climbing habit of a sterile individual. E. Rhizome in cross section, showing five large meristeles surrounded by a dark sclerenchymatous sheath, and many smaller vascular traces of the leaves. F. High-climbing rhizome of P. goyazensis attached firmly to a phorophyte by means of clasping roots produced on the ventral surface. (All photos by Bianca K. Canestraro.) 

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Polybotrya (Dryopteridaceae) is a genus of Neotropical endemic ferns that exhibits diverse growth forms and often plays important ecological roles in tropical forests. The genus is composed of about 35 species, yet the taxonomy of some groups of species in the genus is poorly understood. In this work we present the first detailed taxonomic treatmen...

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... status.-Least Concern (LC): found in numerous locations and widespread in the Brazilian Atlantic Forest (IUCN ...
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... status.-Near Threatened (NT): geographic range severely fragmented and re- stricted and occurence in more than 10 locations and in the four southeastern Brazilian states (IUCN 2001). Polybotrya pilosa can be recognized by its dark brown, appressed, bicolorous scales with erose to lacerate margins; lamina with long hairs, distrib- uted only on the veins (usually) (1.1 mm to 1.6 mm, 8-13-celled); unbranched paraphyses (usually); and lamina margins pilose (Fig. 7C). A few specimens, including the type (Goes & Dionisio 1086, RB), have some paraphyses with a unicellular minute ramification. Polybotrya pilosa resembles P. tomentosa due to the pubes- cence of the abaxial surfaces of the lamina; how- ever, P. tomentosa has shorter hairs on the veins and the laminar tissue between the veins (0.5 mm long, 3-celled; Fig 10). Polybotrya pilosa is also similar to P. osmundacea; whereas the latter has the pinnules of the proximal pinnae anadromic and pinnules of the medial pinnae usually catadromic and lamina margins glabrous (Fig. 8 H-J Plants scandent or terrestrial. Rhizomes long- creeping, 0.7-1.1 cm thick; scales 0.7-1.3×0.07- 1.4(-0.3) cm, deltate linear to lanceolate linear, appressed, soft or stiff (rarely), ligth to dark brown, translucent, bicolorous, the base curved and truncate to ovate or auriculate, margin erose to lacerate toward the apex, denticulate when young; petioles 13-30×0.4 cm. Sterile leaves 50-102 cm long, ovate, 2-pinnate to 2-pinnate- pinnatisect (rarely), the apex pinnatifid; laminae 37-70×20-60 cm, chartaceous; pinnae 10-15(- 30)×4-7(-16) cm, petiolate, ovate to deltate, the base obtuse to truncate, the apex acute; pinnules 3-7(-10)×1.2-2.5(-4) cm, petiolate or sessile, ovate to deltate, the base obtuse to truncate and asymmetric, the base of the distal side of pinnule more divided than the rest, usualy auriculate, the apex acute to acuminate or obtuse, the proximal side of pinnae and pinnule reduced, all pinnules anadromic; segments 1.2(-2.6)×0.8-1 cm, ob- long to ovate (rarely), the base obtuse, the apex obtuse to round; pinnules margin crenate or entire and glabrous; adaxial surface subglabrous, hairs 0.2(-0.5) mm long, 2-celled, on sulcus only; ab- axial surface subglabrous, hairs 0.2 mm long, 1- 3-celled, on the midrib only, with microscales; axes sparsely to moderately pilose, hairs 0.2(- 0.5) mm long, on the sulcus, with scales and microscales; veins free. Fertile leaves 35-50(- 73) cm long, ovate, 3-pinnate; pinnae 8-15×3- 6.5(-10) cm, deltate, the base truncate, the apex acute to cuneate; pinnules 2-3.5(-6)×(0.2-)1(-2) cm, deltate to linear toward the apex, the base truncate and asymmetric, the apex cuneate; paraphyses unbranched or with a minute bifurca- tion (rarely); sori coenosoric; spores (47-)53- 65(-70)×36-46(-50) ...
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... tomentosa is distinct based on its reddish brown, patent, concolorous scales with den- ticulate margins; conspicuously branched paraphy- ses; and hairs 0.2 mm to 0.6 mm long and 3-6- celled, covering both surfaces of the laminae on the veins and between the veins (Fig. 10). Polybotrya tomentosa was considered a synonym of P. speciosa by Moran (1987). However, P. tomentosa has hairs all over the laminar surfaces, whereas P. speciosa has hairs only on the veins or on the leaf sulcus ( Fig. 9) (see P. speciosa ...
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... speciosa is recognized by its red- dish brown, patent, concolorous, soft scales with denticulate margins; adaxial surface of the leaves subglabrous, with hairs only on the sulcus on the main axes; abaxial surface sparsely to moderately pilose, with hairs restricted to the veins (0.1- 0.4 mm, 2-4-celled); and by its conspicuously branched paraphyses (Fig. 9). Polybotrya speciosa can be confused with P. tomentosa by its similar scales, paraphyses and sterile and fertile leaf divi- sions; however, P. tomentosa differs by the hairs 0.6 mm long and 3-6-celled distributed on both lamina surfaces and between the veins (Fig. ...

Citations

... The first contributions to the knowledge of this family in the country were given by Raddi (1819), Presl (1822), Schrader (1824), Martius (1828Martius ( -1834, Fée (1869Fée ( , 1873, Baker (1870), and Christensen (1913Christensen ( , 1920. More recently, there were the contributions by Brade (1961Brade ( , 1972, Sundue et al. (2013), Prado et al. (2014), Canestraro & Labiak (2015), Engels & Canestraro (2017), Schwartsburd et al. (2018), Bohn et al. (2020), and -in addition to the monographs already mentioned in the previous paragraph and some regional floras (e.g., Brade 1946;Sehnem 1979;Salino & Carvalho 2005;Garcia & Salino 2008;Prado et al. 2017). Garcia & Salino (2008) studied the species of Dryopteridaceae from Minas Gerais but adopted a different circumscription for the family (i.e., that of Moran & Riba 1995, which included Didymochlaena, but excluded Bolbitis, Ctenitis, Elaphoglossum, Lastreopsis, Megalastrum, Mickelia, and Parapolystichum). ...
... Polybotrya is neotropical, consisting of 36 species (Moran 1987;Canestraro & Labiak 2015). In Brazil, there are 16 species, seven of which are endemic (Canestraro 2022). ...
... Polybotrya tomentosa further differs from P. speciosa by having these hairs also between the veins (vs. hairs restricted to veins and major axes in P. speciosa) (Moran 1987;Canestraro & Labiak 2015). ...
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We present a taxonomic treatment for the species of Didymochlaenaceae and Dryopteridaceae occurring in the region of Viçosa, Minas Gerais, Brazil. This study was based on 50 herbarium gatherings from IAN, K, NY, PACA, RB, SP, UPCB, US, and VIC. Didymochlaenaceae is represented by a single species, Didymochlaena truncatula, whereas Dryopteridaceae has 11 infrageneric taxa in seven genera: Ctenitis (four taxa, including two varieties), Megalastrum (two species), Mickelia, Parapolystichum, Polybotrya, Rumohra, and Stigmatopteris (one species each). Among the 12 infrageneric taxa here recognized, seven are endemic to the Brazilian Atlantic Forest. Ctenitis distans var. isabellina, Megalastrum connexum, and Polybotrya speciosa are here reported for the first time in Viçosa. Identification keys, descriptions, illustrations, geographical distribution, and comments are presented to all taxa.
... It was the only scandent species found in the CSP. Besides its long-creeping, scandent rhizomes and dimorphic fronds, it is also distinct due to its appressed, rigid and dark brown rhizome scales (Canestraro & Labiak 2015 Copel but differs in that the rachis are densely coated by orange scales and have strongly revolute margins (vs. glabrous or sparsely scaly rachis with flat margins in S. gracilis). ...
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We present a checklist of the ferns and lycophytes from the Cerrado State Park, Jaguariaíva, Paraná, Brazil. This region represents the southernmost limit of Cerrado vegetation in Brazil. The park is located along the Devonian Escarpment, one of the most iconic geological formations in southern Brazil. The remnants of Cerrado in Paraná are relics of a colder and drier climate that occurred during the Last Glacial Maximum, about 5,600 years ago. They are now surrounded by natural grasslands and Araucaria Forest, which is typical of southern Brazil. We recorded 112 species (102 ferns and ten lycophytes), from 51 genera and 16 families. Ten species were first recorded in the Devonian Escarpment, and Ctenitis bigarellae and Tryonia areniticola are rare. None of the species are endemic to the Cerrado and most are typical of the humid forests of Eastern Brazil. We provide notes and illustrations for all species.
... According to Moran (1987), Polybotrya occurs preferably in shaded areas in mature forests, rarely growing in disturbed sites. In Brazil, P. cylindrica is mainly found inside mature Atlantic forest formations (Canestraro and Labiak 2015), being less common along trail margins (Hirai and Prado 2012) and in restinga (Nóbrega et al. 2011;Gonzatti et al. 2014). ...
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Ferns are an important component in the understorey of tropical forests and their distribution is influenced by several biotic and abiotic factors. At a regional scale, soil characteristics and canopy openness play an important role in fern species composition and richness, as well as in the abundance of individuals. Our objective was to compare the influence of edaphic conditions and vegetation structure on the abundance and distribution of fern communities in Atlantic forest and restinga forest. Our hypotheses were that fern species richness and diversity are higher in Atlantic forest than in restinga due to limiting conditions in this habitat and the composition of fern species in Atlantic forest differs from restinga , especially due to differences in edaphic conditions. A principal coordinates analysis was applied to ordinate sampling units in relation to the environmental variables and a permutational multivariate analysis of variance was used to test that environmental variables did not differ between the two vegetation types. Species richness was compared using rarefaction curves. The influence of abiotic variables in species composition and abundance was verified using canonical correspondence analysis. No differences were observed in species richness, diversity or dominance between vegetation types, although abundance was higher in restinga . Fern communities respond to edaphic conditions and vegetation structure variations between vegetation types, the soil playing a major role. A greater variety of habitats resulting from differences in soil drainage in restinga facilitates the co-existence of species with different ecological tolerance, increasing local diversity and compensating for limiting conditions in restinga .
... Polybotrya caudata e P. osmundacea possuem caule longo-reptante e trepador, e possuem a lâmina estéril 2-pinado-pinatífida ou mais dividida. Costa Rica, Guiana Britânica, Suriname, Venezuela, Colômbia e Brasil: AL, GO, MG, MT, PA, PE, RR, SP (Canestraro & Labiak 2015). Serra dos Carajás: Serra da Bocaina, Serra Norte: N1, e Serra Sul: S11-B, S11-D. ...
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This study addressed the Dryopteridaceae taxa recorded in ferruginous formations of Serra dos Carajás, Pará state, bringing descriptions, illustrations, geographical distribution, and comments. In the study area four genera and four species were recorded: Ctenitis nigrovenia, Dryopteris huberi, Elaphoglossum glabellum and Polybotrya sorbifolia.
... by Moran (1986) and Sánchez et al. (1991). Polybotrya was studied by Moran (1987a), with updates by Rojas (2007) and Canestraro and Labiak (2015). ...
... Be-cause the climbing rhizome's connection to the soil is maintained, it is termed a terrestrial root climber (Canestraro et al. 2014). (Hemiepiphytes sensu Benzing [1990] eventually lose their connection to the ground.) ...
... The clade formed by Polybotrya espiritosantensis Brade to Polybotrya speciosa Schott ( fig. 1) is endemic to the Atlantic forest of southeastern Brazil (Moran 1987a;Canestraro and Labiak 2015). Although highly supported by our molecular results (PP p 1.0, ML-BS p 100%), the clade does not exhibit any exclusive morphological characters of which we are aware. ...
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Premise of research. The polybotryoid fern clade is completely Neotropical and consists of Cyclodium, Maxonia, Olfersia, Polybotrya, and Polystichopsis. It has never received a detailed phylogenetic analysis. We performed such an analysis to examine the relationships among species and genera and to map the evolution of their morphological and anatomical characters. Methodology. Our study included 46 (77%) of the 60 species in the clade. It also included 37 outgroup species from 19 genera. We sequenced four plastid DNA markers (rbcL, rps4-trnS, trnG-trnR, and trnL-trnF) and analyzed the data with maximum likelihood and Bayesian inference. One anatomical and 11 morphological characters were mapped on the resulting phylogenetic trees using the criterion of maximum parsimony. Pivotal results. The polybotryoid clade was strongly supported as monophyletic, as were its component genera. Nearly all its species have long-creeping rhizomes. Polystichopsis was resolved sister to the other polybotryoid genera. Its monophyly is supported by the morphological synapomorphies of distichous phyllotaxy, long straightish white hairs on the leaves, and tuberculate perines. Two species currently classified in Arachniodes (Arachniodes macrostegia and Arachniodes ochropteroides) form a clade with Olfersia. No known morphological characters support this clade. Olfersia, however, is highly distinct from all other polybotryoids by the combination of its imparipinnate laminae, submarginal connecting vein, strong sterile-fertile leaf dimorphy, loss of indusia, and evolution of acrostichoid sori. Maxonia is defined by its terrestrial root-climbing habit and dimorphic sterile and fertile leaves. Cyclodium and Polybotrya were resolved as sister. The presence of peltate indusia is synapomorphic for Cyclodium. Polybotrya is defined by several morphological synapomorphies: a rhizome anatomy unique among dryopteroid ferns (each individual meristele is surrounded by a dark sclerenchymatous sheath), strong sterile-fertile dimorphy, and loss of indusia. A possible synapomorphy for Polybotrya is the terrestrial root-climbing habit. Within Polybotrya, anastomosing veins and round discrete sori have evolved more than once. Conclusions. This is the first phylogenetic analysis of the polybotryoid ferns. The clade was resolved as monophyletic, as were its genera. An unexpected result was that two species currently classified in Arachniodes (A. macrostegia and A. ochropteroides) were resolved sister to Olfersia. Most of the main clades of polybotryoids were supported by morphological and/or anatomical characters.
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We present a taxonomic monograph for Dryopteridaceae in Brazil. This is the second largest fern family (after Pteridaceae) in the country, comprising 16 genera and 191 species. An online version is available at: http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB90950
Article
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Polybotrya (Dryopteridaceae) is a genus of Neotropical endemic ferns that exhibits diverse growth forms and often plays important ecological roles in tropical forests. The genus is composed of about 35 species, yet the taxonomy of some groups of species in the genus is poorly understood. In this work we present the first detailed taxonomic treatment of Polybotrya in the Brazilian Atlantic Forest. We provide a key, descriptions, comments, conservation status, distribution maps, and illustrations for the ten species found in the Atlantic Forest. Seven species are endemic (P. cylindrica, P. espiritosantensis, P. matosii sp. nov., P. pilosa, P. semipinnata, P. speciosa and P. tomentosa) and three are widespread in Brazil or in the Neotropics (P. goyazensis, P. osmundacea and P. sorbifolia). A new species, Polybotrya matosii, is described, and the name P. tomentosa is reconsidered in this study. Lectotypes are also designated for P. incisa and P. semipinnata.