WOOD ANATOMYOF THE

NEOTROPICAL SAPOTACEAE

XX. MANILKARA

RESEARCH PAPER FPL 371

FOREST PRODUCTS LABORATORYFOREST SERVICE

U.S. DEPARTMENT OF AGRICULTUREMADISON, WIS.

JUNE 1981

Abstract

Consisting of about 32 neotropical species, the genus Manilkara has a wide-spread distribution in the Americas ranging from southeastern Brazil andnorthern Bolivia as far north as southern Florida. It is probably the mostimportant of the American genera, being the source of commercial timbers,fruits, and gums. Manilkara, as constituted here, consists of a genericcomplex that includes Achras, Muriaeanthe, Schaferodendron, Manilkariopsis,and Chiclea. This "complex" exhibits remarkable anatomical uniformity; thevariability exhibited between "genera" is no greater than that which may beencountered within a single species.

Preface

The Sapotaceae form an important part of the ecosystem in the neotropics; forexample, limited inventories made in the Amazon Basin indicate that thisfamily makes up about 25 percent of the standing timber volume there. Thiswould represent an astronomical volume of timber, but at present only a verysmall fraction is being utilized. Obviously, better information would helputilization-- especially if that information can result in clear identificationof species.

The Sapotaceae represent a well-marked and natural family, but the homogeneousnature of their floral characters makes generic identification extremelydifficult. This in turn is responsible for the extensive synonomy. Unfor-tunately, species continue to be named on the basis of flowering or fruitingmaterial alone, and this continues to add to the already confused state ofaffairs.

This paper on Manilkara is the twentieth in a series describing the anatomy ofthe secondary xylem of the neotropical Sapotaceae. The earlier papers, allby the same author and under the same general heading, include:

I.II.

III.IV.V.

VI.VII.

VIII.IX.X.

Bumelia--Res. Pap. FPL 325Mastichodendron--Res. Pap. FPL 326Dipholis--Res. Pap. FPL 327Achrouteria--Res. Pap. FPL 328Calocarpum--Res. Pap. FPL 329Chloroluma--Res. Pap. FPL 330Chrysophyllum--Res. Pap. FPL 331Diploon--Res. Pap. FPL 349Pseudoxythece--Res. Pap. FPL 350Micropholis--Res. Pap. FPL 351

XI.XII.

XIII.XIV.xv.

XVI.XVII.

XVIII.XIX.

Prieurella--Res. Pap. FPL 352Neoxythece--Res. Pap. FPL 353Podoluma--Res. Pap. FPL 354Elaeoluma--Res. Pap. FPL 358Sandwithiodoxa--Res. Pap. FPL 359Paralabatia--Res. Pap. FPL 360Gambeya--Res. Pap. FPL 361Gomphiluma--Res. Pap. FPL 362Chromolucuma--Res. Pap. FPL 363

Publication in this manner will afford interested anatomists and taxonomiststhe time to make known their opinions, and all such information is herebysolicited. At the termination of this series the data will be assembled intoa single comprehensive unit.

WOOD ANATOMY OF THE NEOTROPICAL SAPOTACEAE

XX. MANILKARA

By

B. F. Kukachka, Botanist1/

Forest Products Laboratory,2/ Forest ServiceU.S. Department of Agriculture

----

Introduction

The genus Manilkara was described by Adanson in 1763 but was generally reducedto the earlier described (1753) Mimusops of Linneaus. Revived by Dubard in1915, the first species assigned to Manilkara was the Asiatic speciesM . kauki (L.) Dub., and this has generally been considered to be the type.Much has been written regarding the relative merits of Mimusops versusManilkara, but it is now generally conceded that the American species onceattributed to Mimusops now properly belong to Manilkara which is pantropicalin distribution. The genus Mimusops is now restricted to Africa and Asia.

The nomenclatural situation with regard to Achras, now usually included inManilkara, is beyond the scope of this paper and the reader is referred for

details to Gilly (7), Monachino (10), and others.3/ The genus Muriea, oncerepresented by the single American species Muriea albescens (Griseb.) Hartogex Baill., is now restricted to south and east Africa, and the M. albescens hasbeen transferred to Murieanthe albescens (Griseb.) Aubr. by Aubréville (1) andaccepted by Baehni (2). However, Cronquist (3) made the new combinationManilkara albescens (Griseb.) Cronq. In 1942 Gilly (6) described two speciesin his new genus Shaferodendron, based on Mimusops mayarensis Ekm. ex Urb.,and both were placed in synonomy under Manilkara mayarensis (Ekm.) Cronq. (3).In his studies of the "Sapodilla-Nispero complex," Gilly (7) created four newsubgenera in the genus Manilkara, one of which, Manilkariopsis, was elevated togeneric status by Lundell (8) in 1975. The latest addition to this alreadyvery complex "complex" was the description of the new genus Chiclea by Lundell(9) in 1976. For the latter, two species are described as new, and Manilkarastaminodella Gilly is reduced to synonomy under Chiclea staminodella (Gilly)Lundell.

1/ Pioneer Research Unit, Forest Products Laboratory.

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

3/ Underlined numbers in parentheses refer to literature cited at the endof this report.

From the anatomical point of view, the genus Manilkara (including the afore-mentioned segregate genera) is remarkably uniform, and the observed differencesappear to be only quantitative. The anatomical differences between thesegregate genera appear to be no greater than those existing between thedifferent species of Manilkara. Cronquist (3) observes, "Our species ofManilkara are in general separated by strong floral characters, although theymay be vegetatively very similar. Sterile specimens may be difficult or evenimpossible to determine with certainty, even after one is familiar with theentities involved, except as vegetatively similar species are eliminated bygeographic considerations." To this must be added the observations of Ducke(5), "The greatest difficulty in studying species of Manilkara on herbariumspecimens is the high variability in size and shape of the leaves which oftenappear successively on the same tree in two very different season-forms. Atthe principal flowering period, usually at the end of the rainy season, branch-lets densely covered with flowers have comparatively small and stiff leaves,often with rounded apex. At all other seasons, however, on the same tree few-flowered branchlets are sometimes observed with leaves similar to those of thesterile branches, namely comparatively large and thin and often a littleacuminate at the apex."

Record (11) provided general and anatomical descriptions of the wood of Achrasand Manilkara, which are practically identical, and at the time did not mentionthe great similarity in the anatomy of the "two" genera. The occurrence ofvascular tracheids was not mentioned for Manilkara, but in the general familydescription their occurrence was noted for Bumelia, Henoonia (Solanaceae), andParalabatia.

Cronquist (3) cites 13 species of Manilkara native to Florida, the Caribbean,Mexico, and-Central America. Three of these are known only from the typelocalities, and several others have rather restricted ranges. Monachino (10)listed 22 species for the South American area, three of which also occur inthe Caribbean and Central America.

Twenty-six named species were available for this study, but two of these(amazonica and huberi) have been reduced to synonomy by Monachino (10), andnitida was reduced to synonomy by Cronquist (3). In addition, four of Gilly'sspecies (catingae, froesii, negrosensis, and solimoesensis) apparently areherbarium names only, and their status is uncertain. For comparative purposesthe wood of the generic type (M. kauki) has been included here. Because ofthe large number of wood specimens examined (254) which came from all parts ofthe known range of Manilkara, it is safe to assume that the examination ofadditional species would not alter to any significant degree the alreadyobserved variability.

Manilkara is probably the most important genus of the neotropical Sapotaceae,providing timbers of commercial significance, delicious fruits, and thebalata gum and chicle.

-2-

Description

This study is based on the examination of 254 specimens representing theentire range of Manilkara in the neotropics. The wood of M. kauki wasincluded here for comparative purposes since it is the generic type.Twenty-six named species plus the southeast Asia kauki were availablefor this research and are given as follows (numbers in parentheses referto the number of wood specimens available for each species): albescens(Griseb.) Cronquist (13), amazonica Chev. (8), bidentata (A. DC.) Chev.(49), catingae Gilly (1), chicle (Pittier) Gilly (11), elata (Fr. Allem.)Monachino (9), excelsa (Ducke) Standley (1), floribunda (Mart.) Dubard(4), froesii Gilly (1), huberi (Ducke) Chev. (13), inundata (Ducke)Ducke (1), jaimiqui (Wright) Dubard (15), kauki (L.) Dubard (2),longiciliata Ducke (1), longifolia (A. DC.) Dubard (2), meridionalisGilly (1), negrosensis Gilly (2), nitida Dubard (1), paraensis (Huber)Standley (1), rufula (Miq.) Lam (3), salzmannii (A. DC.) Lam (3),sideroxylon (Griseb.) Dubard (1), solimoesensis Gilly (6), staminodellaGilly (1), surinamensis (Miq.) Dubard (7), williamsii Standley (1),zapotilla (Jacq.) Gilly, and unassigned (48). Table 1 lists the specimensthat are backed or believed to be backed by herbarium material at someherbarium. Specimens from commercial or trade sources, even though named,have been eliminated from this listing because their specific identity willalways remain in doubt.

General: Heartwood a drab red-brown when freshly cut, soon changing toshades of brown or dark brown to almost black (superficially) in oldspecimens. Sapwood appreciably lighter in color and usually separatedfrom the heartwood by a transitional zone. Growth rings vague andindistinct. Wood hard, heavy, straight-grained (occasionally curlyfigure) with little if any luster. Specific gravity (at a moisturecontent of 6-7 pct) ranges from 0.73 to 1.23, the lowest values found injuvenile wood or specimens free of heartwoods. Species averages rangefrom 0.91 to 1.13 with an overall average of 1.03. Heartwood of albescensand jaimiqui frequently have an oily appearance and feel somewhat oily.

Anatomical:

Pores essentially diffuse in most species (figs. 1,3); tending toechelon arrangement (fig. 5) or a clustered-echelon arrangement(fig. 7). Pores commonly in radial multiples of 2 to 4 andoccasionally to 6; infrequently longer. Maximum pore diameter ofindividual specimens ranges from 89 to 197 µm; smallest in thegeneric type kauki (aver. 83 µm), largest in specimens of huberiand surinamensis (aver. 181 and 1.97 µm, respectively). Genericaverage is 122 µm.

-3-

Vessel member length averages 690 µm) for all species; shortest inwilliamsii (480 µm) and longest in nitida (890 µm). Theseparticular species were represented by a single specimen andhence the values cannot be considered representative. A morerepresentative range of averages may be attributed to jaimiqui(with the shortest average of 550 µm) and huberi (with thelongest aver. of 760 µm). Tyloses commonly thin-walled butfrequently thick-walled or sclerotic in the denser specimens.Very large crystals were observed frequently in the tyloses ofalbescens and jaimiqui, in most instances would be classified astwo-sized; lacking or very sporadic occurrence in the other speciesexamined. Intervessel pitting of 6 to 8 µm diameter was common toall specimens, a most unusual situation considering the number ofspecies and specimens examined. Perforation plates simple.

Axial parenchyma typically banded (figs. 1-8), variably spaced,frequently discontinuous; locally diffuse. The individual bandsirregularly 1 to 3 seriate. Many to all of the heartwood cellsfilled with brown organic deposits. Crystalliferous strands commonin the North American specimens and frequently two-sized, particu-larly in albescens and jaimiqui; in most South American specimensthe cells may contain 2 to 4 rhombic crystals or occur in shortmultiples thereof; crystals were found to be completely lackingin a few South American specimens (at least in the materialavailable for study). Silica and microcrystals not observed.

Wood rays 1 to 2 seriate and may be, in part, 3 to 4 seriate;heterocellular, vertical fusions frequent. The maximum bodyheight of the 1 to 4 seriate portion ranges from 79 to 552 µm;very inconsistent between and within species and of no diagnosticvalue. Vessel-ray pitting irregular in shape and size. Browndeposits very common. Rhombic crystals commonly present in thesquare or erect cells of albescens and jaimiqui and frequentlytwo-sized; occurring in vertical files of four crystals in thetall erect marginal cells and as a cluster of four crystals in thesquare marginals. Crystals generally much less abundant in theother species. Short, horizontal files of small crystals wereobserved in the tabular cells of several specimens. Lateral wallsof square and erect marginal cells conspicuously pitted, occasion-ally approaching the disjunct condition. Silica and microcrystalsnot observed.

Wood fibers thick walled; the fiber length averages for all specimensranging from 1.07 mm to 186 mm with anof 1.51 mm. Vascular tracheids common

For the species summary, see table 2.

overall species average(paratracheal).

Diagnostic features: Heartwood dull red-brown to brown and dark brown;majority of heartwood specimens (thoroughly dry) will sink in water;parenchyma banded; pores diffuse to echelon arrangement; axial parenchyma

-4-

with few to many crystals; silica absent. The only possible confusioncould be with Dipholis and Mastichodendron both of which have crystalstrands and are silica free but differ from Manilkara in that both havereticulate parenchyma.

Notes

Manilkara kauki, the generic type from southeast Asia, is practicallyidentical with specimens of jaimiqui from southern Florida. The growthrings are relatively distinct in kauki, and if this is a constant featureit would represent the only character that would separate these species.

During the sectioning of Manilkara wood blocks, it was observed thatsome of the blocks were apparently "overdone" when utilizing the time-temperature schedule for softening in 4 percent ethylenediamine, particularlythose that were dark colored and with specific gravities in excess of1.00. In these instances it was observed that the ethylenediaminesolution would become opaque after the sectioning blocks had been soakingfor a 24-hour period. It is well known that extractives contribute tothe specific gravity value as does also the ash content. To determineprecisely the amount of extractive and ash, two samples were selected ofthe same specific gravity. The specimens so tested were a dark heartwoodsample of albescens and a lighter colored heartwood sample of bidentata,each with a specific gravity of 1.07. Sawdust prepared from thesespecimens was extracted with a solution of 4 percent ethylenediamine andthe following results were obtained: bidentata produced an extractivecontent of 14.05 percent and albescens 22.10 percent. These results, ineffect, reduced the specific gravity of bidentata to 0.92 and that ofalbescens to 0.83. The total ash content determined for bidentata was0.63 percent and 4.29 percent for albescens, findings which furtherreduced the specific gravity values to 0.91 and 0.80 respectively.Subsequently, the heating time in ethylenediamine was reduced by approxi-mately one-half to compensate for the lowered or "true" specific gravity,producing a marked increase in the quality of the transverse sectionscut on the microtome.

-5-

Table l.--Herbarium-backed wood specimens of Manilkara

utilized in this study

Collector WoodSpecies and Source collection

number and numbers1/-----------------------------------------------------------------------------

albescens Fors 35Graves s.n.Scarff 3Scarff 9Scarff 18 ESchiffino 2Schiffino A 2

amazonica Bastos s.n. BrazilDucke 88 BrazilKrukoff 1440 BrazilOliveira 3121 BrazilPires et al. 51770 BrazilRodrigues and Coehlo 1936 Brazil

bidentata BAFOG 258BAFOG 263BAFOG 1240BAFOG 1272Beard (318) 511Bertin 3039Bertin s.n.BW 36 Uw2262Conservator of ForestsCox 2Englerth-GoytiaForest Department 54Forest Department 2936Goytia 161Goytia 162Goytia 172Holdridge 6208Kluge 55Maguire 23506Miller 1622Navy-Yale 163Navy-Yale 204Navy-Yale 205Navy-Yale 206Navy-Yale 242Pittier 2699Pittier 4318Pittier 11848Rose 18Smith, A.C. 3204Stahel 4U.S. Trop. Station 21Woodworth 206

Cuba SJR 13366Cuba SJR 4996Dominican Republic SJR 32186Dominican Republic SJR 32192Dominican Republic SJR 35359Dominican Republic SJR 35213Dominican Republic SJR 35136

French GuianaFrench GuianaFrench GuianaFrench GuianaSt. LuciaFrench GuianaFrench GuianaSurinamGuyanaPanamaPuerto RicoGuyanaGuyanaPuerto RicoPuerto RicoPuerto RicoPanamaPanamaGuyanaPuerto RicoPuerto RicoGuyanaGuyanaGuyanaSurinamPanamaPanamaVenezuelaVenezuelaGuyanaSurinamPuerto RicoVirgin Islands

RB 3086SJR 21347MAD 32865

MAD 21463INPA 829

SJR 50882SJR 50886SJR 32964MAD 32966SJR 49521SJR 12733SJR 6401MAD 32931SJR 32838SJR 6734MAD 20007SJR 5104MAD 4188MAD 23165MAD 23166MAD 23176MAD 24806SJR 7345MAD 11912MAD 20851SJR 45556SJR 45565SJR 45566SJR 45567SJR 45573MAD 19266MAD 5803SJR 7945SJR 2673

SJR 35858SJR 41084SJR 50524SJR 40173

(Page 1 of 4)

Table 1 .--Herbarium-backed wood specimens of Manilkara

utilized in this study--con.

Collector WoodSpecies and Source collection

numberand number1/

-----------------------------------------------------------------------------

catingae

chicle

elata

excelsa

floribunda

froesii

huberi

inundata

Froes 437

Conservator of Forests 229Dugand 535 (181)Dugand 772 (343)Dugand 1042 (490)Forgeson 67 AStevenson 54Stevenson 184Whitford and Stadtmiller 75Williams 9432Williams 9574

Filho and MagnaniFilho and RizziniFroes 834Froes 1079Servico Florestal

Krukoff 5496

Froes 1031Froes 1032Froes 1044Froes 1055

Froes 1068

Bastos s.n.Black 47-963Capucho 367Dahlgren 11Ducke 140Krukoff 1723Michigan 3926Michigan 3929Monteira da Costa

42

319Nagib Saddi M-16 (898)Oliveira 3107Silva 3374

Krukoff 4745

Brazil

BelizeColombiaColombiaColombiaPanamaBelizeBelizeGuatemalaMexicoMexico

BrazilBrazilBrazilBrazilBrazil

Brazil

BrazilBrazilBrazilBrazil

Brazil

BrazilBrazilBrazilBrazilBrazilBrazilBrazilBrazilBrazilBrazilBrazilBrazil

Brazil

A 27502

SJR 37257SJR 27081SJR 29620SJR 33781SJR 50969SJR 8822SJR 35106MAD 10848SJR 34855MAD 16146

RB 2957RB 5473A 27536A 28018

SJR 44736

MAD 18711

A 4561A 18223A 28001A 28006

A 28012

RB 3107SJR 45799SJR 21670SJR 16790SJR 22600MAD 32867SJR 21082SJR 21083MAD 23664RB 6243

MAD 18543

(Page 2 of 4)

Table 1 .--Herbarium-backed wood specimens of Manilkara

utilized in this study--con.

Collector WoodSpecies and Source collection

numberand number1/

-----------------------------------------------------------------------------

Caldwell 8748Conservator of Forests 66Fors 55Gill and Whitford 81Marts and Smith s.n.Stern 112Stern and Brizicky 239Stern and Brizicky 295Stern and Brizicky 503

Forest Department 4463Smith, A. C. 1450 (3373)

Azevedo 2025

Filho and Rizzini s.n.Froes 1080

Fors 105

Froes 833Krukoff 8691

Bernardi

Rosa, N. A. 1384

Froes 1042Froes 1052Froes 1056

Curran 17Souza 33

IICA J-14

Froes 247Froes 250Froes 251Froes 253Froes 276Krukoff 8628Krukoff 8643

Stevenson 5

FloridaJamaicaCubaCubaFloridaFloridaFloridaFloridaFlorida

SJR 49275SJR 47996SJR 13371SJR 9092SJR 45000SJR 49438SJR 51082SJR 51124SJR 51285

jaimiqui

JavaFiji

SJR 22417SJR 28229

kauki

SJR 47874Brazillongiciliata

longifolia BrazilBrazil

RB 5488A 28019

Cuba MAD 13806meridionalis

negrosensis BrazilBrazil

A 27535MAD 32851

Venezuela MAD 24286nitida

paraensis

rufula

Brazil

BrazilBrazilBrazil

A 24658A 28005A 28007

salzmannii BrazilBrazil

SJR 4687SJR 36087

Jamaica MAD 33912sideroxylon

solimoesensis A 26367A 27421A 27422A 27424A 27432

MAD 32852MAD 36348

BrazilBrazilBrazilBrazilBrazilBrazilBrazil

Belize SJR 8940staminodella

(Page 3 of 4)

Table 1. --Herbarium-backed wood specimens of Manilkara

utilized in this study--con.

Collector WoodSpecies and Source collection

numberand number1/

-----------------------------------------------------------------------------------

surinamensis Lao 56 and 90Williams 893Williams 2260Williams 5735Wurdack and Adderley 42737

williamsii Williams 11860

zapotilla Caldwell 8753Conservator of Forests 50Conservator of Forests s.n.Conservator of Forests s.n.Durland 35Fors 71Haufe et al. 34Kluge 27Maina 111MEXF 115Stern 34Stern 119Stern and Brizicky 309Stevenson s.n.Stevenson 183Steyermark 44636Whitford and Stadtmiller 86Williams 8260Wilson F-25

unassigned Breteler 5046Cuatrecasas 16707Froes 1091Froes 2000Irmay 124Irmay 131Krukoff 4681Krukoff 6292Krukoff 6624Shank 10Stevenson 8Williams 14489Williams 15640

PeruPeruPeruPeruVenezuela

Venezuela

FloridaBelizeBelizeBelizeDominican RepublicCubaHondurasMexicoMexicoMexicoFloridaFloridaFloridaBelizeBelizeGuatemalaGuatemalaMexicoFlorida

VenezuelaColombiaBrazilBrazilBoliviaBoliviaBrazilBrazilBrazilNicaraguaBelizeVenezuelaVenezuela

MAD 22227SJR 17455SJR 17853SJR 18221SJR 54116

MAD 32888

SJR 49280MAD 32882MAD 7398SJR 7401SJR 5055MAD 13781MAD 23103SJR 6192MAD 25228MAD 25220SJR 49393SJR 49445SJR 51134SJR 8828SJR 35105MAD 7659SJR 3745SJR 34543MAD 15968

SJR 55685SJR 43098

A 28023A 28024

SJR 47774SJR 47781MAD 18531MAD 12401MAD 12575SJR 46804SJR 8943SJR 41591SJR 42229

1/ A = Harvard University, Cambridge, Mass.; INPA = Instituto National dePesquisas da Amazonia, Manaus, Brazil; MAD = Forest Products Laboratory,Madison, Wis.; RB = Jardim Botanico do Rio de Janeiro, Brazil; SJR = SamuelJ. Record Memorial Collection, formerly at Yale University but now housed atMadison, Wis.

(Page 4 of 4)

Table

2.--Ma

nilk

ara spe

cies

su

mmar

y

Spec

ies

VML1

/Av

erag

eFL

Aver

age

MPAv

erag

eMB

SP GR

rang

era

nge

rang

era

nge

rang

eAverage SP

----

----

----

----

----

----

----

----

----

----

----

----

----

----

----

----

----

----

----

----

----

----

----

----

----

----

--

albe

scen

sam

azon

ica

bide

ntia

cati

ngae

chic

leel

ata

exce

lsa

flor

ibun

dafr

oesi

ihu

beri

inun

data

jaim

iqui

kauk

ilo

ngic

ilia

talo

ngif

olia

meri

dion

alis

negr

osen

sis

niti

dapa

raen

sis

rufu

lasa

lzma

nnii

side

roxy

lon

soli

moes

ensi

sst

amin

odel

lasu

rina

mens

iswi

llia

msii

zapo

till

aun

assi

gned

µm

460-

740

580-

800

600-

880

570-

820

620-

760

580-

700

680-

850

430-

660

620-

730

660-

730

650-

760

580-

710

700-

760

520-

870

580-

860

510-

870

600-

980

630

690

720

640

680

700

750

640

690

760

770

550

675

710

695

560

705

890

790

650

730

690

740

610

690

480

640

740

mm

1.07

-1.6

31.

27-1

.56

1.25

-1.8

0

1.52

-1.8

41.

35-1

.68

1.43

-1.5

5

1.43

-1.7

5

1.10

-1.4

11.

29-1

.47

1.41

-1.6

0

1.33

-1.6

3

1.50

-1.6

01.

54-1

.70

1.36

-1.6

7

1.30

-1.8

6

1.17

-1.7

71.

33-1

.73

1.42

1.42

1.50

1.42

1.66

1.54

1.64

1.50

1.52

1.61

1.76

1.22

1.38

1.36

1.50

1.32

1.48

1.74

1.67

1.55

1.61

1.66

1.50

1.55

1.53

1.29

1.49

1.57

µm

102-

142

87-1

5879

-158

102-

134

79-1

58

87-1

10

118-

181

87-1

3579-

87

110-

134

142-

142

87-1

3411

0-14

2

110-

165

95-1

97

79-1

7379

-165

119

129

125

102

119

131

142 97 110

149

165

111 83 102

122

126

142

110

110

110

131

142

135

126

130 95 117

130

µm

118-

355

134-

323

118-

434

102

197-

394

165-

433

394

252-

331

276

181-

418

433

134-

355

118-

236

220

236-

355

236

173-

236 95 220

213-

276

394-

489

276

158-

315

197

173-

552

315

95-3

9479

-512

0.96

-1.1

60.

94-1

.12

0.88

-1.1

9

0.96

-1.0

70.

89-1

.10

0.95

-1.0

6

0.98

-1.1

1

0.85

-1.0

90.

98-1

.12

0.96

-0.9

8

1.02

-1.0

7

0.97

-1.0

20.

96-1

.11

0.93

-1.1

2

0.90

-1.0

6

0.73

-1.2

30.

85-1

.16

1.07

131.

038

1.05

491.

091

1.01

110.

979

0.99

11.

014

0.91

11.

0513

1.04

10.

9915

1.05

21.

111

0.97

21.

091

1.04

21.

001

1.00

11.

003

1.02

31.

081

1.03

60.

971

0.98

71.

131

1.05

471.

0448

-------------------------------------------------------------------------------------------------------------

Genu

s43

0-98

069

01.

07-1

.86

1.51

79-1

9712

279

-552

0.73

-1.2

31.

03

1/

VML

= vessel member length;

FL =

fiber

length;

MP =

max

imum

tan

gent

ial

pore

dia

mete

r; M

B =

maxi

mum

height of 2

to 4

seriate

portion

of wood ray;

SP GR =

specific gravity

based

on weight and

volume at

approximately

6 to 7

pct; SP =

number of specimens

examined.

Literature Cited

1. Aubréville, Andre.1964. Sapotacées. Adansonia, Mémoire No. 1:1-157.

2. Baehni, Charles.1965. Mémoires sur les Sapotacées. III. Inventaire des genres.Boissiera II:1-262.

3. Cronquist, Arthur.1945. Studies in the Sapotaceae. IV. The North American species ofManilkara. Bull. Torrey Bot. Club 72(6):550-562.

4. Cronquist, Arthur.1946. Studies in the Sapotaceae. VI. Miscellaneous notes. Bull.Torrey Bot. Club 73(5):465-471.

5. Ducke, Adolpho.1942. New and noteworthy Sapotaceae of Brazilian Amazonia. Trop.Woods 59:38-44.

6. Gilly, Charles L.1942. Studies in the Sapotaceae: A new genus from eastern Cuba and anew species from Haiti. Trop. Woods 71:3-6.

7. Gilly, Charles L.1943. Studies in the Sapotaceae. II. The Sapodilla-Nispero complex.Trop. Woods 73:1-22.

8. Lundell, Cyrus L.1975. Studies of American plants. X. Sapotaceae. Wrightia5(6):167-196.

9. Lundell, Cyrus L.1976. Chiclea, a new genus of gum yielding trees in the Sapotaceae.Wrightia 5(7):221-225.

10. Monachino, J. V.1952. The South American species of Manilkara. Phytologia 4:94-

11. Record, Samuel J.1939. The American woods of the family Sapotaceae. Trop. Woods59:38-44.

Figure 1. --Manilkara rufula, pore and Figure 2. --M. rufula, detail ofparenchyma arrangement (Froes 1052) parenchyma variationsx 30. (Froes 1052) X 110.

Figure 3. --M. bidentata, pore and Figure 4. --M. bidentata, detail ofparenchyma arrangement parenchyma variations(Goytia 163) X 30. (Goytia 163) X 110.

Figure 5.--M. jaimiqui, pore and Figure 6.--M. jaimiqui, detail ofparenchyma arrangement (Marts parenchyma variations (Martsand Smith) X 30. and Smith) X 110.

Figure 7.--M. sp., pore andparenchyma arrangement(Froes 1091) X 30.

Figure 8.--M. sp., detail ofparenchyma variations(Froes 1091) X 110.

U.S. Forest Products Laboratory

Wood Anatomy of the Neotropical Sapotaceae: XX.Manilkara, By B. F. Kukachka, Madison, Wis., FPL

14 p. (USDA For. Serv. Res. Pap. FPL 371).

Consisting of about 32 neotropical species, the genusManilkara has a widespread distribution in the Americasranging from southeastern Brazil and northern Bolivia asfar north as southern Florida. It is probably the mostimportant of the American genera, being the source ofcommercial timbers, fruits, and gums. Manilkara, as con-stituted here, consists of a generic complex that includesAchras, Muriaeanthe, Schaferodendron, Manilkariopsis, andChiclea. This "complex" exhibits remarkable anatomicaluniformity; the variability exhibited between "genera" isno greater than that which may be encountered within asingle species.

U.S. GOVERNMENT PRINTING OFFICE: 1981-752-161