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The Occurrence of Scalariform Perforation Plates and Helical Vessel Wall Thickenings in Wood of Myrtaceae

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The Occurrence of Scalariform Perforation Plates and Helical Vessel Wall Thickenings in Wood of Myrtaceae IAWA Bulletin n.s., Vol. 5 (3), 1984 197 THE OCCURRENCE OF SCALARIFORM PERFORATION PLATES AND HELICAL VESSEL WALL THICKENINGS IN WOOD OF MYRTACEAE by Rudolf Schmid and Pieter Baas Department ofBotany, University ofCalifornia, Berkeley, CA 94720, U.S.A. and Rijksherbarium, P. O. Box 9514, 2300 RA Leiden, The Netherlands Summary Introduction The occurrence of multiple perforation The Myrtales, as recently defined, have either plates and helical wall thickenings in vessel ele­ eleven or twelve families (respectively, Dahlgren ments of 144 species (plus 12 varieties and 2 & Thorne, 1984, and Cronquist, 1981, the lat­ hybrids) in 53 genera of Myrtaceae was exten­ ter including Thymelaeaceae). The Myrtaceae sively explored. Scalariform perforation plates represent the only myrtalean family in which occur in 40 species (plus I hybrid), in Luma, some species have wood with scalariform per­ Myrceugenia, Myrteola, Ugni, and in the mono­ foration plates (Metcalfe & Chalk, 1950; Van typic Myrtastrum rujo-punctatum, Neomyrtus Vliet & Baas, 1984). As summarised in Table I, pedunculata, and Tepualia stipularis. Ugni can­ there are vague reports of such in the old litera­ dollei also has foraminate (i.e., sieve-like) per­ ture, but the first specific record is Reiche's foration plates. Helical wall thickenings occur (1897). Without giving details, he attributed in 33 species (plus 1 hybrid), in Acmena, Aus­ scalariform perforation plates to the Chilean­ tromyrtus, Myrceugenia, Myrcia, Myrcianthes~ Argentinean Eugenia apiculata, also known as Psidium, Xanthomyrtus, and in Myrtus com­ Myrceugenia apiculata, Myrceugenella apiculata, munis. Most of these records are new. The spe­ or, most properly sensu Landrum (1981, pers. eies with exclusively scalariform perforation comm. 1983) and McVaugh (1968), Luma api­ plates (in Luma, Myrteola, Neomyrtus, and culata. Record and Hess (1943) and others have Ugni) are from cool mesic habitats; those with confirmed this re cord (Tables 1, 2). mixed simple and multiple perforation plates Metcalfe and Chalk (1950: 625) attributed are also largely cool mesic but show a some­ scalariform perforation plates to Myrceugenia, what greater diversity of habitats. Myrtaceae Myrtus communis, and an unspeeified species with exclusively simple perforation plates pre­ of Eugenia 'cultivated at Kew' (Table 1). No dominate in all habitat types. Helical wall thick­ species of Myrceugenia, a genus of 38 species enings occur sporadically throughout the dis­ (Landrum, 1981), was indicated, but perhaps tributional range ofthe family. However, tropi­ this is the aforenoted Luma apiculata. The re­ cal species tend to have weaker helical thicken­ ports for simply 'Eugenia' (Table 1) are mean­ ings than the subtropical and temperate species ingless as records because the genus in its exhibiting them. The possible functional signi­ broadest sense incIudes some 70 genera and ficance of these ecological tendeneies is discus­ over 2800 binomials (Schmid, 1972). Presum­ sed. It is hypothesised that multiple perfora­ ably Metcalfe and Chalk's 'cultivated' Eugenia tion plates were retained in some cool mesic is one of the commonly cultivated South Amer­ Myrtaceae because of a lack of strong selective ican species of 'Eugenia' or 'Myrtus' (including pressure to eliminate them from this type of Luma apiculata and Ugni molinae), which are environment, rather than that they were re­ hardy in England (Chittenden, 1956), which tained because of adaptive significance in trap­ have scalariform perforation plates (Table 2), ping embolisms. The systematic and diagnostic and which are properly referable to other genera value of multiple perforation plates and helical (McVaugh, 1968). As discussed below, the re­ wall thickenings is also discussed. Scalariform port of scalariform perforation. plates for Myr­ plates are largely confined to related genera in tus communis is erroneous, as is Fasolo's (1939- Myrtoideae; Tepualia is the only representative 40) report for Syzygium guineense. from Leptospermoideae. Helical wall thicken­ Arecent and fully documented account of ings are only of limited diagnostic and systema­ scalariform perforation plates in Myrtaceae is tic value above the species level. Butterfieid and Meylan's (1974, 1980; Meylan Key words: Scalariform perforation plates, fo­ & Butterfield, 1975, 1978a) for the monotypic raminate perforation plates, helical wall thick­ Neomyrtus pedunculata of New Zealand. How­ enings, Myrtaceae, ecological and functional ever, only simple perforation plates occur in the wood anatomy, systematic wood anatomy. other four New Zealand genera of Myrtaceae of Downloaded from Brill.com10/10/2021 12:05:33PM via free access 198 IAWA Bulletin n.s., Vol. 5 (3),1984 Table 1. Literature reports of scalariform perforation plates and helical wall thickenings in vessel elements of wood of Myrtaceae l REPORTS OF SCALARIFORM PERFORATION PLATES Niedenzu 1893: 58: Myrtaceae with vessel elements 'meist einfach, zuweilen aber, wie bei manchen Eugenia-Arten, auch leiterförmig perforiert'; this report, which perhaps refers to Syzygium, apparently is based on original observations. Reiche 1897: 89: Luma apiculata ('Eugenia apiculata '), but no details are given. Solereder 1899: 400: Myrtaceae with vessel elements 'entweder einfach oder ausschliesslich leiter- förmig (Eugenia apiculata DC. nach Reiche) perforiert.' Solereder 1908: 354: English translation of preceding. Brown 1922: 318: Myrtaceae 'with simple or scalariform perforations.' Hegi 1925-26: 774: Myrtaceae 'mit einfachen oder ausschliesslich leiterförmigen Durchbrechungen.' Fasolo 1939-40: Syzygium guineense, totally scalariform, with many bars per perforation plate. Record & Hess 1943: 404: Myrtaceae with perforation plates simple, except in Luma apiculata ('Myrceugenia apiculata '), 'which has many-barred scalariform plates.' Wagemann Wiedenbrug 1949: 342: Luma apiculata ('Myrceugenella apiculata '), with numerous bars per perforation plate. Metcalfe & Chalk 1950: 625: Myrtaceae with 'perforation plates simple, except in Myrceugenia and Myrtus communis L., in which they are exclusively scalariform with about 15-25 fine bars. Scalariform perforation plates also observed in a young stern of Eugenia sp. cultivated at Kew.' Tortorelli 1956: 570: Luma apiculata ('Myrceugenella apiculata '), with numerous bars per perfora­ tion plate. Emberger 1960: 1436: 'perforations scalariformes chez certaines especes.' Butterfieid & Meylan 1974: Neomyrtus pedunculata, with vestures and microfibrillar webs in some of the perforations (SEM study). Meylan & Butterfieid 1975: 3, 1978a: 100: same information as preceding. Ragonese, pers. comm. Oct.1978 to Schmid: in the 9 species of Myrtaceae she studied (see Ragonese, 1976) scalariform perforation plates observed only in Luma apiculata ('Myrceugenella apicu­ lata'), with '20-30 fine bars' per perforation plate. Butterfieid & Mey1an 1980: 66: same information as their 1974 paper above. Schmid (980: 568: Luma apiculata and L. chequen, but not in Myrtus communis. Schmid, pers. comm. in Landrum (1981: 18): Myrceugenia, Luma, Myrteola, Ugni, and Neomyrtus, but not in Myrtus communis. Landrum 1981: 17: Luma apiculata and 6 species of Myrceugenia: M. alpigena var. alpigena (but not in var. ruta), M. lanceolata (SEM photographs of both simple and sca1ariform perforation plates), M. ovata var. acutata and var. gracilis, M. planipes, M. pilotantha var. pilotantha (2 sampies), M. reitzii. Metcalfe & Chalk 1983: 202: in some Myrtaceae. Van Vliet & Baas 1984: Luma (= probably L. apiculata), and citations for some earlier literature. REPORTS OF HELICAL WALL THICKENINGS IN VESSEL ELEMENTS Sanio 1863: 402: Myrtus communis, but helices absent from tracheids. Moll & Janssonius 1918: 389: Acmenaacuminatissima ('Eugeniaacuminatissima'), with 'spiralige Streifung auf der Innenseite der Gefässwände vorhanden'; on pp. 443-444 noted that helices are present only when bordering on parenchyma and only in I of 2 sampies; on pp. 444-445 two other varieties of E. acuminatissima (var. angustitolia, var. oblonga) said to have very similar wood, but helices are not mentioned. This is a completely reliable report and needs no independent verification. Record & Hess 1943: 404: absent except in Myrceugenia schultzei and M. ternandeziana. Wagemann Wiedenbrug 1949: 343: Myrceugenia exsucca, with delicate helices. Metcalfe & Chalk 1950: 625: 'in some species of Eugenia and Myrceugenia'; report of Moll & Jans­ sonius (1918) noted. Greguss 1959: 232: Myrtus communis, in some of the vessel elements, especially narrow ones (heli­ ces also in tracheids and fibre-tracheids). Downloaded from Brill.com10/10/2021 12:05:33PM via free access IAWA Bulletin n.s., Vol. 5 (3),1984 199 Landrum 1981: 17: 9 species of Myrceugenia: helices weakly present in vessel elements of M. acuti­ flora, M. glaucescens var. latior, M. lanceolata, M. miersiana (only in 1 of 2 sampies); helices conspicuously present in M. alpigena var. alpigena and var. rufa, M. euosma (SEM photograph), M. ovata var. acutata (but not in var. gracilis), M. oxysepala, M. rufescens. Baas et al. 1983: 146: Myrtus communis. Metcalfe & Chalk 1983: 203: in Myrtaceae. 1 Only positive literature records are noted; a few authors (e.g., Ingle & Dadswell, 1953: 388 - see Introduction, Landrum, 1981) have noted the absence of scalariform perforation p1ates and(or helical wall thickenings. The records for scalariform perforation plates in Syzygium guineense and Myrtus communis are erroneous - see Discussion. Bierhorst & Zamora (1965:
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