Cunoniaceae from Caparaó National Park, Mantiqueira Mountain Range, Brazil

Abstract A floristic treatment is presented for Cunoniaceae in the Caparaó National Park (PNC), located in the Mantiqueira Mountain Range, on the border of the Minas Gerais and Espírito Santo states. This conservation unit comprises the highest mountain peak in southeastern Brazil, Bandeira Peak, at 2,890 m of elevation. Its vegetation encompasses different phytophysiognomies, including Dense Ombrophilous Forests, Montane Semi-deciduous Seasonal Forests, and High-Altitude Grasslands. Cunoniaceae was represented in the PNC by four species: Lamanonia ternata, L. ulei, Weinmannia humilis, and W. paulliniifolia. To contribute to further taxonomic studies and species conservation in the Mantiqueira Mountain Range, morphological descriptions, identification keys, photographs in vivo, and comments on the taxonomy, geographic distribution of the recorded species are provided.


Introduction
The Atlantic Forest is a biodiversity hotspot (Myers et al. 2000) that comprises some conservationrelevant mountain ranges.Mountainous habitats from this phytogeographic domain are important relicts of the original biodiversity, considering that most of the lowland vegetation has been degraded and a large portion of it has been completely lost due to human disturbance (Ribeiro et al. 2009;Guedes et al. 2020).One of these regions is the Mantiqueira Mountain Range which extends for ca.1,000 km within four Brazilian states (Espírito Santo, Minas Gerais, Rio de Janeiro, and São Paulo).Usually, it is delimited in the meridional and septentrional portions (Machado-Filho et al. 1983).In addition to typical rainforest vegetation, this montane range also includes High-Altitude Grasslands and grass-dominated formations found mostly above elevations of 1,500 m (i.e., campos de altitude, sensu Vasconcelos 2011).
Rodriguésia 74: e01182022.2023 The largest vegetation remnants in the Mantiqueira Range are mostly found in small disjunct conservation units (CUs).According to Gonzaga & Menini (2017), there are five protected areas in the meridional portion (Itatiaia National Park, State Park (P.E, acronym in Portuguese) Campos do Jordão, P.E.da Serra do Papagaio, P.E.do Ibitipoca, and P.E.Serra Negra da Mantiqueira) and two protected areas in the septentrional portion (P.E. Serra do Brigadeiro and Caparaó National Park).Recent studies have indicated that most plant records are collected inside these CUs.However, it is notable that many areas of the Mantiqueira Mountain Range still lack botanical collections, and anthropogenic impacts on non-protected areas potentially threaten their poorly known plant diversity (Gonzaga & Menini 2017;Gonzaga et al. 2019;Santos-Silva et al. 2019;Moraes et al. 2020;Pereira et al. 2021).
Cunoniaceae (Oxalidales) is an angiosperm family commonly found in montane habitats in the Southern Hemisphere (Bradford & Barnes 2001;Bradford et al. 2004;Pillon et al. 2021).It currently comprises 27 genera and approximately 355 species of trees and shrubs that inhabit both tropical and extratropical environments (Bradford & Barnes 2001;Bradford et al. 2004;Pillon et al. 2021).Two genera occur in Brazil: Lamanonia and Weinmannia, with six species each.Lamanonia is widely distributed in this country, except in the Amazon, and has higher diversity in the southeastern mountains (Santos-Silva et al. 2020a).On the other hand, most Weinmannia species are restricted to the southern and southeastern regions and have been recently recorded for the Amazonian highlands (Barbosa-Silva et al. 2016;Santos-Silva et al. 2020a).
Cunoniaceae was subject to a few regional and local flora in Brazil (Bastos 1966;Cuatrecasas & Smith 1971;Mesquita et al. 2003;Pirani & Castro 2011;Santos-Silva et al. 2017;Oliveira et al. 2019), of which only one was focused on a Mantiqueira Range protected area (Santos-Silva et al. 2017).Therefore, the objective of this study was to present a taxonomic treatment of Cunoniaceae from the PNC, contributing to further taxonomic studies and species conservation in the Mantiqueira Mountain Range.Here, we provide descriptions, identification key, photographs in vivo, and comments regarding the taxonomy, and geographical distribution of these species.

Material and Methods
Caparaó National Park (PNC) is located in the east of Minas Gerais and southeast of Espírito Santo states (20º18'-20º37'S and 41º42'-41º52'W) (Fig. 1a).This conservation unit (CU) currently comprises an area of 31,853 hectares with altitudes ranging from 997 to 2,890 m.The highest altitude is the Bandeira Peak, the third highest Brazilian peak.The seasons in the PNC region are divided into rainy summers (from November through March) and wet winters (from April through October).The PNC vegetation comprises different phytophysiognomies, which include Dense Ombrophilous Forests (Montane and High montane) (Fig. 1b,d), Montane Semi-Deciduous Seasonal Forest (Fig. 1c), and High-Altitude Grasslands (Fig. 1e-f).Dense Ombrophilous Forest occurs only in the Espírito Santo portion of the park (ICMBio 2015).

Results and Discussion
In PNC, Cunoniaceae is represented by four species comprising two genera: Lamanonia ternata Vell., L. ulei (Engl.)L.B.Sm., Weinmannia humilis Engl.and W. paulliniifolia Pohl ex Ser, which corresponds to ca. 30% of the species richness in the Brazilian flora currently recorded (Santos-Silva et al. 2020a).In addition to these four species, W. pinnata L. was recorded in the list of terrestrial plants from the PNC, voucher (L.Krieger s.n.CESJ 22323), by Carrijo et al. (2020) and Moreira et al. (2020).However, the occurrence of W. pinnata is not confirmed here, given that none of the examined specimens (including the specimen cited above) presented suborbicular stipules and leaflets with toothed margins, the diagnostic characteristics of W. pinnata (Santos-Silva et al. 2020a).This specimen was identified in our study as W. paulliniifolia.
In PNC, the four Cunoniaceae species can be distinguished by vegetative characteristics such as leaf type, relative color between their surfaces, indumentum, number and dimension of leaflets, presence or absence of axis, and vein appearance on the adaxial surface.Some reproductive characteristics can complement the identification of taxa, such as the number of perianth whorls and stamens, indumentum on the ovary and seeds, and seed type.
Regarding the geographical distribution of Cunoniaceae species present in the PNC, Lamanonia ternata presents the widest geographical distribution in Brazil, and are the only non-endemic species from Brazil.On the other hand, L. ulei shows the most restricted distribution and is endemic to southeastern Brazil (Santos-Silva et al. 2020a).Both Weinmannia species are endemic to Brazil and are distributed along the northeastern (only Bahia state), southeastern, and southern regions (Santos-Silva et al. 2020a).
Previous taxonomic treatments of Cunoniaceae for different CUs located in Minas Gerais and Espírito Santo states recorded three species of Serra do Cipó National Park (Pirani & Castro 2011) and Ibitipoca State Park (Santos-Silva et al. 2017), two species of Grão-Mogol State Park (Mesquita et al. 2003), and one species of Mestre Álvaro Environmental Protection Area (Valadares et al. 2022) Lamanonia ternata is the only species that occurs in all four of these areas.Weinmannia paulliniifolia is also present in Ibitipoca State Park.The other two species (L.ulei and W. humilis) are not shared between the previously studied areas and the PNC.

Cunoniaceae.
Trees Lamanonia ternata occurs in Argentina, Brazil, and Paraguay (Zickel & Leitão-Filho 1993;Pirani & Castro 2011).This species is widespread in the Brazilian Cerrado and Atlantic Forest domains (Santos-Silva et al. 2020a).Lamanonia ternata displays the highest morphological variation within the genus, even in relation to important characters for taxa delimitation (e.g., the number of leaflets and indument) (Zickel & Leitão-Filho 1993).Among the species that occur in the PNC, L. ternata presents morphological similarities with L. ulei and can be differentiated from it by the presence of concolorous or slightly discolorous leaflets, glabrous or puberulent abaxially, with nonprominent veins (vs.markedly discolorous leaflets, densely tomentose abaxially, and prominent veins in L. ulei).It was collected with flowers in January, July, and August, and with fruits in January and April.This species has its IUCN conservation status assessed as LC (Least Concern) according to Botanic Gardens Conservation International (BGCI) & IUCN SSC Global Tree Specialist Group (2019b).
Lamanonia ulei is endemic to the Brazilian Atlantic Forest and occurs in the high-altitude  surface, indumentum-type in leaflets.This species has been collected with flowers in March and June and fruits in April.It is worth to mention that this species has its IUCN conservation status assessed as EN, "Endangered" (Fernandez et al. 2020) 3. Weinmannia humilis Engl., Linnaea 36: 620 (1870).Type: BRASIL.MINAS GERAIS: without further locality.P. Clausen 328, Type: G n.s, Possible isotype: K image!(barcode K000486179).Fig. 2e Trees or shrubs 0.5-2.5 m height, young branches sparsely to densely incanous-tomentose.Leaves opposite, petiolate, composite, pinnate, 9-17-foliolate; lateral leaflets 0.5-1.3× 0.3-0.5 cm, median leaflets 0.6-1.5 × 0.3-0.5 cm, sessile, obovate-elliptic or oblong-elliptic, discolorous, apex obtuse, acute in the median leaflet, base obtuse or rounded, entire margin until the median region, serrate towards the apex, adaxial and abaxial surfaces glabrous, veins glabrous, rarely tomentose, slightly prominent on the abaxial surface; axis present, winged; stipules circular.Pseudoracemes apical, 2 per apex of the branch, axis 2.6-7.3 cm long, densely tomentose.Flowers actinomorphic, dichlamydeous, white; pedicel 3-5 mm long, tomentose; sepals 5-6 mm long, ovatelanceolate, united at the base, apex acute, abaxial surface puberulous, petals ca. 1 cm long, ovate to oblong, apex obtuse, abaxial surface glabrous, deciduous; stamens 10; ovary glabrous.Septicidal capsule 3.7-4 × 1-1.5 mm, oblong, castaneous to pinkish.Seeds numerous, not winged, pilose.Weinmannia humilis is endemic to the Brazilian Atlantic Forest, occurring in all states of the southeast and south regions, as well as in Bahia (Santos-Silva et al. 2020a).In PNC, W. humilis presents morphological similarities with W. paulliniifolia and can be distinguished mainly by its smaller leaflet dimensions, having lateral leaflets with 0.3 to 0.5 cm width, median leaflets 0.6 to 1.5 cm length and 0.3 to 0.5 cm width (vs.lateral leaflets 0.6 to 1 cm wide, median leaflets 2 to 3.1 cm length and 0.8 to 1.6 cm width in W. paulliniifolia).The species was collected with flowers in February and March, and from January through April and October.This species has its IUCN conservation status assessed as LC (Least Concern) according to Botanic Gardens Conservation International (BGCI) & IUCN SSC Global Tree Specialist Group (2019b).
Weinmannia paulliniifolia is endemic to the Brazilian Atlantic Forest and Cerrado domains throughout the south region and in São Paulo, Minas Gerais, and Bahia (Santos-Silva et al.Rodriguésia 74: e01182022.2023 2020a).Specimen L. Krieger s.n.(CESJ 22323) was identified as W. pinnata by Carrijo et al. (2020).However, the diagnostic characteristics of this species (suborbicular stipules and leaflets with toothed margins) were not observed in any of the analyzed specimens of Weinmannia from the PNC, as mentioned in the Results and Discussion section.
Among the Cunoniaceae species that occur in PNC, Weinmannia paulliniifolia presents morphological similarities with W. humilis and can be distinguished by its leaflet dimensions (as mentioned above).This species has been collected with flowers in May, August, and October, and with fruits in August, September, and November.This species does not have its conservation status assessed by IUCN.
, in addition to the aforementioned literature.Phenology was based on data gathered from herbarium specimens.