Guatteria hirsuta Ruiz & Pav.

Maas, P. J. M., Westra, L. Y. T., Guerrero, S. Arias, Lobão, A. Q., Scharf, U., Zamora, N. A. & Erkens, R. H. J., 2015, Confronting a morphological nightmare: revision of the Neotropical genus Guatteria (Annonaceae), Blumea 60 (1), pp. 1-219 : 95-97

80. Guatteria hirsuta Ruiz & Pav. — Fig. 36d, e , 43 ; Plate 4g ; Map 18

Guatteria hirsuta Ruiz & Pav. (1798) 146; R.E.Fr. (1939) 369, f. 9b. — Type: Ruiz s.n., B100242419 (lecto B, here selected; isolecto B, BM, BR, F, G 3 sheets, HAL, P, W), Peru, Huánuco, Chinchao (‘in nemoribus versus Chinchao vicum’).

Guatteria excelsa Poepp. ex Mart. (1841) 38, syn. nov.; R. E.Fr. (1939) 371, f. 9c, d. — Type: Poeppig 139 (1467) (holo W; iso B, BM, F, G, HAL, P), Peru, Huánuco, Pampayacu (‘in submontanen Wäldern bei Pampayaco’), Oct. 1832 .

Guatteria pilosula Triana & Planch. (1862) 31, syn. nov.; R. E.Fr. (1939) 369. — Type: Linden 767 (holo P; iso BR, G), Colombia, Santander (‘ Las Lajitas, entre Pie de Cuesta et los Santos, prov. del Socorro’), 1300 m, Nov. 1842 .

Guatteria cestrifolia Triana & Planch. (1862) 32; R. E.Fr. (1939) 366, f. 9a, syn. nov. — Type: Triana s.n. (holo P; iso BM, F, G, K, P, S, fragment), Colombia, Meta, Apiay (‘Prov. de Bogotá, basin du Meta, Apiai’ ), 300 m.

Guatteria laurina Triana & Planch. (1862) 32; R. E.Fr. (1939) 366, t. 23, syn. nov. — Type: Goudot s.n. (holo P; iso K), Colombia, Meta, Irica (‘haut Orénoque, pueblo de Iraca’), Dec. 1844 .

Guatteria longipes Triana & Planch.(1862) 32; R. E.Fr. (1939) 516,t. 35,syn. nov. — Type: Triana s.n. (holo P; iso BM, COL, F, G, K), Colombia,Cundina- marca (‘Prov. de Bogotá, Biota, Andes de Bogotá’ ), 2000 m, Sept. 1853 .

Guatteria persicifolia Triana & Planch. (1862) 33, non J. D.Hook. & Thomson 1855; R.E. Fr. (1939) 367, syn. nov. — Type: Schlim 700 (holo P; iso BM, BR 2 sheets, G 2 sheets, P), Colombia, Norte de Santander, Enllanada, 1200 m (‘prov. de Ocaña, Enllanada , 3500 ped., June 1846 –1852’).

Guatteria alutacea Diels (1905) 126, syn. nov. — Type: Ule 6427 (holo B; iso F, G, K, L, MG), Peru, San Martín , Tarapoto, Oct. 1902.

Guatteria juruensis Diels (1905) 126, syn. nov. — Type: Ule 5010 (holo B; iso F, G, K, L, MG, S), Brazil, Amazonas , Rio Juruá, Marary, Sept. 1900.

Guatteria augusti Diels (1924a) 51,syn.nov. — Type: Weberbauer 7062 (holo B; iso F, S, US), Peru, La Libertad, Prov. Patáz, Río Mixiollo, left branch of Río Huallaga, below Ongón , 1200–1300 m, 7 Aug. 1914.

Guatteria guentheriana Diels (1931) 75; R. E.Fr.(1939) 361, syn.nov. — Type: Buchtien 1773 (holo B 2 sheets; iso F, NY, US), Bolivia, La Paz, Mapiri Region , San Carlos, 850 m, 2 May 1927 .

Guatteria alutacea Diels var. angustifolia R.E.Fr. (1938) 710, syn. nov. — Guatteria alutacea Diels f. angustifolia (R.E.Fr.) R.E.Fr.(1939) 362. — Type: Spruce 4270a (holo B; iso BM,BR, E, G, K, NY,P, S, W), Peru, San Martín , Tarapoto, 1855–1856.

Guatteria klugii R.E.Fr. (1938) 710, syn.nov. — Type: Klug 3299 (holo S; iso F, G, K, MO, NY, S, US), Peru, San Martín , Zepalacio, near Moyobamba, 1100–1200 m, Oct.–Nov. 1933.

Guatteria brevipedicellata R.E.Fr. (1939) 359, syn. nov. — Type: Lawrance 771 (holo S; iso E, F 2 sheets,G, K 2 sheets,MO, US 2 sheets), Colombia, Boyacá, El Humbo, high forested fronts, 1250 m, 27 Apr. 1933.

Guatteria alutacea Diels var. steinbachii R.E.Fr. (1939) 362,syn.nov. — Type: Steinbach 6486 (holo S; iso BM, E, F, G, K, MO, NY, PH), Bolivia, Santa Cruz, prov. Sara, Buenavista , 450 m, 27 Sept. 1924.

Guatteria ecuadorensis R.E.Fr. (1939) 364, syn. nov. — Type: Mexia 7111 (holo S; iso F, K, NY,US), Ecuador, Napo, Zatzayacu , 400–500 m, 22 Mar. 1935.

Guatteria melinii R.E.Fr. (1939) 364, f. 26d, e, syn. nov. — Type: Melin 122 (holo S), Peru, San Martín , San Roque , SE of Moyobamba, 9 May 1925.

Guatteria jamundensis R.E.Fr. (1939) 365, f. 8b, syn. nov. — Type: Ducke MG 11780 = RB 35322 (holo S; iso RB, S), Brazil, Pará , Lower Rio Ja- mundá, 18 May 1911.

Guatteria latisepala R.E.Fr. (1939) 368, syn. nov. — Type: Lehmann K 27 (holo S;iso F, GH,K), Colombia, Cauca,Popayán, El Hatico, 1500–1800 m, 20 Mar. 1905.

Guatteria curvipetala R.E.Fr. (1939) 526,f. 36a, b, syn.nov. — Type: Krukoff 6600 (holo S; iso BM, BR, F, G, K, MO, NY, RB 2 sheets, U, US), Brazil, Amazonas , Basin of Rio Madeira , Mun. Humaitá, Livramento, on Rio Livramento, 12 Oct. 1934.

Guatteria metensis R.E.Fr. (1948b) 6, pl. 2 a, b, syn. nov. — Type: Haught 2470 (holo S 2 sheets; iso COAH, COL, NY, S, US 2 sheets), Colombia, Meta, Llanos and along Río Ocoa , 400 m, 19 Dec. 1938.

Guatteria chocoensis R.E.Fr. (1950a) 333, syn. nov. — Type: Cuatrecasas 16592 (holo S 2 sheets; iso COL, F, US), Colombia, Valle del Cauca, Río Calima (región del Chocó), La Trojita, 5–50 m, 29 Feb. 1944.

Guatteria sp. 13 Chatrou et al. (1997) 111.

Tree or rarely a shrub 2.5–30(–42) m tall, 5–60 cm diam; young twigs densely to sparsely covered with erect to appressed, brown hairs 1.5–2 mm long, soon glabrous or long-persisting. Leaves: petiole 2–5(–10) mm long, 0.5–2 mm diam; lamina narrowly elliptic to narrowly oblong-elliptic, sometimes narrowly obovate, 7–22 by 1.5–6 cm (leaf index (2.3–)3.3–4.8(–8)), chartaceous, not verruculose, shiny or dull above, greyish, greyish black to brown above, brown to greyish brown, rarely greenish below, sparsely covered with erect to half-appressed hairs to glabrous above, the primary vein to densely so, becoming glabrous with age, densely to sparsely covered with erect to appressed hairs 1– 2 mm long, primary vein to densely so below, becoming glabrous or almost so with age, base acute to obtuse, rarely cordate or attenuate, apex acuminate (acu- men 5–20 mm long), primary vein flat, impressed or slightly raised above, secondary veins distinct, 8–15 on either side of primary vein, raised or sometimes impressed above, smallest distance between loops and margin 1–3 mm, tertiary veins raised above, reticulate. Flowers in 1(–3)-flowered inflores- cences in axils of leaves or on leafless branchlets; pedicels 10–40(–80) mm long, 1–1.5 mm diam, fruiting pedicels 2–3 mm diam, densely to sparsely covered with erect to appressed, brown hairs, articulated at 0.2–0.5 from the base, with the part above the articulation sometimes thicker than the part below, bracts 5–7, soon falling, basal ones broadly elliptic to elliptic, 1– 2 mm long, the 2 upper ones and sometimes also a third bract below them elliptic to broadly elliptic, 4–10 mm long, oc- casionally foliaceous, to c. 45 mm long; flower buds depressed ovoid, pointed; sepals free, broadly ovate-triangular, 3–12 by 3–9 mm, appressed, finally reflexed, outer side densely to sparsely covered with erect or appressed hairs; petals green, maturing yellowish green, yellow, or white in vivo, ovate to obovate or narrowly so, 8–35 by 3–15 mm, outer side densely covered with appressed or erect hairs, particularly at the base; stamens 1–1.5 mm long, connective shield papillate to glabrous. Monocarps 10–100, green, maturing purplish to black in vivo, black in sicco, ellipsoid, 8–13 by 3–7 mm, sparsely covered with appressed hairs, soon glabrous, apex apiculate (apiculum 0.1–2 mm long), wall 0.1–0.3 mm thick, stipes 8–25 by 0.5–1 mm. Seed ellipsoid, 6–10 by 3–6 mm, dark, shiny brown, pitted, sometimes also transversely grooved, raphe raised.

Distribution — Colombia (Amazonas, Antioquia, Cauca, Chocó, Huila, Meta, Norte de Santander, Santander, Valle del Cauca), Venezuela (Apure, Barinas, Falcón, Lara, Me- rida, Portuguesa, Táchira, Trujillo, Zulia), Ecuador (Azuay, Morona-Santiago, Napo, Pastaza, Zamora-Chinchipe), Peru (Amazonas, Cajamarca, Loreto, Madre de Dios, Pasco, Puno, San Martín ), Brazil (Acre, Amazonas, Pará), Bolivia (Beni, Cochabamba, La Paz, Santa Cruz).

Habitat & Ecology — In non-inundated, rarely periodically inundated (várzea) lowland, montane or submontane forest, sometimes in savannas, on sandy to limestone soil. At eleva- tions of 0–2000(–2400) m. Flowering: throughout the year; fruiting: August to April.

Vernacular names — Bolivia: Negrillo (Steinbach 6486), Pi- raquina macho (Zonta et al. 104), Piraquina negra (Arrazola 16, Del Aguila et al. 364, Meneces & Zabala 356, Moraes R. 940, Zabala & Terceros s.n.). Brazil: Envira-caju ( Daly et al. 8325), Envira-tambaqui (Sothers & Araujo 18). Colombia: Anona de monte (Little 8731), Cargadero (Cuatrecasas 16592), Garrapato (Cárdenas L. & Ramirez 2654, Zarucchi 3378), Suta (Lawrance 771). Peru: Carahuasca (Croat 19216, Vásquez 6686, Vásquez & Jaramillo 10321), Espintana (J. Ruíz & Murphy 276). Ven- ezuela: Laurel (Veillon 85), Magua negra (Breteler 3922).

Field observations — Nee 37977 (U) from Bolivia has ‘spicy fragrant flowers’. Amaral et al. 457 (U) from Brazil has immature fruits emitting a scent of ripe bananas (‘frutos imaturos exalando cheiro de banana madura’).

Notes — The main features of G. hirsuta , belonging to sect. Trichoclonia , are narrow, often greyish black coloured leaves, an indument of erect to appressed, brown hairs on the leafy twigs, relatively long pedicels and pitted seeds.

We decided to unite here four species described by Triana and Planchon in the same paper (1862). These species differ from each other only in minor aspects (among others, leaf base, leaf indument, shape of petals), but examination of more specimens than must have been at the disposal of Triana and Planchon makes obvious that there are many intergradations.

It is with some reluctance that we united G. latisepala with G. hirsuta . Guatteria latisepala is found mostly at higher elevations (c. 800 m and up), and many specimens look quite distinct by a more dense indument and by markedly thickened pedicels above the articulation point. There are too many transitional forms, though, to justify keeping the two species apart.

Guatteria alutacea as described at first seems well distinct by the greyish to greyish black colour of the upper side of the leaves, the mostly obtuse to acute leaf base, and often the presence of rather long appressed and forward-pointing hairs on the lower side of the lamina. However, there are also many intergradations here that make it impossible to uphold G. alutacea as a separate species.

Forms described as G. alutacea var. angustifolia are noticeable because of very narrow and attenuate leaves with a leaf index up to 8 and generally long pedicels, but otherwise are not clearly distinct.

Guatteria longipes , known only from the type collected in the Colombian state of Cundinamarca (‘Prov. de Bogotá, Biota, Andes de Bogotá’ ), at an elevation of 2000 m, differs by pedicels 50–90(–120) mm long !

Some forms, all from the Colombian state of Chocó, are note- worthy because of their tendency towards longer-than-average pedicels: Cuatrecasas & Llano 24070 (to 80 mm long!), Gentry et al. 17844 and 30242. Apart from that, they match well with the overall description.

Figures

Plate 4 a. Guatteria ferruginea A.St.-Hil. Stem with flowers and fruit. – b, d, e. Guatteria goudotiana Triana & Planch. b. Fruit; d. flowering branch; e. flower. – c, f. Guatteria heteropetala Benth. c. Fruiting branch; f. flowering branch. – g. Guatteria hirsuta Ruiz & Pav. Fruiting branch. – h, i. Guatteria inundata Mart. h. Flowering branch; i. fruiting branch (a: Amorim et al. 818; b, d, e: Sánchez S. et al. 4776; c: Stevenson et al. 1115; f: Berry s.n.; g: Clark et al. 5291; h: Croat 19234; i: Maas et al. 6288). — Photos: a, i: P.J.M. Maas; b, d, e: D. Sánchez S.; c: D.W. Stevenson; f: P.E. Berry; g: J.L. Clark; h: T.B. Croat.

Fig. 36 a. Guatteria elata R.E.Fr. Flower. – b. Guatteria fractiflexa Maas & Westra. Flower. – c. Guatteria herrerana N.Zamora & Maas. Fruit on stem. – d, e. Guatteria hirsuta Ruiz & Pav. d. Flowers; e. fruit. – f, g. Guatteria maguirei R.E.Fr. f. Flowers; g. fruit. – h. Guatteria maypurensis Kunth. Fruit (a: Daly et al. 5797, U; b: Vásquez 27596, U; c: G. Herrera C. 8401, holotype part CR; d: Killeen 2802, U; e: W. Palacios 7526, U; f: O. Huber 5130, U; g: Huber & Tillett 2888, U; h: Zarucchi & Barbosa 3828, U).

Fig. 43 Guatteria hirsuta Ruiz & Pav. Flowering branch (Nee 37977, U).