Heteranthera pumila M.Pell. & C.N.Horn, 2017

Pellegrini, Marco O. O. & Horn, Charles N., 2017, Two peculiar new species of Heteranthera Ruiz & Pavon (Pontederiaceae) from Brazil, with notes on inflorescence architecture in the family, PhytoKeys 82, pp. 35-56 : 35

2. Heteranthera pumila M.Pell. & C.N.Horn sp. nov. Figs 3 , 4 , 5

Diagnosis.

Similar to H. reniformis Ruiz & Pavón due to its petiolate leaves with blades two or more times wider than long, reniform to broadly cordate, cincinnus enclosed by the basal bract, glandular-pubescent cincinnus axis, perianth lobes with a 5+1 arrangement and acute to acuminate at apex, filaments straight, and intrusive-parietal placentation. It differs due to its diminute petiolate leaves [3.5-11.8-(13.2) × 3.2-12.1 mm], inflorescences 1-2-(3)-flowered, peduncle densely glandular-pubescent, basal bract glandular-pubescent at base, apex aristate, flowers pale lilac to lilac or light pink, seeds smooth or with 7-9 inconspicuous longitudinal wings.

Type.

BRAZIL. São Paulo: Piraju, várzea do rio Paranapanema , na divisa com o município de Manduri, 23°07'50"S 49°19'32"W, fl., fr., 10 Oct 2016, M.O.O. Pellegrini & R.F. Almeida 495 (holotype: RB!; isotypes: NBYC!, SPF!, US!).

Description.

Herbs annual or short-lived perennials. Roots thin, delicate, unbranched, white. Stems repent on the substrate or floating in shallow water, delicate, spongy, rooting at the nodes; internodes 1.7-64.1 mm long, glabrous. leaves not seen. Petiolate leaves distichously-alternate, distributed along the stem, floating to emergent; sheaths 2.8-7.5 mm long, glabrous, covered with mucilage, longitudinally split and light green when mature, ligule 2-parted, surpassing the sheath, 0.2-0.8 mm long, membranous, light green, glabrous, apex triangular; petiole 8.5-82.9 mm long, not inflated, glabrous; blades 3.5-11.8-(13.2) × 3.2-12.1 mm, cordate to broadly cordate to reniform, rarely narrowly cordate, membranous, glabrous, base cordate, margins glabrous, apex acute to obtuse. Inflorescences axillary or apparently terminal, reduced to a solitary pedunculate cincinnus; peduncle 0.5-3.4 cm long, deflexed and submerged in fruit, densely glandular-pubescent; basal bract (spathe) 0.9-1.9 × 0.4-0.8 cm, spathaceous, broadly elliptic, conduplicate, green, glandular-pubescent at base, margins hyaline, apex aristate; cincinnus bract absent; cincinnus 1-2-(3)-flowered, all flowers included in the basal bract, when present the third flower always exerted, axis 0.2-1.8 mm long, densely glandular-pubescent. Flowers bisexual, tubular, chasmogamous, sessile, enantiostylous; floral buds narrowly ovoid, light green to lilac or pink, densely glandular-pubescent; perianth tube 4.9-7.3 mm long, light green, densely covered with glandular hairs, lobes 5 superior and 1 inferior, pale lilac to lilac or light pink, lateral superior lobes 3.6-5 × 0.8-1.4 mm, elliptic, base cuneate, apex acute to acuminate, central superior lobe 3.6-4 × 1.7-2.1 mm, ovate to broadly ovate, base obtuse to rounded, slightly involute, apex acute, with a nectar guide at base, yellowish green to green with an upper vinaceous to brown spot, inferior lobe 4.2-4.9 × 0.5-0.8 mm, narrowly elliptic to linear elliptic, base cuneate, apex acuminate; stamens 3, lateral stamens with filaments straight, 1.6-1.8 mm long, pale lilac to light pink, not inflated, apically barbate with eglandular, multi-celled, lilac to pink hairs, anthers 0.4-0.6 × 0.3-0.5 mm, broadly oblongoid to quadrangular, yellow, central stamen with filament straight, 2-2.3 mm long, lilac to pink, not inflated, medially sparsely villose with eglandular, white hairs, anthers 1.2-1.6 × 0.3-0.5 mm, ellipsoid, greyish blue to greyish mauve; ovary 3.1-3.5 × 1-1.2 mm, linear ovoid, glabrous, 1-locular, placentation intrusive-parietal, style gently sigmoid, 4.2-5.1 mm long, lilac to pink, terete, densely villose in the distal portion with eglandular, white hairs, stigma unevenly trilobate, purple to pink, densely glandular-pubescent. Capsule 5.3-7.2 × 1.1-1.9 mm, linear ovoid, glabrous, smooth, light green when immature, light to medium brown when mature; persistent perianth base (anthocarp) smooth, medium to dark brown. Seeds 0.5-0.7 × 0.2-0.3 mm, oblongoid, light brown to yellowish brown, testa smooth, sometimes with 7-9 inconspicuous longitudinal wings; hilum punctate; embryotega dorsal, inconspicuous, without a prominent apicule.

Specimens seen

(paratypes). BRAZIL. Minas Gerais: São Sebastião do Paraíso, Fazenda Fortaleza, fl., 20 Apr 1945, A.C. Brade & A. Barbosa 17846 (RB, SP, UNA). Paraná: Guaratuba, Boa Vista, fl., 28 Jan 1964, G. Hatschbach 11078 (MBM); Rio da Divisa, fl., fr., 16 Dec 1971, G. Hatschbach 28523 (MBM, UPCB). Rio Grande do Sul: Bom Jesus, Rio Socorro, fl., 19 Feb 2008, Grupo de Estudos Reófitas UHBG 2116 (MBM). Vacaria, vale do Rio Ibitíria, ca. 30 km NE de Vacaria, fl., s.dat., J.C. Lindeman et al. s.n. (ICN9466). Santa Catarina: Lages, Santo Antônio, near Passo de Socorro, estrada de rodagem Federal km 67-71, south of Lages, fl., 14 Jan 1957, L.B. Smith & R. Reitz 9959 (HBR, RFA, US). São Paulo: Americana, Praia Azul, fl., 2 Mar 1993, Faria 96/16 (UEC). Bálsamo, estrada sentido Bálsamo-Mirassolândia, fl., 30 Jan 1997, A.D. Faria et al. 97/350 (UEC). Dracena, margem do Rio do Peixe, fl., fr., 7 Sep 1995, L.C. Bernacci et al. 2124 (IAC, SP, SPF, UEC). Estrela D’Oeste, SP-320, lago localizado na Fazenda Santo Antônio, lado direito da pista no sentido Estrela D’Oeste-Jales, fl., fr., 30 Jan 1997, L.Y.S. Aona et al. 97/167 (UEC). Igarapava, lagoa localizada na Fazenda Flor das Frutas, lado direito da pista no sentido Igarapava-Rifaina, na altura do km 16, fl., 15 Jan 1997, A.D. Faria et al. 97/102 (UEC). Ouro Verde, SP-563, km 113, Ponte Nova, Rio do Peixe, fl., 10 Jul 1996, A.D. Faria et al. 96/122 (UEC); loc. cit., fl., fr., 10 Jul 1996, A.D. Faria et al. 96/130 (BOTU, IAC, SP, SPF, UEC). Paulo de Faria, fl., Oct 1994, V.C. Souza et al. 12294 (ESA, IAC, UEC). Pedregulho, rodovia Antônio Giolo, acesso à Estreito, solo encharcado próximo à uma cachoeira, fl., fr., 14 Jan 1997, A.D. Faria et al. 97/64 (UEC). Piraju, várzea do Ribeirão São Bartolomeu, fl., fr., 15 May 1996, E.L.M. Catharino et al. 2090 (PMSP). Riolândia, brejo localizado em estrada de terra no sentido Riolândia-Paulo de Faria, fl., 29 Jan 1997, L.Y.S. Aona et al. 97/152 (UEC). Santa Rita do Passa Quatro, rodovia Anhanguera, km 239, Sítio Aubiri, fl., 13 Jan 1997, A.D. Faria et al. 97/20 (UEC). São José do Rio Preto, represa, fl., 25 Nov 1965, G. Marinis & E.M.P. Martins 20 (FUEL, SJRP, SP); Estação Experimental de Zootecnia de São José do Rio Preto, fl., 28 Dec 1977, M.A. Coleman 220 (SP). São Pedro do Turvo, 8 km da estrada em direção à Marília, desvio em estrada de terra ca. 3.5 km, 49°70'W 22°48'S, est., 9 Dec 1994, M.C.E. Amaral & V. Bittrich 94/48 (UEC). Sud Mennucci, distrito de Bandeirantes D’Oeste, fl., 4 Aug 1995, M.R. Pereira-Noronha et al. 1552 (SP). Teodoro Sampaio, margem do lago ao lado da estrada Teodoro Sampaio-Planalto, ca. Km 11.5, fl., Oct 1997, L.Y.S. Aona et al. 97/241 (UEC).

Etymology.

The epithet means “small”, making allusion to the small stature of the new species, especially its diminute leaf blades.

Distribution, habitat and ecology.

Heteranthera pumila is endemic to the Paraná, Uruguay, and Southeastern Atlantic watersheds, in the Atlantic Forest domain. It is restricted to Brazil, in the states of Minas Gerais, São Paulo, Paraná, Santa Catarina and Rio Grande do Sul (Fig. 2 ), growing on open marshy areas and slow water environments along the Paraná, Paranapanema and Rio Grande rivers (and their respective tributaries), from 700 to 1,800 meters above the sea level. It is very likely that H. pumila also reaches the state of Mato Grosso do Sul. Nonetheless, we have been unable, so far, to find any vouchers from this state in the visited herbaria.

Phenology.

Heteranthera pumila blooms throughout the year, with flowering peaks during the wet season, and was found in fruit from September to October and from January to March.

Conservation status.

Heteranthera pumila is widely distributed across the upper Paraná, Uruguay, and Southeastern Atlantic watersheds, with a wide EOO (ca. 318,815.754 km2) which would render this species as Least Concern. On the other hand, its AOO is considerably smaller (ca. 88.000 km2), which would render H. pumila as Endangered. The Paraná, Uruguay, and Southeastern Atlantic watersheds cover eight Brazilian states (Distrito Federal, Goiás, Mato Grosso do Sul, Minas Gerais, Rio Grande do Sul, Santa Catarina, São Paulo, and Paraná), embedded in the Atlantic Forest and Cerrado domains. Its main tributaries are the Iguaçu, Paranaíba, Paranapanema, Rio Grande and Tietê rivers. It possesses the greatest energy generation potential in Brazil, with 176 active hydropower plants, the largest being Itaipu, Furnas, Porto Primavera and Marimbondo. Nonetheless, all the major rivers are currently saturated with hydropower plants, and new projects aim to occupy the smaller tributaries, in order to fulfil the growing energy demand in the region ( ANA 2002). Almost all the known subpopulations of H. pumila coincide with areas currently flooded, and might already have gone extinct, due to the construction of the aforementioned water dams. The few extant subpopulations vary from medium to large, with many clones and mature individuals. Nonetheless, they are currently strongly threatened due to pollution, deforestation, and by ongoing and future constructions of new hydropower plants. Thus, following the IUCN recommendations ( IUCN 2001), H. pumila should be considered as Critically Endangered [CR, A2acd+B1b(ii, iii, iv)+B2ab(ii, iii, iv)+C1+E].

Morphological notes.

Extensive morpho-ecological studies ( Horn 1983, 1988) have shown that Heteranthera species are highly polymorphic vegetatively, as an adaptation to submersion and variations in water level. The same can be observed in the new species herein described, that despite the diminute general stature, may sometimes possess extremely long petioles and peduncles. Heteranthera pumila has been kept in cultivation by the senior author, and even under different environmental conditions, little change was observed in the species’ vegetative morphology. Nevertheless, when cultivated in aquariums with different water depths, the change in the length of petioles and peduncles could be observed in less than a week. The already existing structures elongated in order to keep the leaf blades floating and flowers emerged, and the subsequently produced petiolate leaves and inflorescences were considerably longer than the previous ones of the same individual.

Affinities.

Heteranthera pumila is morphologically similar to H. reniformis due to its petiolate leaves with blades two or more times wider than long, cordate to reniform, rarely narrowly cordate, cincinnus enclosed by the basal bract, glandular-pubescent cincinnus axis, perianth lobes with a 5+1 arrangement with acute to acuminate apex, filaments straight, lateral stamens apically barbate, central stamen basally sparsely villose, and intrusive-parietal placentation ( Horn 1985). It is also similar to H. multiflora due to its petiolate leaves with blades two or more times wider than long, cordate to broadly cordate to reniform, rarely narrowly cordate, perianth lobes with a 5+1 arrangement and acute to acuminate apex, and straight filaments ( Horn 1985). Nonetheless, it can be easily differentiated from all remaining species of Heteranthera by its petiolate leaves with diminute blades [i.e. 3.5-11.8-(13.2) × 3.2-12.1 mm], inflorescences 1-2-(3)-flowered, peduncle densely glandular-pubescent, basal bract basally glandular-pubescent with aristate apex, and seeds smooth or with 7-9 inconspicuous longitudinal wings (Fig. 5 ). The only other species in Heteranthera that possesses seeds not conspicuously winged is H. gardneri , in which the wings are very short, giving the seeds a striate appearance. Nevertheless, in H. pumila , the testa is almost smooth, with the stripes representing only pigmentation. All the remaining species of Heteranthera possess seeds with 8-19 conspicuous longitudinal wings ( Horn 1985; Table 1 ).

Figures

Figure 2. Line drawing of Heteranthera catharinensis C. N. Horn & M. Pell. A Detail of the leaf blade B Detail of the apex of the stem, showing the ligule and an inflorescence at anthesis C Detail of the basal bract, showing the spatulate-mucronate apex D Glandular hair from the cincinnus axis and floral tube E Dissected perianth lobes, showing the 5 + 1 arrangement F Lateral stamen G Uniseriate hair from the lateral stamen H Central stamen I Gynoecium, showing the glabrous style and unevenly trilobate stigma. Illustration by M. O. O. Pellegrini, based on the paratype (Smith & Reitz 9103, US).

Figure 3. Distribution map. ■ Heteranthera catharinensis C. N. Horn & M. Pell. ● H. pumila M. Pell. & C. N. Horn. Green- Parana watershed; Yellow- Uruguay watershed; Red- Southeastern Atlantic watershed; following ANA - Agencia Nacional de Aguas (2002).

Figure 4. Field photos of Heteranthera pumila M. Pell. & C. N. Horn. A Bog at the Paranapanema river, Piraju, Sao Paulo, Brazil B Habit, showing the dense subpopulation at the muddy shore of the bog C Leaf D Detail of the apex of the stem, showing the ligule and the inflorescence E Detail of the inflorescence, showing the glandular hairs at the peduncle, base of the basal bract and cincinnus F Front view of the flower, the shape of the perianth lobes and the color of the nectar guide. Photographs A-E by M. O. O. Pellegrini, F by V. Bittrich.

Figure 5. Line drawing of Heteranthera pumila M. Pell. & C. N. Horn. A Habit B Detail of the apex of the stem, showing a petiolate leaf, the ligule and a pre-anthesis 2 - flowered inflorescence C Glandular hair from the inflorescence, perianth tube and lobes D Dissected perianth lobes, showing the 5 + 1 arrangement E Lateral stamen. F Uniseriate hair from the lateral stamen G Central stamen H Eglandular hair from the central stamen I Gynoecium, showing the stigma J Eglandular hair from the style K Detail of the inconspicuously winged seed, showing the persistent funiculus with raphid crystals. Illustration by M. O. O. Pellegrini, based on the holotype.