Tovomita cornuta (Clusiaceae): a new and threatened species from the Amazonian campinaranas revealed by ecological field research

Layon Oreste Demarchi Maria Teresa Fernandez Piedade Lucas Cardoso Marinho About the authors

ABSTRACT

High species diversity, large area, and taxonomically under-studied groups are some of the explanations for the frequent discovery of new species of plants during ecological field campaigns in the Amazon region. Some of these new species are already being threatened from unplanned urban expansion associated with deforestation and habitat loss. Here we describe Tovomita cornuta, a new species discovered during ecological studies in the white-sand campinaranas of the Amazon, which is also threatened. The species is endemic to the state of Amazonas, Brazil, where it occurs in the understory of forested campinaranas physiognomies and is easily recognized by its narrowly obovoid fruits with dilated free styles. We provide a description, taxonomic and ecological comments, assessment of conservation status, geographic distribution, phenology, and illustrations for the new species.

Keywords:
Central Amazonia; conservation; IUCN Red List; new species; sampling gaps; threatened species; Tovomita; white-sand vegetation

Introduction

The demand to solve the main issues that deal with environmental degradation and the conservation of biodiversity is hampered by a lack of knowledge in relation to biodiversity. This paradox is partly caused by the demand to produce general environmental standards and related solutions, which require large data sets. As a result, the proportion of ecological investigations based on field research has declined in recent decades. For example, Ríos-Saldaña et al. (2018Ríos-Saldaña CA, Delibes-Mateos M, Ferreira CC. 2018. Are fieldwork studies being relegated to second place in conservation science? Global Ecology and Conservation 14: e00389. doi: 10.1016/j.gecco.2018e00389
https://doi.org/10.1016/j.gecco.2018e003...
) analyzed the related environmental literature between the 1980s and 2014 and found a 20 % decrease in publications based on fieldwork and an increase of 600 % and 800 % in modeling and meta-analysis studies, respectively, for the same period. On the other hand, new species are still being found almost every day, indicating that the diversity of animal and plant species is still poorly known due to sampling gaps in large areas (Hopkins 2007Hopkins MJG. 2007. Modelling the known and unknown plant biodiversity of the Amazon Basin. Journal of Biogeography 34: 1400-1411. ) and this diversity is underestimated even for well-studied taxa (Ceballosa & Ehrlichb 2009Ceballosa G, Ehrlichb PR. 2009. Discoveries of new mammal species and their implications for conservation and ecosystem services. Proceedings of the National Academy of Sciences 106: 3841-3846.).

In the Amazon region, the large area, the high diversity of species and, in some cases, taxonomically understudied groups are some of the reasons for the frequent discovery of new species of plants (Sobral & Stehmann 2009Sobral M, Stehmann JR. 2009. An analysis of new angiosperm species discoveries in Brazil (1990-2006). Taxon 58: 227-232.; Cardoso et al. 2017Cardoso D, Särkinen T, Alexander S, et al. 2017. Amazon plant diversity revealed by a taxonomically verified species list. Proceedings of the National Academy of Sciences 114: 10695-10700.; Hopkins 2019Hopkins MJG. 2019. Are we close to knowing the plant diversity of the Amazon? Anais da Academia Brasileira de Ciências 91: e20190396. doi:10.1590/0001-3765201920190396
https://doi.org/10.1590/0001-37652019201...
). In this sense, long-term ecological studies are important because they generally conduct frequent field campaigns and collaborate in the discovery of new species (Lindenmayer et al. 2012Lindenmayer DB, Likens GE, Andersen A, et al. 2012. Value of long-term ecological studies. Austral Ecology 37: 745-757.). A good example is the genus Tovomita, for which most of the new specimens collected in the Amazon have come from research in ecology (e.g., Marinho et al. 2016Marinho LC, Fiaschi P, Santos FAR, Amorim AM. 2016a. Three new species of Tovomita (Clusiaceae) from the Amazon River basin and first record of papillae for Tovomita. Plant Systematics and Evolution 302: 1121-1134.a; 2019Marinho LC, Fiaschi P, Amorim AM. 2019a. A gold mine: four more new species of Tovomita (Clusiaceae: Clusieae) from Amazonia. Willdenowia 49: 343-350. a). The large trees, which are common to the genus, usually have small, greenish flowers and are hardly seen during rapid collection expeditions, but will likely be found during collections in systematic monitoring studies (Marinho & Beech 2019Marinho LC, Beech E. 2019. Red List of Tovomita. Richmond, UK, Botanic Gardens Conservation International.).

Tovomita comprises 53 exclusively Neotropical species belonging to the Clusiaceae family (Marinho et al. 2020Marinho LC, Luján M, Fiaschi P, Amorim AM. 2020. Tovomita nebulosa (Clusiaceae), a new species from Cerro de la Neblina, Venezuela. Acta Amazonica 50: 149-154. ). These species occur more frequently in humid forests, with only a few representatives in areas of seasonal semi-deciduous forest in eastern Brazil (Marinho et al. 2016Marinho LC, Fiaschi P, Gahagen B, Santos FAR, Amorim AM. 2016b. Tovomita (Clusiaceae) from the Brazilian Atlantic Forest: taxonomy and utility of leaf venation characters at the species level. Systematic Botany 41: 758-774. b). In the Amazon region, they occupy areas of upland forests (terra-firme), white- and black-water floodplain forests (várzea and igapó respectively) and white-sand forests (campinaranas). In the Brazilian Amazon, Tovomita is sympatric to Arawakia and Chrysochlamys, two other genera of the Clusieae tribe that are commonly confused with each other. Tovomita can be distinguished by the acute or cuneate to attenuate leaf base (vs. leaf base gradually narrowed, ultimately shortly truncate or abruptly rounded in Arawakia) and its floral buds enclosed by the outer sepals (vs. floral buds not enclosed by the outer sepals in Chrysochlamys) (Marinho et al. 2019Marinho LC, Cai L, Duan X, et al. 2019b. Plastomes resolve generic limits within tribe Clusieae (Clusiaceae) and reveal the new genus Arawakia. Molecular Phylogenetics and Evolution 134: 142-151. b).

The present study describes a new species of Tovomita collected within the scope of the PELD-MAUA project (Brazilian Long-term Ecological Research Network - Ecology, monitoring and sustainable use of wetlands). Illustrations, a distribution map and an assessment of the conservation status of the new species are provided. Taxonomic characters are discussed and compared with those of other closely related species.

Materials and methods

The descriptions and photos of the species in its habitat were carried out in the field within the scope of the doctoral thesis of the first author, and collections were made monthly from July 2017 to March 2020. Specimens were collected and processed following the usual procedures for botanical samplings (Fidalgo & Bononi 1989Fidalgo O, Bononi VLR. 1989. Técnica de coleta, preservação e herborização de material botânico. São Paulo, Instituto de Botânica.). The description of dried material was complemented by collections from the INPA herbarium (acronym according to Thiers 2021Thiers B. 2021, continuously updated. Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. http://sweetgum.nybg.org/science/ih/. 25 Sep. 2020.
http://sweetgum.nybg.org/science/ih/...
, continuously updated) and their respective duplicates. Flower measurements were taken from rehydrated flowers and floral buds; colors of all the structures described were recorded in the field and on the herbarium labels. Leaf terminology follows Ellis et al. (2009Ellis B, Daly D, Hickey LJ, et al. 2009. Manual of leaf architecture. Ithaca, Cornell University Press.), and flower and fruit terminology follows Radford et al. (1974Radford AE, Dickison WC, Massey JR, Bell CR. 1974. Vascular plant systematics. New York, Harper & Row.). The geographic distribution map was created using the website SimpleMappr (Shorthouse 2010Shorthouse DP. 2010. SimpleMappr, an online tool to produce publication-quality point maps. https://www.simplemappr.net. 26 Sep. 2020.
https://www.simplemappr.net...
) with subsequent style modifications. The conservation status of the new species was assessed using the software GeoCAT (Bachman et al. 2011Bachman S, Moat J, Hill AW, Torre J, Scott B. 2011. Supporting red list threat assessments with GeoCAT: Geospatial conservation assessment tool. ZooKeys 150: 117-126. ) and discussed based on the IUCN (2012)IUCN 2012. IUCN Red List categories and criteria, version 3.1. Second Edition. https://portals.iucn.org/library/efiles/documents/rl-2001-001-2nd.pdf.
https://portals.iucn.org/library/efiles/...
criteria.

Results and discussion

Taxonomic treatment

Tovomita cornuta Demarchi & L. Marinho, sp. nov. Type: BRAZIL. Amazonas: São Sebastião do Uatumã, Uatumã Sustainable Development Reserve (SDR), PELD-MAUA plots, understory of white-sand forest campinarana with high groundwater level, 2°11’47” S, 59°00’53” W, 35 m a.s.l., 10/III/2020, fr., L.O. Demarchi 1674 (holotype: INPA; isotype: MAR; RB).

(Figs. 1, 2, 3, 4).

Figure 1
Tovomita cornuta. A. Branch with staminate inflorescence. B. Leaf abaxial surface. C. Staminate floral bud. D. Staminate flower. E. Stamen with detail of anther. F. Staminodes detail in the pistillate flower (from floral bud), sepals and petals removed. G. Pistillate flower. H. Style (from pistillate floral bud). I. Branch with closed fruits. J. Closed fruit. (A-E from L.O. Demarchi 1558; F-G from W. Thomas et al. 5330; H from L.O. Demarchi & Andriolli 1691; I-J from L.O. Demarchi 1674). Illustration by Marcos Melo Corrêa.

Figure 2
Details of Tovomita cornuta structures in sicco. A. Leaf abaxial surface with detail showing venation. B. Staminate inflorescence. C. Staminate floral bud. D. Staminate flower. E. Stamens with detail of anther. F. Pistillate floral bud, sepals and petals removed. G. Staminode. H. Closed mature fruit. (A-E from L.O. Demarchi 1558; F-G from W. Thomas et al. 5330; H from L.O. Demarchi 1674).

Figure 3
Tovomita cornuta, geographic distribution with conservation assessment and habitat. A. Distribution map of T. cornuta in the state of Amazonas, Brazil, with the detail of the state’s location in South America; black circles indicate the locations where the plants were collected. B. The polygon represents the extent of occurrence of T. cornuta indicated by the GeoCAT software. C. Habitat of T. cornuta in São Sebastião do Uatumã, Amazonas, Brazil. D. Detail of the shallow water table forming small “puddles” during the wet season in São Sebastião do Uatumã, Amazonas, Brazil. Photos by Layon O. Demarchi.

Figure 4
Tovomita cornuta, live plants. A. Staminate inflorescence with senescent stamens. B. Staminate flower, top view of the pistillode in detail (arrow). C. Pistillate inflorescence with floral buds, flower, and immature fruits (arrow). D. Pistillate 5-branched pleiochasium with immature fruits. E. Closed fruit, arrow pointing to the yellow exudate. F. Open fruit, lateral view. Photos by Layon O. Demarchi. (A from L.O. Demarchi 1558; B from L.O. Demarchi 1599; C from L.O. Demarchi 1692; D from L.O. Demarchi 1691; E-F from L.O. Demarchi 1674).

Tovomita cornuta can be differentiated from T. umbellata Benth. by the number of secondary veins (25-28 pairs vs. 10-15 in T. umbellata) and by the narrowly obovoid fruit with dilated styles (vs. pyriform with non-dilated styles in T. umbellata), from T. amazonica (Poepp.) Walp. by the number of secondary veins (25-28 vs. 13-18 in T. amazonica), floral bud length (3-4.3 vs. 5-7 mm long in T. amazonica), and fruit epicarp (smooth vs. rugose in T. amazonica).

Trees or treelets 3-8 m alt., dbh < 9 cm, prop roots inconspicuous, not exceeding 30 cm in height; outer bark gray, smooth, inner bark reddish; exudate yellow, orangish when oxidized, scarce; proximal internodes 3-11 cm long, distal internodes 3-20.5 mm long. Petioles 8.6-17.2 mm long, green, slightly canaliculate, not striated, lenticels absent. Leaf blades 3.4-15.1 × 1.4-6.1 cm, greenish adaxially and light brownish abaxially in sicco, black dots absent, subcoriaceous, brittle, shiny, oblong to obovate, base convex to rounded, apex acute to cuneate, margin entire and slightly revolute, exudate canals conspicuous, parallel to secondary veins, visible only abaxially in sicco. Venation brochidodromous, midvein prominent on both surfaces, more so abaxially, secondary veins in 25-28 pairs, 1.5-3.5 mm apart, forming an angle 75-80º to the midvein, slightly prominent adaxially, prominent abaxially; intersecondary veins present, one or rarely two per intercostal area, similar to the secondary veins, parallel to major secondaries and reticulating near the margin, visible only abaxially in sicco; tertiary veins percurrent sinuous to reticulate; intramarginal vein present. Staminate inflorescences a lax 5-6-branched pleiochasial cyme, ranging from 20-47 flowers, with primary flower, sometimes with reduced dichasia lacking the central flower. Pistillate inflorescences a lax 5-branched pleiochasial cyme or a dichasial cyme, ranging from 7-8 flowers, usually with reduced dichasia lacking the central flower. Pedicel 3-5.5 mm long on the flower and strongly dilated and elongated on the fruit with 7.1-10 mm long, whitish green, proximally articulated in the lateral flowers of the dichasia; lenticels present when in fruit; bracteoles 2, non-calyptrate, apex rounded to acute. Floral buds 3-4.3 mm long, ovoid to oblong, apex rounded, slightly apiculate, lenticels absent. Sepals 2, 3.3-4.1 × 1.7-1.9 mm, oblong to ovate, apex rounded, greenish to white; petals 4, 6.2-6.7 × 2.5-2.6 mm, oblong, reflexed, apex acute, greenish to white. Staminate flowers with 23-28 stamens, 1.5-3.5 mm long, heterodynamous; filaments terete, white becoming yellowish when senescent; anthers 0.3-0.4 mm long, slightly wider than filaments, connective not exceeding the thecae; pistillode white, diminute. Pistillate flowers with 20-26 staminodes, 1-1.6 mm long, rudimentary anther present, ca. 0.2 mm long; ovary ca. 1 mm long, not costate or lobed, 4-locular, white-yellowish; styles 0.8-1 mm long, free from each other, curved in the floral bud, erect on anthesis; stigmas 4, 0.5-0.8 mm width, capitate, white. Capsules fleshy, 1.5-2 × 0.8-1.1 cm, 4-septate, narrowly obovoid when closed, not costate, not lobed, epicarp smooth, green when immature; when mature green with visible yellowish resin canals, mesocarp purplish-red; pedicel dilated, styles strongly dilated; sepals, petals and staminodes caducous, styles and stigmas persistent; free styles 6.3-8.3 × 1.6-2.8 mm, conical to oblong. Seeds ca. 12.5 × 6 mm, ellipsoid, aril bright orange.

Paratypes: BRAZIL: Amazonas, Manaus, Rio Negro, Ponta Negra, 11-14/IV/1972, fr., R.E. Schultes & W. Rodrigues 26191A (GH, INPA, NY); Tupé Sustainable Development Reserve, Comunidade Agrovila, campinarana forest on sand-soil, 02°58’55’’ S, 60°14’26’’ W, 9/VI/2013, fb. ♂, L.O. Demarchi et al. 192 (EAFM, INPA); 9/VI/2013, fb. ♂, L.O. Demarchi et al. 193 (EAFM, INPA); 28/IX/2013, fb. ♂, L.O. Demarchi et al. 194 (EAFM, INPA); 4/X/2020, fl. ♀, fr., L.O. Demarchi & F. Andriolli 1691, 1692, 1693 (INPA); 04/X/2020, fl. ♂, L.O. Demarchi & F. Andriolli 1694, 1695 (INPA); Presidente Figueiredo, represa de Balbina on Rio Uatumã, ca. 4 km NW of dam on D-1 road, campinarana on sand-soil, 1°50’ S, 59°32’ W, 4/VII/1986, fb. ♀, fr., W. Thomas et al. 5330 (INPA, NY); São Sebastião do Uatumã, Uatumã Sustainable Development Reserve, PELD-MAUA plots, understory of forested campinarana on sand-soil, 2°11’47’’ S, 59°00’53’’ W, 35 m a.s.l., 4/IX/2018, fb. ♂, L.O. Demarchi 1300 (INPA); 11/VIII/2019, fl. ♂, L.O. Demarchi 1558 (INPA); 10/X/2019, fl. ♂, L.O. Demarchi 1599 (INPA).

Distribution: Tovomita cornuta is presumably endemic to the state of Amazonas, Brazil, with occurrence records in the region of Manaus and the neighboring municipalities of Presidente Figueiredo and São Sebastião do Uatumã (Fig. 3A).

Habitat and ecology: All known specimens were collected in white-sand forests (campinarana) (Fig. 3C). The species occurs in the understory in dense phytophysiognomies with a shallow water table that forms small “puddles” during the wet season (Fig. 3D), and does not occur in open areas with direct sunlight.

Phenology: Tovomita cornuta can be seen with flowers from June to October and with fruits in March, April, and July. In the specific case of the Sustainable Development Reserve (SDR) Uatumã, phenological monitoring over 24 months showed the species with flower buds in August; flowers from September to October, which is the middle of the dry season; and immature fruits from November to February and the dehiscence of fruits and release of seeds in March, which is the wet season (Demarchi et al. unpublished data).

Etymology: The specific epithet refers to the conspicuously prominent styles observed on the fruit. It is derived from the Latin word cornu, which means horn.

Conservation status: Based on the GeoCAT software (Bachman et al. 2011Bachman S, Moat J, Hill AW, Torre J, Scott B. 2011. Supporting red list threat assessments with GeoCAT: Geospatial conservation assessment tool. ZooKeys 150: 117-126. ) Tovomita cornuta was classified as Endangered (EN) according to the area of occupancy (AOO, B2a) (Fig. 3A, B). However, in accordance with the IUCN criteria (2012)IUCN 2012. IUCN Red List categories and criteria, version 3.1. Second Edition. https://portals.iucn.org/library/efiles/documents/rl-2001-001-2nd.pdf.
https://portals.iucn.org/library/efiles/...
, other factors and threats discussed herein further allow us to classify the species as critically endangered (CR). Among the four locations of known occurrence of the proposed new species, two are outside protected areas and under strong threat. The Ponta Negra district in the municipality of Manaus (Schultes & Rodrigues 26191A) has been strongly impacted by unplanned urban expansion associated with deforestation and habitat loss (B2bi, ii, iii); Balbina, a district in the municipality of Presidente Figueiredo (Thomas et al. 5330), was affected in the 1980s by the construction of the Balbina dam, which resulted in large areas being flooded in order to form a reservoir for power generation (Fearnside 1989Fearnside PM. 1989. Brazil's Balbina Dam: Environment versus the legacy of the pharaohs in Amazonia. Environmental Management 13: 401-423. ) and has probably affected populations of this species (A1a + B2cii). The other two locations are in protected areas under the category of Sustainable Development Reserves (SDR), and some minor threats may also occur such as selective logging (Demarchi et al. 2019Demarchi LO, Scudeller VV, Moura LC, Lopes A, Piedade MTF. 2019. Logging impact on Amazonian white-sand forests: perspectives from a sustainable development reserve. Acta Amazonica 49: 316-323. ) and suppression of forest areas for small-scale agriculture and cattle ranching (IDESAM 2009IDESAM. 2009. Série Técnica Planos de Gestão: Reserva de Desenvolvimento Sustentável do Uatumã. Volume 1 e 2. Itapiranga, São Sebastião do Uatumã - Amazonas. Manaus, IDESAM - Conservação e Desenvolvimento Sustentável.) (B2biii). The species appears to be naturally rare, since, despite occurring in the surrounding areas of Manaus, one of the most sampled areas in the Amazon, only a few specimens could be found. In the two protected areas, 21 individuals were recorded in the Tupé SDR (Demarchi et al. 2018Demarchi LO, Scudeller VV, Moura LC, et al. 2018. Floristic composition, structure and soil-vegetation relations in three white-sand soil patches in central Amazonia. Acta Amazonica 48: 46-56. , supplementary material mistakenly identified as T. acutiflora), and less than 10 individuals were recorded in the Uatumã SDR (Demarchi pers. obs.) (C2ai). (A1a + B2a + B2bi, ii, iii, cii + C2ai).

Taxonomic notes and comments: Two specimens of Tovomita cornuta kept at the INPA herbarium were mistakenly identified as T. acutiflora and T. umbellata. In the field, T. cornuta can be easily differentiated from both by its inconspicuous prop roots. The prop roots are an important vegetative characteristic to differentiate it from other genera of Clusiaceae in the field, but it is not a characteristic that is shared by all species of Tovomita. The T. cornuta leaves resemble those of both related species, since they have numerous secondary veins that are close to each other (Figs. 1B, 2A, see Table 1). Although the fruits are quite characteristic, they were initially interpreted by Marinho (2019Marinho LC. 2019. Sistemática de Tovomita Aubl. (Clusiaceae) e gêneros relacionados. PhD thesis, Universidade Estadual de Feira de Santana, Feira de Santana.) as possible galls or a malformation (Figs. 1J, 4E). This was mainly due to the absence of specimens with floral buds or flowers. The free and dilated style is the most striking feature for differentiating T. cornuta. Other species, such as Tovomita stylosa and Tovomita turbinata, share such characteristics, T. cornuta does not have the same geographic distribution (Colombia, Costa Rica, and Panama for T. stylosa; Colombia, Trinidad, and Venezuela for T. turbinata) and can be easily differentiated from both by the number of secondary veins (25-28 pairs vs. 8-9 in T. stylosa; 12-15 in T. turbinata) and epicarp texture (smooth vs. asperous in T. stylosa and T. turbinata).

Tovomita cornuta can be differentiated from T. acutiflora by its smaller floral buds (3-4.3 vs. 12-13 mm long in T. acutiflora), smaller number of stamens (23-28 vs. 50-70 in T. acutiflora) and, especially, by its four carpels and dilated styles on the fruit (vs. five-carpellate fruits with non-dilated styles) (Barros & Mariz 1982Barros MS, Mariz G. 1982. Tovomita acutiflora Barros et G. Mariz espécie nova do Amazonas. Acta Amazonica 12: 291-292. ). In addition to the arrangement of the secondary veins, T. cornuta shares with T. umbellata the ovoid floral buds (Figs. 1C, 2C), the four-carpellate fruits with a smooth epicarp, the distal portion of the pedicel dilated in the fruit (Figs. 1J, 4F) and, finally, the number of stamens (23-28 vs. 20-35 in T. umbellata). The gynoecium of T. cornuta is composed of long styles and tiny stigmas (Figs. 1H, 2F, 4C) that will give rise to a fruit with persistent dilated styles that are similar to horns (Figs. 1J, 2H, 4F), while the T. umbellata gynoecium is composed of sessile and wide stigmas (ca. 2 mm width) that will give rise to a fruit without apparent styles, with only the conspicuous stigmas at the apex.

Due to the dilated distal portion of the pedicel and the shiny leaves in vivo, Tovomita cornuta can also be compared to the sympatric species T. spruceana. Tovomita cornuta can be differentiated from T. spruceana by the ovoid floral buds (vs. oblong in T. spruceana), heterodynamous stamens (vs. isodynamous in T. spruceana), 23-28 stamens (vs. 30-40 stamens in T spruceana) and finally, the narrowly obovoid fruit with dilated styles (vs. pyriform with non-dilated styles).

The specimens Demarchi et al. 192, 193, 194, which belongs to T. cornuta, were mistakenly included by Marinho et al. (2016Marinho LC, Fiaschi P, Santos FAR, Amorim AM. 2016a. Three new species of Tovomita (Clusiaceae) from the Amazon River basin and first record of papillae for Tovomita. Plant Systematics and Evolution 302: 1121-1134.a) as paratypes of T. trachycarpa, now synonymized under T. amazonica (sensuMarinho 2020Marinho LC. 2020. Tovomita. In: Flora do Brasil 2020. Jardim Botânico do Rio de Janeiro. http://reflora.jbrj.gov.br/reflora/floradobrasil/FB6883. 20 Sep. 2020.
http://reflora.jbrj.gov.br/reflora/flora...
). The characters differentiating these three and other related species of Tovomita are presented in Table 1.

Table 1
Morphological comparison of Tovomita cornuta and related Amazonian species.

Considering the large estimated number of plants not yet described, especially in the Amazon Biome, the only way to bring this neglected biodiversity to light is through intensive collections followed by detailed taxonomic studies (Hopkins 2019Hopkins MJG. 2019. Are we close to knowing the plant diversity of the Amazon? Anais da Academia Brasileira de Ciências 91: e20190396. doi:10.1590/0001-3765201920190396
https://doi.org/10.1590/0001-37652019201...
). From this point of view, long-term ecological research is an appropriate investment option since it requires a multidisciplinary approach with frequent field expeditions (Lindenmayer et al. 2012Lindenmayer DB, Likens GE, Andersen A, et al. 2012. Value of long-term ecological studies. Austral Ecology 37: 745-757.). The discovery of T. cornuta is an example of the relevance of field work, and highlights how ecological and taxonomic studies can interact in order to reveal Amazonian biodiversity.

Acknowledgements

We thank the CNPq for the scholarship to LOD and for financing the project CNPq/PELD-MAUA, Process: 441590/2016-0 and FAPEAM/PELD-MAUA, Process: 403792/2012-6; INPA/MAUA Group, PPI: 1090-5; and the Project ADAPTA (CNPq/FAPEAM/INPA, Process: 465540/2014-7). We also thank the SEMA and the project ATTO for fieldwork logistics. We are grateful to Valdeney Azevedo and Fernando Andriolli for assistance in fieldwork; Mariana Mesquita for herbarium procedures; Marcos Melo Corrêa for the illustration; Fernanda Cabral, Pedro Fiaschi and an anonymous reviewer for their constructive suggestions.

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Publication Dates

  • Publication in this collection
    16 Feb 2022
  • Date of issue
    Oct-Dec 2021

History

  • Received
    12 Nov 2020
  • Accepted
    11 Jan 2021
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