Guatteria punctata (Aubl.) R.A.Howard

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 : 138-141

130. Guatteria punctata (Aubl.) R.A.Howard — Fig. 63 ; Map 28

Guatteria punctata (Aubl.) R.A. Howard (1983) 260. — Annona punctata Aubl. (1775) 614, t. 247. — Type: Aublet s.n. (lecto BM, selected by Howard 1983), French Guiana, Sinnamary (‘Habitat in sylvis Sinemariensibus & prope amnem Galibiensem’).

Guatteria glauca Ruiz & Pav.(1798) 145; R.E.Fr.(1939) 384,f. 13a, b; Maas & Westra (2010) 267, pl. 1, syn. nov. — Type: Ruiz L. s.n. (syn B, BR, F, G, HAL, K, MA, NY, P), Peru, Huánuco, ‘ Habitat in Peruviae in nemoribus versus Cochero et Chinchao vicos’, anno 1798.

Annona axilliflora DC. (1824) 86. — Guatteria axilliflora (DC.) R.E.Fr. (1939) 427, f. 15a. — Type: Martin 151 (holo G; iso C, K 2 sheets, P 4 sheets), French Guiana (‘Cayenne’), without location.

Guatteria caniflora Mart. [ var. caniflora ] Mart.(1841) 37,syn.nov. — Guatteria caniflora Mart.var. angustifolia Mart.(1841) 37;R. E.Fr. (1939) 442. — Type: Martius s.n. (holo M), Brazil, Amazonas , Rio Solimões (‘prov. Rio Negro, in sylvis inundatis ad fluv. Solimões’), Nov.–Dec. 1819 .

Guatteria caniflora Mart. var. latifolia Mart. (1841) 37; R.E.Fr. (1939) 442, syn. nov. — Type: Martius s.n. (holo M, not seen), Brazil, Amazonas , Rio Solimões , Dec. 1819 (‘prov. Rio Negro, in sylvis ad fluvium Solimões’).

Guatteria poeppigiana Mart. (1841) 37 (as ‘ pÖppigiana ’); R. E.Fr. (1939) 424, syn. nov. — Type: Poeppig D 2963 (lecto W, selected by Fries 1939, not seen), Brazil, Pará , Colares (‘in silvis ad collares’), June 1838 .

Annona chrysopetala Steud.(1843) 754. — Guatteria chrysopetala (Steud.) Miq. var. chrysopetala (1849) 466; R. E.Fr. (1939) 434. — Type: Hostmann & Kappler 1295, ed. Hohenacker = Hostmann 1295 (holo P; iso B, BM, G 3 sheets, GH, K, MO, P, S, U, UPS, W), Suriname, Suriname River .

Guatteria pteropus Benth. (1853) 8, syn. nov. — Type: Spruce 1680 (holo K; iso BM, E, G 3 sheets, K, M, NY, P), Brazil, Amazonas , mouth of Rio Negro, Aug. 1851.

Guatteria cargadero Triana & Planch.(1862) 34; R. E.Fr. (1939) 380,syn.nov. — Type: Triana s.n. (holo P; iso B, BM, BR, G, W), Colombia, Valle del Cauca, Anserma , 1000 m .

Guatteria platyphylla Triana & Planch.(1862) 35;R.E.Fr. (1939) 449,syn.nov. — Type: Triana s.n. (holo P; iso BM, COL), Colombia, Cundinamarca, Servita, 1000 m, (‘ Servita , versant oriental des Andes de Bogota, alt. 800 mêtres’).

Guatteria sylvicola S. Moore (1895) 298, syn. nov. — Type: Spencer Moore 142 (holo BM; iso B), Brazil, Mato Grosso, Serra da Chapada , 600 m, Aug. 1891.

Guatteria coeloneura Diels (1906) 408; R. E.Fr. (1939) 387, f. 13c, syn. nov. — Type: Weberbauer 3548 (holo B 3 sheets; iso F, G, S), Peru, Huánuco, SW of Monzón , 1600 m, 8 Aug. 1903 .

Guatteria pleiocarpa Diels (1906) 409; R. E.Fr. (1939) 387, f. 12e, syn. nov. — Type: Weberbauer 4530 (holo B 3 sheets; iso F, G 4 sheets, GH), Peru, Loreto, Moyobamba , 800–900 m, 17 Aug. 1904 .

Guatteria guentheri Diels (1927) 169; R. E.Fr. (1939) 433, f. 14b, c, syn. nov. — Type: Tessmann 4387 (holo B 2 sheets; F fragment, NY, S fragment), Peru, Loreto, Upper Río Marañon, near mouth of Río Santiago , 160 m, 27 Oct. 1924 .

Guatteria atra Sandwith (1930) 468, syn. nov. — Type: Sandwith 406 (holo K; iso B, FDG, K 2 sheets, NY, P, RB, U, US), Guyana, Moraballi Creek , Essequibo River, 9 Oct. 1929.

Guatteria acutissima R.E.Fr. (1938) 712;(1939) 525,t. 37, syn. nov. — Type: Klug 1268 (holo F; iso NY, US), Peru, Loreto, Mishuyacu, near Iquitos , 100 m, Apr. 1930.

Guatteria calliantha R.E.Fr. (1938) 715; (1939) 429,syn. nov. — Type: Tessmann 5164 (holo B; iso B,NY,S), Peru, Loreto, Upper Amazon River, Lower Itaya River, Soledad , 110 m, 6 June 1925.

Guatteria juninensis R.E.Fr.(1938) 716;(1939) 387,f. 12d,syn.nov. — Type: C. Schunke 267 (holo F 2 sheets), Peru, Junín, Schunke Hacienda,above San Ramón , 1400–1700 m, 8–12 June 1929.

Guatteria ovalifolia R.E.Fr. (1939) 328, syn. nov. — Type: A.C. Smith 3449 (holo S; iso B, F, G, K, MO, NY, P, U), Guyana, NW slopes of Kanuku Mts, in drainage of Moku-Moku Creek (Takutu tributary), 150–400 m, Apr.1938.

Guatteria collina R.E.Fr. (1939) 381, syn. nov. — Type: Goudot s.n. (holo K; iso P), Colombia, ‘ Savana grande’.

Guatteria buchtienii R.E.Fr. (1939) 388, syn. nov. — Type: Buchtien 699 (holo S; iso NY, US), Bolivia, La Paz, Mapiri Region , San Carlos, 850 m, 18 Dec. 1926.

Guatteria lasiocalyx R.E.Fr. (1939) 388, syn. nov. — Type: Bang 583 (holo UPS; iso BM, BR, C, E, F, G, K, L, M, MO, NY 2 sheets, S, U, US, W), Bolivia, La Paz, Yungas, anno 1890.

Guatteria rhamnoides R.E.Fr. (1939) 389, syn. nov. — Type: Krukoff 5707 (holo S; iso BM, F, G, K, M, MO, NY, RB, U, US), Brazil, Acre, Basin of Rio Purus, near mouth of Rio Macauã (‘Macauhan’), tributary of Rio Yacu , 27 Aug. 1933.

Guatteria pteropus Benth.var. angustior R.E.Fr.(1939) 420,syn.nov. — Type: Spruce 1342 (holo K; iso F 2 sheets, K, P), Brazil, Amazonas , Manaus (‘Barra’), Feb. 1851.

Guatteria pteropus Benth.var. cinerea R.E.Fr. (1939) 420, syn. nov. — Type: Ducke RB 29018 (holo S; iso MO, RB), Brazil, Amazonas, Parintins, be- tween Paraná de Ramos and Tracajá, 11 Jan. 1936.

Guatteria olivacea R.E.Fr. (1939) 423; Maas et al. (2007) 641, syn. nov. — Type: Krukoff 6853 (holo S; iso BM, BR, F, G, K, MO, NY, U, US), Brazil, Amazonas , Basin of Rio Madeira,Mun. Humaitá,near Livramento, 12 Oct.– 6 Nov. 1934

Guatteria obliqua R.E.Fr. (1939) 424, f. 14e, syn. nov.. — Type: J.G. Kuhlmann 643 = RB 24280 (holo S; iso F, INPA, RB, SPF), Brazil, Rondônia, Rio Abunã (‘Abunam’), 17 Oct. 1923.

Guatteria occidentalis R.E.Fr. (1939) 430, f. 15b, syn. nov. — T ype: Rose et al. 23429 (holo S; iso F, GH, NY), Ecuador, El Oro, Portovelo , ‘984 m’, 6–15 Oct. 1918.

Guatteria chrysopetala (Steud.) Miq.var. major R.E.Fr. (1939) 435,f. 16a, b. — Type: Ducke RB 19612 (holo S, iso RB), Brazil, Pará , Ôbidos, 19 Sept. 1927.

Guatteria chrysopetala (Steud.) Miq. var. tenuipes R.E.Fr. (1939) 435. — Type:A nonymous collector 239, 249 (syn UPS,not seen), French Guiana, without location.

Guatteria sagotiana R.E.Fr. var. gracilior R.E.Fr.(1939) 437. — Type: Poiteau s.n. (holo K), French Guiana, without location, July 1824.

Guatteria sagotiana R.E.Fr. [ var. sagotiana ] (1939) 437. — Type: Sagot 1263 p.p. (holo K 2 sheets), French Guiana, Roura , anno 1859.

Guatteria gracilipes R.E.Fr. (1939) 438.— Type: Krukoff 1156 (holo S; iso G, NY, P, U), Brazil, Pará , Upper Cupari (‘Cupary’) River, between Xingu and Tapajós Rivers, 16 Sept. 1931.

Guatteria longestipitata R.E.Fr. (1939) 438, f. 17a, b, syn. nov. — Type: Ducke RB 19613 (holo S; iso RB), Brazil, Pará , Juruti Velho, 23 Dec.1926.

Guatteria parviflora R.E.Fr. (1939) 440, f. 18a, syn. nov. — Type: J.G. Kuhlmann RB 24263 (holo S), Brazil, Pará, Santa Júlia, Serra de Parintins , 18 Mar. 1924.

Guatteria lanceolata R.E.Fr. (1939) 443, f. 18b, c, syn. nov. — Type: Krukoff 5950 (holo S; iso BM,F,G,K,MO,NY,S,U, US 2 sheets), Brazil, Amazonas, Mun. Borba, near Urucurituba, 4–6 Sept. 1934.

Guatteria elliptica R.E.Fr. (1939) 445, f. 19c, d, syn. nov. — Type: Glaziou 9605 (holo C; iso K, P 2 sheets), Pará, Santarém (?) (‘ Rio de Janeiro, São Fidelis’ ), 18 Feb. 1876.

Guatteria umbonata R.E.Fr. (1939) 447, f. 16e, g, syn. nov. — Type: Ducke RB 19614 (holo S), Brazil, Pará , Santarém, ‘loco Mahicá’, 25 Mar. 1916.

Guatteria gamosepala R.E.Fr. (1939) 528, f. 37a–d, syn. nov. — Type: Krukoff 6047 (holo S; iso BM, BR, F, G, K, MO, NY, U, US), Brazil, Amazonas , Basin of Rio Madeira, Mun. Manicoré, near Santa Fé, 8–11 Sept. 1934.

Guatteria leiocarpa R.E.Fr. (1939) 530, f. 36c, d, syn. nov. — Type: Krukoff 8995 (holo S; iso BM, BR, F, G, K, MO, NY, P, U, US), Brazil, Amazonas, Mun. São Paulo de Olivença, Basin of Creek Belém , 26 Oct.–11 Dec.1936.

Guatteria macrocalyx R.E.Fr.(1939) 530,syn.nov. — Type: Mutis 3705 (holo US), Colombia, without location and date.

Guatteria calimensis R.E.Fr. (1950a) 332, syn. nov. — Type: Cuatrecasas 16566 (holo S 2 sheets; iso F 3 sheets,U, US), Colombia, Valle del Cauca, La Trojita, 5–50 m, 19 Feb.–10 Mar. 1944.

Guatteria latipetala R.E.Fr. (1957b) 602, syn. nov. — Type: Schultes 5512 (holo US; iso US), Colombia, Vaupés, Río Macaya, near Cachivera del Diablo and mouth of river, 300 m, May 1943.

Guatteria asplundiana R.E.Fr. (1959a) 24; Erkens et al. (2008) f. 3, pl. 4, syn. nov. — Type: Asplund 19673 (lecto S, selected by Maas et al. 1994; isolecto G, P), Ecuador, Pastaza, Mera , near Manguyacu , 14 Dec. 1956.

Guatteria wessels-boerii Jans.-Jac.(1970) 336,pl. 1. — Type: Wessels-Boer 1302 (holo U; iso K, NY,WAG,Z), Suriname,near confluence of Paloemeu and Tapanahoni Rivers, 15 Apr. 1963.

Guatteria sp. 10 Chatrou et al. (1997) 111.

Tree or shrub 1–45 m tall, 5–75 cm diam. Young twigs densely to sparsely covered with appressed (or rarely erect) hairs, soon glabrous. Leaves: petiole 4–30 mm long, 1–6 mm diam; lamina chartaceous to coriaceous, narrowly elliptic, sometimes elliptic or narrowly obovate, 7–35(–45) cm long, 3–14(–18) cm wide (leaf index 1.5–3.5(–5)), not verruculose, dull to shiny, grey to brown above, greyish brown to brown below, glabrous or primary vein variably covered with appressed or rarely erect hairs above, sparsely to rather densely (rarely densely) covered with appressed hairs below, base acute, shortly to long-attenuate, sometimes obtuse or even cordate, apex acuminate (acumen 5–25 mm long), rarely acute, primary vein slightly impressed to flat above, sometimes keeled below, secondary veins distinct, impressed to flat above, 10– 25 on either side of primary vein, smallest distance between loops and margin 1–5 mm, tertiary veins flat, slightly raised or slightly impressed above, reticulate to percurrent. Flowers in 1–3(–5)-flowered inflorescences in axils of leaves or on leafless branchlets; pedicels 10–35(–55) mm long, 1–2 mm diam, fruiting pedicels 2–4 mm diam, densely to sparsely covered with appressed (to semi-erect or erect) hairs, articulated at 0.2–0.5 from the base, bracts 5–7, soon falling, the 2 upper ones elliptic to broadly elliptic, to 6(–10) mm long (one upper bract exceptionally foliaceous and c. 40 by 10 mm), middle bract elliptic, c. 5 mm long, lower bracts elliptic to ovate or broadly so, 1–2 mm long; flower buds de- pressed ovoid, apex obtuse or acute; sepals free, rarely basally connate, broadly to shallowly ovate-triangular, 4–10 by 4–10 mm, reflexed or appressed, outer side densely covered with appressed to erect hairs; petals green, yellow, cream or white, sometimes glaucous in vivo, oblong-elliptic, 10–40{–50} by 5–20 mm, outer side densely covered with appressed hairs; stamens 1–2 mm long, connective shield hairy, papillate, or glabrous, umbonate to flat. Monocarps 5–100, green, maturing black to purple-black in vivo, black to brown, sometimes glaucous in sicco, ellipsoid to subglobose, 7–20 by 4–15 mm, sparsely covered with appressed hairs, soon glabrous, apex rounded to apiculate (apiculum <1 mm long), wall 0.1–1{–4} mm thick, stipes 6–30 by 0.5–2{–3} mm. Seed ellipsoid to subglobose, 6–12 by 4–8 mm, pale to dark brown, pitted to smooth, sometimes rugose, raphe raised.

Distribution — Colombia (Amazonas, Antioquia, Boyacá, Cauca, Chocó, Guainía, Meta, Valle del Cauca, Vaupés, Vichada), Venezuela (Amazonas, Bolívar, Delta Amacuro, Zulia), Guyana, Suriname, French Guiana, Ecuador (Azuay, Carchi, Esmeraldas, Morona-Santiago, Napo, Pastaza, Santiago-Zamora, Sucumbios, Zamora-Chinchipe), Peru (Ama- zonas, Cajamarca, Cusco, Huánuco, Loreto, Madre de Dios, Oxapampa, Pasco, San Martín , Ucayali), Brazil (Acre, Amapá, Amazonas, Maranhão, Mato Grosso, Pará, Rondônia), Bolivia (Beni, Cochabamba, La Paz, Pando, Santa Cruz).

Habitat & Ecology — In non-inundated lowland, premontane or montane forest, often on clayey soil. At elevations of 0–2800 m. Flowering: throughout the year; fruiting: throughout the year.

Vernacular names — Bolivia: Chia (de Michel et al. 590), Chirimoya del monte (de Michel et al. 590), Chocolatillo negro (D.N. Smith et al. 13252, 13749), Laurel macho (deWalt et al. 121), Maurel canelón (Endara s.n.), Midha dhahua (Tacana) (Bourdy 1740, de Walt et al. 121, Serato 58), Palo pancho verde (Serato 58), Piraquina (Guillén & Roca 2494), Piraquina de barbecho (Meneces 2087), Piraquina negra (Quevedo et al. 90, D.N. Smith et al. 14128). Brazil: Envira (Krukoff 6411, Pacheco et al. 120), Envira-cajú (Figueiredo 803), Envira-flor-grande (Fróes 11638), Envira-fofa ( Daly et al. 6878), Envira-manga-de-anta ( Pardo et al. 106), Envira-preta ( Daly et al. 4331, Fróes 11787, M.J.P. Pires et al. 868, Rosa et al. 1750, 2768), Envireira (H. C. Lima et al. 2716, Rodrigues 233), Envireira-da-birida (Sobel et al. 4615), Invira (Krukoff 6047), Invireira (Campbell et al. 9694), Seiseiunahi (Yanomami name) (Milliken 1706, 2089), Taiwi’i (Balée & Ribeiro 1742). Colombia: Cargadero (Cuatrecasas 16253, Faber-Langendoen & Rentería A. 1368, Gentry et al. 53678, Monsalve B.1482, 1500, Triana s.n.), Cargadero negro (Cuatrecasas & Patiño 27452), Garrapato (Tick tree) (Dawe 946, Fonnegra et al. 3135, 3140), Guasco dulce (Zarucchi 3271), Kïbojïu dujeko (Muinane name) (Murillo A. & Rodríguez 561), Zuto (Betancur et al. 6811). Ecuador: Cargadera negra (C. & E. Aulestia 899), Chiwiachim (Shuar name) (Aulestia & Gonti 1915), Fandicho (Cofán name) (Cerón 20720, 41738), Fanicho (Cofán name) (Cerón 20901), Gañitahue (Huaorani name) (Aulestia & Gonti 1795), Mucataremon (Huaorani name) (Aulestia et al. 1044), Neayatio (Secoya name) (Freire et al. 2861), Oñitahua (Huaorani name) (Aulestia & Bainca 3088), Oñitahue (Huaorani name) (Aulestia et al. 1965, 3206), Oñita- huemo (Huaorani name) (Aulestia & Quihuiñamo 3173), Shapa- ttovo (Cofán name) (Cerón 20895), Ucucha anona (Quichua name) (Gudiño & Andi 2040). French Guiana: Iliwa (Wayana name) (Fleury 1475), Iwi (Wayãpi name) (Prévost & Grenand 1020), Iwilusi (Wayãpi name) (de Granville 2633), Mamayavé/ Maman yawée (Creole name) (de Granville B.4614, B.5125, Oldeman 1579, B.3142, B.3477), Mamayawé commun (Creole name) (Oldeman 2841), Pina ? ipinu (Wayãpi name) (de Granville B.5156), Pinaou (Galibi Carib name; see Aublet (1775) 615). Guyana:Arara (Arawak name) (Tutin 360), Black Kuyama (Arawak name) (Sandwith 406), Black maho (Creole name) (Van Andel et al. 666), Kuyama (Carib name) (Van Andel et al. 666), Yarayara (Carib name) (Van Andel et al. 1127). Peru: Amarillo (D.N. Smith & Pretel 1491, 1549), Anonilla (D. Smith 2088), Anonilla blanca (Pariona et al. 23, 941), Auca hicoja (Schunke V. 14100), Carahuasca ( Angulo 5a, Ellenberg 2459, Hartshorn 1683, KrÖll Saldaña 674, Reynel R. 3, D.N. Smith et al. 1203), Carahuasca negra (Gutiérrez R. et al. 206), Caravasca (Tessmann 3235), Muraya (Shuar name) ( Neill & Manzanares 13144), Wámpuyais (Shuar name) (Kayap 395), Yaïs (Shuar name) (C. Díaz et al. 7230, 7237, R. Rojas et al. 393), Yanahuasca (Schunke V. 2065), Yana huasca (Graham 2578), Zoro caspi (Spichiger et al. 1773), Zorro caspi ( Daly et al. 5727). Suriname: Arara (Arawak name), Baaka pau, mamaai (Saramaccan name), Blaka paw (Saramaccan name) (van Donselaar 2078), Boesi-soensaka (S), Boszuurzak (Sranang name) (Elburg, LBB 9881, Lindeman 3646, 4982, 6151, 6759, Schulz, LBB 8250, Vreden, LBB 13711), Koeli koejokoe (Arawak name) (Stahel s.n.), Krabietakaka (Sranang name) (BW 470), Panta (Sranang name) (van Donselaar 1806, Lindeman 6809), Pedre- koe pisi (Sranang name). Venezuela: Annoncillo (Steyermark 88270), Majagua (Delgado 836, Velazco 1969), Majagua blanca (Marcano-Berti 68-979), Majagua hoja larga (Liesner 6188), Majagua negra (Marcano-Berti et al. 39-1-77), Majagua verde (Aymard C. 9793, Liesner 7055), Wosewayek (Hernández et al. 29, 118), Yarayara amarilla (Blanco 332A), Yarayara morada (Marcano-Berti 422, 622).

Uses — Suriname: Edible fruits, leaves in herbal bath (Van Andel 5354). Wood used for boards (Van Andel & Poeketi 4807).

Notes — In 1939 Robert Fries described sect. Pteropus as one of the 30 sections he then recognized in the genus Guatteria . As the name ‘Pteropus’ (‘winged foot’) already suggests, one of the features of this section is the attenuate and almost winged leaf base (‘Blätter an der Basis zugespitzt, meist nach unten keilförmig verschmälert’; in Latin: ‘Folia vulgo decur- rentia’). Another main feature of sect. Pteropus consists in the indument, which is almost always composed of appressed hairs (except for G. sylvicola in which the hairs are often erect). The section is mainly distributed in the Guianas and the Amazon Region.

Section Pteropus originally included 16 species, increased after that to 20 species ( Fries 1959b). The species were keyed out by Fries (1939) almost exclusively by leaf features such as leaf shape, leaf base, leaf apex, leaf indument and in a few cases pedicel length.

After extensive comparison of much more material than Fries had at his disposal as late as 1959, we can now accept G. elata , G. elongata (only known from the type) and G. modesta (includ- ing 4 of Fries’ species: G. chlorantha , G. geminiflora , G. glaberrima and G. tessmannii ). The remaining species together with a number of other species not classified by Fries in sect. Pteropus all form the G. punctata complex which will be discussed now. This complex includes most of the species of sect. Pteropus , namely G. atra , G. axilliflora , G. calliantha , G. chrysopetala , G. guentheri , G. obliqua , G. occidentalis , G. olivacea , G. ovalifolia , G. poeppigiana , G. pteropus and G. sylvicola . We checked the material of all these above mentioned species and tried to find some distinguishing features. Despite many trials we were unable to delimit these species convincingly. The differences found in the leaves, like shape of the leaf base, did not hold: the variation in the shape of the leaf base simply is too variable in this complex.

1. Typically, G. pteropus and G. olivacea show a strongly atte-

nuate leaf base, but in much material of Brazil, Peru, Ecuador

and Venezuela one can observe attenuate, acute, to obtuse

or even cordate leaf bases. The extremes, in other words,

may look very different, but they are connected by all kinds

of intermediate forms (see also Fig. 63 ), and we could not

maintain them as distinct species. 2. Leaf colour in dried material in this complex is also quite vari-

able: brown leaves are most commonly found, but greyish

colours in various shades as well as transitional colours are

also encountered. 3. Leaf size is highly variable in this complex ( Fig. 63 ), large

leaves often abound (among others in Peru, the Brazilian

state of Acre and elsewhere), vs very small and narrow lea-

ves (among others in the Brazilian state of Pará and in some

parts of Suriname). We also checked features of inflorescence, flowers and fruits of this complex, only to find great variation, too, and hardly useful differences. It is worth noting that seeds of all species involved in this com- plex appear remarkably similar, being of almost equal size and mostly smooth to only slightly pitted. Fries (1939) distinguished in Guatteria a sect. Tylodiscus with as main feature an umbonate connective shield (‘Staubblattschilde mit einem centralen Umbo’). In that section he included 20 different species, several of which occur in the Guianas and adjacent Brazil, namely G. chrysopetala , G. gracilipes and G. sagotiana . After an intensive study of Guianan Annonaceae we came to the conclusion that these species are part of one complex species, namely G. punctata . The next logical step for us then was to compare the concepts of G. pteropus and G. punctata more closely. The feature of the presence of an umbo on the connective shield, striking though it may be, is not constant, the centre of the connective shield varying from distinctly elevated (= umbonate) to merely a non-elevated area which still might stand out as such to not at all. This being so, the main distinguishing feature of sect. Tylodiscus appeared not to be constant. Furthermore, we did not find good differences in leaves, flowers and fruits to be able to maintain the two entities as separate species. Therefore we propose herewith to unite both species under the oldest name, namely G. punctata . In a late stage of our revision we compared G. punctata with species of sect. Guatteria . In an earlier publication ( Maas & Westra 2010), we already united all species of this section, namely G. buchtienii , G. coeloneura , G. juninensis , G. lasiocalyx , G. pleiocarpa and G. rhamnoides under one species, namely G. glauca . Although G. glauca shows some slight differences with G. punctata in a narrower sense, namely an indument often composed of erect hairs and often basally connate sepals, we found material with erect hairs and slightly connate sepals in G. punctata as well. For us there remained little else, and much to our reluctance, than to unite G. glauca , mainly restricted to Bolivia and Peru, with G. punctata . Then, during the last phase of our revisionary work, we had to deal with a group of species mainly restricted to the Pacific coast of Colombia and Ecuador (among others, G. cargadero and G. asplundiana ). Notable features in this group are often some- what larger monocarps (in G. asplundiana ) and relatively broad and leathery leaves (in G. cargadero ). Here, again, distinctions are not as sharp as they seem at first, with both monocarps and leaves being variable, but as in all other features of leaves and flowers there were no great differences with G. puncata we decided, albeit again with much hesitation, to include both concepts in G. punctata . The outcome of this study for the present is: one large spe- cies, namely G. punctata , covering most parts of tropical South America (except for SE Brazil). We are fully aware that this decision is quite tentative and we hope that additional studies (including intensive field work and molecular studies) will contribute to a final unraveling of this complex.

Figures

Fig. 63 Guatteria punctata (Aubl.) R.A.Howard. Analysis of leaf shapes.