WOOD ANATOMY OF THE NEOTROPICAL SAPOTACEAE XXXI. POUTERIA RESEARCH PAPER FPL 419 FOREST PRODUCTS LABORATORY FOREST SERVICE U.S. DEPARTMENT OF AGRICULTURE MADISON, WIS. OCTOBER 1982
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WOOD ANATOMYOF THE
NEOTROPICAL SAPOTACEAEXXXI. POUTERIA
RESEARCH PAPER FPL 419
FOREST PRODUCTS LABORATORYFOREST SERVICE
U.S. DEPARTMENT OF AGRICULTUREMADISON, WIS.
OCTOBER 1982
A b s t r a c t
As c i r cumsc r ibed by Baehn i t he genus Pou te r i a c o n s i s t s o f a h e t e r o g e n e o u sa s s e m b l a g e o f o v e r 3 0 0 s p e c i e s p a n t r o p i c a l i n d i s t r i b u t i o n . The wood anatomya l s o r e f l e c t s t h e t r e m e n d o u s h e t e r o g e n e i t y a n d m a k e s i d e n t i f i c a t i o n b a s e d o nt h e w o o d a l l b u t i m p o s s i b l e . T h i s a u t h o r h a s e l e c t e d t o f o l l o w A u b r é v i l l e ' ss y s t e m t h a t r e d u c e s P o u t e r i a t o l e s s t h a n 5 0 s p e c i e s c o n f i n e d t o t h e A m e r i c a nt r o p i c s . A s a r e s u l t , t h e A m e r i c a n s p e c i e s o f P o u t e r i a compr i se a ve ry homoge-n o u s a n a t o m i c a l g r o u p t h a t i s r e a d i l y i d e n t i f i a b l e f r o m i t s w o o d a n a t o m y .
P re f ace
T h e S a p o t a c e a e f o r m a n i m p o r t a n t p a r t o f t h e e c o s y s t e m i n t h e n e o t r o p i c s ; f o re x a m p l e , l i m i t e d i n v e n t o r i e s m a d e i n t h e A m a z o n B a s i n i n d i c a t e t h a t t h i s f a m i l ymakes up about 25 percent of the s tanding t imber volume there . This wouldr e p r e s e n t a n a s t r o n o m i c a l v o l u m e o f t i m b e r b u t a t p r e s e n t o n l y a v e r y s m a l lf r a c t i o n i s b e i n g u t i l i z e d . O b v i o u s l y , b e t t e r i n f o r m a t i o n w o u l d h e l pu t i l i z a t i o n - - e s p e c i a l l y i f t h a t i n f o r m a t i o n c a n r e s u l t i n c l e a r i d e n t i f i c a t i o no f t h e s p e c i e s .
The Sapotaceae represent a wel l -marked and natural family but the homogeneousn a t u r e o f t h e i r f l o r a l c h a r a c t e r s m a k e s g e n e r i c i d e n t i f i c a t i o n e x t r e m e l y d i f f i -c u l t . T h i s i n t u r n i s r e s p o n s i b l e f o r t h e e x t e n s i v e s y n o n o m y . Unfo r tuna t e ly ,s p e c i e s c o n t i n u e t o b e n a m e d o n t h e b a s i s o f f l o w e r i n g o r f r u i t i n g m a t e r i a la l o n e a n d t h i s c o n t i n u e s t o a d d t o t h e a l r e a d y c o n f u s e d s t a t e o f a f f a i r s .
T h i s p a p e r o n P o u t e r i a i s t h e t h i r t y - f i r s t i n a s e r i e s d e s c r i b i n g t h e a n a t o m yof t he s econda ry xy l em o f t he neo t rop i ca l Sapo t aceae . T h e e a r l i e r p a p e r s , a l lby t he s ame au tho r and unde r t he s ame gene ra l head ing , i nc lude :
I .I I .
I I I .IV.V.
VI.V I I .
V I I I .IX.X.
XI.X I I .
X I I I .XIV.
XV.
Bumelia--Res. Pap. FPL 325 XVI. Paralabatia--Res. Pap. FPL 360Mastichodendron--Res. Pap. FPL 326 XVII. Gambeya--Res. Pap. FPL 361Diphol i s - -Res . Pap . FPL 327 XVII I . Gomphiluma--Res. Pap. FPL 362Achrouter ia - -Res . Pap . FPL 328 XIX. Chromolucuma--Res. Pap. FPL 363Calocarpum--Res. Pap. FPL 329 XX. Manilkara--Res. Pap. FPL 371Chloroluma--Res. Pap. FPL 330 XXI . Barylucuma--Res. Pap. FPL 372Chrysophyllum--Res. Pap. FPL 331 XXII . Pradosia--Res. Pap. FPL 373Diploon--Res. Pap. FPL 349 XXII I . Gayella--Res. Pap. FPL 374Pseudoxythece--Res. Pap. FPL 350 X X I V . Ecclinusa--Res. Pap. FPL 395Microphol i s - -Res . Pap . FPL 351 X X V . Ragala--Res. Pap FPL 396P r i e u r e l l a - - R e s . P a p . F P L 3 5 2 X X V I . Myrtiluma--Res. Pap. FPL 397Neoxythece--Res. Pap. FPL 353 X X V I I . Sarcaulus--Res. Pap. FPL 398Podoluma--Res. Pap. FPL 354 X X V I I I . Labatia--Res. Pap. FPL 416Elaeoluma--Res. Pap. FPL 358 X X I X . Eglerodendron--Res. Pap. FPL 417Sandwithiodoxa--Res. Pap. FPL 359 XXX. Pseudocladia--Res. Pap. FPL 418
P u b l i c a t i o n i n t h i s m a n n e r w i l l a f f o r d i n t e r e s t e d a n a t o m i s t s a n d t a x o n o m i s t st h e t i m e t o m a k e k n o w n t h e i r o p i n i o n s a n d a l l s u c h i n f o r m a t i o n i s h e r e b ys o l i c i t e d . A t t h e t e r m i n a t i o n o f t h i s s e r i e s t h e d a t a w i l l b e a s s e m b l e d i n t oa comprehensive uni t .
WOOD ANATOMY OF THE NEOTROPICAL SAPOTACEAE
XXXI. POUTERIA
By
B . F . K u k a c h k a , B o t a n i s t1/
F o r e s t P r o d u c t s L a b o r a t o r y ,2 / F o r e s t S e r v i c eU.S. Depar tment of Agr icul ture
- - - -
I n t r o d u c t i o n
T h e g e n u s P o u t e r i a w a s d e s c r i b e d b y A u b l e t i n 1 7 7 5 a n d c o n s i s t e d o f t h e s i n g l especies known to him, Pouter ia guianensis f rom French Guiana. Unfo r tuna t e ly ,t h e f r u i t d e s c r i b e d b y A u b l e t i s k n o w n n o w t o b e l o n g t o S l o a n e a o f t h eElaeocarpaceae and, a s a r e s u l t o f t h i s e r r o r , t h e n u m b e r o f s p e c i e s w i t h i nthe genus remained very smal l . On the o ther hand, Lucuma descr ibed by Molinai n 1 7 8 2 d e v e l o p e d i n t o a r a t h e r l a r g e g e n u s d e s p i t e t h e f a c t t h a t M o l i n a ' so r i g i n a l d i a g n o s i s c o n t a i n e d t w o s p e c i e s b e l o n g i n g t o f a m i l i e s o t h e r t h a n t h eSapotaceae. Lucuma is now regarded as a "nomen nudum" and the major i ty of i t sspecies have been reduced to synonyms of Pouter ia .
The nomenc l a tu r a l h i s t o ry o f t he Pou t e r i a -Lucuma c o m p l e x i s r a t h e r i n t e r e s t i n ga n d , a l t h o u g h i t h a s b e e n a r g u e d b y e v e r y s t u d e n t o f t h e f a m i l y , t h e p r e c i s e
c i r c u m s c r i p t i o n o f P o u t e r i a s t i l l r e m a i n s c o n t r o v e r s i a l . Baehni (4 )3/ i n h i s"Monograph of Pouteria" c i t e s 3 1 8 s p e c i e s d i s t r i b u t e d a m o n g 1 5 s e c t i o n s t h a ta r e p a n t r o p i c a l i n d i s t r i b u t i o n . F o r t h e N e o t r o p i c s h e c i t e s 1 7 8 s p e c i e s w i t ha n a d d i t i o n a l 2 3 v a r i e t i e s a n d f o r m s . I n h i s " I n v e n t o r y o f t h e G e n e r a " ( 5 ) heci tes for Pouter ia 24 gener ic synonyms and an addi t ional 23 probable synonyms.A l l i n a l l , a v a s t a n d h e t e r o g e n o u s a s s e m b l a g e o f p l a n t s .
A t t h e o t h e r e x t r e m e , A u b r é i l l e ( 1, 2) restricts Pouteria to a much smallern u m b e r o f s p e c i e s t h a t a r e c o n f i n e d t o t h e A m e r i c a n T r o p i c s a n d a r e c h a r a c t e r -i z e d b y t h e i r 4 - m e r o u s f l o w e r s , s u b u l a t e s t a m i n o d e s , s t a m e n s w i t h f i l a m e n t sin se r t ed nea r t he midd l e o f t he co ro l l a t ube , a f ou r - chambe red ova ry , andl e a v e s w i t h a d i s t i n c t n e t w o r k o f v e i n s a p p a r e n t f r o m b o t h s u r f a c e s .
F r o m t h e a n a t o m i c a l s t a n d p o i n t , B a e h n i ' s P o u t e r i a i s u n t e n a b l e , i m p r a c t i c a l ,a n d c o n s i s t s o f a n e x t r e m e l y h e t e r o g e n e o u s a s s e m b l a g e ; a n a n a t o m i c a l
1 / P i o n e e r R e s e a r c h U n i t , F o r e s t P r o d u c t s L a b o r a t o r y .
2 / Mainta ined a t Madison, Wis . , i n coope ra t i on w i th t he Un ive r s i t y o fWisconsin.
3 / U n d e r l i n e d n u m b e r s i n p a r e n t h e s e s r e f e r t o l i t e r a t u r e c i t e d a t t h ee n d o f t h i s r e p o r t .
descr ipt ion would near ly encompass a l l of the American Sapotaceae. A u b r é v i l l e ' sPou t e r i a f o r m s a c l o s e l y a l l i e d g r o u p i n g o f s p e c i e s c h a r a c t e r i z e d b y b a n d e dp a r e n c h y m a a n d t h e p o r e s i n d i f f u s e a r r a n g e m e n t a n d , f u r t h e r , c a n b e s e p a r a t e di n t o t w o g r o u p s o n t h e s i z e o f t h e i n t e r v e s s e l p i t t i n g .
Reco rd ( 1 3 ) p r o v i d e d a v e r y b r i e f a n d g e n e r a l i z e d d e s c r i p t i o n o f t h ePou te r i a -Lucuma c o m p l e x t h a t w a s s i m i l a r i n m a n y r e s p e c t s t o t h e o t h e r g e n e r ahe had de sc r ibed . H i s d e s c r i p t i o n s w o u l d n o t p e r m i t g e n e r i c i d e n t i f i c a t i o nu n d e r a n y c i r c u m s t a n c e b u t i t s h o u l d b e r e m e m b e r e d t h a t a t t h e t i m e h i sd e s c r i p t i o n s w e r e b e i n g p r e p a r e d , t he Ya le wood co l l ec t i on con ta ined a r e l a -t i ve ly sma l l number o f nomenc l a tu r a l l y con fused Sapo taceae . However , i t i sr a t h e r s u r p r i s i n g t h a t n o m e n t i o n w a s m a d e o f t h e p r e s e n c e o f v a s c u l a rt r a c h e i d s a n d t h e o c c u r r e n c e o f s i l i c a i n t h e w o o d r a y s . I n f a c t , s i l i c a i sn o t m e n t i o n e d i n a n y o f h i s d e s c r i p t i o n s a n d t h i s i s n o d o u b t d u e t o t h e " t h e ns t anda rd p rocedu re" o f soak ing s ec t i on ing b locks i n hyd ro f luo r i c a c id wh ichw a s q u i t e e f f e c t i v e i n r e m o v i n g s i l i c a a n d c r y s t a l l i n e c o n t e n t s .
K r i b s ( 1 0 ) d e s c r i b e d t w o s p e c i e s o f P o u t e r i a and , a s Reco rd , made no men t iono f t h e p r e s e n c e o f v a s c u l a r t r a c h e i d s a n d s i l i c a . H i s d e s c r i p t i o n o f P o u t e r i ac a r a b o b e n s i s i s n o w k n o w n t o b e f o r a s p e c i e s o f P r a d o s i a , a n d P o u t e r i agu i anens i s i s d e s c r i b e d a s h a v i n g i n t e r v e s s e l p i t t i n g 4 - 5 µ m i n d i a m e t e r w h e na c t u a l l y i t i s 8 - 1 0 µ m i n d i a m e t e r . I t i s e v i d e n t t h a t t h e P o u t e r i a g u i a n e n s i sd e s c r i b e d b y K r i b s b e l o n g s i n a n o t h e r g e n u s .
D e s c r i p t i o n
U p t o t h i s p o i n t , many spec i e s o f Pou t e r i a ( s e n s u B a e h n i ) h a v e b e e n r e f e r r e dt o o t h e r g e n e r a t h a t h a v e b e e n d e s c r i b e d i n p r e c e e d i n g m o n o g r a p h s o f t h i ss e r i e s . T h e r e m a i n i n g s p e c i e s , n o t a b l y t h o s e w h i c h A u b r é v i l l e ( 1,2) declareda s b e l o n g i n g t o t h e t r u e P o u t e r i a a n d a f e w n e w s p e c i e s t h a t h a d b e e n s u b s e -quen t l y de sc r ibed by h im , compr i s e a g roup o f c lo se ly r e l a t ed spec i e s hav ingin common the ana tomica l cha rac t e r i s t i c s o f t he gene r i c t ype Pou te r i ag u i a n e n s i s . T o t h i s b a s i c g r o u p h a v e b e e n a d d e d s p e c i e s o f P o u t e r i a t h a t o nt h e b a s i s o f t h e i r a n a t o m i c a l a n d p h y s i c a l c h a r a c t e r i s t i c s b e l o n g h e r e .Pou t e r i a i s h e r e i n d e s c r i b e d o n t h e b a s i s o f 2 9 n a m e d s p e c i e s r e p r e s e n t e d b y111 wood specimens and 21 unassigned specimens. R e f e r t o t a b l e 1 f o r p e r t i n e n td a t a r e l a t i v e t o e a c h s p e c i m e n . T h e l a s t p o r t i o n o f t a b l e 1 l i s t s s p e c i m e n so f n a m e d s p e c i e s o f P o u t e r i a t h a t h a v e b e e n e x c l u d e d f o r v a r i o u s r e a s o n s ( s e eN o t e s f o r d e t a i l s ) .
General : Sapwood light brown to brown; h e a r t w o o d a p p a r e n t l y l a t e i n f o r m a t i o nand, when present , i s usual ly dark brown or reddish brown and sharply demar-ca t ed f rom the l i gh t e r co lo r ed s apwood . Wood hard and heavy, wi th a specif icg r a v i t y r a n g e o f 0 . 6 0 ( b r a n c h w o o d ) t o 1 . 3 0 ( h e a r t w o o d ) ; t h e o v e r a l l a v e r a g ef o r a l l s p e c i m e n s w a s 0 . 9 1 a n d t h e m a j o r i t y o f t h e s e w e r e s a p w o o d o r i n - p a r thear twood. The va lue fo r a l l hea r twood spec imens a lways exceeds t he 1 .00 l eve l .G r o w t h r i n g s u s u a l l y i n d i s t i n c t o r a b s e n t e x c e p t i n t h e s o u t h e r n m o s t s p e c i e s( f i g . 3 ) w h e r e t h e y a r e f a i r l y d i s t i n c t . W o o d s t r a i g h t - g r a i n e d ; w i t h l i t t l eo r n o l u s t e r . F r o t h t e s t n e g a t i v e .
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Anatomical:
P o r e s i n d i f f u s e a r r a n g e m e n t ; o c c u r r i n g a s s o l i t a r y p o r e s a n d i n r a d i a lm u l t i p l e s o f 2 - 5 p o r e s o r i n f r e q u e n t l y l o n g e r w i t h a n o b s e r v e d m a x i -m u m o f 9 p o r e s i n s a l i c i f o l i a ( f i g s . 1 , 3 ) . Maximum tangential pored i a m e t e r o f i n d i v i d u a l s p e c i m e n s r a n g e s f r o m 7 9 µ m i n s a l i c i f o l i a t o244 µ m in an unassigned specimen; overal l average is 157 µ m.
Vessel member length averages range f rom 340 µ m in sal ic i fol ia to 920 µ min be l i z ens i s w i t h a n o v e r a l l a v e r a g e o f 6 6 0 µ m . I n t e r v e s s e l p i t t i n g6 µ m i n d i a m e t e r i n t h e s o u t h e r n s p e c i e s ( f r a g r a n s, gardnerana, andsal ic i fo l ia) and commonly 6-8 µ m or 8-10 µ m in the o ther species .P e r f o r a t i o n p l a t e s s i m p l e . T y l o s e s t h i n - w a l l e d , t h i c k - w a l l e d , a n dcommonly sclerot ic in the dense hear twood.
Ax ia l pa r enchyma typ i ca l l y banded ( f i g . 1 ) ; t h e i n d i v i d u a l b a n d s i r r e g u -l a r l y 1 - 3 s e r i a t e t o i n f r e q u e n t l y i n - p a r t 4 - 5 s e r i a t e . The bands( f i g . 3 ) a r e n o t d i s t i n c t b e c a u s e o f t h e m a g n i f i c a t i o n a n d a b u n d a n c eo f p o r e s b u t a r e q u i t e d i s t i n c t w h e n o b s e r v e d w i t h a h a n d l e n s o f15X. Pa renchyma ce l l s f r equen t ly w i th b rown depos i t s i n wh ich asma l l pa r t i c l e o f s i l i c a may be embedded . Rhombic and microcrysta lsl a c k i n g .
W o o d r a y s h e t e r o c e l l u l a r ; c o m m o n l y 1 - 2 s e r i a t e a n d o c c a s i o n a l l y 3 - s e r i a t ein some species ; m u l t i s e r i a t e p o r t i o n v e r y v a r i a b l e i n h e i g h t b u tg e n e r a l l y n o t e x c e e d i n g 5 0 0 µ m i n h e i g h t ( f i g s . 2 , 4 ) . Ray ce l l st a b u l a r , s q u a r e , t h e m a r g i n a l s u s u a l l y e r e c t a s v i e w e d f r o m r a d i a ls e c t i o n s . Brown deposits common although some cells may appear tob e d e v o i d o f a n y c o n t e n t s . V e s s e l - r a y p i t t i n g i r r e g u l a r i n s h a p eand s ize but commonly obovoid to l inear . P i t t i n g o n l a t e r a l w a l l so f s q u a r e a n d e r e c t c e l l s a b u n d a n t a n d d i s t i n c t . Rhombic crysta lsa n d m i c r o c r y s t a l s l a c k i n g b u t s i l i c a p a r t i c l e s a r e c o m m o n t o v e r yabundant . S i l i c a p a r t i c l e s s p h e r o i d a l , a t t a i n i n g d i a m e t e r s o f 2 0 µ mto 30 µ m and conf ined to those ce l ls wi th brown contents . S i l i c am o s t a b u n d a n t i n t h e t a b u l a r a n d s q u a r e c e l l s ; w h e n i t o c c u r s i n t h ee r e c t m a r g i n a l s i t i s u s u a l l y i n t h o s e c e l l s a d j a c e n t t o t h e a x i a lparenchyma.
Wood f i be r s t h i ck -wa l l ed ; f i b e r l e n g t h a v e r a g e s f o r a l l s p e c i m e n s r a n g i n gfrom 0.69 mm to 1 .75 mm with an overa l l average of 1 .30 mm. Vasculartracheids common.
S i l i c a con t en t de t e rmined by chemica l ana ly s i s r anged f rom 0 .03 pe r cen t( o v a t a ) t o 3 . 1 0 p e r c e n t ( n e g l e c t a ) w i th an ove ra l l ave rage o f0 . 7 8 p e r c e n t f o r t h e 9 5 s p e c i m e n s a n a l y z e d . S i l i c a ana ly se s we reperformed by Martin F. Wesolowski, Chemist, FPL.
D i a g n o s t i c f e a t u r e s : Wood l ight brown, dark brown, or red brown and most lyheavy to very heavy (hear twood specimens always s ink in water) . Parenchymabanded. P o r e s i n d i f f u s e a r r a n g e m e n t ; wood rays commonly 1-2(3) ser ia te ;u n i s e r i a t e r a y s f r e q u e n t l y a b u n d a n t ; t h e m u l t i s e r i a t e b o d y g e n e r a l l y l e s s t h a n
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500 µ m high. I n t e r - v e s s e l p i t t i n g 6 - 8 µ m o r 8 - 1 0 µ m i n d i a m e t e r . T y l o s e s ( i nhea r twood) commonly th i ck -wa l l ed o r s c l e ro t i c . S i l i c a pa r t i c l e s common in t hew o o d r a y s a n d g e n e r a l l y c o n f i n e d t o t h e t a b u l a r a n d s q u a r e c e l l s . N o t r e a d i l ycon fused w i th o the r gene ra w i th d i f fu se pa renchyma and s i l i c a i n t he wood r aysb e c a u s e i n t h e s e i n s t a n c e s t h e p o r e s a r e i n r a d i a l - e c h e l o n a r r a n g e m e n t o r t h ev e s s e l p i t t i n g i s 3 - 4 µ m o r 4 - 6 µ m i n d i a m e t e r . Eglerodendron might be con-fused wi th those Pouter ia wi th red-brown hear twood but in the former the woodr a y s a r e m o s t l y 2 - 4 s e r i a t e a n d t h e m u l t i s e r i a t e b o d y p o r t i o n i s 0 . 6 m m t o1.0 mm in height.
Notes
1. Lucuma amygdalina Standley (14) was descr ibed in 1935 on the bas is ofl e a v e s a n d s e e d s c o l l e c t e d b y D u r l a n d ( s . n . ) i n E l P e t e n , G u a t e m a l a . In 1942Baehn i ( 4 ) t r a n s f e r r e d t h i s s p e c i e s m a k i n g t h e n e w c o m b i n a t i o n P o u t e r i aamygda l ina ( S t a n d l e y ) B a e h n i a n d a d d e d a n i n c o m p l e t e f l o r a l d e s c r i p t i o n t h a ts t a t e d i n - p a r t " s e p a l s 5 , c o r o l l a u n k n o w n . " A p p a r e n t l y t h e f l o r a l d e s c r i p t i o nwas de r ived f rom Lunde l l LP 14 co l l e c t ed i n Be l i ze t ha t S t and l ey had de sc r ibedas Bumelia laur i fol ia but Baehni had reduced to a synonym of Pouter ia amygdal ina .I n 1 9 4 6 C r o n q u i s t ( 7 ) amp l i f i ed t he de sc r ip t i on by t he add i t i on o f e l even co l l ec -t i o n s f r o m B e l i z e a n d G u a t e m a l a a n d i n h i s f l o r a l d e s c r i p t i o n s t a t e d , i n - p a r t"sepals commonly 4 , somet imes 5 ."
Three wood specimens were avai lable for th is s tudy of which Stevenson 2 wasci ted by Cronquis t (7 ), Aguilar 11 was determined at the Field Museum, andL u n d e l l 5 w a s c i t e d b y C r o n q u i s t a s P o u t e r i a u n i l o c u l a r i s . Standley hadre f e r r ed Lunde l l 5 t o h i s new spec i e s S ide roxy lon meyer i a n d t h i s s p e c i e s , i nt u r n , w a s t r a n s f e r r e d t o P o u t e r i a lunde l l i i b y L o u i s W i l l i a m s . Our other woods p e c i m e n s o f P o u t e r i a un i locu l a r i s a r e ve ry d i f f e r en t f r om Lunde l l 5 andseeming ly be long t o F ranche t e l l a .
2 . Lucuma b e l i z e n s i s S t and l ey ( 1 4 ) was de sc r ibed f rom Kluge 41 co l l ec t ed i nB e l i z e a n d b a s e d o n s t e r i l e , l e a f y t w i g s . Baehn i ( 4 ) exc luded t h i s spec imenf r o m h i s P o u t e r i a a n d r e f e r r e d i t t o S ide roxy lon s p . Cronquis t (7 ) made thet r a n s f e r t o P o u t e r i a b e l i z e n s i s ( S t a n d l e y ) C r o n q u i s t a n d a d d e d a f l o r a ld e s c r i p t i o n b a s e d on S ide roxy lon lunde l l i i S t a n d l e y t h a t h e h a d r e d u c e d t osynonomy. However, h e q u a l i f i e d h i s d e s c r i p t i o n w i t h t h e s t a t e m e n t " T h e t y p e( K l u g e 4 1 ) i s c o m p l e t e l y s t e r i l e , a n d t h e r e i s s o m e p o s s i b i l i t y t h a t t h i s i sno t t he s ame spec i e s a s t he f l ower ing ma te r i a l t o wh ich S t and l ey gave t he nameS i d e r o x y l o n l u n d e l l i i."
The wood specimen from the type t ree (Kluge 41) and an associa ted specimenK l u g e s . n . d i f f e r f r o m t h e o t h e r t w o s p e c i m e n s t h a t h a v e a n a p p r e c i a b l y h i g h e rs i l i c a c o n t e n t a n d t h e w o o d r a y s a r e 1 - 3 s e r i a t e . Anatomical ly, a l l f o u r s p e c i -m e n s a r e a c c e p t a b l e a s P o u t e r i a b u t m a y r e p r e s e n t t w o s p e c i e s .
3 . The Pe ruv i an wood spec imens o f Pou te r i a c a i m i t o h a v e a s p e c i f i c g r a v i t yr a n g e o f 0 . 6 0 t o 0 . 7 8 , a v e r a g i n g 0 . 7 0 ; t h e o t h e r s p e c i m e n s a v e r a g e 0 . 9 5 .W i l l i a m s 4 8 0 2 , o r i g i n a l l y r e c e i v e d a s Lucuma h u a l l a g a e S t a n d l e y , h a s a s p e c i f i c
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g r a v i t y o f 0 . 9 4 a n d w a s c i t e d i n " F l o r a o f P e r u " ( 6 ) a s a synonym o f P . ca imi to .The wood specimens of A. C. Smith 2848 and 3245 have a s i l ica content of1 . 8 1 p e r c e n t a n d 2 . 3 3 p e r c e n t , r e s p e c t i v e l y , wh ich s eem r a the r h igh fo r t h i ss p e c i e s . The same may be said for Williams 171 and 472 from Peru.
4 . Pou t e r i a ca l i s t ophy l l a S t and l ey ( 1 5 ) w a s d e s c r i b e d f r o m C o o p e r 4 8 1 w h i c hc o n s i s t e d o f s t e r i l e l e a f y t w i g s a n d i s k n o w n o n l y f r o m t h e t y p e l o c a l i t y ,Province of Bocas de1 Toro, Panama. S t a n d l e y q u a l i f i e d h i s s p e c i e s b y t h ef o l l o w i n g s t a t e m e n t " B e c a u s e o f t h e a b s e n c e o f f l o w e r s a n d f r u i t s , t h e g e n e r i cp o s i t i o n o f t h i s t r e e i s , o f c o u r s e , u n c e r t a i n . I t s e e m s d e s i r a b l e t o g i v ei t a name , b e c a u s e i t i s c l e a r l y d i s t i n c t i n f o l i a g e c h a r a c t e r s f r o m a l l t h eS a p o t a c e a e h i t h e r t o r e p o r t e d f r o m C e n t r a l A m e r i c a . The t r e e i s we l l ma r ked byt h e b r o a d c o r i a c e o u s l e a v e s . d e n s e l y b r o w n - s e r i c e o u s b e n e a t h . " I n 1 9 4 2 B a e h n i( 4 ) t r a n s f e r r e d t h i s s p e c i e s t o P o u t e r i a c a l i s t o p h y l l a ( S t a n d l e y ) B a e h n i b u tp l a c e d i t i n h i s " s p e c i e s o f u n c e r t a i n s t a t u s . " T h i s s p e c i e s h a s b e e n r e t a i n e db y C r o n q u i s t ( 7 ) a n d P i l z ( 1 2 ) b u t i s e x c l u d e d h e r e o n t h e b a s i s o f i t s v e r ys m a l l i n t e r v e s s e l p i t t i n g ( 3 - 4 µ m ) a n d t e n t a t i v e l y p l a c e d i n L a b a t i a .
5 . Lucuma chir icana Standley (15) w a s d e s c r i b e d o n t h e b a s i s o f l e a v e s a n dseeds f rom the Panama col lec t ions of Cooper & Sla ter 254 ( type) and 230. Baehni( 4 ) c i t e d b o t h o f t h e s e n u m b e r s a n d m a d e t h e c o m b i n a t i o n P o u t e r i a ch i r i cana( S t a n d l e y ) B a e h n i b u t p l a c e d t h i s s p e c i e s i n h i s " s p e c i e s o f u n c e r t a i n s t a t u s . "Cronquis t (7 ) c i ted the Cooper & Sla ter number and added Cooper numbers 445and 457, also from Panama. H i s f l o r a l d e s c r i p t i o n w a s a p p a r e n t l y b a s e d o nCooper 457. A f i f t h s p e c i m e n , a l s o f r o m P a n a m a , F o r g e s o n 6 9 A , i s n o t c i t e di n t h e l i t e r a t u r e b u t t h e s h e e t w a s a n n o t a t e d b y C r o n q u i s t a s b e l o n g i n g h e r e .Very adequate wood specimens with accompanying herbar ium specimens are in theM a d i s o n c o l l e c t i o n s . The wood anatomy of Cooper & Slater 230, 254, andF o r g e s o n 6 9 A a r e v e r y s i m i l a r b u t t h e i r p l a c e i n P o u t e r i a a p p e a r s d o u b t f u l .Furthermore, Cooper 445 and 457 are very d i f ferent f rom each other and f romthe p r eceed ing t h r ee spec imens ; t hey do no t be long i n Pou te r i a and t he i rp r o p e r g e n e r i c d i s p o s i t i o n h a s n o t y e t b e e n d e t e r m i n e d . I t t h u s a p p e a r s t h a tt h e f l o r a l d e s c r i p t i o n p r o v i d e d b y C r o n q u i s t i s n o t a p p l i c a b l e . The wood ofC o o p e r 4 5 7 i s v e r y d i s t i n c t i v e b e c a u s e t h e s m a l l p o r e s a r e i n c l u s t e r e d - e c h e l o na r r a n g e m e n t ; t h e i n t e r v e s s e l p i t t i n g i s 6 - 8 µ m i n d i a m e t e r a n d t h e w o o d h a s av e r y h i g h s i l i c a c o n t e n t , 3 . 4 5 p e r c e n t ; d e f i n i t e l y n o t a P o u t e r i a . C o o p e r 4 4 5a l s o h a s s m a l l p o r e s b u t i n d i f f u s e a r r a n g e m e n t a n d t h e i n t e r v e s s e l p i t t i n g i s3 - 4 µ m i n d i a m e t e r ; s p e c i f i c g r a v i t y i s 0 . 5 7 ; n o t a P o u t e r i a .
6 . Pouter ia cooper i Cronquis t (7 ) was descr ibed f rom Cooper 499 and is knownon ly f rom the t ype l oca l i t y i n Panama . Th i s spec imen was o r ig ina l l y i den t i f i edb y S t a n d l e y a s C a l o c a r p u m vi r ide P i t t i e r b u t C r o n q u i s t d i d m a i n t a i n h i s n e ws p e c i e s i n t h e " C a l o c a r p u m g r o u p " u n d e r P o u t e r i a . C r o n q u i s t s t a t e d " T h i ss p e c i e s i s d i s t i n g u i s h e d f r o m o t h e r s o f t h e ' C a l o c a r p u m g r o u p " ' b u t i t s r e l a -t i v e l y c o a r s e a n d i r r e g u l a r v e n a t i o n , f l e s h y g l a b r o u s s e p a l s , a n d w o o d y f r u i t .Lundel l (11) make the new combinat ion Calocarpum cooper i (Cronquis t ) Lundel l .Ana tomica l l y , t he wood f rom the t ype t r ee i s ve ry d i f f e r en t f rom C a l o c a r p u mmammosum which is character ized by re t icula te parenchyma and the radia la r r a n g e m e n t o f t h e p o r e s ; i n t y p i c a l P o u t e r i a a n d i n t h i s s p e c i m e n t h ep a r e n c h y m a i s b a n d e d a n d t h e p o r e s a r e i n d i f f u s e a r r a n g e m e n t . I t would ber a the r d i f f i cu l t t o r e conc i l e t h i s spec imen w i th Ca loca rpum.
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7. Pou t e r i a c u a t r e c a s a s i i A u b r é v i l l e ( 3 ) was de sc r i bed f rom Pac i f i c Co lombiao n t h e b a s i s o f l e a v e s a n d f r u i t s o f C u a t r e c a s a s 1 6 3 9 6 ( h o l o t y p e ) a n d a naddi t ional spec imen, Cuat recasas 15749. Both numbers were col lected with woodspec imens and t he i r ana tomy i s t y p i c a l l y P o u t e r i a .
8 . Pou t e r i a dasys ty l a R i z z i n i a p p e a r s t o b e a d o u b t f u l Pou t e r i a o n t h e b a s i so f i t s s m a l l p o r e s w h i c h a r e a r r a n g e d i n r a d i a l f i l e s . The only specimena v a i l a b l e f o r t h i s s t u d y w a s F i l h o & R i z z i n i 5 5 7 c o l l e c t e d i n t h e R i o d eJaneiro area and is represented by wood specimen 6062 of the Jardim Botanico(Rio) and corresponding herbar ium sheet 155277.
9 . Pou t e r i a d e n t a t a ( P i e r r e ) G i l l y i s r e p r e s e n t e d b y F r o e s s p e c i m e n s 9 6 , 2 0 7 ,213, and 279. The wood specimens exhibi t considerable var ia t ion and may repre-s e n t t w o o r p e r h a p s t h r e e s p e c i e s o f P o u t e r i a .
10. Lucuma i z a b a l e n s i s S t and l ey ( 1 4 ) w a s d e s c r i b e d o n t h e b a s i s o f a s t e r i l ecollection from Guatemala made by Whitford & Stadtmiller No. 35. S t a n d l e yremarked, " T h e l e a v e s o f t h i s t r e e a r e s t r i k i n g l y s i m i l a r t o t h o s e o f L u c u m as c l e r o c a r p a . " Baehn i ( 4 ) m a d e t h e t r a n s f e r t o P o u t e r i a i z a b a l e n s i s (S t and ley )Baehni c i t ing the type specimen (Whit ford & Stadtmi l le r 35) and J . Record 8841.The S. J . Record herbar ium number is ac tual ly G-10 and the accompanying Yalewood col lec t ion number i s 8841. The Reco rd co l l e c t i on appa ren t l y p rov idedm a t e r i a l f o r B a e h n i ' s f l o r a l d e s c r i p t i o n . C r o n q u i s t ( 7 ) i den t i f i ed some add i -t i o n a l c o l l e c t i o n s w i t h t h i s s p e c i e s a n d i n c l u d e d E n g l e s i n g 4 6 , R e c o r d G - 1 0 ,and t he t ype , all of which have accompanying wood specimens. C r o n q u i s t ' sd e s c r i p t i o n s o f i z a b a l e n s i s a n d s c l e r o c a r p a a r e v e r y s i m i l a r a s i s a l s o t h ewood anatomy of the wood specimens avai lable for s tudy. Anatomical ly , the woods p e c i m e n s a r e n o t P o u t e r i a o r C a l o c a r p u m b u t m o s t l i k e U r b a n e l l a ( s e e a l s oNote 15) .
11. Pou t e r i a k rukov i i ( A . C . S m i t h ) B a e h n i ( 4 ) w a s o r i g i n a l l y d e s c r i b e d a sLucuma krukovi i A. C. Smith based on Krukoff 5700 col lected in Acre Terr i tory ,B r a z i l . T h e s m a l l i n t e r v e s s e l p i t t i n g , 4 - 5 µ m , e x c l u d e s t h i s s p e c i e s f r o mP o u t e r i a . F r o e s 2 6 2 r e c e i v e d a s P . k r u k o v i i h a s v e r y s m a l l i n t e r v e s s e l p i t t i n g ,3-4 µm, a n d u n d o u b t e d l y r e p r e s e n t s a n o t h e r s p e c i e s b u t n o t a P o u t e r i a .
12. Pou t e r i a l au r i fo l i a ( G o m e s ) B a e h n i , a c c o r d i n g t o t h e l i t e r a t u r e i s l i m i t e dt o s o u t h e a s t e r n B r a z i l . Al l of our specimens are Amazonian and hence mayr e p r e s e n t s o m e o t h e r s p e c i e s o f P o u t e r i a . The wood specimens represented byL u c i v a l 2 2 9 h a s v e r y s m a l l i n t e r v e s s e l p i t t i n g , 3 - 4 µ m , a n d h a s b e e n t e n t a -t i v e l y r e f e r r e d t o L a b a t i a .
13. Pou t e r i a me lanopoda Eyma ( 9 ) w a s d e s c r i b e d i n - p a r t " 5 s e p a l s a n d 5 p e t a l s ;t h e l e a v e s w i t h p e t i o l e s 1 - 1 / 2 t o 4 c m l o n g ; a n d t h e f r u i t u n k n o w n . " Thep e t i o l e l e n g t h g i v e n i n c e n t i m e t e r s ( c m ) i s a t y p o g r a p h i c a l e r r o r a n d s h o u l dr e a d a s m i l l i m e t e r s ( m m ) . T h i s e r r o r m a y h a v e b e e n r e s p o n s i b l e f o r t h e r a t h e rodd assemblage which has accumulated under this name. T h e f i r s t f o u r w o o dspec im e ns wh ich became ava i l ab l e f o r t h i s s t udy had been i den t i f i ed f roms t e r i l e h e r b a r i u m m a t e r i a l a n d a n a t o m i c a l l y b e l o n g e d t o f o u r d i f f e r e n t g e n e r a .T h e s p e c i m e n s l i s t e d i n t a b l e 1 m a y o r m a y n o t b e l o n g t o t h i s s p e c i e s b u ta p p e a r t o b e t y p i c a l P o u t e r i a .
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14. Pou t e r i a neg lec t a C r o n q u i s t ( 7 ) i s r ep r e sen t ed he r e by a s i ng l e spec imencol lec ted by Cooper s .n . (Oct . 1926) in Costa Rica and accompanied by woodspecimen SJR 9712. C r o n q u i s t i n d i c a t e d t h a t s t e r i l e s p e c i m e n s c o u l d b e c o n -fused wi th Calocarpum but the banded parenchyma and other fea tures make th isn u m b e r a t y p i c a l P o u t e r i a . T h e s i l i c a c o n t e n t o f t h i s s p e c i m e n i s 3 . 1 0 p e r c e n ta n d i s t h e h i g h e s t v a l u e o b t a i n e d f o r a n y s p e c i e s o f P o u t e r i a .
15. P o u t e r i a s a l i c i f o l i a ( S p r e n g . ) R a d l k . i s o n e o f t h e m o s t d i s t i n c t i v e o ft h e g e n u s , b e i n g c h a r a c t e r i z e d a s a s m a l l t r e e w i t h S a l i x-like leaves nativeto the temperate zone of South America . A n a t o m i c a l l y i t i s a l s o v e r y d i s t i n c ta n d r e a d i l y s e p a r a b l e b y i t s a b u n d a n t p o r e s i n d i f f u s e a r r a n g e m e n t w h i c h a t t a i na maximum diameter of 87 µ m (fig. 3 ) ; t h e v e s s e l s a n d p o r e s a r e t h e s h o r t e s tencoun te red in P o u t e r i a ; w o o d r a y s 1 - 3 s e r i a t e ; i n t e r v e s s e l l p i t t i n g 6 µ m i nd i a m e t e r ; a n d w i t h l o w s p e c i f i c g r a v i t y , a v e r a g i n g 0 . 6 3 . Baehn i ( 5 ) p l acedt h i s s p e c i e s i n h i s c o n g l o m e r a t e g e n u s R i c h a r d e l l a .
16. P o u t e r i a s c l e r o c a r p a ( P i t t i e r ) C r o n q u i s t ( 7 ) w a s o r i g i n a l l y d e s c r i b e d b yP i t t i e r a s Lucuma s c l e r o c a r p a . Th i s spec i e s i s known de f in i t e ly on ly f rom thet y p e c o l l e c t i o n , P i t t i e r 4 3 5 7 f r o m P a n a m a , and is accompanied by a wood speci-men. C r o n q u i s t m a d e t h e t r a n s f e r t o P o u t e r i a a n d r e t a i n e d t h i s s p e c i e s i n h i s"Calocarpum group." I n 1 9 7 6 L u n d e l l t r a n s f e r r e d t h i s s p e c i e s t o C a l o c a r p u ms c l ë r o c a r p u m ( P i t t i e r ) L u n d e l l ( 1 1 ) . The wood anatomy is nei ther Pouter ia orC a l o c a r p u m b u t i s v e r y s i m i l a r t o U r b a n e l l a .
17. P o u t e r i a t o r t a ( M a r t . ) R a d l k . i s s a i d t o b e v e r y v a r i a b l e i n s i z e f r o m " al i t t l e s h r u b t o a t r e e 2 5 m e t e r s h i g h " a n d a c c o r d i n g t o " F l o r a o f P e r u " ( 6 )includes Lucuma dolichophylla Standley represented by Ll . Wil l iams 886. Thef i v e s p e c i m e n s a v a i l a b l e f o r t h i s s t u d y e x h i b i t e n o u g h a n a t o m i c a l v a r i a t i o n t os u g g e s t t h a t t w o o r t h r e e s p e c i e s o f P o u t e r i a may have been inc luded he re .
18. P o u t e r i a t r i l o c u l a r i s C r o n q u i s t ( 8 ) w a s d e s c r i b e d o n t h e b a s i s o fAdams 2070 (accompanied by wood specimen W-11) from Acre Territory, Brazil.Also c i ted were Krukoff 5283, 5338, and 5531 a lso f rom Acre Terr i tory whichh a d p r e v i o u s l y b e e n d i s t r i b u t e d a s S ide roxy lon b o l i v i a n u m Rusby . To the abovec o l l e c t i o n s , the au thor has added Capucho 371, Froes 164 , Krukoff 5435, 8109,and M. G. Silva 3242 and 3362. A n a t o m i c a l l y t h i s s p e c i e s i s b r i e f l y d e s c r i b e da s f o l l o w s : wood l ight brown; larges t pores averaging 214 µ m in d iameter andin r ad i a l - eche lon a r r angemen t ; i n t e r v e s s e l p i t t i n g 6 - 8 µ m i n d i a m e t e r ; v e s s e lmember length averages 765 µ m; w o o d r a y s e s s e n t i a l l y u n i s e r i a t e ; p i t t i n g o fl a t e r a l w a l l s o f t h e e r e c t m a r g i n a l s p r o m i n e n t ; s i l i c a p a r t i c l e s o c c u r i n t h et a b u l a r , s q u a r e , a n d e r e c t m a r g i n a l s ; a n d w o o d w i t h a h i g h s i l i c a c o n t e n t ,a v e r a g e o f a l l s p e c i m e n s 2 . 0 8 p e r c e n t . A n a t o m i c a l l y t h i s s p e c i e s i s n o t aPou t e r i a a n d s u p e r f i c i a l l y w i t h h a n d l e n s e x a m i n a t i o n a p p r o a c h e s R a g a l a a n dS a r c a u l u s . T h e c o r r e c t p l a c e m e n t o f t h i s s p e c i e s r e m a i n s t o b e d e t e r m i n e d .
19. Pou t e r i a v e s t i t a Baehn i ( 4 ) w a s d e s c r i b e d f r o m K r u k o f f 4 7 4 3 ( t y p e ) a n dKrukoff 5013 f rom the s ta te of Amazonas , Braz i l . A n a t o m i c a l l y , t h i s s p e c i e sc o u l d b e r e g a r d e d a s a s m a l l e r - p o r e d v e r s i o n o f P . t r i l o c u l a r i s . T h e s i l i c ac o n t e n t o f t h e s e t w o s p e c i m e n s a v e r a g e d a n e x c e p t i o n a l l y h i g h 4 . 0 7 p e r c e n t .P . v e s t i t a a n d t r i l o c u l a r i s c o u l d b e t h e b a s i s f o r a n e w g e n u s .
- 7 -
L i t e r a t u r e C i t e d
1. Aubrév i l l e , Andre .1960. Notes sur les Sapotacées de L 'Afr ique Equator ia le , XV. Le Genre
P o u t e r i a A u b l e t ( s . s . ) n ' e x i s t e p a s e n A f r i q u e . Adansonia 16(3-4) :278-279.
2 . Aubrév i l l e , Andre .1961. N o t e s s u r d e s P o u t é r i é e s A m é r i c a i n e s . Adansonia 1 (2) :155-159 .
3 . Aubrév i l l e , Andre .1967. Sapotacées Nouvel les de la Côte Colombienne du Pacif ique.
Adansonia 7 (2) :141-143 .
4 . Baehni , C h a r l e s .1942. M i m o i r e s s u r l e s S a p o t a c é e s , I I . L e G e n r e P o u t e r i a . Candol lea
9:194-475.
5 . B a e h n i , C h a r l e s .1965. Mémoires sur les Sapotacées , I I I . I n v e n t a i r e d e s g e n r e s .
B o i s s i e r a 1 1 : 1 - 1 5 0 .
6 . Baehni , Char les and Luciano Bernardi .1970. F l o r a o f P e r u , S a p o t a c e a e . F i e l d M u s . N a t . H i s t . B o t . X I I I :
Par t V-A, No. 3 ; 151-170 .
7 . C r o n q u i s t , A r t h u r .1946. S t u d i e s i n t h e S a p o t a c e a e , I I . Su rvey o f t he Nor th Amer i can
Genera. Lloydia 9(4) :257-292 .
8 . C r o n q u i s t , A r t h u r .1946. S t u d i e s i n t h e S a p o t a c e a e , V I . M i s c e l l a n e o u s N o t e s . B u l l . T o r r e y
Bot . C lub 73(5) :469-470 .
9 . Eyma, P . J .1936. Notes on Guiana Sapotaceae . R e c . T r a v . B o t . N e e r l . 3 3 : 1 7 0 - 1 9 2 .
10. Kribs , David A.1968. Commercial foreign woods on the American market , 2nd ed. ,
Dover Publ ica t ions , New York, N.Y. pp . 150-151.
11. Lundel l , Cyrus L.1976. S t u d i e s o f A m e r i c a n P l a n t s , X I I . W r i g h t i a 5 ( 7 ) : 2 5 2 - 2 5 3 .
12. P i l z , G e o r g e E .1981. Sapotaceae of Panama. Ann. Missour i Bot . Gard . 68(1) :172-203 .
13. Record, Samuel J .1939. American woods of the family Sapotaceae. Trop. Woods 59:36-38.
- 8 -
14. S t a n d l e y , P a u l C .1925. An enumerat ion of the Sapotaceae of Centra l America . Trop. Woods
4 : 5 - 7 .
15. S t a n d l e y , P a u l C .1929. S t u d i e s o f A m e r i c a n p l a n t s , I I . F i e l d M u s . N a t . H i s t . B o t .
IV:8;252.
- 9 -
Tab
le 1
. --P
ou
teri
a:
Sel
ecte
d
par
amet
ers
of
spec
imen
s ex
amin
ed1
/
(sp
ecim
ens
mar
ked
(T
) ar
e fr
om
ty
pe
tree
)
Sp
ecie
sC
oll
ecto
r an
d n
um
ber
Sp.
gr.
SiP
MPD
V
ML
FL
R
MB
H
IVS
ourc
e
%µm
INT
ER
VE
SSE
L P
IT D
IAM
ET
ER
6 µ
m
Wil
liam
s,
Ll.
1
10
11
0.8
60.
87
D
126
µmmm
µmµm
740
1.54
2
095
6V
enez
uel
a
No
ver
ras
39
.62
.51
D
118
570
1.22
335
56
Arg
enti
na
Cu
rran
3
21
.62
1.04
D
95
600
1.2
22
355
6A
rgen
tin
a
Se
rvic
e F
ore
sta
l s.
n.
.61
.09
D87
Lin
dem
an &
de
Haa
s 34
94.6
1.3
2D
79P
atte
rso
n
81
35
.66
.34
D87
INT
ER
VE
SSE
L P
IT D
IAM
ET
ER
6-8
µm
430
.88
339
46
Uru
guay
420
.80
323
66
Bra
zil
340
.69
321
36
Arg
enti
na
Au
gil
ar 1
1.9
31
.90
D13
464
01
.30
223
6L
un
del
l5
.93
.54
D10
260
01
.29
215
8S
tev
enso
n2
.89
1.0
1D
134
580
1.25
356
8
Co
ns.
F
ore
sts
49.9
31
.58
D12
673
01
.46
Klu
ge
s.n
..9
6.1
3D
118
770
1.5
6K
luge
41
(T)
1.0
3.1
2.8
6D
126
920
1.7
5S
tev
enso
n 1
85
.95
D12
661
01
.33
3
276
2
339
2
291
3
315
be
no
isti
i A
ub
r.M
elin
on
108
1.0
0D
11
879
01
.37
207
9
6-8
6-8
6-8 6-8
6-8
6-8
6-8
6-8
Gua
tem
ala
Be
liz
eB
eli
ze
Be
liz
eB
eli
ze
Be
liz
eB
eli
ze
Fr.
G
uia
na
ca
rab
ob
en
sis
Pit
tie
r
fra
gra
ns
(Pie
rre
)D
ubar
d
gar
dn
eran
a (A
.DC
.)R
adlk
.
sali
cif
oli
a
(Sp
ren
g.)
Rad
lk.
amy
gd
alin
a
(Sta
nd
ley
)B
aeh
ni
be
liz
en
sis
(Sta
nd
ley
)C
ron
q.
(Pag
e 1
of
8)
Ta
bl
e
l.-
-Pou
teri
a:S
elec
ted
p
aram
eter
s o
f sp
ecim
ens
exam
ined
1/
(sp
ecim
ens
mar
ked
(T
) ar
e fr
om
ty
pe
tree
)--c
on
.
Sp
ecie
sC
oll
ecto
r an
d n
um
ber
Sp
. g
r.S
iP
MPD
V
ML
FL
R
MB
H
IVS
ou
rce
caim
ito
(R.&
P.)
Rad
lk.
his
pid
a E
ym
a
%µm
µm
m
mµm
INT
ER
VE
SSE
L P
IT D
IAM
ET
ER
6-8
pm
--co
n.
Fo
r.
Dep
. (G
uy
ana)
9
19
0.9
10
.18
D15
868
01
.30
207
9F
or.
D
ep.
(Gu
yan
a)
36
52
.95
.68
D14
265
01
.14
213
4K
ruk
off
6
37
71
.00
.59
D14
266
01
.26
213
4L
anje
uw &
Lin
dem
an 1
576
.88
.85
D11
873
01
.27
206
3M
agu
ire
et
al.
51
69
6.9
6.5
5D
9563
01
.32
204
2S
ilva
&Ja
ngou
x 33
21
.00
.68
D11
855
01
.05
215
8S
mit
h,
A.
C.
28
48
.92
1.8
1D
158
670
1.2
62
315
Sm
ith
, A
. C
. 3
24
5.9
72
.33
D11
066
01
.20
215
8W
illi
ams,
L
l.
17
1.7
81
.64
D14
270
01
.27
215
8W
illi
ams,
L
l.
47
2.7
01
.21
D14
248
00
.94
209
5W
illi
ams,
L
l.
17
89
.68
D16
553
01
.23
227
6W
illi
ams,
L
l.
17
92
.70
.18
D14
259
01
.09
219
7W
illi
ams,
L
l.
19
83
.60
D13
446
01
.14
231
5W
illi
ams,
L
l.
26
83
.64
D15
056
01
.09
216
5W
illi
ams,
L
l.
39
90
.77
.40
D16
573
01
.44
231
5W
illi
ams,
L
l.
47
17
.72
.58
D14
245
01
.03
227
6W
illi
ams,
L
l.
48
02
.94
.72
D15
861
01
.25
212
6
Cap
ucho
512
LB
B 1
0835
Old
enbe
rger
&N
orde
43
4S
ilv
a,
N.
T.
39
47
jen
ma
nii
(P
itti
er)
San
dw.
Fo
r.
Dep
. (G
uy
ana)
3
11
2
1.0
11
.05
1.0
2.9
0
1.0
9
.45
D13
467
01
.30
239
4.3
3D
126
800
1.5
53
512
1.0
4D
102
700
1.4
02
158
D13
450
0.9
33
347
.51
D
158
840
1.60
2 11
8
µm 6-8
6-8
6-8
6-8
6-8
6-8
6-8
6-8
6-8
6-8
6-8
6-8
6-8
6-8
6-8
6-8
6-8
6-8
6-8
6-8
6-8
6-8
Guy
ana
Guy
ana
Bra
zil
Sur
inam
eB
raz
ilB
raz
ilG
uyan
aG
uyan
aP
eru
Per
uP
eru
Per
uP
eru
Per
uP
eru
Per
uP
eru
Bra
zil
Sur
inam
eS
urin
ame
Bra
zil
Guy
ana
(Pag
e 2
of
8)
Tab
le 1
.--P
ou
teri
a:
Sel
ecte
d
par
amet
ers
of
spec
imen
s ex
amin
ed1
/
(sp
ecim
ens
mar
ked
(T
) ar
e fr
om
ty
pe
tree
)--c
on
.
Sp
ecie
sC
oll
ecto
r an
d n
um
ber
Sp.
gr.
SiP
MPD
V
ML
FL
R
MB
H
IVS
ou
rce
lau
rifo
lia
(G
om
es)
Bae
hn
i
ov
ata
A.
C.
Sm
ith
sch
rop
hy
lla
(A
uth
.?)
tric
ho
po
da
B
aeh
ni
coo
per
i C
ron
qu
ist
cu
atr
ec
asa
sii
Au
br.
dem
erar
ae
San
dw.
Fo
r.
Dep
. (G
uy
ana)
s.
n.
.94
ech
ino
carp
aW
. R
odri
gues
%µ
m
µm
INT
ER
VE
SSE
L P
IT D
IAM
ET
ER
6-8
pm
--co
n.
Fro
es 1
85
0.9
20
.21
D18
180
01.
31
2 39
4M
ag
uir
e e
t a
l.51
767
.99
.55
D12
663
01.
19
2 15
8O
liv
eira
2
96
6D
126
720
1.28
2
095
Oli
vei
ra
41
72
D11
864
01.
14
2 23
6
Kru
ko
ff 1
84
1 (
T)
.78
.03
D
134
610
1.38
3
433
Wil
liam
s,
Ll.
1
89
2.6
7D
15
8 50
0.9
3 2
197
Kru
ko
ff 6
34
4 (
T)
.93
.27
D
150
720
1.57
2
331
INT
ER
VE
SSE
L P
IT D
IAM
ET
ER
8-1
0 µ
m
Co
op
er
49
9
(T)
Cu
atre
casa
s 1
57
49
Cu
atre
casa
s 1
63
96
de
nta
ta
(Pie
rre
) G
illy
F
roe
s 9
6F
roe
s 2
07
Fro
es
21
3F
roe
s 2
79
Ro
dri
gu
es 8
77
9
.95
1.0
41
.02
.85
.84
.71
1.1
2
1.3
0
mm
µmµm
.24
D15
875
01
.53
206
3
.24
D14
266
01
.32
219
7.3
0D
181
680
1.4
23
473
.14
D19
753
01
.07
231
5
.96
D14
265
01
.34
218
9.9
2D
205
890
1.6
02
079
1.3
7D
158
700
1.4
72
189
.80
D11
066
01
.28
207
9
1.2
0D
181
710
1.2
32
434
6-8
6-8
6-8
6-8
6-8
6-8
6-8
Bra
zil
Bra
zil
Bra
zil
Bra
zil
Per
u
Bra
zil
8-1
0Pa
nam
a
8-1
0C
olom
bia
8-1
0C
olom
bia
8-1
0
8-1
08
-10
8-1
08
-10
Guy
ana
Bra
zil
Bra
zil
Bra
zil
Bra
zil
8-1
0B
raz
il
(Pag
e 3
of
8)
Tab
le 1
. --P
ou
teri
a:
Sel
ecte
d p
aram
eter
s o
f sp
ecim
ens
exam
ined
l/
(sp
ecim
ens
mar
ked
(T
) ar
e fr
om
ty
pe
tree
)--c
on
.
Sp
ecie
sC
oll
ecto
r an
d
nu
mb
erS
p.
gr.
Si
P M
PD
VM
L
FLR
M
BH
IV
So
urc
e
%µm
µm
m
mµ
m
µm
INT
ER
VE
SSE
L P
IT D
IAM
ET
ER
8-1
0 pm
--co
n.
gu
ian
ensi
s A
ub
l.
gu
tta
(Du
cke)
B
aeh
ni
lasi
oc
arp
a
(Ma
rt.)
Rad
lk.
mal
anop
oda
Em
ya
BA
FOG
129
31
.07
BB
S 10
681
.21
BW
27
1.0
8C
apuc
ho 3
941
.03
Cap
ucho
50
51
.03
Cap
ucho
576
1.0
6F
or.
Dep
. (G
uy
ana)
33
46
1.1
0F
roes
15
21
.05
Ko
ols
10
67
1.2
4M
agu
ire
et a
l. 5
18
31
1.1
3O
liv
eira
2
95
91
.00
D17
367
01
.13
231
51
.03
D16
565
01
.44
227
6D
165
710
1.3
22
394
.92
D16
570
01
.35
233
9.6
7D
165
540
1.1
02
394
1.0
6D
181
670
1.2
52
443
.38
D15
872
01
.68
217
3.8
8D
236
740
1.2
52
236
.50
D18
980
01
.52
235
5.6
7D
142
690
1.3
72
173
D19
771
01
.31
231
5
D10
278
01
.44
255
0.9
2D
220
570
1.2
02
197
D18
966
01
.30
239
4.2
3D
213
690
1.3
42
315
.51
D14
279
01
.49
247
3
8-1
0F
r. G
uia
na
8-1
0S
urin
ame
8-1
0S
urin
ame
8-1
0B
raz
il8
-10
Bra
zil
8-1
0B
raz
il8
-10
Guy
ana
8-1
0B
raz
il8
-10
Sur
inam
e8
-10
Bra
zil
8-1
0B
raz
il
Oli
vei
ra
29
98
.90
Plow
man
&R
osa
9420
1.0
0R
osa
, N
. A
. 1
12
2.9
5S
tah
el
91
.10
Sta
hel
2
72
1.0
8
2/6
-88
-10
8-1
08
-10
8-1
0
Bra
zil
Bra
zil
Bra
zil
Sur
inam
eS
urin
ame
Bla
ck
47
-10
92
1.0
71
.25
D18
172
01
.47
239
48
-10
B
raz
ilM
on
teir
o d
a C
ost
a 2
83
1.0
7.5
9D
158
520
1.3
22
433
8-1
0
Bra
zil
Kru
ko
ff
13
22
.96
.71
D21
370
01
.30
207
98
-10
B
raz
il
Cap
ucho
564
Fro
es 1
92
Fro
es
31
4
.91
.97
.95
1.62
D
19
7 59
0 1.
17
3 39
4
.81
D18
167
01
.24
226
0.9
3D
150
710
1.3
12
197
8B
raz
il
3/6-
8 B
razi
l8
-10
B
raz
il
(Pag
e 4
of
8)
Tab
le 1
. --P
ou
teri
a:
Sel
ecte
d
par
amet
ers
of
spec
imen
s ex
amin
ed1
/
(sp
ecim
ens
mar
ked
(T
) ar
e fr
om
ty
pe
tree
)--c
on
.
Sp
ecie
sC
oll
ecto
r an
d n
um
ber
Sp.
gr.
SiP
MPD
V
ML
FL
R
MB
H
IVS
ou
rce
%µm
µm
m
mµ
m
µm
INT
ER
VE
SSE
L P
IT D
IAM
ET
ER
8-1
0 µ
m--
con.
mal
ano
po
da
Ey
ma-
-co
nti
nu
ed Fro
es 3
22
0.9
1F
roes
4
08
1.0
6F
roes
1
07
1
(Bah
ia)
.96
Ro
sa,
N.
A.
1.0
0
0.6
7D
181
680
1.3
02
213
.27
D17
374
01
.31
219
7.4
5D
142
700
1.4
72
213
D13
463
01
.22
215
8
8-1
0B
raz
il8
-10
Bra
zil
8-1
0B
raz
il
Bra
zil
3/ 6
-8
.96
3.10
D
18
9 63
0 1.
28
2 43
38
-10
Co
sta
Ric
aC
oo
per
s.n
.n
egle
cta
Cro
nq
.
ob
iden
sis
Hu
ber
soli
mo
esen
sis
Au
br.
&P
ell
eg
r.
tem
are
(H.B
.K.)
A
ub
r.
tort
a
(Ma
rt.)
R
ad
lk.
Fro
es 5
76
.99
.36
D
165
610
1.26
2
055
8-1
0B
raz
il
Kru
ko
ff 8
61
8K
ruk
off
86
49
(T
).9
8.3
0D
142
640
1.3
32
197
8-1
0.9
6.7
1D
126
880
1.6
52
079
8-1
0B
raz
ilB
raz
il
.26
D
158
570
1.14
2
244
8-1
0W
illi
ams,
L
l.
14
30
0.7
3V
enez
uel
a
Kru
ko
ff
12
01
Kru
ko
ff
64
15
Kru
ko
ff
66
36
Wil
liam
s,
Ll.
8
86
Wil
liam
s,
Ll.
4
95
1
.96
1.5
0D
181
580
1.2
42
355
8-1
0.8
92
.72
D18
969
01
.48
218
98
-10
.94
1.1
3D
150
680
1.3
72
173
4/ 6
-8
.75
.31
D19
772
01.
252
118
4/ 6
-8
.71
1.1
4D
165
560
1.2
92
158
4/ 6
-8
Bra
zil
Bra
zil
Bra
zil
Per
u
Pe
ru
Sch
unke
46
26W
illi
ams,
L
l.
89
9.7
51
.07
D17
358
01
.15
210
28
-10
.29
D16
567
01
.25
224
48
-10
Per
uP
eru
wu
rdac
kii
Au
br.
(Pag
e 5
of
8)
Tab
le 1
.--P
ou
teri
a:
Sel
ecte
d
par
amet
ers
of
spec
imen
s ex
amin
ed1
/
(sp
ecim
ens
mar
ked
(T
) ar
e fr
om
ty
pe
tree
)--c
on
.
Sp
ecie
sC
oll
ecto
r an
d n
um
ber
Sp.
gr.
SiP
MPD
V
ML
FL
R
MB
H
IVS
ou
rce
%µm
µmm
mµm
µm
ca
list
op
hy
lla
(Sta
nd
ley
) B
aeh
ni
BW
47
Cu
rran
9
5C
urr
an
34
2C
urr
an
45
0F
roes
3
7F
roes
4
5F
roes
7
5F
roes
1
60
Fro
es 2
97
Fro
es 3
09
Fro
es
41
2II
CA
20
5K
ruk
off
1
08
11
Kru
ko
ff
10
92
4L
isb
oa
16
95
Lis
bo
a 1
72
2M
onte
ath
TS
14
Pit
tie
r 4
1R
osa
, N
. A
. 5
72
Sil
va,
N
. T
. 5
11
4T
S
(SJR
w
5534
)
UN
ASS
IGN
ED S
PEC
IMEN
S
1.0
0D
165
860
1.6
02
158
61
.02
0.3
4D
158
740
1.4
12
315
8-1
0.9
7.8
3D
165
730
1.3
82
260
8-1
01
.02
.38
D15
880
01
.53
226
08
-10
.96
1.3
1D
181
640
1.4
92
079
6-8
1.0
31
.06
D21
351
0.9
62
276
8-1
0.9
2.7
0D
158
690
1.3
53
552
8-1
0.9
3.7
0D
220
640
1.4
22
039
8-1
01
.02
.33
D17
358
01
.08
217
38
-10
1.2
0.2
0D
189
730
1.3
92
236
8-1
0.9
7.6
3D
173
590
1.1
62
236
8-1
0.8
2.6
5D
244
690
1.5
52
394
8-1
0.9
51
.16
D17
361
01
.23
223
68
-10
1.0
1.8
2D
158
590
1.1
62
095
8-1
0.9
2.9
2D
189
620
1.2
82
236
8-1
01
.02
.65
D17
366
01
.41
239
48
-10
.76
1.7
0D
165
650
1.3
82
095
6-8
.84
1.2
5D
197
620
1.2
12
276
8-1
0D
181
560
1.1
32
079
8-1
01
.00
.18
D16
581
01
.48
371
08
-10
.95
D18
173
01
.42
239
48
-10
Nam
ed
spec
ies
of
Po
ute
ria
ex
clu
ded
fr
om
d
escr
ipti
on
.
Co
op
er 4
81
(T
).9
3.9
3 D
11
8 73
0 1.
72
2 27
6 3-
4
Sur
inam
eB
raz
ilB
raz
ilB
raz
ilB
raz
ilB
raz
ilB
raz
ilB
raz
ilB
raz
ilB
raz
ilB
raz
ilPa
nam
aB
oli
via
Bo
liv
iaB
raz
ilB
raz
ilP
eru
Ven
ezu
ela
Bra
zil
Bra
zil
Bra
zil
Pan
ama
(Pag
e 6
of
8)
Tab
le1
.--P
ou
teri
a:
Sel
ecte
d
par
amet
ers
of
spec
imen
s ex
amin
ed1
/
(sp
ecim
ens
mar
ked
(T
) ar
e fr
om
ty
pe
tree
)--c
on
.
Sp
ecie
sC
oll
ecto
r an
d n
um
ber
Sp
. g
r.S
iP
MPD
V
ML
FL
R
MB
H
IVS
ou
rce
ch
iric
an
a
(Sta
nd
ley
)B
aeh
ni
cra
ssin
erv
ia
En
gle
r
Coo
per
230
0.8
1C
oo
per
25
4 (
T)
.76
Coo
per
445
.57
Coo
per
457
.73
Fo
rges
on
6
9A
.96
Fro
es 1
63
.98
da
syst
yla
R
izz
ini
Fil
ho&
Riz
zin
i 5
57
iza
ba
len
sis
(Sta
nd
ley
)B
aeh
ni
En
gle
sin
g
46
Rec
ord
G-1
0W
hit
ford
&S
tad
tmil
ler
35(T
)
kru
ko
vii
(A
. C
. S
mit
h)
Bae
hn
i
lau
rifo
lia
(G
om
es)
Bae
hn
i
mel
anop
oda
Eym
a
scle
roc
arp
a
(Pit
tie
r)C
ron
q.
%µm
µm
m
mµm
µm
Nam
ed
spec
ies
of
Po
ute
ria
ex
clu
ded
fr
om
d
escr
ipti
on
--co
n.
.81
.82
.88
Fro
es
21
6.8
5F
roes
26
2.8
5K
ruk
off
57
00
(T
).8
5
Lu
civ
al
22
9
Sch
ulz
7
45
3
Pit
tie
r 4
35
7
(T)
.90
.79
.77
1.2
0D
158
710
1.3
52
158
3.8
0D
134
660
1.5
53
291
1.7
8D
9559
01
.24
335
53
.45
R95
550
1.2
22
158
1.6
1D
158
700
1.5
22
118
1.3
9D
134
630
1.1
22
276
.36
R87
690
1.3
32
236
1.5
4D
134
710
1.5
43
355
2.1
0D
158
820
1.5
83
315
1.9
9D
134
710
1.5
23
473
.43
R11
067
01
.35
207
9.8
7R
126
770
1.3
82
276
.30
D10
268
01
.39
211
0
.65
D13
462
01
.25
226
0
.19
R16
590
02
.10
209
5
1.7
2D
142
620
1.1
82
134
5-6
5-6
3-4
6-8
5-6
4-5
6-8
Pana
ma
Pana
ma
Pana
ma
Pana
ma
Pan
ama
Bra
zil
Bra
zil
4-6
Nic
arag
ua
4-6
Gua
tem
ala
4-6
Gua
tem
ala
4-6
Bra
zil
3-4
Bra
zil
4-6
Bra
zil
3-4
Bra
zil
8-1
0S
urin
ame
4-5
Pana
ma
(Pag
e 7
of
8)
Tab
le
1. -
-Po
ute
ria
:S
elec
ted
p
aram
eter
s o
f sp
ecim
ens
exam
ined
1/
(sp
ecim
ens
mar
ked
(T
) ar
e fr
om
ty
pe
tree
)--c
on
.
Sp
ecie
sC
oll
ecto
r an
d n
um
ber
Sp.
gr.
SiP
MPD
V
ML
FL
R
MB
H
IVS
ou
rce
%µm
µm
m
mµ
m
µm
Nam
ed
spec
ies
of
Po
ute
ria
ex
clu
ded
fr
om
d
escr
ipti
on
--co
n.
tril
oc
ula
ris
Cro
nq
uis
tA
dam
s 20
70/W
-11
(T)
0.8
11
.68
R22
074
01
.43
207
96
-8C
apuc
ho 3
71.7
2.6
92
.57
R18
968
01
.39
211
86
-8F
roes
1
64
.75
R24
482
01
.56
207
98
-10
Kru
ko
ff
52
83
.77
2.2
5R
220
670
1.4
92
134
6-8
Kru
ko
ff
53
38
.81
1.5
3R
205
790
1.4
72
055
6-8
Kru
ko
ff
54
35
.74
2.8
6R
220
630
1.3
72
126
6-8
Kru
ko
ff
55
31
.75
.81
3.0
1R
236
810
1.3
22
158
6-8
Kru
ko
ff
81
09
1.9
6R
205
870
1.5
82
134
6-8
Sil
va,
M
. G
. 3
24
2R
228
790
1.2
92
079
6-8
Sil
va,
M
. G
. 3
36
2R
173
850
1.4
92
095
6-8
Bra
zil
Bra
zil
Bra
zil
Bra
zil
Bra
zil
Bra
zil
Bra
zil
Bra
zil
Bra
zil
Bra
zil
ve
stit
a
Ba
eh
ni
Kru
ko
ff 4
74
3 (
T)
.71
3.0
8R
118
840
1.2
91
000
6-8
Bra
zil
Kru
ko
ff 5
01
3.6
85
.20
R15
073
01
.15
207
96
-8B
raz
il
1/
Sp.
gr.
=sp
ec
ific
gra
vit
y;
Si
= s
ilic
a c
on
ten
t; p
= p
ore
arr
an
ge
me
nt,
D =
dif
fuse
, R
--ra
dia
l;M
PD
= m
axim
um p
ore
diam
eter
;V
ML
=
v
esse
l m
emb
er
len
gth
; F
L
=
fib
er
len
gth
;R
=
m
axim
um
ra
y
seri
atio
n;
IV
=
inte
r v
ess
el
pit
d
iam
ete
r.2
/ S
ee
no
te_
.3
/ S
ee n
ote
13
.4
/ S
ee
no
te
17
.
Err
ata,
Tab
le 1
: 'S
urin
ame'
sho
uld
be s
pelle
d 'S
urin
am.
Als
o, f
ootn
ote
2 sh
ould
app
ly t
o th
e en
tire
spec
imen
Oliv
iera
299
8, a
nd s
houl
d re
ad,
'Thi
s is
dub
ious
.
(Pag
e 8
of
8)
F i g u r e 1 . - -Pouteria guianensis, trans-ve r se s ec t i on showing l a rge po re sa n d m u l t i p l e s i n d i f f u s e a r r a n g e m e n tparenchyma narrow banded X 30( M a g u i r e e t a l . 51831).
F i g u r e 2 . - - S a m e a s f i g u r e 1 , t a n g e n t i a ls e c t i o n s h o w i n g c h a r a c t e r o f 1 a n d 2s e r i a t e r a y s X 1 1 0 .
F i g u r e 3 . - -Pouteria salicifolia, trans-verse sect ion showing abundance ofp o r e s o f s m a l l d i a m e t e r i n d i f f u s ea r r a n g e m e n t ; g r o w t h r i n g s p r e s e n tX 30 (Lindeman & de Haas 3494).
F i g u r e 4 . - - S a m e a s f i g u r e 3 , t a n g e n -t i a l s e c t i o n s h o w i n g r a y s m o s t l y2 - 3 s e r i a t e X 1 1 0 .
U.S. GOVERNMENT PRINTING OFFICE: 1982—654-254
U . S . F o r e s t P r o d u c t s L a b o r a t o r y
Wood anatomy of the neotropical Sapotaceae: XXXI.Pouter ia . by B. F . Kukachka. FPL.
18 p: (USDA For. Serv. Res. Pap. FPL 419).
As c i r cumsc r ibed by Baehn i t he genus Pou te r i a c o n s i s t so f a h e t e r o g e n e o u s a s s e m b l a g e o f o v e r 3 0 0 s p e c i e s p a n t r o p i -c a l i n d i s t r i b u t i o n . The wood anatomy also ref lects thet r emendous he t e rogene i t y and makes i den t i f i c a t i on ba sed ont h e w o o d a l l b u t i m p o s s i b l e . T h i s a u t h o r h a s e l e c t e d t of o l l o w A u b r é v i l l e ' s s y s t e m t h a t r e d u c e s P o u t e r i a t o l e s st h a n 5 0 s p e c i e s c o n f i n e d t o t h e A m e r i c a n t r o p i c s . As ar e s u l t , t he Amer i can spec i e s o f Pou t e r i a c o m p r i s e a v e r yh o m o g e n o u s a n a t o m i c a l g r o u p t h a t i s r e a d i l y i d e n t i f i a b l efrom its wood anatomy.
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