IAWA Journal, Vol. 18 (3),1997: 247-259

WOOD ANATOMY OF SOME ANAUERIA AND SPECIES () 1 by Catia H. Callado & Cecilia G. Costa

Anatomy Sector, Rio de Janeiro Botanical Garden, Rua Jardim Botänico, 1008 Jardim Botänico, Rio de Janeiro, RJ / CEP 22400-000, Brazil

SUMMARY

The wood anatomy of the species Anaueria brasiliensis Kosterm., Beilschmiedia emarginata (Meissn.) Kosterm., B. rigida (Mez) Kosterm. and B. taubertiana (Schw. et Mez) Kosterm. (Lauraceae) is described. The and ecology of these species, important components of the Amazonian forest or Atlantic forest of southeastern Brazil, are discussed as related to wood anatomy. The main anatomical differences are: presence, type, arrangement and location of inorganic inclusions and secretory cells, and the arrangement of the axial parenchyma. Key words: Wood anatomy,Anaueria. Beilschmiedia, Lauraceae, taxon­ omy, ecology.

INTRODUCTION

Anaueria Kosterm. is amonotypic genus, with the single speciesA. brasiliensis Kosterm. (Kostermans 1938; Van der Werff 1991).1t was once (1957) included in Beilschmiedia by Kostermans, but is anatomically closer to Mezilaurus Taubert (Rohwer 1993). The genus Beilschmiedia Nees consists of about 250 species and is found through­ out the tropics (Rohwer 1993); six species occur in the Atlantic forest of southeastern Brazil: Beilschmiedia angustifolia. B. emarginata. B. fluminensis. B. rigida. B. stricta and B. taubertiana. Three of these species were examined in this study. They were selected on the basis of availability of botanical material, whether fresh from the forest or through the exchange of wood sampies from collections in Brazil. Beilschmiedia rigida (Mez) Kosterm. and B. taubertiana (Schw. et Mez) Kosterm. are among the most important species at the Macae de Cima Munü;:ipal Ecological Reserve and the Parafso State Ecological Station (Relat6rio Tecnico do Programa Mata Atläntica do Jardim Botänico do Rio de Janeiro 1990). Beilschmiedia emarginata (Meissn.) Kosterm. is very similar morphologically to B. rigida, thus making species identifica­ tion difficult. Anaueria brasiliensis and the three Beilschmiedia species are important compo­ nents of two of the most endangered Brazilian ecosystems and are very difficult to identify.

1) Part of the first author' s MSc thesis.

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This study is part of aseries on the anatomy of the most important Atlantic forest families (Plant Anatomy Project, Atlantic Forest Program, Rio de Janeiro Bo­ tanical Garden).

MATERIALS AND METHODS

Characteristics of the individual used in the study are given in Table 1. Wood sampies were treated as recommended in Coradin and Mufiiz (1991), using techniques appropriate for the study of wood anatomy (lAWA Commitee 1989; Burger & Richter 1991). Generally, classification, measurements and counts of cell elements, as well as termino10gy, follow the recommendations of the JAWA Committee (1989) and Coradin and Mufiiz (1991). Description of si1ica bodies and prismatic crystals is based on Rich­ ter (1980); silica bodies were subjected to X-ray microanalysis (Wi1cox 1994). Lengths of oil/ mucilage cells were measured on radial or tangential longitudinal sections, or on dissociated material.

Table 1. Tree measurements and characteristics of the wood and collection site of each tree sampled.

Q 0 o '.:;::: a ... ü ü .~ ., .~ ~ g g 0 Q " ~= '"0 0 ]' ., ....," ü 0 ~§ .E ~ 'B G .,"0 0 Q E .~ ..!l § '"o 0 ~ "0 § '§ 11 .~ ~ ..c: 11 ü ., ~ ~- .~ <=I Q 1;l~ ~ lZl ., e on <=I • '+< ., ~ 0'" ., .. -d '§ § ... ~ Cl'" § '" ... @ ~ '" 0 .~ a ... - ';:::l o 0 Q ..... 'E: ., ~ " ü lS" o Q Q 0 ~ '" " ., E- ... 0 a <=1- E ~ ~~ .g "<=I .-f3 'B ~ §8 ,,~ o '" '" Q ~ ::l '" ",..c: '"B Q '" :i§ § 0, 11 '" ., "..c: '" '" ~ 0 ::a " '" ~ "" 11 Species ~~ Ci ~:r: u ~ ~ ~., ~1Zl

Anaueria

brasiliensis 6843 45 H Barreirinha-AM 150 25 0 2500 S brasiliensis 7324 61 H Tefe-AM 150 25 0 2500 S

Beilschmiedia

emarginata 7347 86 H S. da Cantareira-SP 750 20 0 1300 M rigida 7277 39 H N. Friburgo-RJ 1100 18 0 1750 M

rigida 7298 63 S N. Friburgo-RJ 1100 18 0 1750 M rigida 7266 29 S N. Friburgo-RJ 1100 18 0 1750 M rigida 7300 30 H N. Friburgo-RJ 1100 18 0 1750 M taube rtiana 7297 26 S Mage-RJ 650 21 0 2250 M

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RESULTS

Anaueria brasiliensis Kosterm. (Fig. 1-7). Synonym: Beilschmiedia brasiliensis (Kosterm.) Kosterm. Common names: Anaueni, Louro Anaueni Branco, Anoeni-ferro. Anatomical descriptions: Growth rings: indistinct (Fig. 1). Vessels: wood diffuse-porous (Fig.l); (6-)8(-10) vessels/mm2; solitary (77%), ra­ dial multiples of 2 (22%) or clusters of 3-4 (1 %); circular to oval outline; element length (396-)646( -835) JlIll; tangential diameter (101-) 130(-152) JlIll; mean wall thick­ ness 4 11m; end walls oblique (30°); exclusively simple perforation plates; large, alter­ nate and circular to polygonal intervessel pits; reduced borders, elongate horizontally and diagonally vessel-ray pits, similar to the vessel-axial parenchyma pits (Fig. 4); tyloses present. Libriform fibres: length (1131-)1521(-1920) 11m; tangential diameter (16-)25 (-33) JlIll; lumina (8-)13(-21) JlIll; thin- to thick-walled; minutely bordered pits, mainly confined to the radial walls; gelatinous fibres present (Fig. 3). Axial parenchyma: paratracheal unilateral (Fig. 1), aliform and confluent, uniting the vessels diagonally, but never forrning concentric bands (Fig. 1); strands of 3-8 cells, (399-)627(-847) JlIll. Rays: (5 -)7 (-8) / mm; uniseriate composed of square to upright cells, 1-7 cells high, (54-)161(-357) 11m (Fig. 2) and multiseriate, 2-3 cells wide, (15-)24(-41) 11m (Fig. 2), composed entirely of procumbent cells, or with the body of procumbent cells with 1 (or 2) marginal rows of square or upright cells (Fig. 5), sometimes only at one end; height (186-)375(-603) JlIll; rough silica grains present (Fig. 6 & 7). Oil/mucilage cells: associated with axial parenchyma, length (125-)198(-271) 11m (Fig. 1 & 5).

~ Fig. 1-5. Anaueria brasiliensis - 1: Transverse section: diffuse-porous; solitary vessels (*), radial multiple of two elements (*); oil/mucilage cells (arrow). Scale bar = 330 iJIll. - 2: Tangential section: vessel elements with slightly oblique end walls (*); uniseriate rays (thin arrow); biseriate rays (thick arrow) and series ofaxial parenchyma (*). Scale bar = 330 iJIll. - 3: Transverse section: detail of gelatinous fibres (thick arrow). Scale bar = 10 iJIll. - 4: Tangential section: detail of strand axial parenchyma (*), vessels parenchyma pits evident (thin arrow). Scale bar = 130 iJIll. - 5. Radial section: rays with square marginal cells (thin arrow) and detail of oil/ mucilage cell in longitudinal section (thick arrow). Scale bar = 130 iJIll.

~~ Fig. 6 & 7. Anaueria brasiliensis, silica bodies in ray cells. - 6: In optical microscope (thick arrow). Scale bar = 10 iJIll. - 7: In scanning electron microscope (*), detail of ray pits (thick ar­ row). Scale bar = 7.5 iJIll. - Fig. 8-10. Beilschmiedia emarginata. - 8: Transverse section: growth ring bounded by marginal parenchyma (black arrow) and axial parenchyma aliform (white arrow). Scale bar = 330 iJIll. - 9: Dissociated vessel element: simple perforation plates and short appendices (thick arrow). Scale bar =75 iJIll. - 10: Tangential seetion: uniseriate rays (thick arrow) and multiseriate rays (ti). Scale bar = 330 iJIll.

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For legend, see page 249.

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For legend, see page 249.

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6: (3),1997 00 18 Vol. Journal, Journal, Downloaded from Brill.com10/07/2021 06:20:25PM IAWA IAWA 252 Callado & Costa - Anatomy of Anaueria and Beilsehmiedia 253

Beilschmiedia emarginata (Meissn.) Kosterm. (Fig. 8 -10) Synonyms: Hufelandia emarginata (Meissn.) Mez; emarginata Meissn. Anatomical descriptions: Growth rings: distinct, marked by marginal parenchyma bands, reduced diameter and increased wall thickness of latewood vessels (Fig. 8). Vessels: wood diffuse-porous (Fig. 8); (3-)4(-5) vessels/mm2; solitary (61 %), ra­ dial multiples of 2-3 (37%) or clusters of 3-4 (2%); circular to oval outline; element length (330-)453(-529) 11m; tangential diameter (122-)143(-162) /lffi; mean wall thickness 311m; end walls oblique (31°); exclusively simple perforation plates (Fig. 9); large, alternate and circular to polygonal intervessel pits; bordered, elongate hori­ zontally and diagonally vessel-ray pits, similar to the vessel-parenchyma pits; tyloses present. Libriformfibres: length (1247-)1542(-1873) /lffi; tangential diameter (24-)30(-39) /lffi; lumina (13-)19(-27) 11m; thin-walled; minutelybordered pits, mainly confined to the radial walls. Axial parenchyma: paratracheal vasicentric, aliform (Fig. 8), confluent and bands of 2-4 cells wide, marginal parenchyma bands up to 5-8 cells (Fig. 8); strands of 2-4 cells, (335-)429(-561) /lffi. Rays: (4-)6(-8)/mm; uniseriate composed of square to upright cells (Fig. 10), 1-5 cells high, (54-)133(-326) 11m, and multiseriate, 2-4 cells wide, (24-)48(-74) 11m (Fig. 10), composed entirely of procumbent cells, or with the body of procumbent cells with one marginal row of square or upright cells, sometimes only at one end; height (143-)303(-462) /lffi.

Beilschmiedia rigida (Mez) Kosterm. (Fig. 11-14) Synonym: Hufelandia rigida Mez ex Taubert. Common name: Canela tapinha. Anatomical descriptions: Growth rings: distinct, marked by marginal parenchyma bands, reduced diameter and increased wall thickness oflatewood vessels (Fig. 13).

Fig. 11-14. Beilschmiedia rigida. - 11: Tangential seetion: detail of pits to parenehyma eells (arrow). Seale bar = 75 ~. - 12: Tangential seetion: detail of oil/mueilage eell in longitudinal seetion (arrow) and multiseriate ray. Seale bar = 75 ~. - 13: Transverse seetion: growth ring bounded by marginal parenehyma (*); axial parenehyma aliform (thick arrow). Scale bar = 330~. - 14: Tangential seetion: uniseriate rays (thin arrows), multiseriate rays (*) and strand axial parenchyma (*). Scale bar = 330 ~. - Fig. 15. Beilschmiedia taubertiana. Transverse seetion: solitary vessels and in radial multiples of two elements; axial parenchyma vasicentric (thick arrow); confluent (*); oil/mucilage cells in transverse section (thin arrows). Scale bar = 130~.

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Vessels: wood diffuse-porous (Fig, 13); (2-)3(-5) vessels/mm2; solitary (68%), radial multiples of 2-4 (30%) or clusters of 3-4 (2%); circular to oval outline; length (239-)445(-564) J.lm; tangential diameter (126-)158(-212) J.lm; mean wall thickness 4 J.lm; end walls oblique (32°); exclusively simple perforation plates; large, alternate

Fig. 16-19. Beilschmiedia taubertiana. - 16: Radial seetion: ray eomposed of proeumbent eells in eentral region with siliea grains (*) and upright at margins (arrow). SeaJe bar = 75 ~. - 17: Crystal sand in ray (arrow). Se ale bar = JO~ . - 18: Overall view of siliea bodies in ray eell in seanning eleetron microseope (arrows). Seale bar = 5 ~. - 19: Tangential seetion: uniseriate rays (thin arrow), multiseriate rays (thiek arrows), oil/mueilage eell in longitudinal seetion (white arrow). Seale bar = 130 ~.

Downloaded from Brill.com10/07/2021 06:20:25PM via free access Callado & Costa - Anatomy of Anaueria and Beilschmiedia 255 and circular to polygonal intervessel pits; bordered, elongate horizontally and diago­ nally vessel-ray pits, similar to the vessel-parenchyma pits (Fig. 11); tyloses present. Librifonnfibres: length (976-)1376(-1713) Jlm; tangential diameter (19-)25(-32) Jlm; lumina (11-) 16(-24) Jlffi; thin-walled; minutely bordered pits, mainly confined to the radial walls. Axial parenchyma: paratracheal vasicentric, aliform (Fig. 13), confluent and in bands of 2-4 cells wide, marginal parenchyma bands up to 5-10 cells (Fig. 13); strands of 2-4 cells, (319-)483(-819) Jlm. Rays: (4-)6(-8)/mm; uniseriate composed ofsquare to upright cells (Fig. 14~, 1-5 cells high, (68-)171(-370) Jlffi, and multiseriate, 2-4 cells wide, (24-)38(-56) Jlffi (Fig. 14), composed of procumbent cells, or the body procumbent cells with one mar­ ginal row of square or upright cells, sometimes only at one end; length (172-)398 (-864) Jlffi. Oil/mucilage cells: extremely rare or absent in some individuals, associated with axial parenchyma, length (119-)169(-202) Jlffi (Fig. 12).

Beilschmiedia taubertiana (Schw. et Mez) Kosterm. (Fig. 15-20) Synonym: Hufelandia taubertiana Schw. et Mez. Anatomical descriptions: Growth rings: distinct, marked by marginal parenchyma bands, reduced diameter and increased wall thickness of latewood vessels. Vessels: wood diffuse-porous; (8-)11(-14) vessels/mm2; solitary (44%), radial mul­ tiples of 2-4 (51 %) or clusters of 3-4 (4%); circular to oval outline (Fig. 15); length (220-)503(-660) Jlffi; tangential diameter (95-)112(-137) Jlffi; mean walls thickness 3 Jlffi; end walls oblique (33°); exclusively simple perforation plates; large, alternate and circular to polygonal intervessel pits; bordered, elongate horizontally and diago­ nally vessel-ray pits, similar to the vessel-parenchyma pits. Librifonnfibres: length (795-)1061(-1452) Jlffi; tangential diameter (19-)23(-29) Jlffi; lumina (11-)15(-18) Jlm; thin-walled; minutely bordered pits, mainly confined to the radial walls; gelatinous fibres present. Axial parenchyma: paratracheal vasicentric (Fig. 15), aliform, confluent (Fig. 15) and bands of 2-4 cells wide, marginal parenchyma bands up to 5-7 cells; strands of 2-5 cells, (412-)541(-643) Jlffi. Rays: (6-)8(-1l)/mm; uniseriate composed of square to upright cells, 1-6 cells high, (85-)181(-322) Jlffi, and multiseriate, 2-5 cells wide, (26-)47(-75) Jlffi (Fig. 19), composed of procumbent cells, or the body procumbent cells with one marginal row of square or upright cells (Fig. 16), sometimes only at one end; length (162-) 384(-600) Jlm; silica aggregate bodies present (Fig. 16 & 18) and crystal sand in the ray cells (Fig. 17). Oil/mucilage cells: associated with axial parenchyma; rather elongate, the length (380-)540(-711) Jlffi (Fig. 19).

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DISCUSSION AND CONCLUSION

Wood structure in the Lauraceae is reportedly very uniform (Richter 1987), yet a number of anatomical characteristics readily differentiate lauraceous wood from that of other families (Stern 1954). Most lauraceous genera, however, are not weIl defined. Some species belonging to the same genus do not always show similar anatomical character­ istics; wood anatomy may differ considerably within a given genus. These differences often are related to erroneous taxonomie identification (Dadswell & Eckersley 1940; Record & Hess 1942; Janssonius 1950). An analysis of the wood anatomy in Anaueria brasiliensis and in the three Beilschmiedia species show uniformity of traits for these genera, in agreement with descriptions by Record and Hess (1942) and Richter (1987). The number of examined samples is small and it is necessary to study a larger number to verify the results presented herein. In the material we examined the ana­ tomical characteristics that separate the different species most effectively are organi­ sation of the axial parenchyma and the presence and type of inorganic inclusions and secretory cells. Anaueria brasiliensis has unilaterial paratracheal axial parenchyma that extends tangentially or obliquely in an aliform or confluent pattern; vasicentric axial paren­ chyma is rare. The three Beilschmiedia species have vasicentric, aliform and confluent axial parenchyma, as weIl as marginal/zonate bands. Richter (1980) investigated the occurrence of inorganic material in the xylem and its diagnostic significance of 750 lauraceous species. Anaueria brasiliensis has rough silica grains (Richter 1980). Beilschmiedia emarginata and B. rigida belong to the 30% of lauraceous species that have no inorganic inclusions, while B. taubertiana has two types of inclusions: aggregate silica grains and calcium oxalate crystals. The former are found mostly in rays and are cited as a frequent inclusion in South American spe­ eies of Beilschmiedia (Richter 1980). Calcium oxalate crystals appear as crystal sand in the single sampie of B. taubertiana, an occurrence that had not previously been reported for this genus. Oil and mucilage cells are common in Lauraceae (Metcalfe 1987). They have been reported for several genera, and are found throughout the plant. Little taxonomie sig­ nificance has been attributed to the location, arrangement, size and content of these cells (Richter 1987). The taxonomie value of oil and mucilage cells at the generic and infrageneric levels has been questioned by several authors, e.g. Bakker & Gerritsen (1992) and Bakker et al. (1992). However, it is possible that these cells may be a useful trait in the species treated here, but additional samples of each species need to be examined. Anaueria brasiliensis has many oil/mucilage cells, usually associated with the axial parenchyma. The one sampie of Beilschmiedia emarginata examined did not have secretory ceIls, while B. rigida has smaIl, very rare oil/mucilage cells embedded in the axial parenchyrna, or these cells rnay be totally lacking. Beilschmiedia taubertiana had elongated oil/mucilage cells (380-711 11m) in the axial parenchyma. In Beilschmiedia, secretory cells may be present or absent in different plants of the same species, and the proportion of these structures may vary significantly from one

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Table 2. Differences in secondary xylem features of Anaueria brasiliensis, Beilschmiedia emarginata, B. rigida, and B. taubertiana.

Features A. brasiliensis B. emarginata B. rigida B. taube rtiana

Growth rings indistinct distinct distinct distinct Vessels - vessels/mm2 range 6-10 3-5 2-5 8-14 - tangential diameter range (flIIl) 101-152 122-162 126-212 95-137 - length range (flIIl) 396-835 330-529 239-564 220-660 Fibres - 1ength range (flIIl) 1131-1920 1247-1873 976-1713 795-1452 Axial parenchyma type unilateral, vasi- vasicentric, ali- vasicentric, ali- vasicentric, a1i- centric (rare) form, confluent form, confluent form, conflu- and bands and bands ent and bands - marginal bands absent present present present Oi1 / mucilage cells present not observed present (rare) present Silica grains rough grains not observed not observed aggregate grains and calcium oxalate crystals Prismatic crystals not observed not observed not observed present Vulnerability 15.33 35.74 52.72 10.16 Mesomorphy 9899 16179 23481 5113

plant to the next (Kosterrnans 1957). Beilschmiedia emarginata and B. rigida are are not readily separated on the basis of wood anatomy, the two species are morphologi­ cally very similar and are separated only by leaf and flower size, and ovary pubes­ cence in B. rigida (Kosterrnans 1938). Kosterrnans noted that B. rigida could be struc­ turally more developed since it is the only American species with trichomes on the ovary. When dealing with wood identification, it is essential to distinguish between those anatomical characters that are relatively constant, and those that vary with grow­ ing conditions (Metcalfe & Chalk 1985). Some characteristics such as the number of vessels/mm2, length and diameter of vessel elements and proportion ofaxial paren­ chyma may be strongly influenced by the environment (Brazier 1976; Metcalfe & Chalk 1985). Plants from mesic habitats have a tendency for vessels to be wider, to have fewer vessels per group and a smaller number of vessels/mm2• Species from xeric habitats have many narrow vessels, a larger number of vessel groups, and also more vessels per group (Starr 1912, apud Vieira 1994). All four species have solitary vessels, or few vessels grouping, with a medium tangential diameter. These characteristics are typical of mesophytes according to Starr (1912, apud Vieira 1994), Baas (1982) and Carlquist (1988).

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Indices of vulnerability and mesomorphy and their mathematical relationships have been labelled by critics as arbitrary in the analysis of those structural variables that are influenced by the environment (Rury 1985; Kuniyoshi 1993). However, in one species these indices have been shown to be effective in the case of xylem cell elements when the habitat is known (Kuniyoshi 1993). According to Baas (1982) and Carlquist (1988), the values expressed by these indices represent a range of ecological factors, for ex am­ pIe, vessel diameter decreases and frequency increases as rainfall diminishes or tem­ perature falls. Indices of vulnerability and mesomorphy (Carlquist 1977) show the species studied here to be mesomorphic (V > 1; M > 200). Beilschmiedia taubertiana has the lowest mesomorphy index and B. rigida has the highest mesomorphy index (Table 2). Indistinct or poorly defined growth rings are characteristic of the wood of tropical lowland species; tropical mountain species, on the other hand, are similar to those of temperate zones as far as structural features ofthe xylem are concemed (Baas 1973). Anaueria brasiliensis does not have distinct growth rings, which is to be expected since this is a tropicallowland species. On the other hand, Beilschmiedia emarginata, B. rigida and B. taubertiana have distinct growth rings, indicating periodic cambial activity. According to Carlquist (1978) and Norton (1979, apud Ogden & West 1981), higher altitudes may favour the appearance of these traits.

ACKNOWLEDGEMENTS

The authors would like to thank the Atlantic Forest Program of the Rio de Janeiro Botanical Garden, technicians Ines C.N. Grillo and Paulo R.R. Dias, CNPq and FAPERJ for the scholarships, and Dr. Raul D. Machado for helpful comments.

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