172 IAWA Bulletin n.s., VoL 1 (4),1980

WOOD ANATOMY OF THE

by

J. T. Quirk Forest Products Laboratory*, Forest Service, P.O. Box 5130, Madison, Wisconsin 53705, U.S.A.

Summary are a11 woody, ranging from a few climbers and A wood anatomy key based on reliable xylem shrubs through sma11 to very large trees. Some features of ray type and proportions separates are among the largest emergent canopy trees the six genera in the family Vochysiaceae. Se• (Huber, 1909) and are characteristic genera of lected anatomical features of 57 are the lowland rain forest of Middle and South summarized. The wood anatomical range en• America (Daubenmire, 1978). countered is presented in descriptions for each . Materials and Methods Woods of the Vochysiaceae differ from clo• Wood sampies were used from the Madison sely related taxa. Vochysiaceae have vestured and S. J. Record co11ections housed at the U.S. vessel pitting and only libriform fibers; banded Forest Products Laboratory; a few sampies are axial parenchyma and traumatic interce11ular from Utrecht. Woody material more than 5 canals occur frequently; heartwoods of a11 centimeters (ern) from the pith was used for Vochysiaceae test positive (turn blue) with the the anatomical descriptions. application of Chrome-Azurol-S. Wood anatomical features were measured from transverse, radial, and tangen tial sections Introduction and from macerations. Where possible, section• For the family Vochysiaceae there is no ana• ing blocks were cut to yield cut sections 1 cen• tomica11y based key with which to separate the timeter square for a maximum survey area. Af• six genera. This study of the anatomy of Vochy• ter final sectioning of the block a 1 mm-thick siaceae species was made with a three way ob• wafer was split from the radial face, macerated jective: to generate a key to separate the genera; using Franklin's method (1946), stained with to determine if anatomical data could be used safranin, and mounted in glycerine for measur• to separate species or species groups similar to ing. One hundred and sixty sampie blocks re• subsections and seetions as in the taxonomie presenting 57 species were prepared and the monographs of the family by Stafleu (1948- following anatomical measurements made. For 1954); to assist in evaluating growth and tissue each sampie, 100 fiber lengths and 50 total ves• patterns by using ecological da ta as did Baas sei-element lengths (as per Chalk and Chatta• (1973) or Carlquist (1977). way, 1934) were measured from the macera• This first report presents data on the six tions. Twenty-five measurements were made genera of the family and a key for their separa• from each prepared slide: maximum vessel dia• tion based on anatomical data. A subsequent meter, fiber diameter, fiber wall thickness, total report will present species descriptions, quanti• height and procumbent cell portion of tallest tative and qualitative anatomical data, and the rays, number of cells high and wide, and dia• implication of ecological factors. meters of intervascular pits and ray-vessel pit The family Vochysiaceae consists of the six size. Ten random fields of 1 mm2 size were genera Erisma, Erismadelphus, Callisthene, counted on the tangential section for the pro• Qualea, Salvertia, and Vochysia, and is biconti• portion of multiseriate to uniseriate rays. I f nental in distribution due to the disjunction of one-half or more of the ray length was within the single genus Erismadelphus in the Camer• the count boundary, alternating halves were oons and Gabon in equatorial West Africa. counted as in or out. Number of pores per Brazil is the distribution center of the five mm 2 were counted in 1 mm 2 grids over the Western Hemisphere genera. Of these five only entire area of the transverse section. Percen• the genus Vochysia has species north of Panama tages by volume of fibers, vessels, parenchyma, to the Yucatan Peninsula of Mexico. The major• and rays were obtained using a Zeiss dot-grid ity of a11 species are in the Amazonian rain for• eyepiece and by counting the points falling on est. Many species are limited, with few excep• a particular tissue type (Quirk, 1975). A polar• tions, to a minor geographical unit. The species izing microscope was used to assess presence

* Maintained at Madison, Wis., in cooperation with the University ofWisconsin.

Downloaded from Brill.com10/01/2021 10:54:35PM via free access IA WABulIetin n.s., Vol. 1 (4), 1980 173 and type of crystals from cut sections and from macerations. Percent silica was assessed from microchemical assay. 000000\0\000 Reliability and comparability of materials A taxonomic monograph of the family Vochysiaceae was published in four parts by Stafleu (1948, 1952, 1953, and 1954), who recognized 183 species distributed over six genera and several sections and su bsections. Since that time approximately 12 new species, mainly of the genus Vochysia, have been de• scribed, implying that the naming of some of our specimens may be incorrect. I am aware that not all our specimens are vouchered and 88~888 Oot-OtnC') that some of the names may not be absolutely MM~M('1')- I 1 1 I 1 1 reliable, but there is no doubt that all of the OtnOOOO tnNtnOOO specimens described here belong to the various N ...... N-• genera of the V ochysiaceae. Wood specimens "-'''-'''-''''-''''-''''-' may have been collected from the bole or branch, near pith or under bark, and these data * are not included on the vouchers. For our pur• *,-..,,-,.-..-..-..-.. poses, sampIes at least 5 cm or more from the ('f)V'lOVJN_ -N pith were considered as mature wood sampIes I 1 I 1 I 1 0-0--0\ for descriptive purposes. '-" '-" '-' '-" "-' "-' C')Otnt-\oOtn "":0i0i0i

General features of the family Vochy siilceae '-o '"::: Authorities such as Cronquist (1968), Hut• ....

Downloaded from Brill.com10/01/2021 10:54:35PM via free access -...l Table 1h. - Fiber data for six genera of Vochysiaceae ~

Genus Percent Diameters Length Wall Septate by (Ilm) (Ilm) thickness volume (Ilm)

Erismadelphus 46 16.3 (14-19)* 1,428 ( 800-2,060) 4.0 (2.5- 5.0) +** Erisma 40 20.5 (16-28) 1,422 (1,000-2,000) 3.1 (1.5- 5.0) + Vochysia 57 19.7 (10-37) 1,445 ( 600-2,400) 3.0 (1.2- 9.0) rare Salvertia 38 17.0(13-20) 1,067 ( 710-1,430) 3.0 (2.0- 4.0) + Qualea 52 16.3 (11-22) 1,330 ( 600-2,300) 4.2 (1.0-11.0) + CaUisthene 38 18.8 (16-20) 906 ( 663-1,350) 3.2 (2.5- 4.0) + Family 45 18.1 1,266 3.4 * Range ofvalues in parentheses. ** +, present. For number of species and specimens studied, see Table IA.

Table Ie. - Ray data for six genera of Vochysiaceae

Genus Number Width Height Percent of rays Number of Ray cell (per mm 2) (Ilm) (Ilm) procum bent ceUs shape h/d ;; Multiseriate ::.!E Downloaded fromBrill.com10/01/2021 10:54:35PM Uniseriate ratio > TaU Wide t:e ;:s:: Erismadelphus 38 (30-40)* 16.8 (14- 19) 586 (400- 700) 100 13 (12- 15) I 2.2 Erisma 45 (30-70) 41.0 (25- 90) 672 (375-1,300) 27 73 19 ( 31) 3 4) 5' 8- (2- 1.3 :3 Vochysia 22 ( 7-43) 79.6 (30-275) 1,480 (650-4, I 00) 27 73 48 (20-170) 5(3-10) 1.6 Salvertia 37 (31-47) 89.5 (70-110) 730 (675- 925) 43 57 19(16- 24) 3 (2- 4) 1.2 r Qualea 22 ( 45.0 (20-110) 678 (300-1,860) 70 30 90) < 8-36) 14(10- 3 (2- 6) 1.3 ~ Callisthene 35 (26-40) 50.0 (40- 60) 480 (420- 570) 75 25 22 (15- 30) 3 (3- 4) 1.1 Family 33 61 771 1.5 ~

via freeaccess * Range ofvalues in parentheses. \0 For number of species and specimens studied, see Table IA. 00 0 IAWA Bulletin n.s., Vol. 1 (4), 1980 175

The numerical data in this study are much Vochysia. Fibers often septate, although this is more comprehensive and detailed for many not common in all specimens and rather rare in anatomical features. Vochysia. Fibers occupy an average 45 percent Tables lA, IB, and lC summarize the anato• of specimen volume and for individual speci• mical data by genus. The average values are mens fiber volume ranges from 35 to 80 per• computed on the basis of each species having cent Dry-site species tend to have shorter equal weight rather than every specimen having fibers, I mm or less on the average. equal weight. The range values are the smallest and largest value encountered for any single Rays (Table IC) specimen. The descriptions follow the termi• Numerous x = 33 per mm 2 but very variable; nology used by Metcalfe and Chalk (1950) for range 7 to 70 per mm2. Rays 2-4 cells wide, lengths and sizes. mostly 3 in Erisma, Salvertia, and Qualea; 4-6, mostly 5, but sometimes up to 8-10 cells wide Grossfeatures (Figs. A-C) in Vochysia. Uniseriate in Erismadelphus. Tall• All woods diffuse porous. Growth zones est rays average 0.77 mm in height and seldom usually abseI'lt or indistinct, but sometimes exceed I mm in all genera except Vochysia present in species from dry habitats. Borders of where tallest rays average over 1 mm in height growth zones sometimes can be distinguished and very frequently exceed 2 to 3 mm. Rays due to a differential in number of vessels per heterocellular (Kribs type HA and B) in Erisma, mm2 (Figs. A and B) or some variation in the Vochysia, and Callisthene with up to 3 to 4 coloration of the fiber tissue. uniseriate marginal rows of square or upright Heartwood variable, generally ranging from cells, but usually only I row in Callisthene. pinkish to reddish brown, not always sharply Rays generally homocellular in Qualea and Sal• differentiated from the sapwood. Woods gener• vertia with some scattered rays sometimes ally lustrous. Texture moderately coarse. Grain having a single marginal row of square cells, es• straight to very roey. Without distinctive odor pecially in Qualea dinizii. or taste when dry. Average basic density is 0.55 Procumbent portion of the tallest multiseria• (34 Ib/ft3) and ranges from 0.26 to 0.74 (16 to tes averages 17 cells in Erisma, Qualea, Salver• 47 Ib/ft3); ranging from light and soft to heavy tia, and Callisthene, and can range up to 30 and hard (Record and Hess, 1943). cells; 90 for Qualea. In Vochysia the tallest multiseriate rays average 50 procumbent cells Vessel elements (Table IA) but can range up to 150-170 cells. Procumbent Generally solitary, some radial multiples of ray cell shape: the ratio of height/diameter 2-4 cells. Diameter: medium (125 JLm) in Cal• (h/d) iselongate (2.2) in Erismadelphus through listhene to large (250 JLm) in the other genera. ovoid (1.5) in the other genera trending toward Pore frequency 9-21/mm2 in Callisthene and circular (1.0) in the dry-site species. 0-12/mm2, usually 2/mm2 in the other genera; end-wall perforations simple, transverse to Parenchyma (Figs. A-D) slightly oblique. Mean vessel-element length Apotracheally banded in Erismadelphus, Eris• medium (510 JLm), range 140 to 1,277 JLm for ma, and Callisthene. Bands moderately long all genera. Intervascular pitting, alternate and and 4-6 cells wide, sometimes appearing to en• vestured (Fig. J), medium (8 JLm), range 3-16 velop the vessels, sometimes only touching on JLm. Vessel-ray pitting generally the same as the adaxial' side. Vessels surrounded by fibers intervessel pitting, but just slightly smaller; in have scanty or no paratracheal parenchyma. Erisma some ray-cell pitting isoblong and sim• Parenchyma predominantly paratracheal in ple with only very narrow borders, even on the Vochysia, Qualea, and Salvertia. Extremely verge of appearing somewhat scalariform. Tylo• variable: from vasicentric, to aliform, to con• ses may be present in any group although they fluent. are most common in Erisma. Good evidence from the data indicates a trend for shorter ves• Included phloem sei elements the drier the site, and a concomi• Found only in Erismadelphus and Erisma; tant association of narrower and more numer• small bundles scattered in the parenchyma ous vessels (frequency). zones, also found in some rays.

Fibers (Table IB) Crystals and siIica (Table IB, Figs. E-H) Libriform fibers of medium length 1,266 Crystals of calcium oxalate are fairly com• (600-2,400) JLm, thin- (1 JLm) to thick- 11 JLm) mon in all genera except Erismadelphus and walled (average 3.4 JLm) with few simple pits, rare in Vochysia. Crystals when present were although some slightly bordered pits found in more common in the axial parenchyma than in

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Fig. A-J. -- A: Growth zone boundaries in Qualea dinizii, x 5. - B: Growth zone boundariesin Callisthene, x 5. - C: Traumatic resin ducts, Qualea acuminata, x 5. - D: Parenchyma band on adaxial side of vessel, Erisma floribundum, x 14. - E: Crystals in axial parenchyma of Qualea rosea, x 50.- F: Silica bodies in procumbent ray cells in Qualea albiflora, x 180. - G and H: Silica bodies in ray parenchyma, SEM, x 450. - J: Vestured vessel wall pits, SEM, x 900.

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the ray tissue. When present in the rays they Parenchyma. Paratracheal scanty to absent, are more common in the marginal cells than some fragmentary vasicentric rings. Apotracheal the procumbent ray parenchyma (Fig. E). bands 4-6 cells wide on adaxial side. On the basis of microchemical analysis of Included phloem. Common axially, also ashed specimens some silica (Si02) was indica• found in rays. ted in all genera except Erismadelphus. Concen• Other. Prismatic crystals rare in rays; no sili- trations never exceeded 0.5 % on a dry-weight ca. basis in any genus. Silica was most plentiful in Qualea and seldom exceeded 0.05 % in the Erisma other genera. Under the microscope silica Sixteen species from the Amazon Basin and bodies (Figs. F, G & H) could be found in the the Guianas. Sapwood greyish, heartwood yel• procumbent ray cells: medium (4-6 pm) in lowish brown. Basic density of 0.41 (0.34- some Qualea specimens and small (1-3 pm) in 0.47). Diffuse porous; growth zones absent to a few specimens of Erisma. Vochysia. Salvertia. somewhat distinct. and Callisthene. Use of the microscope occa• Vessels. Mostly solitary, some radial multiples sionally revealed a few fine silica bodies in the 2 to 4, x = 2/mm2 (I to 5). Diameters x = 223 axial parenchyma in Erisma and Vochysia, but Jlm (125 to 300). Length x = 603 Jlm ± 77 only in proximity to the vessels. Silica bodies (225 to 950). Vessel-vessel pitting x = 10 Jlm were found in all three locations only in Calli• (8 to 13). Vessel-ray pitting, similar. Some• sthene: procumbent and marginal ray cells, and times elongated with simple margin and appear• axial parenchyma. In a good share of the speci• ing somewhat reticulate or scalarifonn. mens where silica was indicated by microchem• Fibers. Septate scanty. Thirty-five to 50 per• ical analysis, no silica could be discemed under cent by volume. Diameters x = 20 Jlm (16 to the microscope. 28). Wall thickness x = 3 Jlm (1.5 to 5.0). Length x = 1,422 Jlm ± 130 (1,000 to 2,000). Descriptions of the individual genera Rays. 45/mm2 (30 to 70), 27 % multiseriate I have adopted the following fonnat for ex• and 73 % iiniseriate. Rays heteroceUular, pro• pressing values. In the example, vessel element cumbent cells marginated by one or more rows length x = 677 pm ± 164 (224 to 1,127), ± 164 of square and upright cells. Uniseriates predom• is the average standard deviation for several inantly of upright cells. Height of taUest rays specimens, not standard deviation of the mean. x = 672 Jlm (375 to 1,300) and x = 19 (8 to This type of data will appear in the subsequent 31) cells tall. Ray width x = 41 Jlm (25 to 90) publication with the individual species descrip• and 3 (2 to 4) cells wide. Ray cell shape, h/d tions. The bracket values (222 to 1,127) are the ratio 1.3. minimum and maximum values encountered Parenchyma. Paratracheal parenchyma scan• for any specimen. Data are based on the 183 ty or in fragmentary rings around pores. Apo• species recognized by Stafleu. tracheal parenchyma banded in lang bands 2 to 6 cells wide, mostlyon the adaxial side of Erismadelphus pores (Fig. D). Two species from west tropical Africa. Sap• Included phloem. Common axially and also wood greyish, heartwood pale reddish brown. found in rays. Basic density 0.57. Diffuse porous; growth Other. Cubic crystals absent in sapwood, zones absent to some;what distinct. common in heartwood and mostly in axial pa• Vessels. Mostly solitary, some radial groups renchyma chains of 2 to 4 crystals, seldom in of 2 to 3, x = 1.3/mm2 (0 to 3). Diameters x = rays. Silica predominantly in axial parenchyma. 284 Jlm (250 to 300). Length x = 677 Jlm ± Traumatic axial gum ducts present. 164 (224 to 1,127). Vessel-vessel pitting x = 10 Jlm (8 to 12). Vessel-ray pitting x = 7 Jlm Vochysia (StolO). Ninety-seven species from southem Mexico Fibers. Some septate. Forty-six percent of (Yucatan Peninsula) south through Central volume. Diameters x = 16.3 Jlm (14 to 19). America, from east of the Andes through Brazil Wall thickness x = 4 Jlm (2.5 to 5.0). Length and Guianas. Sapwood off white to greyish, x = 1,428 Jlm ± 185 (800 to 2,060). heartwood pinkish to reddish brown. Basic Rays. 38/mm2, all uniseriate. Rays hetero• density 0.51 (0.26 to 0.74). Diffuse porous; cellular, bordered by single row of square or growth zones (rings) absent to somewhat dis• upright cells. Height of tallest rays x = 586 pm tinct. (400 to 700), having 12 to 15 cells. Ray width Vessels. Mostly solitary, some radial groups x = 17 pm (14-19). Procum bent ray cell shape, 2 to 3, x = 2.5/mm2 (0 to 10). Largest vessel h/d ratio 2.2. diameters x = 269 Jlm (150 to 475). Length x

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= 515 pm ± 46 (150 to 1,277). Vessel-vessel Qualea pitting x = 8 pm (5 to 16). Vessel-ray pitting, Sixty species from Amazon Basin and Guia• similar. nas. Sapwood greyish to yellowish, heartwood Fibers. Rarely septate, some fibers with pinkish to reddish brown. Basic density 0.60 slightly bordered pitting. Fifty-seven percent (0.48 to 0.74). Diffuse porous; growth zones by volume. Diameters x = 19.7 pm (10 to 37). (rings) absent to distinct in some of the drier Wall thickness x = 3 pm (1.2 to 9.0). Length site species. x = 1,445 pm ± 140 (600-2,400). Vessels. Mostly solitary, some radial groups Rays. 22/mm2 (7 to 43), 27 % muItiseriate, 2 to 3, x = 4.6/mm2 (1 to 12). Largest vessel 73 % uniseriate. Rays heterocellular, procum• diameters x = 220 pm (100 to 350). Length bent cells marginated by one or more rows of x = 490pm± 76 (140 to 1,175). VesseJ-vessel square and upright cells. Uniseriates predomi• pitting x = 6 pm (3 to 12). Vessel-ray pitting, nantly of upright cells. Height of tallest rays similar. x: = 1,480 pm (650 to 4,100) and 48 (20 to Fibers. Some septate. Fifty-two percent by 170) cells tall. Ray width x = 80 pm (30 to volume, 25 to 80 percent. Diameters x = 16.3 275) and average 5 (3 to 10) cells wide. Ray pm (11 to 22). Wall thickness x = 4.2 pm (I to cell shape, h/d ratio 1.6. I I). Length x = 1,330 pm ± 132 (600 to Parenchyma. Paratracheal vasicentric to ali• 2,300). form and aliform confluent forming strong Rays. 22/mm2 (8 to 36), 70 % multiseriate, ban ding in a few cases. Apotracheal parenchy• 30 % uniseriate. Rays typically homocellular, ma in some groups diffuse-in-aggregates to procumbent cells marginated by one or two some short and narrow bands. rows of square or upright ceIIs in some rays of Other. No crystals found. Silica common in Q. albiflora, Q. dinizii, and Q. acuminata only. a few specimens, mostly in the ray tissue. Trau• Uniseriates with some procumbent, mostly matic axial gum ducts common. square and upright ceIIs. Height of taUest rays x = 678 pm (300 to 1,860) and x = 14(10 to 90) cells taU. Ray width x = 45 pm (20 to 110) Salvertia and 3 (2 to 4 [rare 1) ceIIs wide. Ray ceII shape, One species from Campos region of eastern h/d ratio I.3. BraziI. Sapwood yellowish, heartwood pinkish Parenchyma. Paratracheal vasicentric to ali• brown. Basic density 0.59. Diffuse porous; form and strongly confluent in some species. growth zones absent to somewhat distinct. Apotracheal parenchyma diffuse to diffuse-in• Vessels. Mostly solitary, some radial groups aggregates and forming some very short bands. of 2 to 3, x = 2.7 /mm2 (1 to 5). Largest vessel Other. Crystals common, either single or in diameters x = 234 pm (200 to 300). Length short to long chains in the axial parenchyma, x = 432 pm ± I 15 (193 to 768). Vessel -vessel few in the rays. Crystals found in fibers in Q. pitting x = 8 pm (6 to 10). Vessel-ray pitting, cymulosa. Silica when present not exceeding similar. 0.5 percent in any case; found in aII parenchy• Fibers. Commonly septate. Thirty~ight per• ma tissues. Traumatic axial gum ducts fairly cent by volume. Diameters x = 17 pm (13 to common. 20). WaU thickness x = 3 pm (2 to 4). Length x = 1,067 pm ± (710 to 1,430). Callisthene Rays. 37/mm2 (31 to 47), 43 % multiseriate, Eight species from southeast Brazil Campos 57 % uniseriate. Rays homocellular, cells typic• and Catingas areas, from northern Paraguay ally procumbent; some scattered rays with one and eastern Bolivia. Wood yeIIowish. Basic den• row of square or upright cells. Upright cells sity 0.59 (0.58 to 0.62). Diffuse porous; growth commonly two chambered if crystals are pres• zones (rings) indistinct to distinct. ent. Uniseriate rays of procumbent and upright Vessels. Mostly soJitary, up to 50 percent in or. all square and upright. Height of tallest rays radial groups 2 to 4, x = 14/mm2 (9 to 21). x = 730 pm (675 to 925) and x = 19 (16 to Largest vessel diameters x = 125 pm (100 to 24) cells tall. Ray width x = 90 pm (70 to 110) 130). Length x = 341 pm ± 62 (141 to 475). and 3 to 4, mostly 3 cells wide. Ray cell shape, Vessel-vessel pitting x = 6 pm (5 to 8). Ves• h/d ratio 1.2. sel-ray pitting x = 5 pm. Parenchyma. Paratracheal scanty. Apotrach• Fibers. Septate common. Thirty-eight per• eal banded, bands 2-6 cells wide. cent by volume. Diameters x = 18.8 pm (16 to Other. Crystals common, found only in the 20). Wall thickness 3.2 pm (2.5 to 4.0). Length axial parenchyma. Silica rare, found in ray tis• x = 906 pm ± 107 (663 to 1,350). sue when present. Tyloses and gum common in Rays. 35/mm2 (26 to 40),75 % muItiseriate, the vessels. 25 % uniseriate. Rays heterocellular with one

Downloaded from Brill.com10/01/2021 10:54:35PM via free access IA WABulIetin n.s., Vol. I (4), 1980 179 row of square or upright cells. Vniseriate rays Diagnostic value of the wood characters with procumbent and upright cells. Height of Only few of the wood anatomical characters tallest rays x = 480 /.Im (420 to 570) and x = are diagnostic at the genus level. 22 (15 to 30) cells tall. Ray width x = 50 /.Im The characteristics which were constant at (40 to 60) and 3 to 4, mostly 3 cells wide. Ray the genus level are the types and proportions of cell shape nearly circular, h/d ratio 1.1. rays. One genus is uniseriate, two are homo• Parenchyma. Paratracheal scanty or incom• geneous, and three heterogeneous. The propor• plete vasicentric rings. Apotracheal parenchy• tion of multiseriate-uniseriate rays varied from ma banded in long bands 2 to 5 cells wide pas• field to field over 10 random counting fields, sing the vessels on the adaxial side. more within a single specimen. than between Other. Crystals common in rays and also specimens. The average value calculated from short to medium chains in the axial parenchy• giving each counting field equal weight com• ma. Silica rare, found in rays and axial paren• pared to the average calculated giving each spe• chyma. cimen equal weight did not differ significantly.

Generic key to the woods of the Vochysiaceae Pits vestured, all heartwood reacts positive to Chrome-Azurol-S treatment. I. Rays all uniseriate ...... Erismadelphus I. Some rays multiseriate ...... 2 2. Multiseriate (M) rays predominant over uniseriate (V) (per mm 2). M:V ratio 3: I ...... 3 2. Vniseriate rays predominant or sometimes nearly equal to multiseriate. M:V ratio 1:3 or I: I .. 4 3. Rays heterocellular ...... Callisthene 3. Rays homocellular ...... Qualea 4. Rays predominantly homocellular, some scattered rays with I row of square or upright cells. M:V ratio average 2:3 ...... Salvertia 4. Rays heterocellular ...... 5 5. Largest rays 3 cells wide, usually I millimeter high and >25 cells tall. Crystals rare, no in- cluded phloem ...... Vochysia

References Baas, P. 1973. The wood anatomical range in cal spot-test for aluminum and its value in Ilex (Aquifoliaceae) and its ecological and wood identification. IA WA Bull. n.s. I: phylogenetic significance. Blumea 21 : 104-109. 193-258. Metcalfe, C.K. & L. Chalk. 1950. Anatomy of Carlquist, S. 1977. Ecological factors in wood the Dicotyledons. Vol. I. Clarendon Press, evaluation: A floristic approach. Am. J. Oxford, V.K. Bot. 64: 887-896. Quirk, J.T. 1975. Dot-grid integrating eyepiece: Chalk, L. & M.M. Chattaway. 1934. Measuring Two sampling techniques for estimating the length ofvessel members. Trop. Woods cell wall areas. Wood Sci. 8: 88-91. 40: 19-26. Record, S.J. & R.W. Hess. 1943. Timbers of Cronquist, A. 1968. The evolu tion and c1assifi• the New World. Yale Univ. Press, New cation of flowering . Houghton Mif• Haven, Conn. flin Co., Boston. Stafleu, F.A. 1948. A monograph of the Vochy• Daubenmire, R. 1978. geography. Acad. siaceae. I. Salvertia and V ochysia. Rec. Press, New York. Trav. Bot. Neerl. 41: 397-540. Also in Franklin, G.L. 1946. A rapid method of soften• Med. Bot. Mus. Vtr. 95: 397-540. ing wood for microtome sectioning. Trop. -- 1952, 1953, 1954. A monograph of the Woods 88: 35-36. Vochysiaceae. Il. Callisthene, IlI. Qualea, Huber, J. 1909. Mattas e madeiras amazonicas. IV. Erisma. Acta Bot. Neerl. I: 222-242, Bol. Mus. Goeldi: (Para') 6: 91-225. 2: 144-217,3: 459-480. Hutchinson, J. 1967. The genera of flowering Takhtajan, A. 1969. Flowering plants and dis• plants. II. Dicotyledons. Oxford, V.K. persal. Smithsonian Institution Press, City Kukachka, B.F. & R.B. Miller. 1980. A chemi- of Washington.

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