Aphelandra martiusii Wasshausen (1975: 108)

Villanueva-Espinoza, Rosa, Azevedo, Igor H. F. & Montenegro, Luis Torres, 2022, Taxonomic update and first record of Aphelandra martiusii (Acanthaceae) in Peru, Phytotaxa 549 (2), pp. 199-208 : 200-205

Aphelandra martiusii Wasshausen (1975: 108) ( Fig. 1–3 )

Replaced name: Lagochilium repandum Nees von Esenbeck (1847a: 86) , not Aphelandra repanda Nees von Esenbeck (1847b: 728) .

Type: — BRAZIL. Pará: “In silvys ad fluvium Xingú”, 10–12 September 1819, C. F. P. Martius s.n. (lectotype designated here: M0186188 [image!]; isolectotypes: M0186187 [image!], M0186190 [image!]) .

Shrub 0.6-3 m tall, erect, cystoliths absent. Stem terete, glabrous. Leaves opposite-decussate, isophyllous; petiole 0.5–1.3 cm long, puberulent; blade submembranaceous, 25–32.5 × 4–9.6 cm, oblong-lanceolate to elliptic, apex acuminate to attenuate, base decurrent, margin entire, venation eucamptodromous, lateral veins (11–) 16–19 pairs, adaxially flat, glabrous, greenish, abaxially puberulent along veins, pale green. Inflorescence a terminal spike, 7–10.7 cm long, quadrangular, congested; peduncle 5 mm long, puberulent; rachis with flowers opposite, glabrous; inflorescence bracts foliose, mid-vein pubescent, green to red. Flowers zygomorphic, imbricate; bracts red to pink, papyraceous, conduplicate, 3.5–3.9 cm long, 2.4–2.6 cm wide, obovate-cuneate, reticulate-nerved, glabrous, apex obtuse-mucronulate; bracteoles pink, papyraceous, (1.8–) 2.3–2.4 × (0.4–) 0.5–0.6 mm, oblong, apex asymmetric, glabrous. Calyx papyraceous, lobes subequal, quincuncial aestivation, oblong, apically acute, margin entire, posterior sepal ca. 15 × 2.5 mm, anterior pair ca. 14 × 2.5 mm, lateral pair ca. 8 × 1.5 mm, glabrous. Corolla bilabiate, (4–) 6.2 cm long, erect, purple, glabrous outside, puberulent inside; tube erect, ca. 3 cm long, ca. 1.5 mm wide at base to ca. 3 mm wide at top, then gradually enlarged to ca. 8 mm wide at mouth; mouth oblique; upper lip (0.8–) 1.2 × (0.8–) 1.2 cm, cucullate, ovate, apex retuse-revolute; lower lip 3-lobed, ca. 1.3 cm long, lobes subequal, ovate, middle lobe (4–) 6 × 6 mm, lateral lobes ca. 5 × 4 mm. Stamens 4, slightly didynamous, included; filaments ca. 2 cm long, puberulent; anthers monothecous, thecae ca. 7 mm long, apically obtuse, a tuft of long hairs on the abaxial surface. Ovary ca. 5 mm long, pyriform, glabrous; style ca. 2.4 cm long, filiform, glabrous; stigma ca. 0.5 mm long, bifid. Capsule slightly obovoid, 14–16 × 6–7 mm, glabrous. Seeds not seen.

Distribution and habitat: — Aphelandra martiusii is an endemic species to South America, distributed in Brazil, Colombia and Peru ( Fig. 4 ). In Peru, it is only known from the northeastern Amazonian Department of Loreto. All collections are from the province of Mariscal Ramón Castilla (Locality of San Juan de Loretoyacu) and Putumayo (Cotuhé Camp), belonging to the territory of the Cotuhé River Conservation Concession.

Aphelandra martiusii grows at floodplain forests on uneven terrain (IH 14380, 14517) and low terrace forest (IRP 2648) mostly undisturbed, at an elevation of 115– 140 m. In the Cotuhé Camp (IH 14380, 14517), this species is found associated with some arboreal species such as Coussapoa trinervia Spruce ex Mildbraed (1928: 416) , Micrandra spruceana ( Baillon 1864: 288) R.E. Schultes (1952: 217) , Mabea nitida Spruce ex Bentham (1854: 367) and Ficus guianensis Desvaux ex Hamilton (1825: 62) ; some ferns from genus Adiantum Linnaeus (1753: 1094) , Lindsaea Dryander ex Smith (1793: 413) and Trichomanes Linnaeus (1753: 1097) in the understory, and some Rubiaceae shrub species ( Pitman et al. 2011).

Phenology: —This species was observed in flowering from March to October, showing some falling flowers in this later month, and fruiting in May.

Conservation status: — Aphelandra martiusii is endemic to the Amazon Forest, growing in Brazil, Colombia and Peru. In Brazil, this species is only known from its type locality (banks of Xingu river, Pará) having no information of its current possible threats. Therefore, this species should be listed as DD. In Colombia, it is widespread, growing in more than 10 localities with a total EOO> 20,000 km ² and an AOO> 50 km ². In Peru, A. martiusii is only known from two localities (San Juan de Loretoyacu and Cotuhé Camp) belonging to the Cotuhé River Conservation Concession. Despite being close to the Yaguas National Park, Cotuhé is currently being highly affected by deforestation for agricultural purposes, leading to the loss of several hectares of land ( Novoa et al. 2021). A negative impact on the habitat of Aphelandra martiusii is therefore expected.The Colombian populations meet the criteria for Near Threatened; however, the species is classified as [EN (B2a+biii)] in Peru. Therefore, we assign this species to the category NT (AOO<2,000 km ², continue declining,> 10 locations) ( IUCN 2012a, 2012b, 2019).

Specimens examined:— BRAZIL. Pará: ‘’ In silvis ad fluvium Xingú. ’’, 10–12 Sep 1819, C . F . P. Martius s. n. ( GZU, M) . COLOMBIA. Amazonas: Tarapacá, ‘’ Río Porvenir Grande cerca de Caño Pajaro. Bosque de zona plana temporalmente inundado.’’, 2°31’24.24”S, 70°11’06”W, 12 Feb 2006, M. Rodríguez & J. Aguila 10 ( COAH); ‘’Corregimiento Puerto Arica, asociación AIZA, comunidad Puerto limón. Bosque de tierra firme ondulado, con transición de llanura aluvial’’, 2°17’50.6”S, 72°01’31.8”W, 97 m, 15 Nov 2017, M . Rodríguez et al. 3530 ( COAH!); ‘’ Zona étnica de conservación de Puerto Caimán, asociación AIPEA , resguardo indígena Curare-Los Ingleses. ’’, 8 Apr 2018, A . Barona-C. 4952 ( COAH); ‘’ Rio Caquetá, km 20 trocha Santa Isabel-Mirití. ’’, 18 May 1980, M . Pabón E . 1000 (COAH!, COL!). Vaupés: “Porto dos Miranhas dit. [ditionis] Japurensis, Prov. Fl. Nigri ”, Dec 1819, C . F . P. Martius s. n. ( GZU, M) . PERU. Loreto: Prov. Mariscal Ramón Castilla, Dist. Caballo Cocha, San Juan de Loretoyacu , 115 m, 03°25’44.22”S, 70°45’30.25”W, 3 Nov 2019 (fl.), I . Revilla Pantigoso 2648 ( MOL). Prov. Putumayo, Dist. Yaguas, Inventario Rápido de las cabeceras del río Cotuhé , NE de Iquitos y Pebas, en la esquina del trapezoide de Colombia. Drenaje del bajo Putumayo , entre los ríos Yaguas y Amazonas. Camp 2, “Cotuhé”, borde sur, 140 m, 3°13’32.4”S, 70°54’93”W, 21 Oct 2010 (fl.), I . Huamantupa, Z . Cordero, N . Pitman & R . García 14380 ( AMAZ, F). Same collection data as preceding, 135 m, 3°11’40.7”S, 70°52’45.9”W, 23 Oct 2010 (fl.), I . Huamantupa, Z . Cordero, N . Pitman & R . García 14517 (AMAZ, F).

Lectotypification: —In the protologue of Lagochilium repandum, Nees von Esenbeck (1847a) did not cite any specimens but mentioned two localities where the species was collected: (1)“ In the woods at the river Xingu, prov . Paraensis ” and (2)“ In the woods at Porto dos Miranhas under the jurisdiction of Japurensis ”. Four duplicates housed at Herbarium M (M0186187, M0186190, M0186188 and M0186189) were traced. The handwriting on the two first sheets “ Brasilia prov . Pará ” and on the third sheet “ Habitat in sylvis ad fluv. Xingú. Provinciae Paräensis ” [Prov. Paraensis] indicating that these materials were collected by Martius in Pará, mainly close to the Xingu river, dating 10-12 September 1819. The another sheet M0186189 has the label “ in sylvis ad Porto dos Miranhas dit. Japurensis. Provinciae R. N. ” [Porto dos Miranhas, prov. Rio Negro, district of Japurensis]. Martius arrived at Rio Negro in December 1819, few months after his visit to Pará ( Spix & Martius 1938); however, this district currently belongs to Vaupés, a department of Colombia ( Wasshausen 1975). Among these four sheets, M0186188 and M0186189 bear the original blue label of Martius, while the sheets M0186187 and M0186190 are duplicates. These specimens may be the original material and they are part of the Martius’ collection which was obtained during his exploration to the Amazonian basis of Brazil between 1817 and 1820 ( Spix & Martius 1938). Additionally, another sheet of L. repandum was found in GZU (GZU000250146). This material was given by Baron von Zschock to the University of Graz and belonged to the personal herbarium of Nees which was split up after his suspension as a professor at the University of Breslau ( Steenis-Kruseman et al. 1950; Stafleu & Cowan 1981). In the sale of his herbarium and library, Nees von Esenbeck (1852) highlighted the historical significance of this material “ it is rendered still more important by its containing the original specimens on which my monographie labours, the dissertations on the Laurineae, Solaneae, Acanthaceae, Hepaticae , Asteraceae , Cyperaceae , Gramineae, and Restiaceae, are founded ”. Accordingly, the sheet housed at GZU is also the original material of Lagochilium repandum . Since there is no holotype indicated by Nees von Esenbeck (1847a), all of these specimens are syntypes according to the Art. 9.6 of Shenzhen Code ( Turland et al. 2018).

Prior to the World Wide II, Francis MacBride travelled to Europe to photograph the type specimens of tropical American plants preserved at several European herbaria, including Munich (M) ( Grimé & Plowman 1986). MacBride achieved to take photos of the syntypes of L. repandum housed at M and these photos are available online, F20492 (https://fm-digital-assets.fieldmuseum.org/288/549/20492.jpg) and F20498 (https://fm-digital-assets.fieldmuseum. org/288/481/20498.jpg). However, these photos cannot be considered as syntypes of L. repandum because these do not include any parts from the original material

Despite being the one specimen with flowers, the duplicate housed at GZU is probably a mixed preparation of both syntypes, as it shows a label of both localities. Additionally, it is uncertain to which syntype belongs every fragment. Consequently, the duplicate GZU000250146 is discarded as lectotype. Among the remaining duplicates, the sheet M0186188 was the only one which bears a blue label of Martius with a note “ Bractea et cor. purpureae ”, the one collected in Pará and the first locality where this sample was collected by Martius. As discussed above, we here designate a duplicate of Martius s.n. at M (barcode 0186188) as the lectotype for the name Aphelandra martiusii .

Discussion: —In Peru, Aphelandra martiusii is currently found in the city of San Juan de Loretoyacu, district of Caballo Cocha, province of Mariscal Ramon Castilla and Camp Cotuhé, district of Yaguas, province of Putumayo, both bordering to Colombia. This might be explained by the fact that Putumayo and Caballo Cocha are close to the department of Amazonas, Colombia, where recent material (Rodríguez et al. 3530) was collected to still confirm its existence in the jungle of this country. On the other hand, collections from Brazil solely date back from Martius’ exploration (1817-1820) when original materials were collected ( Zanatta et al. 2020). This only one occurrence in Brazilian territory could owe to the missing of botanical explorations in the zone or otherwise, its extinction. The long distance between the known populations of A. martiusii might be related to a water dispersal through the Amazonas River. Similar situation has been observed in the genus Suessenguthia Merxmüller (1953: 178) , with a sole collection in Pará ( Brazil), and the others individuals over 2,000 km westward ( Schmidt-Lebuhn 2003).

The Peruvian species of Aphelandra may be separated into two groups: species with entire leaves and those with spiny leaves. The species in former group can be distinguished by the spiny or entire floral bracts. A. martiusii along with Aphelandra plowmanii Wasshausen (2013a: 109) and Aphelandra latibracteata Wasshausen (1973: 468) shares the presence of leaves and floral bracts entire and the cucullate corolla. Aphelandra latibracteata and A. plowmanii , endemic species to Peru, are known in Huánuco, San Martin and Ucayali, and Loreto and San Martin, respectively. These three species can be confusable by their inflorescence spike and corolla shape, however, A. martiusii differs by having obovate-cuneate floral bracts, larger bracteoles and a pink corolla. More detail of the diagnostic characters is summarized in Table 1 .

Figures

FIGURE 1. Aphelandra martiusii. A. Habit. B. Detail of the elliptic leaves, without cystolith. C. Terminal quadrangular spike. D. Young bilabiate flower. From I. Huamantupa s.n. (B) y 14517 (A, C, D). Photos by Isaú Huamantupa (A-D).

FIGURE 2. Aphelandra martiusii. A. Fertile branch with terminal inflorescence. B. Bract. C. Flower basally enclosed by a pair of bracteoles. D. Bracteoles E. Calyx. F. Open corolla, containing the androecium and gynoecium. G. Stamen. H. Stigma. I. Capsule. From I. Revilla 2648 (MOL) and M. Pablón 1000 (COAH). Drawing by Igor Azevedo.

FIGURE 3. Lectotype of Aphelandra martiusii Wasshausen at M (barcode M0186188). [© Reproduced with the consent of the herbarium of the Botanische Staatssammlung Münche, Munich].

FIGURE 4. Distribution map of A. martiusii in South America.