Leaf Morphology of the Pejibaye Palm (Bactris Gasipaes H.B.K.)
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Rev. Biol. Trap., 31(1), 103·112, 1983 Leaf morphology of the Pejibaye palm (Bactris gasipaes H.B.K.) Charles R. Clement Instituto Nacional de Pesquisas da Amazonia, Manaus, Brazil Jorge Mora Urpí Escuela de Biología, Universidad de Costa Rica, San José, Costa Rica (Received fOI publication December 7, 1982) Abstraet: This is a preliminar y study of the Pejibaye (Bactris gasipaes) frond and begins with a general morphological description of the Icaf, complemcnting those already in existence. The data ohtained fr om Central American Pejibayes show that lhe rachis length of their fronds is considerably ¡onger lhan those recorded from Surinam populations and it is assumed that al least part of this difference has a genotypic background. It was also fa und, contrary to expectations, that the older fronds llave a longer rachis than younger mature ones fr om the same trunk. This difference could be due to seasonal environmental factors such aS variations in rainfall. The association of leaflets in groups, each leaflet having a diffe rent angle of insertion on the raehis wi thin the group, gives a many-planed le aflet arrangement that may inerease light-use efficieney. The groups are fo rmed by a variable number of Icaflets and conscquently arc not a reUable eharacter for use as a descriptor. The many-planed arrangement fades in oJd fr onds when drooping of the leaflets inereases, obscuring the cffeet of the different angles of insertion. lt was also found that as.<;ymetry exists·between opposite sides of the same frond in regard to leaflet number and length. An assymetrical fr ond outline is evident on both sides of the frond. A larger leaflet number was found in the younger fr onds when comparcd wi th the older ones from the same plant in spite of the longer rachis ofthe latter ones. This may indieate that even 12 year-old paIms have not reached the maximum development of their fronds since it is assumed that leaflet number is less affected by environmental variations tban raehis length. The frond of a palm is its principal this species various morphological charac photosynthetic organ and is ultimately teristics of the leaf are constant enough to be responsible for bio-mass increment. For this used in the description of varieties (Nixon, reason a knowledge of the frond is essential for 1950). For the Pejibaye palm (Bactris ga sipaes understanding many gene tic and agronomic H.B.K.) very few bibliographic references on data of a given species. the fronds are found. In the oil palm (E/aeis guineensis Jacq.) these studies were initiated in 1925 General morphology: Among the various (Broekmans, 1957) and have led to an extensive articles by taxonomists who have described the knowledge of the frond and 'its relation to Pejibaye palm, the one by Wessels·Boer (1965) growth and production. In Malaysia Hardon et is one of the most complete in terms of frond al. (I969) faund in the oil palm .a positive description: " .. .10-20 contemporaneous leaves correlation between leaf area and production, in a plt¡mose crown, leaves curling and although this was less evident in other studies in drooping, up to 3 m long of which the rachis West Africa (¡-Iartley, 1977). The study of the about 2 m long, more or less sparsely armed, fron d has led to the calculation of various with up to 120 pairs of pinnae clustered in to parameters such as the loaf Area Index, loaf groups of usually 4 inserted in various direc Area Ratio, Relative loaf Area Growth Rate, tions, 50·60 cm long, about 3 cm wide". The which permit a more detailed analysis of the works of Humboldt, Bonpland and Knuth growth of the palm (Hartley, 1977). (1816), Martius (1824) and Barbosa·Rodrigues The fr ond of the date palm (Phoenix dac ty· (1903) contain sorne information bu t give a less lifera L.) has also be en intensively studied. In complete description of the fr onds. 103 104 REVISTA DE BIOLOGIA TROPICAL In general, the aboye descriptions serve well Thus to distinguish between genotypes, a for the Pejibaye from Surinam and perhaps qualitative and quantitative analysis of the from the Amazon Basin as a whole. However, different characters is required. Prelirninary the plants studied in Costa Rica and reported information on sorne characters is here pres here show generally larger dimensions. Sorne ented, based on six different plants (B/3, 4/1, additions to this description that are worth 9/1, 16/1, 1/8 and 1/10). One -B/3- from mentioning come from Tomlinson (1961): Panamá, planted in 1973 (Morera Monge, " .. .leaflets narrowly lanceolate, reduplica te, 1981), and the other five from Costa Rica each with a prominent adaxial midrib, often planted in 1968; both collections at CATIE, setose. Rachis and petiole densely spinous Turrialba. The older frond was chosen between below". From MacBride (I960) : " ...midner ve positions 12 and 16 and the newer between 8 prominent aboye, the 3 or 4 secondaries on and 12, thus both were probably mature. both sides, margins, remotely setulose or mi nutely aculeate ..." From Martius (1824) (trans Rachis length: The rachis lengths observed lated by W. Rodriguez, pers. com.): " ...petiole are presented in Table l. One notes imme base convex vaginate lanceolate ... rachis bifacial diately that these fronds are indeed much larger on dorsal part, on ventral side convex, spiny ... than those described from Surinam (Wessels pinnae ... the more apical and basal crispa te, Boer, 1965), the smallest being more than 50 linear-lanceolate acuminate ... " cm longer and the largest 130 cm longer. As With these additions and the observations mentioned aboye this may be due to the genetic gathered fo r this article, a more detailed variation between the populations of Pejibaye description may be presented: 10 to 25 (rarely from Costa Rica and those of Surinam, or to reduced to 4 or 5, or up to 36) contempora the richer soils of Costa Rica, or to a combi neous fronds in a plumose crown, fronds cur nation of both. Further study is necessary to ving, when old even curling, and drooping; 300 determine the cause of these differences. One to 450 cm in length of which the rachis is 200 also notes that the younger group of fronds is to 330 cm long; pe tiole sheath involves one half srnaller. This result was not expected to be so to two thirds trunk circumference, margins bro constant as is shown here. In the oil palm, the ken somewhat fibrous, con cave aboye, convex size of the leaf increases during the juvenile below (Fig. 1); rachis concave becoming bifacial stage, until it reaches fu ll maturity, after which aboye, convex beneath ; spinyness absent to it stabilizes fo r a while and then starts to extremely dense on trunk and frond, usually diminish slightly (Hardon, et al. 1969; Corley more or less aligned on petiole, less so on and Gray, 1976), although the age factor alone rachis, occasionally stattered; leaflets in groups cannot explain these results in Pejibaye. The of to 9 with lower leaflet of group more :2 difference in size too large between fron ds perpendicular to the rachis dorsel surface, close in age. In addition,is on e of the palms -B/3 following leaflets less perpendicular, giving a from Panamá- also presents a rachis length many-planed arrangement; 110 to 140 leaflets reduction which is not expected to occur an per side, arranged in sup-opposite pairs or immature plant. Because all the palms showin the alternately on rachis with more one side 011 same size reduction one must assume that there than on the other, basal and apical leaflets is an environmental factor, such as rainfall, that frequently fu sed with grouping less pronounced influences rachis length and frond size. Such or a bsent ; leaflets linear-lanceolate changes must consequently be seasonal and bi-acuminate , length 20 to 80 cm, width 3 to superimposed on the change which occurs with 5.8 cm; midnerve prominent aboye, with 3 or 4 age. This subject obviously requires more study. secondaries on both sides; midvein, secondaries and/or margin of leaflets may present small Leaflet number: The number of leaflets per spines; leaflet base convex vaginate-lanceolate frond and per frond sides are presented in Table at insertion on rachis; leaflets glossy dark green 2. In all the fr ortds studied there was a notable a boye , opaque green below with more spacial difference in the point of insertion of trichomes and stomata below; fr ond outline the first basal leaflets between the two sides of irregular to occasionally ovoid or obovoid. the fr ond. It will be noted that plants 9/1 and Although this description is more detailed 16/1 have more leaflets on the right side, while than the others given here, it is still general. B/3, 4/ 1, 1/8 and 1/10 ha ve more on the left CLEMENT & MORA URP/: Leaf morphology of Bactris gasipaes 105 fronds. This is to be expected on immature plants where leaflet number is assumed to increase with age until maturity and then remain relatively constant. The diffe rence is only 1.67% and so statistically not significant; and consequently, it may be attributed to environmental influence . However, comparing Tables 1 and 2, the younger fronds are shorter and have more leaflets than the older, so that one finds more leaflets per meter on the younger fronds. Thus the older fronds have an average of 86.8 leaflets per meter while the younger have 96.6 (11.3% more per meter).