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WOOD ANATOMY of FOUR MYRTACEAE GENERA in the SUBTRIBE MYRCIINAE from SOUTH AMERICA By

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WOOD ANATOMY of FOUR MYRTACEAE GENERA in the SUBTRIBE MYRCIINAE from SOUTH AMERICA By IAWA Journal, Vol. 16 (I), 1995: 87-95 WOOD ANATOMY OF FOUR MYRTACEAE GENERA IN THE SUBTRIBE MYRCIINAE FROM SOUTH AMERICA by Claudia Luizon Dias-Lerne!, Peter Gasson2 & Eirnear Nie Lughadha2 SUMMARY The wood anatomy of 31 species representing four genera of subtribe Myrciinae, Myrtaceae, Calyptranthes, Gomidesia, Marlierea and Myrcia is described. In general the wood of subtribe Myrciinae can be char­ acterised by solitary vessels, simple perforations, alternate vestured pits, fibres with bordered and/or vestured pits (fibre-tracheids), parenchyma scanty paratracheal, diffuse and/or diffuse-in-aggregates forming com­ plete or interrupted bands, and heterocellular rays with disjunctive cell walls. The four genera share all these features, but exhibit considerable variation in axial parenchyma patterns and crystal occurrence and distri­ bution. No anatomical features serve to separate the genera, but some, e. g., helical thickenings and crystals, appear to be restricted to species in one genus or another. All wood sampies were obtained from wood collections and the exact environmental conditions in which the trees grew are unknown. For this reason, it is impossible to assess the relative contributions of ecological and taxonomic factors to the observed anatomical variation. Key words: Myrtaceae, Calyptranthes, Gomidesia, Marlierea, Myrcia, South America. INTRODUCTION The Myrtaceae is a large family of trees and shrubs consisting of about 3850 species in 127 genera (Brummitt 1992, Mabberley 1987). The family is mainly tropical and subtropical worldwide and is well-represented in tropical South America. All but one of the South American species belong to the Myrteae, the baccate-fruited tribe. The most recent comprehensive revision of this tribe is that of Berg (1855/56), in which 3 subtribes are recognised within the Myrteae. The present study deals with the subtribe Myrciinae in which Berg distinguished 11 genera. Of these, only 5 (Calyptranthes, Gomidesia, Marlierea, Myrceugenia and Myrcia) are now recognised, the others being relegated to synonymy. Myrceugenia is very different from these Myrciinae and is postulated to be the primitive genus within the subtribe (Landrum 1981). This arrange­ ment was first suggested by McVaugh in his re-alignment of the American Myrteae (1968). He regarded the myrcioid genera as "perhaps the most readily compre- 1) Depto de Botänica, lust. Biociencias, Universidade de Silo Paulo, Silo Paulo, Brazil. 2) Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, United Kingdom. Downloaded from Brill.com10/02/2021 06:58:30AM via free access 88 IAWA Journal, Vol. 16 (1),1995 ----- Table 1 Species Character (see key below): 2 3 4 5 6 7 8 9 10 11 12 13 Calyptranthes jasciculata Berg + + + + + >5 grandifolia Berg + + + + >8 lucida Mart. ex DC. + + + + + >8 pileata D. Legrand + + + + + + - 4-8 pulchella DC. + + + + + >8 speciosa Sagot + + + + + >8 widgreniana Berg + + + + >8 Gomidesia affinis (Cambess.) D. Legrand + + + - 5-8 anacardiifolia Berg + + + + + + + 4-8 sellowiana Berg + + + + + + - 4-8 tijucensis (Kiaersk.) D. Legrand + + + + + + + 4-8 Marlierea montana (AubI.) Amshoff + + + + + >8 parvijIora Berg + + + + + >8 scytophylla Diels + + + + + >8 spruceana Berg + + + + + + >5 sylvatica (Gardner) Kiaersk. + + + + + - 5-8 umbraticola (Kunth) Berg + + + + + + >8 Myrcia amazonica DC. + +/- +/- + + +/- - +/- - >8 bracteata (Rieh.) DC. + + + + + + 3-8 deflexa DC. + + +/- +/- + + - 3-8 jallaxDC. + + +/- +/- + + + +/- - 3-8 grandis McVaugh + + + + + + >5 guianensis DC. + + + + + >5 inaequiloba (DC.) D. Legrand + + + + + + >5 multijIora (Larn.) DC. + + + + + + - 3-8 pyrifolia (Desy. ex Harn.) Nied. + + + + + + + >5 servata MeVaugh + + + + - 3-8 splendens (Swartz) DC. + + + + + + - 3-8 subobliqua (Benth.) Nied. + + + + + >5 sylvatica (G. Meyer) DC. + + + + + >5 tomentosa (AubI.) DC. + + + + + - 4-10 1. Growth rings 8. Axial parenchyma in bands more than three 2. Fibres thick-walled cells wide 3. Fibres thin-walled 9. Axial parenehyma with erystals 4. Axial parenchyma diffuse 10. Dark contents 5. Axial parenchyma diffuse in aggregates 11. Pith flecks 6. Axial parenchyma scanty paratracheal 12. Helical wall thickenings in fibres 7. Axial parenchyma in bands up to three 13. Axial parenchyma strand length (number of cells wide cells) Downloaded from Brill.com10/02/2021 06:58:30AM via free access Dias-Lerne, Gasson & Lughadha - Wood anatorny of S. Arnerican Myrciinae (Myrtaceae) 89 ---~ hensible of the groups of Myrteae." He also highlighted the absence of sharp dis­ tinctions between the genera and suggested that if all historical considerations could be set aside, Calyptranthes, Marlierea and Myrcia might be combined in one vast genus. As currently recognised Gomidesia is distinct by virtue of its 4-locular anthers, the other 3 genera having bilocular, longitudinally dehiscent and explanate anthers. Both Gomidesia and Myrcia have more or less regularly 5-lobed calyces, while Mar­ lierea has an irregularly splitting 3- to 5-lobed calyx and Calyptranthes has a closed circumscissile calyx. However, both anther and calyx characters vary continuously and, as Mc Vaugh emphasised, there are many borderline species which have to be assigned arbitrarily to one genus or another. These problems of generic delimitation prompted the present study, which com­ prises a survey of the wood anatomy of 31 species representing the spectrum of diver­ sity of the myrcioid genera and a review of previously published wood anatomical data for this group. There are no detailed comparative works on the wood anatomy of these four genera, although they are all included in the general description of Myrtaceous wood by Metcalfe and Chalk (1950). Detienne et al. (1982) and Detienne and Jacquet (1983) include representatives of Calyptranthes, Marlierea and Myrcia. Ilic (1991) illustrates Calyptranthes and Myrcia and Schmid and Baas (1984) describe vessel characters of Myrcia. The only description of Gomidesia wood anatomy (Barbosa et al. 1977/78) appears to be based on a collection of Myrcia pubipetala Miq. errone­ ously identified as Gomidesia crocea (Vell.) Berg. We have examined 7 species of Calyptranthes, 4 of Gomidesia, 6 of Marlierea and 14 of Myrcia (Table 1) to provide the most detailed wood anatomical account of the genera to date. MATERIAL AND METHODS Wood samples and slides of the 31 species in 4 genera studied in this paper were ob­ tained from: Institute of Systematic Botany, University of Utrecht, The Netherlands (Uw); Museum Collections, Royal Botanic Gardens, Kew (Kw) and Rijksherbarium, Leiden, The Netherlands (all slides in Leiden were originally from the Oxford For­ estry Institute, FHOw). The abbreviations for wood collections follow those in Stern (1988). Material studied: Calyptranthes Jasciculata Berg: Guyana, Uw 997 and FHOw 14782. - C. grandifolia Berg: Uw 6382. - C. lucida Mart. ex DC.: Suriname, Uw 2522. - C. pileata D. Le­ grand: Brazil, Uw 14519. - C. pulchella DC.: Brazil, Uw 19683. - C. speciosa Sagot: Suriname, Uw 340. - C. widgreniana Berg: Brazil, Uw 6383. - Gomidesia affinis (Carnbess.) D. Legrand var. catharinensis D. Legrand: Brazil, Uw 12946 and Uw l34l3. - G. anacardiifolia Berg: Brazil, Uw 6381, Uw 6903 and Kw 22082. - G. sellowiana Berg: Brazil, Uw 14028. - G. tijucensis (Kiaersk.) D. Legrand: Brazil, Uw 14341. - Marlierea montana (AubI.) Amshoff: Guyana, Uw 1003 and FHOw 14786. - M. parviflora Berg: Brazil, Uw 6923. - M. scytophylla Diels: Brazil, Uw 8069. - M. spruceana Berg: Brazil, Uw 7864. - M. sylvatica (Gardner) Kiaersk.: Brazil, Uw 12536. - M. umbraticola (Kunth) Berg: Brazil, Uw 7502 and Uw 7792. - Myrcia amazonica DC.: Suriname, Uw 2l3. - M. bracteata DC.: Surinam, Uw 1630. - M. deflexa DC.: Surinarne, Uw 1744 and Uw 2529. -M. JaUax DC.: Surinarne, Uw 1744 and Venezuela, Uw 1887 and Uw Downloaded from Brill.com10/02/2021 06:58:30AM via free access 90 IAWA Journal, Vol. 16 (I), 1995 ---------------- 12195, - M. grandis McVaugh: Guyana, Uw 21510, - M. guianensis DC. var. guianensis: Gu­ yana, Uw 21634. - M. inaequiloba (DC.) D. Legrand: Guyana, Uw 21514 and Uw 21540. -M. multiflora (Larn.) DC.: Brazil, Kw 22097. -M. pyrifolia (Desv. ex Harn.) Nied.: Surinarne, Uw 335. -M. servata McVaugh: Guyana, Uw 1006. -M. cf. splendens (Swartz) DC.: Kw 71381 and Uw 1466. - M. subobliqua (Benth.) Nied.: Guyana, UwlO04 and FHOw 14789. - M. sylvatica (G.Mey.) DC.: Surinarn, Uw 2528. - M. tomentosa (AubI.) DC.: Guyana, Uw 32229. The wood sampies were boiled in water. Transverse and longitudinal (tangential and radial) sections were then cut at 20 - 25 /JIll on a Reichert OME sliding microtome, stained in 1% safranin in 50% alcohol and 1% aqueous alcian blue, taken through an alcohol series to 100% ethanol, cleared in histoclear and mounted in euparal. For scanning electron microscopy (SEM), wood sections were split longitudinally and air-dried. Dried specimens were then sputter-coated with gold and viewed in radial longitudinal section using a Cambridge 240 SEM at 18 K v. Photomicrographs were taken at various magnifications using SEM and light mi­ croscopy. The IAWA List of Features for Hardwood Identification (IAWA Committee 1989) was used to interpret the microscopical observations. GENERAL DESCRIPTION OF THE SUBTRIBE Wood: diffuse porous. Vessels (Figs. 1-4,9-10): typically solitary (Figs. 1-4), perforations simple (Fig. 9), intervessel pits alternate, vestured (Fig. 10); vessel to ray pitting of same type as the intervascular pitting. Fibres (Figs. 11-12): mostly thick-walled with bordered pits in both
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