Tovomita manauara A. Nobre & L. Marinho, 2022

Marinho, Alexandra Monteiro Nobre Lucas Cardoso, Cabral, Fernanda Nunes, Demarchi, Layon Oreste & Sousa, Juliana Hipólito De, 2022, Tovomita manauara (Clusiaceae): a new species revealed by fruit morphology and leaf anatomy, Phytotaxa 536 (3), pp. 270-278 : 271-276

publication ID 10.11646/phytotaxa.536.3.7


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Tovomita manauara A. Nobre & L. Marinho

sp. nov.

Tovomita manauara A. Nobre & L. Marinho   , sp. nov. ( Figs 1 View FIGURE 1 , 2 View FIGURE 2 )

Type:— BRAZIL. Amazonas: Manaus, ca. 90 km N de Manaus, Distrito Agropecuário da Suframa , rodovia BR 174, km 72, depois 6 km oeste da BR, Fazenda Dimona, mata de terra-firme sobre latosolo amarelo, árvore n°3362, 02°19’S, 60°05’W, 50–125 m. a.s.l., 16 July 1992, fr., M. Nee 42991 (holotype INPA barcode INPA0096785 View Materials !; GoogleMaps   isotypes GH!, MG!, NY barcode NY01417188 !, UEC!) GoogleMaps   .

Diagnosis: — Tovomita manauara   is morphologically similar to T. caloneura   , and can be differentiated by the amount of secondary veins (25–36 pairs vs. up to 26 pairs in T. caloneura   ), fruit morphology (4-septate and turbinate vs. 5-septate and fusiform) and anatomical aspects—secretory canals larger (> ~40 µm vs. <~20 µm), presence of collenchyma in the petiole (vs. absent), and the absence of druses in the midvein (vs. presence).

Description: —Trees 15–18 m alt., dbh ca. 30 cm; prop roots conspicuous with 1 m in height; outer bark gray, smooth, inner bark reddish; exudate yellow; proximal internodes 2.5–8.2 cm long, distal internodes 1.2–3.4 cm long. Petioles 1–2 cm long, green, slightly canaliculate, not striated, lenticels absent. Leaf blades 4.7–14.2 × 1.9–7.1 cm, brownish-green and shiny in sicco, the adaxial surface darker than the abaxial, black dots present on both surfaces, subcoriaceous, brittle, oblong or obovate, base convex or rounded, apex acute or apiculate, margin entire and slightly revolute, exudate canals conspicuous, parallel to secondary veins, visible only abaxially in sicco as blackish lines. Venation brochidodromous, midvein immersed adaxially and prominent abaxially; secondary veins in 25–36 pairs, 1–6.2 mm apart, forming an angle ca. 70º to the midvein, immersed adaxially and slightly 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. Capsules fleshy, 2.5–5.3 × 0.9–2.4 cm, green when immature, epicarp smooth; mesocarp purplish-red, pleasantly aromatic, 4-septate, turbinate to napiform when closed, not costate, 4-lobed; pedicel slightly dilated 1–1.5(–4.4) cm long; sepals, petals and staminodes caducous, 4–stigmas persistent 2–6 mm. Seeds 4, often one is aborted, ellipsoid, aril light orange.

Additional Specimens Examined (Paratypes): — BRAZIL. Amazonas : Manaus , estrada Manaus-Porto Velho , estrada entre o lago do Castanho e o rio Araçá, em direção ao Careiro, 12 July 1972, fr., M. F   . Silva et al. 564 ( INPA barcode INPA0096784 View Materials ); Distrito Agropecuário da Suframa, rodovia BR 174   , km 64, depois 23 km leste na ZF3, Fazenda Esteio , mata de terra-firme sobre latosolo amarelo, árvore n°2803, 02°23’S, 59°51’W, 27 August 1986, fr., M. J. R GoogleMaps   . Pereira s/n ( INPA barcode INPA0096786 View Materials ); ca. 90 km N de Manaus, Distrito Agropecuário da Suframa, rodovia BR 174 , km 72, depois 6 km oeste da BR   , Fazenda Dimona , mata de terra-firme sobre latosolo amarelo, árvore n°3362, 02°19’S, 60°05’W, 28 June 1989, fr., E GoogleMaps   . Palheta s/n ( INPA n° 175768, NY barcode NY01417090 ); ca. 90 km N de Manaus, Distrito Agropecuário da Suframa, rodovia BR 174   , km 64, depois 23 km leste na ZF3, Fazenda Esteio , mata de terra-firme sobre latosolo amarelo, árvore n°2803, 02°23’S, 59°51’W, 26 July 1989, fr., S. S GoogleMaps   . da Silva s/n ( INPA barcode INPA0096787 View Materials , NY barcode NY01417091 )   .

Etymology: —Manauara (Portuguese) is a person who is born in the municipality of Manaus, Amazonas state, Brazil, where all the specimens came from. Besides the species, the first author is also a manauara.

Distribution, Habitat and Conservation Status: —All the known specimens of Tovomita manauara   were collected in dense ombrophilous forest situated in low altitudes (between 60 and 120 m. a.s.l.), known as terra-firme forest (see physiognomic descriptions in Ribeiro et al. 1999). The area of occurrence is located on extensive tertiary sediments within the lowland plains of the Amazon, with predominantly clayey latosols (ferralsols) that are generally poor in nutrients due to long periods of leaching ( Oliveira & Mori 1999). Tovomita manauara   grows in the sub-canopy stratum (15 to 20 m tall) of forests that are in a good state of conservation and, apparently do not occur in secondary forests.

Due to the few available records, which are concentrated in only two nearby locations, T. manauara   is considered as having deficient data (DD) following what was adopted by Marinho & Beech (2019). The exsiccates/specimens of T. manauara   were collected in the study sites of the world’s largest and longest-running experimental study of habitat fragmentation, the Biological Dynamics of Forest Fragments Project (BDFFP). This project was created in the 70’s, after the Brazilian government announced the deforestation of a large area for agricultural development in the region. As a result, forest patches of different sizes were maintained for the study of the impacts caused by the fragmentation ( Laurance & Luizão 2007). The BDFF project, in partnership with government agencies, in order to protect its study sites, implemented a protected area of 3,288 ha (Area of Relevant Ecological Interest - Biological Dynamics of Forest Fragments Project AREI – BDFFP) in 1985, which covered the known area of occurrence of T. manauara   . However, nowadays, the protected area and the entire region suffer from threats due to illegal farming, logging, and hunting ( Laurance & Luizão 2007). This situation has become further aggravated by the urban expansion of the suburbs of Manaus, and the construction of small roads or trails that facilitate the illegal occupation of public lands ( Esteves 2007).

Taxonomy notes and comments: —Almost 30 years after the collection of the type materials, specimens of Tovomita manauara   with flowers have not yet been collected. Despite the BDFF project operating in the area of occurrence for more than 40 years, the absence of materials with flowers may be a reflection of the lack of scheduled collection in the area. In this sense, we recommend carrying out systematic fieldwork, followed by detailed taxonomic studies to bring this and other species not yet described to the fore ( Hopkins 2019).

Tovomita manauara   can be recognized among other species of the genus by a large number of secondary veins (25–36 pairs vs. up to 26 pairs in T. caloneura   , Fig. 1B View FIGURE 1 ), and the presence of 4-septate fruits, which are turbinate to napiform and smooth. Tovomita manauara   is sympatric and morphologically similar (leaf shape, petiole, venation type) to T. caloneura   . However, it has 4-carpellate and turbinate fruit (vs. 5-carpellate and fusiform in T. caloneura   ) ( Fig. 1D View FIGURE 1 ), and the number of carpels is one of the most important characteristics for infrageneric delimitation ( Marinho et al. 2021). Turbinate fruits also occur in T. albiflora A.C. Smith (1940: 298)   and T. turbinata Planchon & Triana (1860: 283)   , though T. manauara   can be easily distinguished from these species by the number of secondary veins (<15 pairs in both). Napiform to turbinate fruits ( Fig. 1C View FIGURE 1 ) also appear in T. colombiana L. Marinho (2018: 406)   , which is a species that is endemic to Colombia and has less than 15 pairs of secondary veins ( Marinho 2018).

Anatomy: —In addition, anatomical characters are also important for distinguishing species ( Fig. 2 View FIGURE 2 ). Tovomita manauara   has petiole with plane-convex contour and presents the vascular cylinder with units of vascular bundles in the medullary portion of the vascular tissue, which is a closed vascular system ( Fig. 2A View FIGURE 2 ), as well as the presence of idioblasts concentrated in the cortex. The petiole has about 25/+ vascular bundles ( Fig. 2B View FIGURE 2 ), the secretory canals are aligned on the periphery of the epidermis, and there is the presence of druses along the parenchyma ( Fig. 2C View FIGURE 2 , 3F View FIGURE 3 ). The midvein presents secretory canals in the medulla ( Fig. 4A,B View FIGURE 4 ) and there is no significant increase in the number of vascular bundles when compared to the petiole. In T. caloneura   the petiole has a plane-convex contour and presents the vascular cylinder with units of vascular bundles in the medullary portion of the vascular tissue, collateral bundles with a ring and an internal arch ( Fig. 2D, E View FIGURE 2 ). The idioblasts and canals are present in the subepidermal region and petiole cortex ( Fig. 2F View FIGURE 2 ). In the midvein ( Fig. 4C View FIGURE 4 ), there is a thick cuticle in the epidermis and, in the vascular system, the vascular bundles are numerous (40/+), form collateral   bundles, and are a ring with an inferior arch, a biconvex arrangement with medullary phloem ( Fig. 4C, D View FIGURE 4 ). The idioblasts are arranged in the subepidermal, perivascular and medulla region of the vascular system. The mesophyll is homogeneous, the palisade parenchyma has three layers and the spongy parenchyma has six layers and canals arranged in the medulla, and both tissues are adorned with idioblasts ( Fig 4E View FIGURE 4 ).

Externally, Tovomita manauara   and T. caloneura   share hypostomatic leaves ( Fig. 3C, E View FIGURE 3 ). The abaxial surface is covered with a waxy cuticle ( Fig. 3A–E View FIGURE 3 ). In some regions, the cuticle covers small papillae, similar to those seen in T. vismiifolia L. Marinho   (see Fig. 1 View FIGURE 1 in Marinho et al. 2016b), but is smaller and less evident ( Fig. 3B, D View FIGURE 3 ). Tovomita manauara   and T. caloneura   also share the collateral vascular bundles and consist of one ring with one inner arch, a plane-convex cross-section on the petiole, in addition to the thick cuticle on the midvein. However, they are clearly distinct when the number of vascular bundles and the presence of druses is considered, especially in the midvein, the number of idioblasts, and the diameter of secretory canals in the petiole. This information is summarized in Table 1 View TABLE 1 .


Instituto Nacional de Pesquisas da Amazonia


Harvard University - Gray Herbarium


Museum of Zoology


William and Lynda Steere Herbarium of the New York Botanical Garden


Universidade Estadual de Campinas


Botanische Staatssammlung München


Field Museum of Natural History, Botany Department


University of the Witwatersrand


Departamento de Geologia, Universidad de Chile


Nanjing University


Royal Botanic Garden Edinburgh


Department of Botany, Swedish Museum of Natural History