IAWA Bulletin n.s., Vol. 5 (3), 1984 197

THE OCCURRENCE OF SCALARIFORM PERFORATION PLATES AND HELICAL VESSEL WALL THICKENINGS IN WOOD OF

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 , as recently defined, have either plates and helical wall thickenings in vessel ele­ eleven or twelve families (respectively, Dahlgren ments of 144 (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 , some species have wood with scalariform per­ , Myrteola, , 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, ~ Argentinean apiculata, also known as Psidium, Xanthomyrtus, and in 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 , 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 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 . 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 ), 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 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

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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: , 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).

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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: 702) observed sca­ lariform perforation plates in primary xylem of sp.

which ten of the 18 species have been studicd Leiden, where some 50 additional collections xylomatically (Table 2, Ingle & Dadswell, 1953; were sampled. We tried to sam pie the young Meylan & Butterfie1d, 1975, 1978a, b). wood of species Iikely to have scalariform per­ It is noteworthy that Ing1e and Dadswell foration plates or helical wall thickenings, or (1953) in a detailed analysis of the wood of both, that is, species occurring either in tropi­ some 354 species (including 177 of , cal montane floras or in temperate, high latitu­ also 108 species of this genus in Dadswell, tude ones (Baas, 1973, 1976, 1982, 1984; Baas 1972) and some 35 genera (many sensu latissi­ et al. , 1983; Carlquist, 1975, 1980; Fahn et al. , mo) of mainly southwest Pacific Myrtaceae in press). could find 'no sign of scalariforrn perforation plates' (p. 395). A review of other works on Materials and Methods wood anatomy of Myrtaceae (e.g., Moll & J ans­ Tables 2 and 3 give voucher details and other sonius, 1918, and references in Gregory, 1980, information for the species anatomically sam­ Metcalfe & Chalk, 1950, Solereder, 1908, Van pled. Generic nomenclature generally follows Vliet & Baas, 1984) revealed no mention of ad­ Briggs and J ohnson (1979), Landrum (1981), ditional species with scalariforrn perforation McVaugh (1968), and Schmid (1972), and sub­ plates. Table I summarises the literature. familial nomenclature follows Briggs and J ohn­ Intrigued by the reports of scalariforrn per­ son (1979) and Schmid (1980). We adopted foration plates in thc temperate Luma apicula­ Landrum's species concepts of Myrceugenia ta and Neomyrtus pedunculata, and aware of and Merrill and Perry's (citations in Schmid, the implications of this from the perspective of 1972) of Syzygium. 'ecologieal wood anatomy' (Baas, 1973, 1976; Most wood sam pies were macerated with Carlquist, 1975, 1977), the senior author in Jeffrey's fluid and stained heavily with safranin 1978 began to survey ChiIean and other tem­ (up to 50 hours, and usually in a 60° C oven). perate, mainly American species of Myrtaceae For details and some innovations on this tech­ for type of perforation plate. Scalariform per­ nique see Schmid (1982b). At Leiden addition­ foration plates were found in a fair number of al wood macerations were prepared by Frank­ species, nearly all , and nearly all lin's method (Jane, 1970) and stained with with wood anatomy previous1y unstudied as astra blue. We examined for each macerate a the species are and small with little minimum of threc slide preparations. or no economic value (Table 2). Independently, As researchers who have had prior experience Landrum (1981) found scalariform perforation mainly with eithcr Franklin's maceration tech­ plates in six species of Myrceugenia (Table I), nique (Baas) or with Jeffrey's (Schmid -' also a genus from temperate and subtropical South Schmid, 1982b), and who at Leiden had the America. opportunity to compare the results (sometimes The initial survey of myrtaceous wood by on the same collection of Myrtaceae) of both Schmid revealed helica1 wall thickenings in techniques, we would like to put in a plca for various species. Landrum (1981) independently much broader use of Franklin's technique. discovered helices in the wood of nine species While Jeffrey's technique can produce cxcel­ of Myrceugenia (Table I). In contrast, helices lent results (references below), Franklin's have been only infrequently noted in the litera­ method comparcd to Jeffrey's is gene rally ture (Table 1). faster, easier to control (Jeffrey's is tricky Although begun in fall 1978 in Berkeley, see Schmid, 1982b), and more consistently this study was completed in spring 1984 at (text continued on page 208)

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SEM photographs of multiple perforation plates in Myrtaceae. - Fig. I & 2. Luma apiculata. - 1: scalariform-reticulate perforation plate, x 970. - 2: scalariform perforation with remnants of primary wall and middle lamella a10ng the margins, x 1940. - Fig. 3 & 4. Ugni candollei, sieve­ like, foraminate perforation plates. - 3: x970. - 4: foraminate plate in vessel membera matching a simple perforation in vessel member b, x 1160.

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SEM photographs of perforation plates and vessel wall thickenings in Myrtaceae. - Fig. 5 & 6. Ugni candollei. - 5: perforation plate foraminate above and simple below, x 1940. - 6: simple per­ foration with vestiges of foraminate end wall pattern, x 1160. - Fig. 7. Tepualia stipularis, simple perforation plate with irregular outline, x 1940. - Fig. 8. Myrceugenia euosma, coarse helical vessel wall thickenings, x 1940. - Fig. 9. Myrcia bracteata, faint irregular helical to wavy wall thickenings, x 580.

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Table 2. Myrtaceae with wood having scalariform perforation plates and/or helical wall thickenings in vessel elements - New data 1

Multiple perforation plates Helical wall thickenings 3

~ ..!l ~ 0.. ..!l " ~ P.." P.. 0.. ""0.." ~ .~ E .§ Vl~,.-., ~ ~ .~ 0..0"'~ ~ ~"'0..00 - " E ! ! I' .0 0 0 "€ 6 " <8 6 § ~ o 6 .~ E ~ b.9 S -2 <8 ~ '"0 ~~ öl ~ .~ ~ '§ ~ "E ~ :;'" t;; ..Q .0 0 '" 00 " ~ öl 6 '- ~ ~ -&~ öl '" e :::l öl " ;:j t ~ 0 "0 .0'- ~ ~ <8 u'" <;::: .~" Speciesl Country of originl Collector I Herbarium '" ~ ~ o 0.. '" '" 0 acmenioides (F. v.Muell.) Burret N. : unvouchered slide (Lw, seetions) (0) + + A. bidwillii (Benth.) Burret N. Australia: unvouchered slide (Lw, seetions) (0) ± + Luma apiculata (DC.) Burret : Mazzucconi 1164 (UC) + 20 (9-28) Argentina: Castellanos 114928 (UC) + 15 (9-20) : Hollermayer 93 (L, UC) (also sections) + 15(11-18) Chile: Schmid 1980·53 (UC)(-) + 18(9-21) Chile: Schmid 1980·35 (UC) + 12 (6-14) Chile: Schmid et al. 1982-134 (UC) (*, macerations, seetions, SEM) + 24 (12-37) L. chequen (Molina) A. Gray California (cult.): Schmid 1978 ·194A (UC) (0) + 15 (12-23) Chile: West 5125 (UC) + 14 (8-20) Chile: Schmid 1980·94 (UC)(-) + 15 (10-19) Chile: Schmid 1980·95 (UC)(*) + 19 (16-22) Chile: Eyerdam 10132 (UC)('Lumagayana') + 14 (8-23) Chile: Hohenacker 448 (L) CLuma gayana ') + 13 (9-19)

Myrceugenia acutif/ora (Kiaerskou) Legrand & Kausei SE. : Glaziou 13894 (UC, isolectotype) + 5 (1-8) M. alpigena (DC.) Landrum var. alpigena SE. Brazil: !rwin 2778 (UC) + 3 (1-6) + var. rofa (Berg) Landrum SE. Brazil: Hatschbach 18158** (UC) + 4(1-11) + SE. Brazil: Hatschbach 3 \065 (UC) + 5 (2-7) + M. bracteosa (DC.) Legrand & Kausei SE. Brazil: Glaziou 13891 (UC) + 5 (1-7) M. campestris (DC.) Legrand & Kausei SE. Brazil: Hatschbach 16423 (UC) + 5 (1-9) M. chrysocarpa (Berg) Kausei Chile: Sparre & Constance \0677 (UC) + 5 (2-13) + ± + Chile: Sparre & Constance \0797 (UC) + 8 (1-12) + ± + M. colchaguensis (Philippi) Navas Chile: Landrum 3454 (NY) + ± + M. correifolia (Hook. & Am.) Berg Chile: Werdermann 900 (UC) + 4 (xxx) M. cucullata Legrand SE. Brazil: Hatschbach 18633 (UC) + 7 (1-12) M. euosma (Berg) Legrand SE. Brazil: Kummrow 355 (UC) + 5(1-10) + + +

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Multiple perforation plates Helical wall thickenings 3

~ ..2 ..2 ..2 ..2 .... 0. 0. "N 0. @ 0. 0." ~ ~ .~ .~ fn~_ ~!! @ ~ ·a ~ " .~ o.c ~ I I - S" ~ @ § § ~ ~ ~ >, .~ ~ " o,2..=.- -.:: 0 ~ ~ ~ c ~~ ca 1:: .~ ~ e ~ ·e"" -300 1:1 " ~ ~ e.Ee "'..::~~ :::l ca'" ca e " ~ ..0'- Species / Country of origin/ Collector / Herbarium ~ ~ ~ ~ ä [~ ä .!::I '" t '" 0 M. euosma (Berg) Legrand x M. glaucescens (Cam bess.) Legrand & KauseI var. glaucescens SE. Brazil: Reitz & Klein 4081 (UC) + 2 (1-4) + ± M. exsucca (OC.) Berg Argentina: de Barba 376 (UC) + Chile: Sparre & Constance 10859 (UC) + Chile: Hastings 474 (UC) + M. [ernandeziana (Hook. & Am.) Johow ('Nothomyrcia [ernandeziana') Chile (insular): Pisano & Montaldo 1421 (UC) Chile (insular): Meyer 9527 (UC) Chile (insular): Meyer 9509 (USw) (*) M. [ranciscensis (Berg) Landrum SE. Brazil: Hatschbach 18950 (UC) + M. glaucescens (Cambess.) Legrand & KauseI var. glaucescens Uruguay: Herter 1610a (UC) + 5 (4-7) ± ± + var. latior (Burret) Landrum SE. Brazil: Reitz & Klein 13528 (UC) + ± M. kleinii Legrand & KauseI SE. Brazil: Klein 1540 (L, UC, isotype) + 9 (xxx) M. lanceolata (Juss. ex J. SI. HiI.) KauseI Chile: Behn s.n. (UC) + 4 (xxx) xxx Chile: Zöllner 3721 (L) (sections) + 4 (2-8) + M. leptospermoides (OC.) KauseI Chile: Aravena 018 (UC) + 6 (1-1\) M. miersiana (Gardner) Legrand & KauseI SE. Brazil: Reitz & Klein 8760** (UC) + 5 (xxx) ± + SE. Brazil: Mexia 4257 (UC) + 5 (3-9) M. myrcioides (Cambess.) Berg var. acrophylla (Berg) Landrum SE. Brazil: Reitz & Klein 10673 (UC) + 7(1-11) ± + var. myrcioides SE. Brazil: Hatschbach 16242 (UC) + 6 (1-9) M. myrtoides Berg SE. Brazil: Rambo 26094 (U) ± + M. obtusa (Oe.) Berg Chile: Morrison 16731 (UC) + 3 (3-4) M. ovata (Hook. & Am.) Berg var. acutata (Legrand) Landrum SE. Brazil: Reitz & Klein 7807 (UC) + 5 (1-10) + var. gracilis (Burret) Landrum SE. Brazil: Hatschbach 18568 (UC) + 4 (1-9) + var. nannophylla (Burret) Landrum Chile: Landrum 3284 (NY) + 6 (4-10) + var. ovata Chile: West 4884 (UC) + 8 (6-20) ± + + Chile: Schmid 1980-38 (UC) + 10 (4-20) ± + +

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(Table 2 continued) Multiple perforation plates Helical wall thickenings 3

,-, ..!l .... 'i5.." 0.. ..!l "N -ä E 0.. 0.." ~ .~ .~ 0 .~ ~]~ ~!l "'o..bI) .~ 0..,:: E ~ I 1:: I .D ,:: ,:: § .!!l Ei " ~ E E ~ '- 0 '" >, o ~ :;; <8 '"~ ,::'" ~ .~~ t!::'d) "öl ·e ~ ·E " .... bI) :;; .0 .D 0 '" ~ ~bI) t=~ ~ "öl'" "öl i ~ Ei '- .... ~- ~ '" 0 " :::: .D '- Species/ Country of origin/ Collector /Herbariurn ~ ~ ~ E <8 S 8. ~ '" Jj '" " .= '" 0 M oxysepa/a (Burret) Legrand & Kausei SE. Brazil: Reitz & Klein 8673 (UC) + 5 (1-10) ± ± + M. parvifolia (DC.) Kausei Chile: Sparre & Constance 10839 (UC) + 10 (1-15) ± + M pilatantha (Kiaerskou) Landrurn var. major (Legrand) Landrurn SE. Brazil: Reitz & Klein 8093 (UC) (also SEM) + 7 (1-12) ± + var. pilatantha SE. Brazil: Reitz & Klein 10992 (UC) + 3 ± ± + M pinifalia (Philippi f.) Kausei Chile: Landrurn 3086 (NY) + I (1-2) ± ± + Chile: Landrurn 3381 (NY) + 6 (4-7) ± ± + M p/anipes (Hook. & Am.) Berg Chile: West 4861 (UC) + 7(1-18) ± + Chile: Schrnid 1980-39 (UC) + 8 (1-22) ± + Chile: Schmid 1980 -40 (UC) + 6 (1-10) ± + M. reitzii Legrand & Kausei SE. Brazil: Cabrera et al. 9973 (L, UC) + 4 (2-5) M. rufa (Colla) Skottsb. ex Kausei Chile: Bertero 1170 (UC) M. rufescens (DC.) Legrand & Kausei SE. Brazil: Hatschbach 18309 (UC) ± ± + M. schultzei Johow Chile (insular): Solbrig et al. 3711 (UC) + 5 (I -8) ± ± + M seriatoramosa (Kiaerskou) Legrand & Kausei SE. Brazil: Hatschbach 1393 (MICH) + 7 (1-13) ± ± + M. smithii Landrurn SE. Brazil: Srnith & Reitz 12411 (UC, isotype) + 1 (xxx) ± ± + M venasa Legrand SE. Brazil: Reitz 5474 (L, iso type) + 7 (2-10) ± +

Myrcia bracteata (L. Richard) DC. Surinarn: Lanjouw & Lindernan 2155 (U, UW) ( *, seetions and SEM) + + M. cf paivae Berg Surinarn: Lanjouw & Lindernan 2712 (Uw) ( *)(seetions ) + + M servata McVaugh British Guyana: Fanshaw, F.D. 4491 (Uw) (*) (sections) + + M sylvatica (G. F. Meyer) DC. Surinam: Maguire 24637 (Uw) (0) (sections) + + Myrcianthus mata (Griseb.) McVaugh Argentin.: Meyer et.1. 23130 (Ue) (also SEM) + Myrtastrum rufa-punctatum (Pancher ex Brongn. & Gris) Burret : Balansa 12°-1484 (NY) + 4(1-5)xxx New Caledonia: Baurnann 8179 (NY) + 9 (6-18)

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Multiple perforation plates Helical wall thickenings 3

,-. ~ ~ ~ ~ <>", -ä <> ~ "" E ""<> "" .~ .~ .~"" ~ ]~ ~!l Ei ..2 "'''''Oll -2 ""<= ~ I ! '5 I .0 <= <= 1;; ..2 § ~ >, Ei " § § öl '- 0 '" ;:: ..2 ~ '" ~ .~~ ~~ ~ ~ ~ '§ <> ~ Oll a .Q .0 0 '" -3Oll c=] ~ öl ~ r:: Ei '- ~ ~~ :::I ~ <= ~ .0'- Species/ Country of origin/ Collector /Herbarium ~ ~ ~ ~ ..2 g & ~ <:::" .!:l '" '" 0 New Caledonia: Franc 40 (NY) + 8 (6-12) New Caledonia: Franc 209 (NY) + 6 (3-9) Myrte% acerosa (Berg) Burret Peru: Ferreyra & Ac1eto 15299 (UC) + 11 (7-15) Peru: Hutchison & Wright 5516 (UC) + 15 (10-18) M. leucomyrtillus (Griseb.) Reiche Chile: Kausei 2408 (STG) + 7 (5-10) Chile: Gunekel 9368 (G) + 7 (3-10) M. microphylla (Humb. & BonpI.) Berg var. glabrata Berg Peru: Steinbach 592 (U) + 11(9-14) : Herzog 2139 (L) (seetions) + 11 (8-13) Bolivia: Herzog 2384 (L) (seetions) + 12 (8-15) Bolivia: Williams 1573 (UC) + 15 (12-10) M. nummularia (Poiret) Berg var. barneoudii (Berg) Kausei Chile: Mexia 8012 (UC) + 10 (2-15) Chile: Schmid 1980-43 (UC) + 9 (4-10) Chile: Zöllner 4870 (L) (sections) + 12 (18-16) Chile: Andreas 399 (U) (seetions) + 10 (4-20) var. nummularia Chile: West 4906 (UC) + 7 (4-9) Argentina: Goodall1781 (UC) + 7 (5-13) M. oxycoccoides (Benth.) Berg Colombia: Fosberg 22366 (UC) + 12 (7-13) Myrtus communis L. California (cult.): Schmid 1978-192 (UC) (.) (also seetions) ± + + Greece: Mattfeld 2149 (UC) ± + California (cult., ex Greece): Schmid 1978-193 (UC) ( .) (also sections) ± + + Portugal: Barcuda 580 (.)(sections) + ± ± Greece: Thessaloniki, Lab. For.Util. 55 (Uw) (.) (seetions) + ± ± Netherlands (cult.): unvouchered slide (Lw) (.) (seetions) + ± ± Neomyrtus pedunculata (Hook. f.) Allan New Zealand: Walker 4438 (UC) + 18 (11-27) Nothomyrcia - see Myrceugenia fernandeziana Psidium longipes (Berg) McVaugh ('Myrtus verrucosa') var. longipes Florida: Stern & Brizicky 205 (Uw) ( .)(seetions ) ± + Tepualia stipu/oris (Hook. & Am.) Griseb. Chile: Morrison 17648 (UC) + 6-8 (xxx) Chile: Schmid 1980-78 (UC) (.) + 7 (xxx) Chile: Schmid et al. 1982-127 (UC) (*, macerations, seetions, SEM) + 5 (2-13)

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(Table 2 continued) Multiple perforation plates Helical wall thickenings 3

..!l ~ '""' c.. -ä -ä E -ä "'''c..", .~ .~ ~ .~ .~'" -~~ c..c E ! I ~ I 5-!i 1;; § ::: 5 ; Ei '" '" '5 .c '§ .0 0 '" eo =~ ~ 'öl 'öl ~ ~ Ei ""' ~ :::I '" '" .0'-~- Species/ Country of origin / Collector / Herbarium ~ ~ ~ ~

1 Only macerations were examined unless noted to the contrary. For ecological and geographical information on taxa see Discussion. Symbols: xxx = insufficient information to express character states from sampie ormethod used; * = mature wood sam pie rather than a twig sampie from an herbarium specimen (NB: Myrteola is a shrublet with mature wood usual­ ly represented on herbarium specimens); ** = this collection was also examined anatomically by Landrum (1981; and Table I). Observational criteria (only positive character states tabulated; for Myrceugenia two species without scalariform perforation plates or helices are included in this table; other such taxa are listed in Table 3): type of perforation plate (a) scalariform (simple) = perforation plates predominantly scalariform (65% or over), with 34% or fewer simple; (b) scalariform-simple = 31-64% of perforation plates scalariform or simple; (c) (scalariform) simple = perforation plates predominantly simple (65% or over), with 5-30% scalariform; (d) rarely scalariform = 4% or fewer of perforation plates scalariform; (e) foraminate-simple =any mixture of foraminate and simple per­ foration plates.

2 Number of bars per scalariform perforation plate usually based on ten counts, oceasionally 20 (e.g, some sampies of Myrteola) or 50 (some sampies of Luma).

3 Expression of helical wall thiekenings on vessel elements: + =eonspicuous and eommon; ± = faint and/or of oeea­ sional occurrence.

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Table 3. Myrtaceae with wood lacking both scalariform perforation plates and helical wall thickenings in vessel elements - New data 1

Amomyrtella güili (= Eugenia güili, Pseudocaryophyllus güili), Argentina. luma, Chile (5 sampies, 2*). - A. meli, Chile (2 sampies). Arillastrum gummiferum (= Spermolepis gummifera), New Caledonia (2 sampies). fascicularis, California (cult., ex Australia). (= Myrtus dulcis), Australia. Baeckea sp., Australia. - B. ericoides, New Caledonia (2 sam pies). - B. virgata, New Caledonia, California (cult., ex Australia) (2 sampies). sparsa, California (cult., ex Australia). cruckshanksii (= Temu cruckshanksii, T. divaricatnm), Chile (3 sampies, 1*). gnidioides, New Caledonia (*). validus, California (cult., ex Australia). Calythrix flavescens, Australia. - C. fraseri, Australia. - C. tetragona, California (cult., ex Australia) (2 sam pies, 1 *). squarrosa, Australia. Decaspermum alpinum, New Guinea. - D. forbesii, New Guinea. - D. nivale, New Guinea. - D. novoguineensis, New Guinea. - D. prostratum, New Guinea (holotype). Eucalyptus globulus, Australia. Eugenia beaurepaireana (= Pilothecium ternatifolium), Brazil (2 sampies). - E. carimbatayensis, Paraguay. - E. cupulata var. macrophylla, Brazil. - E. glazioviana (= Pilothecium glaziovianum), Brazil. - E. hassleriana, Paraguay. - E. herbacea, Brazil. - E. oerstedeana, British Honduras. - E. pyriformis (= Pseudomyrcianthes pyriformis), Brazil. - E. uniflora (= Stenocalyx michelii), Bermuda, Costa Rica (2 sam pies, 1*). - E. winzerlingii, British Honduras. Heteropyxis dehniae, southern Africa. edulis (= Myrcianthes edulis, Eugenia edulis), Paraguay. - H. handroi, Brazil. - H. humilis (= Eugenia anomala), Brazil. Kania eugenioides (= Metrosideros eugenioides), New Guinea (cotype). Kjellbergiodendron celebicum, New Guinea, Celebes (2 sam pies). Legrandia concinna, Chile (2 sampies). Leptospermum scoparium var. scoparium, Australia. Lindsayomyrtus brachyandrus, Australia. bullata, New Zealand. - t. bullata x L. obcordata (= Myrtus ralphii), New Zealand. - L. obcordata, New Zealand, California (cult., ex New Zealand) (2 sampies). (= Tristania conferta), California (cult., ex Australia). gnidioides, New Caledonia. - M. quinquenervia, New Caledonia. Meteoromyrtus wynaadensis (= Eugenia wynaadensis), India (holotype). Metrosideros angustifolia, southern Africa (2 sampies). - M. operculata, New Caledonia. Myrceugenia fernandeziana (see Table 2). - M. rufa (see Table 2). var. fragrans, Mexico. - M. pungens (= Acreugenia pungens, Eugenia pungens), Argentina. delicatula (= Eugenia delicatula, Paramyrciaria delicatula var. linearifolia), Brazil. - M. floribunda, British Honduras. Nothomyrcia fernandeziana (see Table 2). Pileanthus filifolius, Australia (2 sam pies). - P. peduncularis, Australia. Psidium pubifolium (= Myrtus ovalis Berg), Uruguay. Psiloxylon mauritianum, (3 sampies, = Psiloxyloideae sensu Schmid, 1980, Psilo- xylaceae sensu Van Vliet & Baas, 1984). Purpureostemon ciliatum, New Caledonia. Reicheia coquimbensis (= Aspidogenia coquimbensis), Chile (2 sampies). Siphoneugenia baporeti (= Myrciaria baporeti, Myrciariopsis baporeti), Argentina. Stereocaryum rubiginosum, New Caledonia. Syzygium adelphicum, New Guinea. - S. alatum (= Aphanomyrtus alata), New Guinea (2 sampies). - S. buxifolium, Japan. - S. coolminianum, California (cult., ex Australia) (2 sampies, 1*). - S. gui­ neense var. macrocarpon, Congo, Dahomey (2 sampies). - S.jambos, Guatemala (cult., ex Indo­ Malaysia). - S. kinabaluensis, Borneo. - S. myrtillus, Borneo. - S. steenisü, Borneo (isotype).

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Temu (see B1epharoca1yx). Tristaniopsis callobuxus (= Tristania callobuxus), New Ca1edonia (2 sampies). - T. 1aurina (= Trista­ nia 1aurina), California (cu1t., ex Austra1ia) (*). archbo1diana, New Guinea. - U. brassii, New Guinea. - U. emarginata, New Caledonia (3 sampies). - U. ngoyensis, New Caledonia. - U. sarawakensis, Sarawak. - U. thymifolia, New Caledonia. monadelpha, Australia (2 sampies). - V. nitens, Australia (2 sampies). Whiteodendron mou1tonianum, Borneo. Xanthomyrtus arfakensis, New Guinea. Xanthostemon francii, New Caledonia (*). - X. rubrum, New Caledonia. - X. sulfureum, New Cale­ donia.

I Voucher information and authorities for scientific names (89 species in 45 genera) are avai1able on request to Schmid. Most vouchers are at Ue. Se1ect synonymy is given for a few species. Countries noted refer to sources of sampies studied. One sampie per species was examined un1ess otherwise noted, and * indicates that mature wood rather than twig wood was samp1ed. A few data are duplicated by previous literature (see References). Except for the species noted in Tab1e 2, the approximate1y 118 species and 43 genera of Myr­ taceae [plus Eucalyptus, which we examined only superficially (for details see Dadswell, 1972, and Ing1e & Dadswell, 1953] , which are represented on slides in the collections at Leiden, Oxford, Richmond (California), Utrecht, and Washington (D.C.), also have wood lacking both scalariform perforation plates and helical wall thickenings in the vessel elements. Most of these species were previously examined by Ingle & Dadswell (1953) and other workers (see References).

produces high quality preparations with a mini­ latter. For example, various taxa, inc1uding mum of undamaged elements. The last factor is Myrtaceae (Tristania - Bierhorst & Zamora, especially important in studying de1icate tissues 1965), have scalariform perforation plates in (primary xylem of sterns and ) as the the primary but not in the seeondary xylem. more potent chemica1s of Jeffrey's fluid can be We attempted methodologieally (see Sehmid, overly (or totally!) destructive. Yet surprising1y, 1982b) to avoid sueh 'contamination' of see­ studies of primary xylem of sterns (Bierhorst & ondary by primary xylem, and in all eases we Zamora, 19155; Muhammad, 1984; Muhammad ignored vessel elements with features of prima­ & Sattler, 1982) have used mainly if not exclu­ ry xylem so as not to eharaeterise falsely the sively Jeffrey's technique. In contrast, our seeondary xylem. macerates of myrtaceous flowers prepared by Terminology for deseriptors involving fre­ Franklin's method are vastly superior to ones queney (e.g., 'often', ete.) folio ws Sehmid prepared by Jeffrey's. (1982a). Other deseriptive criteria are indieated In addition, we examined some sampIes of in note I to Table 2. myrtaceous wood with the scanning electron microscope or, after hand or microtome sec­ Results tioning, w!th the light microseope. We also sur­ Tables 2 and 3 summarise the oeeurrenee of veyed all the available microscope slides of sealariform perforation plates and helical wall sectioned mature woods of Myrtaceae in the thickenings in the wood of our sampie of 144 collections at Leiden, Oxford, Richmond (Cali­ speeies (plus 12 varieties and 2 hybrids) and 53 fornia), Utrecht, and Washington (D.C.). genera of Myrtaceae. For Myrceugenia we sam­ For the macerations mainly wood from pled 33 of the 38 species and one of the three sm all-diameter sterns, especially of herbarium hybrids reeognised by Landrum (1981). specimens, was available. This approach has The morphology of the sealariform perfora­ two main disadvantages: (I) Such juvenile and tion plates in Myrtaecae varies from quite regu­ twig wood is not typica1 of adult wood from lar to very irregular with many forked bars, in the main trunk, particularly in size criteria both cases with low to high numbers of bars (Jane, 1970; Panshin & Oe Zeeuw, 1980). per perforation plate. Excessive branehing Hence we do not report cell dimensions, al­ (' forking') of bars may result in a sealariform­ though we did make Iimited length measure­ reticulate (Fig. 1) or even a honeycomb-likc ments of vessel elements. (2) Intermixing of pri­ retieulate type ofperforation plate that is c1early mary and secondary xylem is common in mace­ derived from a scalariform type. In some spe­ rations of twig wood, such that characterisa­ eies, Ugni selkirkii (only Morrison 17338) being tion of the former could falsely represent the the most notable example, many of the pcrfo-

Downloaded from Brill.com10/10/2021 12:05:33PM via free access IAWA Bulletin n.s., Vol. 5 (3),1984 209 ration plates are scalariform-reticulate or highly in a number of other taxa with either scalari­ reticulate and ordinary scalariform perforation form or simple perforation plates. plates are rather uncommon. In genera with Apart from scalariform and scalariform-reti­ both scalariform and simple perforation plates culate perforation plates, Ugni candollei has (Myrceugenia, Myrtastrum, Tepualia) there of­ sieve-Iike foraminate perforation plates (Figs. 3 ten occur irregularities such as incomplete bars, & 4). Here the individual perforations resemble or bars confined to one side of the perforation large pits without pit membranes. Both forami­ area. Generally all patterns of variation as de­ nate and simple perforation plates occur in the scribed for several dicotyledons by Muhammad same . Intermediates between smooth­ and Sattler (1982), Muhammad (1984), and rimmed simple perforations and complete fora­ others were represented in our sam pIe of Myr­ minate ones are shown in Figures 5 & 6. In one taceae. Table 2 does not give such deviations specimen of U. candollei (Table 2) foraminate from regular scalariform perforation plates. perforations were found to intergrade with sca­ In our entire sam pIe of Myrtaceae (Table 2) lariform-reticulate plates. Moreover, in Ugni the number of bars per scalariform perforation selkirkii which has 100% multiple perforations, plate ranges from one bar (Myrtastrum rufo­ some of the scalariform-reticulate plates inter­ punctatum and 16 species and I hybrid of graded with reticulate plates very c10sely re­ Myrceugenia) to 37 (Luma apiculata). This sembling those of U.candollei. These transi­ contrasts with the value of '15-25 fine bars' tions suggest that in Ugni scalariform and fo­ given by Metcalfe and Chalk (1950), the only raminate perforation plates are related (cf. specific figure for this feature noted in the lit­ Muhammad & Sattler, 1982, who advocate a erature. Generic summaries can be extracted relationship between scalariform and circular from Table 2 as folIows: perforations in angiosperms and Gnetum). Some authors would prefer the term 'reticulate' Genus Number bars: for the type ofperforations iIIustrated in Figures (and number of species) average (range) 3 & 4 (e.g., Gray & De Zeeuw, 1974). It has been reported for a number of unrelated taxa, Luma (2) 16.2 (6-37) e.g. Rosaceae (Metcalfe & Chalk, 1983) and Ru­ Myrceugenia (26) 5.4 (1-22) biaceae (see below), both of wh ich are chiefly Myrtastrum (I) 6.8 (1-18) characterised by simple perforation plates. As Myrteola (5) 10.5 (2-20) with Coprosma of the Rubiaceae (ButterfieId & Neomyrtus (I) 18.0 (11-27) Meylan, 1980; Metcalfe & Chalk, 1983; Meylan Tepualia (I) 6.3 (2-13) & Butterfield, 1975, 1978a), in Ugni the fora­ Ugni (4) 12.6 (2-26) minate perforation plate may be contiguous with a simple perforation plate (Fig. 4). Usually the bars are quite thin (less than I The simple perforation plates characterising J.Lm) but in several sampies thick bars occur (up almost all Myrtaceae have smooth or, as docu­ to 2.5 J.Lm in Myrteola nummularia var. num­ mented by scanning electron microscopy, ves­ mularia; up to 2.1 J.Lm in Ugni myricoides var. tured (Baas, 1977; Kucera et al., 1977), border­ oerstedii). Thick and thin bars may occur even ed or non-bordered, circular to oval perforation within a single vessel element, either in the rims (Meylan & Butterfield, 1978a). In many same perforation plate or at opposite ends of species with admixed simple and scalariform the element. plates, some simple ones have irregular outlines The rims of scalariform perforation plates of (Fig. 7) that suggest intergradations between Myrtaceae may be both vestured and have mi­ simple and scalariform perforation plates (see crofibrillar webs. Both features have been shown Muhammad, 1984, for similar transitions in for Neomyrtus pedunculata (Bu tterfield & Mey­ Comptonia of the ). lan, 1974, 1980; Meylan & Butterfield, 1978a), Helical wall thickenings in the vessel ele­ the only previous member of Myrtaceae with ments of secondary xylem of Myrtaceae can be scalariform perforation plates to be examined arbitrarily c1assified as (I) conspicuous and in detail with the scanning electron microscope. coarse, the thickenings here always widely The few species we studied with the scanning spaced (Fig. 8), (2) conspicuous and fine, with electron microscope had unvestured perfora­ closely spaced helices typically thinner than in tion rims (Figs. I & 2). Structures reminiscent 'I', and (3) faint, with irregular helical to wavy of microfibrillar webs but much coarser in tex­ sculpturing (Fig. 9, which is quite inconspicu­ ture (obviously remains of incompletely hydro­ ous in the light microscope). These categories Iysed primary wall and middle lamella) were include intergradations from weak expression observed with the scanning electron microscope to total absence, i.e., to smooth walls. Table 2 in Luma (Fig. 2) and with the light microscope summarises the various character states, but it

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should be realised that, depending on the quali­ calfe and Chalk's (1950) record of scalariform ty of the macerates and the intensity of their perforation plates for Myrtus communis is cer­ staining, identical types of helical wall thicken­ tainly erroneous. It is contrary to the observa­ ing may appear conspicuous or rather faint in tions of Baas et al. (1983). Ingle and Dadswell the light microscope. Therefore no great impor­ (1953), Sanio (1863), Schmid (1980), Solereder tance should be attached to subtle differences (1885, 1899, 1908, by implication), Van Vliet in the table. In addition, our descriptors for and Baas (1984), the present study (Table 2), helices are less elaborate than those of Meylan and several detailed accounts of the species and Butterfieid (l978b), which are strictly (Fahn et al., in press; Greguss, 1959; Huber & based on SEM observations. Rouschal, 1954). (4) We can also confirm In quite a number of sampies vessel elements Landrum's (1981) finding of scalariform per­ with helices occur with ones lacking helices. In foration plates in six species of Myrceugenia some species helices occur only sporadically (Tables I, 2). and are then often restricted to the narrow tails The present study thus ex tends the known of the vessel elements, which in many species occurrence of scalariform perforation plates can be very long. In some species with scalari­ in wood of Myrtaceae from eight species in form perforation plates and coarse helices, three genera to 40 species in seven genera, thickenings ne ar the former may be continuous namely, Luma, Myrceugenia, Myrtastrum, with the bars. Finally, individual sampies may Myrteola, Neomyrtus, Tepualia and Ugni exhibit the gamut from helices absent to obscure (Table 2). Except for Myrceugenia, which has or faintly expressed to conspicuous. Table 2 38 species (Landrum, 1981), the other genera summarises some of the possibilities. are either monotypic (Myrtastrum, Neomyrtus, In passing, it must be warned that uncritical Tepualia) or very small. The low percentage observations can give the impression of helical (below 5 %) of scalariform perforation pI at es in wall thickenings or even of scalariform perfora­ the Chilean Tepualia stipularis was simply over­ tion plates. Vessel to parenchyma pit-pairs in looked by Record and Hess (1943) and Wage­ Myrtaceae are commonly large and scalariform mann Wiedenbrug (1949). (lngle & Dadswell, 1953). Such pitting can re­ Similarly, prior to 1980 five species of Myr­ semble helical wall thickenings. In addition, taceae were reported to have wood with helical such pitting or helical wall thickenings on the wall thickenings on the vessel elements (Table wall of avessei element opposite a simple per­ 1 - we ignore here literature reports of genera foration plate can give the impression of a sca­ with unspecified species because such are near­ lariform perforation plate, especially as many Iy impossible to verify): (I) We can confirm species of Myrtaceae have simple perforation the reports of such for Myrceugenia exsucca, plates that are rather oblique or that occur at M. schultzei, and Myrtus communis. (2) Moll appreciable distances from the tips of the ele­ and Janssonius's (1918) report for Acmena ments. acuminatissima is highly reliable and needs no further comment. (3) Record and Hess's (1943) Discussion record of helical wall thickenings for Myrceu­ genia fernandeziana is not confirmed by our Systematic occurrence of scalariform perfora­ data (Table 2) or by Landrum's (1981). (4)We tion plates and helical wall thickenings in Myr­ can also confirm Landrum's (1981) finding of taceae helical wall thickenings in eigh t species of Myrc­ Prior to 1980 of Myrtaceae were eugenia (Table I) bu t differ from hirn in not see­ reported to have wood with scalariform perfo­ ing such in M. acutij70ra (Table 2). As this situa­ ration plates (Table I): (I) We can confirm the tion may be similar to our results with two sam­ reports of such for Luma apiculata and Neo­ pIes of M. miersiana (Table 2), it seems reason­ myrtus pedunculata (Table 2). (2) Fasolo's rec­ able to accept Landrum's positive record for ord (1939-40) of such for Syzygium guineense M. acutij7ora. is based on misidentified material, which is not The present study thus extends the known myrtaceous (compare his description with those occurrence of helical wall thickenings in vessel in Metcalfe & Chalk, 1950, and Van Vliet & elements of wood of Myrtaceae from 13 species Baas, 1984). Our material of this species (Table in three genera to 33 species in eight genera, viz. 3) has simple perforation plates. Perhaps signi­ Acmena, Austromyrtus, Myrceugenia, Myrcia, ficantly all xylotomatically known species of Myrcianthes. Myrtus. Psidium. and Xanthomyr­ Eugenia and Syzygium (by modern concepts - tus. Except for Myrtus, which is currently re­ Briggs & Johnson, 1979; McVaugh, 1968; garded as having two species (M. communis and Schmid, 1972) have strictly simple perforation the North African M. nivellii), the remaining plates (Tables 1-3; other literature). (3) Met- genera are of moderate to very large size.

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Summarising the above information for sca­ and countries of origin] : Luma (with 2 species, lariform perforation plates and helical wall L. gayana inc1uded in L. chequen following thickenings in vessel elements, there are in our Landrum, pers. comm., 1980) occurs in Chile sam pie ofMyrtaceae (I) 24 species and 7 genera and adjacent Patagonian Argentina from about with only scalariform perforation plates, (2) 17 30° latitude south in mesic or fairly mesic sites; species and 8 genera with only helical wall Myrteola (ca. 7 species) has (I) two species in thickenings (inc1uding the re cord for Acmene Chile and adjacent Argentina (plus the Malvinas acuminatissima by Moll & Janssonius) and (3) Islands) from 36° latitude south growing at 17 species, all in Myrceugenia, with both fea­ low elevations in mesic forests or in heaths or tures. Myrceugenia is the only genus with all bogs and (2) five tropical species in Venezuela, three possibilities. Colombia, Ecuador, Peru, and Bolivia restricted to mesic high montane sites at elevations from Ecological and functional considerations 2200 to 4000 m (Landrum, pers. comm., is re­ The large family Myrtaceae covers a very vising Myrteola and believes there are only wide and diverse ecological range. Most of the three specie's, M. nummularia with distribution approximately 3700 species and 150 genera land 2, and M. acerosa and M. microphylla (Briggs & Johnson, 1979; Schmid, 1980) occur with distribution 2); Neomyrtus (I species) is in the tropics. However, the family is also quite endemie to New Zealand where it occurs in weil represented in the subtropics, though rath­ coastal to lower montane forest and shrubland er poorly so in temperate zones (temperate from 35° latitude south; Ugni (4 species) has Australia, New Zealand, , and temperate (I) three species in Chile (U selkirkii endemic North and ). In the tropics most to the Juan Fernandez Islands) growing at low species are mesic rainforest elements, but in the elevations between 33° and 45° south latitude su btropics, especially in Australia, Myrtaceae in forest habitats and (2) one polymorphic spe­ range from mesic to quite xeric in their habitat cies, U myricoides, which occurs between preference. Some of the tropical species occur 1500 and 3700 m from Mexico to Peru and the only in cool high montane sites from approxi­ Guyana Highlands. Typically the varieties of mately 1500 to 4000 m elevation. this species are from moist vegetation types, At the onset of this discussion of ecological but two varieties (v ar. longipes and var. oer­ and functional aspects of the occurrence of sca­ stedii in Central America) mayaiso occur on lariform perforation plates and helical wall dry mountain hillsides. The Chilean U can­ thickenings in vessel elements of Myrtaceae it dollei, the only species with predominantly should be stressed that the overwhelming ma­ simple perforation plates and a varying propor­ jority of Myrtaceae with simple perforation tion of foraminate and scalariform perforation plates and vessel elements lacking helical wall plates, appears to grow in cooler and wetter thickenings covers the complete ecological places than U molinae (Landrum, pers. comm.), range of the family. This suggests that these which has exclusively scalariform perforation features are not detrimental (or in a more posi­ plates. tive vein: are even adaptive) for the vital func­ The other genera of Myrtaceae with scalari­ tions of xylem sap conduction in all their di­ form perforation plates also have a varying, verse environments. usually large proportion of simple perforation The restricted num ber of genera of Myrta­ plates. The ecology of these genera is more di­ ceae with scalariform perforation plates and/or verse than that of the genera with exc1usively helical wall thickenings cover only a rather (or mainly) scalariform perforation plates and small part of the ecological range. These genera is summarised below. Tepualia stipularis, which are typically of temperate to subtropical distri­ is monotypic and the only New World represen­ bution, and in the latter case mainly from cool tative of Leptospermoideae, occurs in Chile montane sites. The ecology of genera with and adjacent Patagonia south of 35° latitude 100 % (or nearly 100 %) scalariform perforation in mesic to semi-mesic forests, wh ich often plates can be characterised as 'cool mesic', with contain species of Luma, Myrceugenia, and details as follows [information from Landrum Ugni. Tepualia has a very low percentage of (1981) and McVaugh (1968) and especially the scalariform perforation plates in its wood (see taxonomic references therein and in Briggs and Table 2). Myrtastrum rufo-punctatum is mono­ Johnson (1979) and Schmid (1972, 1980), typic and endemic to New Caledonia (ca. 21 ° with ecological data gleaned especially from south latitude), where it occurs in mesic up­ Landrum (1981) and extended by field obser­ land forests but also in rather dry lowland for­ vations by Schmid in New Caledonia in July ests on serpentine soils. This species also has a and August 1978 and in Chile in Dec. 1980 low percentage of scalariform perforation plates and March 1982 - see also Table 2 for species in its wood (Table 2).

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Myrceugenia, a genus of 38 species, was ex­ In Myrceugenia the considerable variation of plored most comprehensively for ineidence of helical wall thickenings (Table 2) is completely scalariform perforation plates in relation to unrelated to the ecology of the species. Among ecology of its taxa becal'se the genus was re­ species with conspicuous helical thickenings cently revised by Landrum (1981), who also there are highly mesic as weil as xeric ones, and gives fairly detailed ecological and distribution­ cold tolerant Chilean and high montane Brazil­ al information. It appears that most of the taxa ian species as weil as species from warm sub­ of Myrceugenia with 31 % or more of their per­ tropical lowland sites of southeastern Brazil. foration plates scalariform (Table 2) have a The group of species of Myrceugenia lacking mesic to highly mesic ecology. These species any apparent helical wall thickenings in their (Table 2) thusmight be c1assified as 'cool mesic' wood covers the same wide range of habitats. in their ecology, although M. euosma, with an The only generalisations to be made regard­ estimated 33% scalariform perforation plates, ing ecological preference in relation to expres­ is an exception, being 'one of the most xero­ sion of helical wall thickenings are (I) that all phytic species of Myrceugenia' (Landrum, records for tropical taxa concern sporadic ob­ 1981: 86). The species with exclusively simple servations of very weak helices (or in the case perforation plates (Table 2) are more diverse of Myrcia and probably also of Acmena - Moll in their ecology, there are three taxa from & J anssonius, 1918 - of weak, irregular stria­ western South America and five taxa from east­ tions), and (2) that among the subtropical to ern South America, mostly from elevations be­ temperate taxa there are several species with low 700 m. Unfortunately, the precise ecology well-developed fine or coarse helices occurring in of most of these species is not very weil known. most or all of the vessel elements (Myrceugenia However, this group contains some very mesic p.p., Myrcianthes mato, Myrtus communis). species, for example, M. exsucca, M. fernandez­ The occurrence of scalariform perforation iana, and M. rufescens. Lastly, most species of plates and helical wall thickenings in vessel ele­ Myrceugenia have a very low percentage of ves­ ments is largely in agreement with general eco­ sei elements with scalariform perforation plates logical trends established for the dicotyledons (30% or less - Table 2). Almost all of these as a whole. Carlquist (1975) and Baas (1976) species have a mesic ecology, but M. obtusa is demonstrated that scalariformperforation plates an exception. This category also includes spe­ are much more common in cool mesic regions eies from the complete altitudinal and latitudi­ than in tropical lowland vegetation or in xeric nal ranges of the genus. floras. In Myrtaceae this trend is also apparen t. The conclusion for Myrceugenia thus is that Baas (1973), Baas et al. (1983), and Van der the type of perforation plate is only weakly Graaff and Baas (1974) found helical wall thick­ related to the ecological ranges of the species: enings to be a dominant feature of subtropical the most primitive condition (a high percentage to temperate, mesic to moderately xeric floras, of scalariform perforation plates) has alm ost with tropical trees or shrubs only rarely show­ only been retained in cool mesic sites, but the ing helices. This trend is hardly retraceable in more derived condition of a low frequency Myrtaceae except for the fact that the subtrop­ of scalariform perforation plates or even their ical to temperate Myrtaceae with helices show total absence occurs in all types of habitats. In a more conspicuous expression of this feature the mesic sites taxa with only simple perfora­ than do the tropical taxa. tion plates are sympatric with taxa with a high In passing, it may be mentioned that weak percentage of scalariform perforation plates. striations, such as occur in some Neotropical The occurrence of helical wall thickenings species of Myrcia, have probably been over­ on vessel elements of Myrtaceae (Table 2) seems looked in most c1assical wood anatomical hardly related to ecological preference. Of our studies (Moll & J anssonius, 1918, are a notable new records combined with those in the litera­ exception - see Table 1). In the light micro­ ture (Table I), four are for taxa of the tropical scope these striations are so weak that confir­ lowland: Acmena p.p. (Java), Austromyrtus rnation by scanning electron microscopy is p.p. (Queensland), Myrcia p.p. (Surinam and needed. This may somewhat have affected the Guyana), and Psidium p.p. (Florida); one rec­ statistics of incidence of helicalor irregular wall ord is for the tropical high montane flora: thickenings in vessel elements as cited from the Xanthomyrtus humilis (New Guinea, at 3000 m literature (Baas, 1973; Baas et al. , 1983; Van elevation); and the remaining records are for der Graaff & Baas, 1974). subtropical to temperate taxa: Myrceugenia A functionally adaptive interpretation of the p.p. (South America - see below), Myrcianthes diversity of patterns in type of vessel perfora­ mato (northwestern Argentina), and Myrtus tion and wall sculpturing in Myrtaceae, like the communis (Mediterranean, in rather xeric sites). general discussions of functional wood anatomy

Downloaded from Brill.com10/10/2021 12:05:33PM via free access IAWA Bulletin n.s., Vol. 5 (3),1984 213 on a comparative basis, can only remain entirely foration plates in Myrtaceae (Tables 2, 3) indi­ speculative. On the assumption that the Myrta­ cates that this feature is the ancestral condition ceae were primitively a scalariform-perforation­ not only for the family but also for the entire plate family (and we believe this to be the case, order Myrtales (V an Vliet & Baas, 1984). Rec­ as there are no reasons why scalariform per­ ognition of scalariform perforation plates as a foration plates would have evolved in Myrta­ primitive (or plesiomorphic) feature does not ceae by reversion from the derived simple per­ necessarily imply close phylogenetic relation­ foration plates), it is obvious that in all present ship of Luma, Myreeugenia, Myrtastrum, Myr­ day habitats of Myrtaceae the scalariform per­ teola, Neomyrtus, Ugni, and Tepualia, the seven foration plate has largely been eliminated in myrtaceous genera retaining this character state. favour of simple plates, and that only in fairly The first six genera are members of subfamily cool mesic sites some Myrtaceae have retained Myrtoideae. Tepualia, in contrast, is the only this primitive feature. Advocates of a function­ representative of subfamily Leptospermoideae ally adaptive significance of scalariform per­ (subfamilies sensu Briggs & Johnson, 1979, foration plates in trapping embolisms due to McVaugh, 1968, Niedenzu, 1893, or Schmid, thawing of columns of frozen xylem sap may 1980) with scalariform perforation plates. And find these data in support of their hypothesis here they are very infrequent (Table 2), and (cf. Zimmermann, 1978). However, if we con­ largely rudimentary. sider not only that the vast majority of Myrta­ Comparison of the incidence of multiple per­ ceae growing sympatrically with or in similar forations (i.e., scalariform perforations as weil habitats as the species with scalariform perfora­ as sieve-like foramina te ones) with taxonomie tion plates have, in contrast, simple perforation groupings in Myrtoideae is interesting. For ex­ plates, but also that some Myrtaceae with scala­ ample, such perforations are largely restricted riform perforation plates are never subjected to (I) to Briggs and J ohnson's (1979) 'Myrtus al­ heavy frosts (though occurring in cool mesic liance' (= tribe), (2) to McVaugh's (1968) Ugni­ sites), the explanation becomes less convincing, Myrteola-etc. phyletic 'trend' and his 'polymor­ at least for this family. The pattern of variation phic', aphyletic group containing Luma and in Myrtaceae is more in support of the idea, ad­ Myrceugenia, or (3) to Berg's classical (see vanced by one of us (Baas, 1984), that the high McVaugh, 1968, and Niedenzu, 1893) subtribes incidence of scalariform perforation plates in Myrciinae and Pimentinae. Especially intriguing cool mesic floras should be explained by a lack is that Ugni and Myrteola have traditionally of strong selective pressures in these ecological been considered closely allied, and that both conditions to eliminate this primitive feature. genera have scalariform perforation plates re­ Such pressures are presumably stronger in en­ presented in all of their species. Even more in­ vironments with a high er demand on efficient triguing is the hypothesis that these genera are sap transport, that is, in the tropical lowland related to the monotypic genera Myrtastrum and xeric regions. and Neomyrtus, as suggested by Landrum (pers. Our data on the occurrence of helical wall comm., 1981, to Schmid) at the start of his re­ thickenings in vessel elements (Tables 2, 3) cer­ vision of these groups. In view of the facts that tainly do not inspire unambiguous interpreta­ most genera in Myrtoideae have exclusively tion in terms of functionally adaptive signifi­ simple perforation plates (Table 3), and that cance. In Myrtaceae helices are hardly restricted within the coherent group Myreeugenia (Lan­ to species that are periodically su bjected to in­ drum, 1981) there is wide variation in occur­ creased risks of em bolism due to frost and sub­ rence and frequency of scalariform perforation sequent thawing, or due to excessive negative plates (Table 2), too much phylogenetic signifi­ pressures in the xylem sap. Carlquist (1982) cance should not be attached to this character. has hypothesised that helices increase the sur­ Nevertheless, it is a convenient diagnostic tool face area of walls of vessel elements and thus to recognise woods of this sm all group of Myr­ contribute to their bonding capacity for xylem taceae. For example, a collection (Sparre & sap, thus decreasing such risks. Were this to Constance 10797) originally identified as Amo­ apply to Myrtaceae, one would have expected myrtus luma was later reidentified as M. chry­ many more species (e.g., of Myreeugenia) from socarpa because Amomyrtus completely lacks xeric localities or from regions with significant scalariform perforation plates (Table 3). Simi­ winter frosts to show vessel elements with heli­ larly, a slide in the Oxford collection (FHOw cal wall thickenings. As outlined above, this is 11684) with mixed simple and scalariform per­ not the case. foration plates and representing an unvouchered collection of M. jernandeziana is highly suspect Phylogenetie and taxonomie signijicanee as all other sampIes of this species (Tables I, 2) The sporadic occurrence of scalariform per- have only simple perforation plates.

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For helical wall thickenings ofvessel elements Bierhorst, D. W. & P.M. Zamora. 1965. Primary the distribution of genera over different allian­ xylem elements and element associations ces and the tremendous infrageneric variation of angiosperms. Amer. J. Bot. 52: 657- suggest an even more Iimited systematic and 710. diagnostic value than for scalariform perfora­ Briggs, B.G. & L. A. S. Johnson. 1979. Evolution tion plates. However, examples can be cited. in the Myrtaceae - evidence from inflores­ Collections (respectively, Hastings 474 and cence structure. Proc. Linn. Soe. New South Sparre & Constance 10859) originally identi­ Wales 102: 157-256. fied as Temu cruckshanksii and T. divaricatum Brown, F.B.H. 1922. The secondary xylem of were later reidentified as Myrceugenia exsucca Hawaiian trees. Occ. Pap. B.P. Bishop Mus. because Temu (regarded as Blepharocalyx by 8: 215-371. Landrum - pers. comm., 1984) completely Butterfield, B. G. & B. A. Meylan. 1974. Ves­ lacks helical wall thickenings in its vessel ele­ tured scalariform perforation plate openings ments (Table 3). And Wagemann Wiedenbrug in Neomyrtus peduneulata. Austr. J. Bot. (1949) used this feature of M. exsucca in a key 22: 425-427. to separate its wood from Tepualia stipularis. ~ & ~ 1980. Three-dimensional structure of wood: an ultrastructural approach. 2nd Ed. Chapman & Hall, London. Acknowledgements Carlquist, S. 1975. Ecological strategies of xy­ This work was supported in part by NSF lem evolution. Univ. California Press, Ber­ grant DEB78-04289 to Schmid and the Com­ keley. mittee on Research, University of California, ~ 1977. Ecological factors in wood evolution: Berkeley. We thank Cynthia Gibbon (Berkeley) a f10ristic approach. Amer. J. Bot. 64: 887- for enduring the preparation of numerous slides 896. of wood macerations and also Bertie van Heuven ~ 1980. Further concepts in ecological wood (Leiden) for SEM assistance and for additional anatomy, with comments on recent work preparations of sections and macerations. We in wood anatomy and evolution. Aliso 9: appreciate the help of Leslie R. Landrum in 499-553. providing determinations of some of the Schmid ~ 1982. Wood anatomy of IIIicium (Illieia­ collections and in supplying various sam pIes. ceae). Phylogenetic, eeological and func­ tional interpretations. Amer. J. Bot. 70: 1578-1598. References Chittenden, F.J. (ed.) 1956. The Royal Horti­ Baas, P. 1973. The wood anatomical range in eultural Soeiety dietionary of gardening. 4 Ilex (Aquifoliaceae) and its ecological and Vois. 2nd Ed. by P.M. Synge (ed.). Claren­ phylogenetic signifieanee. Blumea 21: don Press, Oxford. (Also 1969 Supp!., 2nd 193-258. ed., by P.M. Synge, ed.) ~ 1976. Some functional and adaptive aspects Cronquist, A. 1981. An integrated system of of vessel member morphology. In: Wood c1assification of f10wering . Columbia structure in biologie al and technologieal Univ. Press, New York. research (eds. P. Baas, A.J. Bolton & D.M. Dadswell, H. E. 1972. The anatomy of euealypt Catling): 157-181. Leiden Bot. Sero 3. woods. CSIRO, Australia; Forest Prod. Leiden Univ. Press, The Hague. Div. App!. Chem. Techno!. Paper no 66: ~ 1977. The peculiar wood structure of Lep­ (1-4),1-28. tospermum crassipes Lehm. (Myrtaceae). Dahlgren, R. & R. F. Thorne. 1984. Circum­ IA WA Bull. 1977/1: 25-30. scription of and variation in the order Myr­ ~ 1982. Systematic, phylogenetie, and ecolo­ tales. Ann. Missouri Bot. Gard. (in press). gical wood anatomy - history and perspec­ Emberger, L. 1960. Les vegetaux vasculaires. tives. In: New perspectives in wood anato­ Vo!. 2 of M. Chadefaud & L. Emberger: my (ed. P. Baas): 23-58. Martinus Nijhoffl Traite de botanique (systematique). 2 Vols. Dr W. Junk Pub!., The Hague. in 3. Masson et Cie, Paris. ~ 1984 (in the press). Ecological patterns in Fahn, A., E. Werker & P. Baas. In press. Wood xylem anatomy. In: On the economy of anatomy and identifieation of trees and plant form and function (ed. T.J. Givnish). shrubs from Israel and adjaeent regions. Cambridge Univ. Press, New York, Cam­ Israel Aead. Sciences, Jerusalem. bridge. Fasolo, U. 1939-40. Atlante micrografico dei ~ , E. Werker & A. Fahn. 1983. Some eeologi­ legni deli' Africa Orientale Italiana. Cartella cal trends in vessel characters. IA WA Bull. I (Introduzione, Tav. 1-12). R. Erbario n.s. 4: 141-159. Coloniale, Firenze.

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