Geonoma maxima (Poit

Geonoma maxima (Poit.) Kunth, Enum. Pl. 3: 229. 1841. Gynestum maximum Poit., Mém. Mus. Hist. Nat. 9: 388. 1822. Type. FRENCH GUIANA. Without locality, no date, A. Poiteau s. n. (holotype: P!).

Plant height 3.2(1.0-9.0) m, CV 0.4, n = 144; stems clustered or rarely solitary; stem diameter 1.3(0.6-3.5) cm, CV 0.3, n = 110; number of leaves per stem 9(4-19), CV 0.4, n = 66. Leaves regularly pinnate or irregularly pinnate with multi-veined pinnae and 1-veined pinnae intermixed, rarely simple; sheath length 12.6(4.0-25.0) cm, CV 0.4, n = 87; petiole length 36.4(1.0-100.0) cm, CV 0.5, n = 97; rachis length 44.6(7.3-120.0) cm, CV 0.6, n = 145; rachis width 3.7(1.4-7.1) mm, CV 0.3, n = 177; number of leaf divisions per side of rachis 8(1-31), CV 1.0, n = 170; basal angle of pinna divergence 46(7-87)°, CV 0.4, n = 174; basal pinna length 45.1(11.5-132.0), CV 0.5, n = 104; basal pinna width 4.3(0.2-34.0) cm, CV 1.3, n = 167; apical angle of pinna divergence 22(7-40)°, CV 0.3, n = 171; apical pinna length 34.6(8.8-78.5), CV 0.4, n = 110; apical pinna width 10.7(0.7-42.5) cm, CV 0.8, n = 157. Inflorescences branched; prophyll length 11.4(4.0-21.8) cm, CV 0.3, n = 47; peduncular bract length 9.8(6.0-13.0) cm, CV 0.3, n = 9; distance between bract scars 0.6(0.2-3.0) cm, CV 0.6, n = 141; peduncle length 8.5(4.3-19.5) cm, CV 0.3, n = 149; peduncle width 4.9(1.5-10.0) mm, CV 0.3, n = 205; orders of inflorescence branching 1-3; rachilla width 2.7(0.7-5.5) mm, CV 0.3, n = 255; rachilla length 11.5(4.8-24.3), CV 0.3, n = 258; number of rachillae 19(4-50), CV 0.5, n = 167; fruit diameter 8.9(5.5-13.0) mm, CV 0.2, n = 96; fruit length 11.9(6.9-18.2) mm, CV 0.2, n = 96.

There is considerable variation in leaf division, as evidenced by the high CV value for this variable. A PCA of 42 specimens (231 specimens excluded because of missing data) and seven log-transformed quantitative variables (petiole length, rachis length, rachis width, number of divisions, basal pinna angle, apical pinna length, apical pinna angle) was carried out. The small sample size here is a result of excluding the variables basal pinna width, basal pinna length, and apical pinna width (because they do not apply to simple leaves) and including petiole length which has many missing values but which is a discriminatory variable. A scatterplot of the first two components (not shown) divides the specimens into five discernible groups. One of these groups can again be divided, on geographical grounds, into two. For these six groups, ANOVA shows that for pairwise comparison probabilities, all vegetative quantitative variables differed significantly (P <0.01) between at least one pair of groups, although no variable differed among all groups. DA, using the same quantitative variables as for PCA, separates the six groups with 98% success in classification matrix and 88% success in jackknifed classification matrix (Wilks´ lambda = 0.005; P <0.01). The most discriminatory variable is number of leaf divisions, followed by petiole length and rachis length. Because of these results, and because several of these groups are separated from each other geographically, these six groups are recognized as subspecies.

Key to the subspecies of G. maxima

1a. Leaves regularly pinnate; pinnae numerous, 4-31 per side of rachis, regularly arranged, usually narrow and equally wide except for apical one; throughout the Amazon region in the Guianas, Venezuela, Colombia, Brazil, Ecuador, Peru, and Bolivia..... G. maxima subsp. maxima.
1b. Leaves irregularly pinnate or blade simple; pinnae few, 1-8 per side of rachis, irregularly arranged, unequally wide, often the apical and basal ones wider than the others. 2.

2a. Leaves with 1-4 pinnae per side of rachis, often blade simple; mean petiole length 13.0 cm. 3.
2b. Leaves with 1-8 pinnae per side of rachis, rarely blade simple; mean petiole length 41.6 cm. 4.

3a. Mean basal pinna width 24.3 cm, or blade simple; basal pinna forming a mean angle of 13°; central-western Amazon region of Brazil and ajdacent Colombia..... G. maxima subsp. spixiana.
3b. Mean basal pinna width 2.6 cm, or blade simple; basal pinna forming a mean angle of 36°; Rio Negro region of Brazil and ajdacent Colombia and Venezuela..... G. maxima subsp. chelidonura.

4a. Mean basal pinnae 77.0 cm long and 9.1 cm wide; southwestern Amazon region of Peru and adjacent areas of Brazil and Bolivia..... G. maxima subsp. peruviana.
4b. Mean basal pinna 49.6 cm long and 5.4 cm wide. 4.

5a. Mean number of divisions per side of rachis 2 (2-5); central-western Amazon region of Brazil and ajdacent Colombia, Peru, and Bolivia..... G. maxima subsp. densiflora.
5b. Mean number of divisions per side of rachis 4 (2-8); Guyana and ajdacent Venezuela and Brazil..... G. maxima subsp. ambigua.

Geonoma maxima subsp. maxima

Geonoma multiflora Mart., Hist. nat. palm. 2: 7. 1823. Type. BRAZIL. Pará: without locality, no date, C. Martius s. n. (holotype: M; F neg. 18516, 18516a).

Geonoma hexasticha Spruce, J. Linn. Soc., Bot. 11: 110. 1871. Type. BRAZIL. Amazonas: near Gama´s sitio, Sâo Gabriel, Mar 1852, R. Spruce 29 (holotype: K!; isotype: NY! F neg. 38649).

Geonoma discolor Spruce, J. Linn. Soc., Bot. 11: 110. 1871. Type. BRAZIL. Pará: Rio Tapajós, no date, R. Spruce 36 (holotype: K).

Geonoma paraensis Spruce, J. Linn. Soc., Bot. 11: 112. 1871. Type. BRAZIL. Pará: near Belém, no date, R. Spruce 69 (holotype: K!).

Geonoma negrensis Spruce, J. Linn. Soc., Bot. 11: 113. 1871. Type. VENEZUELA. Amazonas: Río Negro, San Carlos, Sep 1853, R. Spruce 70 (holotype: K!).

Geonoma speciosa Barb. Rodr., Enum. palm. nov. 9. 1875. Lectotype (Wess. Boer, 1968). Barb. Rodr., Sert. palm. brasil. 1: t. 18. 1903.

Geonoma capanemae Barb. Rodr., Enum. palm. nov. 9. 1875. Lectotype (Wess. Boer, 1968). Barb. Rodr., Sert. palm. brasil. 1: t. 29. 1903.

Geonoma palustris Barb. Rodr., Enum. palm. nov. 11. 1875. Lectotype (Wess. Boer, 1968). Barb. Rodr., Sert. palm. brasil. 1: t. 27. 1903.

Geonoma uliginosa Barb. Rodr., Enum. palm. nov. 11: 1875. Lectotype (Wess. Boer, 1968). Barb. Rodr., Sert. palm. brasil. 1: t. 28. 1903.

Geonoma spruceana subsp. intermedia var. compta Trail, J. Bot. 5: 329. 1876. Type. BRAZIL. Amazonas: Barcellos, 30 Jun 1874, J. Trail 997/LXXXIV (holotype: K!).

Geonoma spruceana subsp. intermedia var. intermedia Trail, J. Bot. 5: 329. 1876. Type. BRAZIL. Amazonas: Lago Cerrado, Rio Juruá, 30 Oct 1874, J. Trail 989/CXLVII (holotype: K!).

Petiole length 41.0(12.0-100.0) cm, CV 0.4, n = 34; number of leaf divisions per side of rachis 18(4-31), CV 0.3, n = 53; basal angle of pinna divergence 60(35-87)°, CV 0.2, n = 63; basal pinna width 0.6(0.2-4.0) cm, CV 0.7, n = 70; basal pinna length 33.0(11.5-52.0) cm, CV 0.3, n = 48.

Distribution. One hundred and one specimens from 5°17´N-11°10´S and 46°30´-77°20´W at a mean elevation of 250 (50-1000) m throughout the Amazon region in the Guianas, Venezuela, Colombia, Brazil, Ecuador, Peru, and Bolivia (map).

There is considerable variation within this subspecies. One specimen from French Guiana (# 3) has only 4 pinnae per side of the rachis, and thus approaches subsp. ambigua. Two specimens (#´s 245, 246) from the eastern Amazon region of Brazil also have few pinnae per side, 4-5, and also approach subsp. ambigua, These are similar to, and come from the same area as, several species described by Barbosa Rodrigues (G. speciosa, G. palustris, G. capanemae; the types of which are illustrations). Some specimens from the Colombian Amazon (#´s 100, 101, 102, 188) have smaller, sigmoid pinnae, narrower rachillae, and larger fruits than usual.

Geonoma maxima subsp. ambigua (Spruce) Henderson, stat. nov. Geonoma maxima var. ambigua (Spruce) Henderson, Palms of the Amazon 278. 1995. Geonoma ambigua Spruce, J. Linn. Soc., Bot. 11: 111. 1871. Type. GUYANA. Without locality, no date, C. Appun 566 (isotype: K!).

Geonoma schomburgkiana Spruce, J. Linn. Soc., Bot. 11: 111. 1871. Type. GUYANA. Without locality, 1837, R. Schomburgk 705 (holotype: K! excluding leaf).

Petiole length 40.9(21.0-80.0) cm, CV 0.4, n = 11; number of leaf divisions per side of rachis 4(2-8), CV 0.3, n = 25; basal angle of pinna divergence 40(20-60)°, CV 0.3, n = 22; basal pinna width 7.9(1.9-22.0) cm, CV 0.7, n = 20; basal pinna length 45.0(28.5-71.0) cm, CV 0.3, n = 12.

Distribution. Fifty-two specimens from 1°50´-8°34´N and 57°52´-62°00´W at a mean elevation of 347 (35-900) m in Guyana and ajdacent Venezuela and Brazil (map).

Geonoma maxima subsp. chelidonura (Spruce) Henderson, stat. nov. Geonoma maxima var. chelidonura (Spruce) Henderson, Palms of the Amazon 279. 1995. Geonoma chelidonura Spruce, J. Linn. Soc., Bot. 11: 111. 1871. Type. BRAZIL. Amazonas: Rio Uaupés, Nov 1852, R. Spruce 73 (holotype: K!).

Geonoma tuberculata Spruce, J. Linn. Soc., Bot. 11: 112. 1871. Type. BRAZIL. Amazonas: Rio Negro, no date, R. Spruce 18 (holotype: K!).

Geonoma parvisecta Burret, Notizbl. Bot. Gart. Berlin-Dahlem 10: 1018. 1930. Type. BRAZIL. Amazonas: Rio Negro, Sâo Pedro do Uaupés, 23 Sep 1928, P. Luetzelburg 22278 (holotype: B, destroyed; isotypes, NY! R; F neg. 18518).

Petiole length 12.8(10.0-18.0) cm, CV 0.2, n = 8; number of leaf divisions per side of rachis 2(1-2), CV 0.3, n = 11; basal angle of pinna divergence 36(14-65)°, CV 0.4, n = 11; basal pinna width 2.6(0.7-4.0) cm, CV 0.5, n = 6; basal pinna length 37.8(19.5-45.5) cm, CV 0.3, n = 6.

Distribution. Twelve specimens from 2°03´N-2°41´S and 60°19´-67°20´W at a mean elevation of 98 (75-140) m in black water regions along the Rio Negro in Brazil and ajdacent Colombia and Venezuela (map).

Geonoma densiflora Spruce, J. Linn. Soc., Bot. 11: 112. 1871. Type. BRAZIL. Amazonas: Sâo Gabriel, near Gama´s sitio, Mar 1852, R. Spruce 30 (holotype: K!).

Geonoma personata Spruce, J. Linn. Soc., Bot. 11: 112. 1871. Type. BRAZIL. Amazonas: Serra of Sâo Gabriel, Jun 1852, R. Spruce 34 (holotype: K!).

Geonoma densiflora var. monticola Spruce, J. Linn. Soc., Bot. 11: 118. 1871. Type. BRAZIL. Amazonas: Sâo Gabriel, Jun 1852, R. Spruce 33 (holotype: K!).

Geonoma brachyfoliata Barb. Rodr., Enum. palm. nov. 10. 1875. Lectotype (Wess. Boer, 1968). Barb. Rodr., Sert. palm. brasil. 1: t. 33. 1903.

Geonoma bijugata Barb. Rodr., Enum. palm. nov. 10. 1875. Lectotype (Wess. Boer, 168). Barb. Rodr., Sert. palm. brasil. 1: t. 16. 1903.

Geonoma falcata Barb. Rodr., Enum. palm. nov. 10. 1875. Lectotype (Wess. Boer, 1968). Barb. Rodr., Sert. palm. brasil. 1: t. 19. 1903.

Geonoma furcifolia Barb. Rodr., Enum. palm. nov. 11. 1875. Lectotype (Glassman, 1972). Barb. Rodr., Sert. palm. brasil. 1: t. 15. 1903.

Geonoma spruceana Trail, J. Bot. 5: 328. 1876. Geonoma spruceana subsp. spruceana var. spruceana Trail, J. Bot. 5: 329. 1876. Type. BRAZIL. Pará: Lago Juriti, 3 Apr 1874, J. Trail 1002/XXIV (holotype: K!).

Geonoma spruceana subsp. spruceana var. heptasticha Trail, J. Bot. 5: 329. 1876. Type. BRAZIL. Amazonas: Rio Negro at Assutuba, 6 Jul 1874, J. Trail 1007/XCIII (holotype: K!)

Geonoma spruceana subsp. spruceana var. micra Trail, J. Bot. 5: 329. 1876. Type. BRAZIL. Amazonas: Lago Juruty, no date, J. Trail 29 (holotype: K).

Geonoma spruceana subsp. intermedia var. tuberculata Trail, J. Bot. 5: 329. 1876. Type. BRAZIL. Amazonas: Rio Marmellos, Rio Madeira, 2 Jun 1874, J. Trail 983/LIV (holotype: K).

Geonoma spruceana subsp. intermedia var. major Trail, J. Bot. 5: 330. 1876. Type. BRAZIL. Amazonas: Rio Solimoes, Coary, 16 Oct 1874, J. Trail 984/CXLIV (holotype: K! isotype: NY).

Geonoma juruana Dammer, Verh. Bot. Vereins Prov. Brandenburg 48: 119. 1906. Type. BRAZIL. Acre: Rio Juruá, Juruá-mirim, Aug 1901 E. Ule 5744 (holotype: B; isotype: MG).

Geonoma longisecta Burret, Bot. Jahrb. Syst. 63: 257. 1930. Type. PERU. Loreto: Iquitos, no date, G. Tessmann 5087 (holotype: B, destroyed; isotype: NY!).

Petiole length 44.5(18.0-73.0) cm, CV 0.4, n = 23; number of leaf divisions per side of rachis 2(2-5), CV 0.3, n = 49; basal angle of pinna divergence 47(28-70)°, CV 0.2, n = 46; basal pinna width 4.5(1.4-11.5) cm, CV 0.5, n = 47; basal pinna length 52.1(20.0-75.5) cm, CV 0.3, n = 26.

Distribution. Sixty-three specimens from 2°24´-11°45´S and 55°00´-75°50´W at a mean elevation of 145 (75-350) m in the central-western Amazon region of Brazil and ajdacent Colombia, Peru, and Bolivia (map).

Several specimens (#s 1, 27, 116) are smaller than usual, and ressemble those of subsp. chelidonura, and may also come from black water regions.

Geonoma maxima subsp. peruviana Henderson, subsp. nov. Type. PERU. Madre de Dios: Río Tambopata, Explorer´s Inn at junction of Río La Torre, 12°50S, 69°17´W, 250 m, 3 Nov 1991, A. Henderson & F. Chávez 1634 (holotype: NY!).

Petiole length 36.8(23.0-64.0) cm, CV 0.3, n = 12; number of leaf divisions per side of rachis 4(1-8), CV 0.5, n = 17; basal angle of pinna divergence 31(19-40)°, CV 0.2, n = 20; basal pinna width 9.1(2.4-19.0) cm, CV 0.6, n = 19; basal pinna length 77.0(53.0-92.0) cm, CV 0.2, n = 7.

Distribution. Twenty-nine specimens from 5°03´-13°45´S and 68°09´-78°22´W at a mean elevation of 367 (135-800) m in the southwestern Amazon region of Peru and adjacent areas of Brazil and Bolivia (map).

Geonoma maxima subsp. spixiana (Mart.) Henderson, stat. nov. Geonoma maxima var. spixiana (Mart.) Henderson, Palms of the Amazon 281. 1995. Geonoma spixiana Mart., Hist. nat. palm. 2: 15. 1823. Type. BRAZIL. Amazonas: Rio Japurá, no date, C. Martius s.n. (holotype: M! F negs. 18525, 18525a).

Petiole length 13.3(1.0-23.0) cm, CV 0.6, n = 9; number of leaf divisions per side of rachis 2(1-4), CV 0.6, n = 15; basal angle of pinna divergence 13(7-20)°, CV 0.3, n = 13; basal pinna width 24.3(10.5-34.0) cm, CV 0.4, n = 5; basal pinna length 91.5(69.0-132.0) cm, CV 0.3, n = 5.

Distribution. Sixteen specimens from 1°18´-7°10´S and 60°00´-72°00´W at a mean elevation of 277 (230-300) m in the central-western Amazon region of Brazil and ajdacent Colombia (map).