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202 IAWA Bulletin N.S., Vol. 8 (3),1987 MORPHOLOGY of the VESSEL

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202 IAWA Bulletin N.S., Vol. 8 (3),1987 MORPHOLOGY of the VESSEL 202 IAWA Bulletin n.s., Vol. 8 (3),1987 MORPHOLOGY OF THE VESSEL ELEMENTS IN THE SECONDARY XYLEM OF THE MYRISTICACEAE FROM BRAZILIAN AMAZONIA by Pedro L. B. Lisboa*, J. Cesar A. da Silva*, A. A. Loureiro**, and Gracielza M. dos A. dos Santos * Summary A morphological study of vessel elements foration plates in some genera, thus making his was carried out in species representing the five key somewhat confusing. genera of Myristicaceae present in the Brazilian In this study, we examined a large quantity Amazon region. The results show that perfora­ of wood specimens obtained from the five tion plates of the scalariform type predominate genera present in the region. Our objective was in Compsoneura, lryanthera and Otoba, wherc­ to document their full range of morphological as perforation plates of the simple type pre­ variability associated with vessel elements. Af­ dominate in Osteophloeum and Virola. The ter characterising these structures in each genus, phylogenetic and taxonomie implications of we consider the implications for the phylogeny these findings are discussed. and taxonomy of the regional genera as a Key words: Compsoneura, lryanthera, Osteu­ group. phloeum, Otoba, Virola, Vessel perforations. Material Introduction In Brazilian Amazonia, the species of Myris­ The family Myristicaceae, widely distributed ticaceae are distributed as folIows: Compsoneu­ in tropical regions, is represented in the Brazilian ra (5), lryanthera (19), Osteophloeum (I), Oto­ Amazon region by five genera (Compsoneura, ba (I) and Virola (29) (Rodrigues, 1978, 1981, Iryanthera, Osteophloeum, Otoba and Virola) 1982). Of the 55 species in the region, 41 were and 55 species. Smith and Wodehouse (1937), examined in this study. In the genus Compso­ Ducke and Black (1954), and Rodrigues (1978) neura, only the wood of C ulei was examined considered Amazonia to be the distribution due to a lack of specimens of other species, centre of this family in the Americas. which are all rare. Specimens were obtained The idea that vessel elements in Angiosperms from the wood collections of the Instituto Na­ originated from more primitive vascular ele­ cional de Pesquisas da Amazönia (INPA) in ments known as tracheids has been weil accept­ Manaus, Brazil; the Centro de Pesquisas Agro­ ed for some time (Frost, 1930; Cheadle, 1943). pecuaria do Tr6pico Omido (lAN) and the Mu­ Through successive morphological modifica­ seu Goeldi (MG) in Belem, Brazil; the Jardim tions, vessel elements transformed into a com­ Botänico (RB) in Rio de Janeiro, Brazil; the pact type with simple perforation plates. This University of Utrecht (U) in Holland; and the general evolutionary sequence gave rise to vari­ Forest Products Laboratory (MAD) and the ous levels of specialisation that are used by Samuel Record Collection (SJR) in Madison, anatomists as a valid parameter for clarifying Wisconsin, U.S.A. the phylogeny and taxonomy of botanical groups. Material studied: Little has been published concerning the Compsoneura Warb.: C ulei Warb.: Amazo­ morphology of vessels in the Myristicaceae of nas, INPA 142, INPA 4799, INPA 8054; Para, Brazilian Amazonia. The most comprehensive MG 2346; Rondönia, MG 4108. publication is by Garrat (1933), who studied lryanthera Warb. - I. campinae W. Rodr.: the wood anatomy of the family and construct­ Amazonas, INPA 6972. - I. coriacea Ducke: ed a key to separate genera on the basis of Amazonas, INPA 786, INPA 7331. - I. crassi­ characters associated with the type of vessel folia A.C. Smith: Amazonas, MAD 15218. - I. perforation plates. Due to a lack of sufficient elliptica Ducke: Amazonas.IAN 101849: INPA material, Garrat was unable to interpret accu­ 2895, INPA 5614; MAD 14105. - I. grandis rately the occurrence of various types of per- Ducke: Amazonas, INPA 5809: Mato Grosso, * Department of Botany, Museu Paraense Emilio Goeldi, Caixa Postal 399, 66.000 Belem, Para, Brazil. ** Instituto Nacional de Pesquisas da Amazonia, Caixa Postal4 78, Manaus, Amazonas, Brazil. Downloaded from Brill.com09/29/2021 09:37:21PM via free access IAWA Bulletin n.s., Vol. 8 (3),1987 203 Uw 19330; Pani, INPA 6052. - 1. hostmanni tions of acetic acid and hydrogen peroxide (Benth.) Warb.: Maranhäo, MG 2358. - 1. inpae (30% vol.) at 60° C for 24 hours. The sampIes W. Rodr.: Amazonas, INP A 5181. - 1. juruen­ were subsequently washed in running water sis Warb.: Amazonas, INPA 5430; MG 1022; and stored in beakers with water and a few Pani, SJR 22074. - 1. laevis Markgraf: Amazo­ drops of 2% formaldehyde. Prior to preparing nas, INPA 4061; Roraima, INPA 4613. - 1. lan­ slides, each sam pIe was stained with a few cifolia Ducke: Amazonas, INPA 5510, INPA drops of a safranin in a water-ethanol solu­ 5529, INPA 5926. - 1. macrophylla (Benth.) tion, and then mounted on a minimum of five Warb.: Amazonas, INPA 1005, INPA 1536, slides in glycerin. A minimum of 25 vessel ele­ INPA 3375. - 1. olacoides A.C. Smith: Rorai­ ments were examined per sampIe. ma, INPA 4401. - 1. paradoxa (Schw.) Warb.: The length of the vessel elements were mea­ Amazonas, INPA 4989, INPA 5118,INPA 5168. sured with the aid of an 01ympus projector un­ - 1. paraensis Huber: Amazonas, INPA 1992, der 100 x magnification and included the entire INPA 4461, INPA 5156. -I. polyneura Ducke: vessel length plus terminal appendices. Diame­ Amazonas, INPA 4915, INPA 5333, IN PA ters were measurcd using cross sections on all 5819, INPA 5986. - 1. sagotiana (Benth.) slides from the Museu Goeldi and INPA collec­ Warb.: Amapa, INPA 1313, INPA 1314. - 1. tions. Counting of the bars in the scalariform tessmannii Markgraf: Amazonas, MAD 22912.- plates was done using a light microscope, which 1. tricornis Ducke: Amazonas, INPA 5506; Ro­ was found to produce more clarity than the raima, INPA 4466. - 1. ulei Warb.: Amazonas, projector. INPA 1360, INPA 5021, INPA 5081. Botanical specimens corresponding with the Osteophloeum Warb. - 1. platyspermum wood sampIes were identified by Dr. William (A.DC.) Warb.: Amazonas, MG 962, MG 2758; Rodrigues of the Botany Department of the INPA 5334, INPA 6192; Para, lAN 136833. Instituto Nacional de Pesquisas da Amazönia Otoba DC. ex Karst. - 0. parvifolia (M.G.F.) (INPA), a specialist on Neotropical Myristica­ A. Gentry: Acre, IAN 133382; Amazonas, INPA ceae. 6231. The terminology utilised in this paper fol­ Virola Aublet. - V. albidiflora Ducke: Ama­ lows that adopted by the Committee on No­ zonas, MAD 14004. -- V. caducifolia W. Rodr.: menclature of the International Association of Amazonas, INPA 5577. - V. calophylla Spr. ex Wood Anatomists (lAWA, 1964). Classification Warb.: Amazonas, INPA 4750, INPA 5027, IN­ of measurements followed the Panamerican PA 6468. - V. carina ta (Benth.) Warb.: Pani, Commission of Rules and Techniques (Comision MG 1673. - V. coelhoi W. Rodr.: Amazonas, Panamericana, 1974). INPA 6200. - V. decorticans Ducke: Amazo­ nas, INPA 15427. - V. divergens Ducke: Ama­ Results and Discussion zonas, INPA 5050. - V. duckei A.C. Smith: Tables 1 and 2 summarise the morphological Amazonas, INPA 5164. - V. elongata (Benth.) characteristics of the vessel elements of the five Warb.: Amazonas, IAN 115634; Para, IAN genera of Myristicaceae examined in this study. 86303, 86353; MG 1814. - V. flexuosa A.C. The morphology reveals an extremely wide Smith: Amazonas, INPA 6189. - V. guggen­ range of forms, including vessels that are short heimii W. Rodr.: Amazonas, IAN 99848; IN PA to 10ng, narrow to wide, and with perforation 5370. - V. lorentensis A.C. Smith: Amazonas, plates located at diverse sites along the vessel RB 2854. -- V. michelii Hecke!: Amapa, MG walls, with their orientation varying from 574; Amazonas, INPA 3259; Pani, IAN 101594, slightly inclined (i. e., almost horizontal) to ver­ IAN 106978. - V. minutiflora Ducke: Amazo­ tical. The perforation plates are likewise of nas, INPA 5390. - V. multicostata Ducke: variable form, not only at the generic but also Amazonas, INPA 3231; Pani, MG 1912. - V. at the specific level. Plates of the simple, scala­ multinervia Ducke: Amazonas, INPA 3855; riform, and reticulate type are common, but Para, IAN 101597. - V. sebifera Aubl.: Rorai­ atypical types occur that appear to be a mix­ ma, INPA 4452. - V. surinamensis (Rol.) Warb.: ture of scalariform and reticulate; these were Amazonas, MG 527; INPA 710; Para, IAN named semi-scalariform. 106872. - V. venosa (Benth.) Warb.: Amazo­ Despite the wide morphological range of per­ nas: INPA 115761. foration plates within each genus, it was pos­ sible to discern apredominant type. Scalari­ Methods form plates are common in all the genera stud­ Sam pIes consisting of longitudinal slices of ied but are especially predominant in Compso­ the heartwood were obtained from each of the neura (Plate1: 1-4) and Otoba (Plate 1: 5-8); wood specimens studied. The sam pIes were although present in all the species of Iryanthera. macerated in a solution containing equal por- they only attain similar predominance in thc Downloaded from Brill.com09/29/2021 09:37:21PM via free access 204 IAWA Bulletin n.s., Vol. 8 (3),1987 Table 1. Characters of vessel elements in Myristicaceae from Brazilian Amawnia. I = inclined, FI = sttongly inclined, SI = slightly inclined, V = vertical. s.d. = standard deviation of the mean. For the characters simple, reticulate, semi-scalariform and scalariform plates, two values indicate the minimum and maximum within the species examined; a single value indicates that only one specimen was examined. ~ ~ '" ""'0 '-' r-.. ~ ~ § '-' r-.. -a 'J:l .~ ~ .~c:: '"0.> '" ~ ~ ~ "'0.> e Kl .,gE .!ii
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