MANIPULUS R U B I A C E A R U M - V . A R E V I S I O N O F T H E G E N U S C a p i r o n a .
J o s e p h H. K i r k b r i d e , J r . (*)
SUMMARY
Tkz -ta.xon.omZc hJjstoty ofa the gcnuA CaptAona ÍÁ ph.eAe,nt.e.d. Complete, òynonymiçj,, ducJuptionò, common noma, dÁMtAtbottonÁ avid dL&uu>Aion& ahc gtvzn ^onCapAAona and iti, &)o òpecttò C. de^contxcanò and C. letophloea. The ge.nejü.c díòtntbution -L& the. Amazon valley and e.o6teAn GuZanaò and coincides with, that o$ the genua HQVQJX. C. dei>coKttcanb Ái known i>iom the. we&teAn and òouthcaòtetn Amazon vatle,y; C. letophloexx. U, h.cpo>vte.d $H0tA
the. eaòteAn Gtiianaò and the, noKthe.at>teAn Amazon valley. The. zone. o£ contact between the. two òpicleA ÍÁ along the. houthetin idge. 0 $ the. Amazon RiveA and the. loweA drainage of,
the Rio Tapajoz In PaAa, Efiaztl.,
(*) Departamento de Biologia Vegetal. Fundação Universidade de Brasília. Caixa Postal 153081, 70.919 Brasília, DF, Brasil. Present address: Plant Exploration & Taxonomy Laboratory,Bldg.265,BARC-East ,U .S .Department of Agriculture,Betsville, Maryland 20705 ,USA.
íaxonomic HistoryCapirona w a s p u b l i s h e d by S p r u c e b a s e d on a c o l l e c t i o n m a d e in the area of Tara -poto, Peru, in 1 8 5 5 . The g e n e r i c name is the common name used in that region for the type species. During 1 8 5 9 , S p r u c e w a s traveling in the central A n d i a n v a l l e y s of E c u a -dor (Spruce, 1970) w h e n C a p i r o n a w a s p u b l i s h e d in London ( S p r u c e , 1 8 5 9 ) . T h e r e f o r e he had recognized his c o l l e c t i o n as a new genus in the field a n d p r e p a r e d the d e s c r i p t i o n in Peru.
A year l a t e r , K a r s t e n ( i 8 6 0 ) presented the same g e n u s again as M o n a d e l p h a n t h u s based on a collection m a d e in the llanos of e a s t e r n Colombia.Hisgeneri c epithet d e s c r i b e s the basal connation of the f i l a m e n t s .
In volume I of Index K e w e n s i s ( J a c k s o n , 1 8 9 3 ) C a p i r o n a b o i v i n i a n a Bail Ion (1880) is listed. Bail Ion (1880) p u b l i s h e d the new g e n u s P l e u r o c o f f e a w i t h a single new species, P. b o i v i n i a n a . A f t e r c o m p a r i n g C a p i r o n a to P l e u r o c o f f e a , h e m e n t i o n s " C . b o i -viniana1 1, an o b v i o u s t y p o g r a p h i c e r r o r , w h i c h was p i c k e d up by the c o m p i l e r s of Index Kewensis as a new s p e c i e s of C a p i r o n a .
In 1912 w h i l e c o n d u c t i n g studies of areas in w h i c h B r a z i l - n u t s w e r e c o l l e c t e d near the Rio T r o m b e t a s , B r a z i l , A d o l p h o D u c k e d i s c o v e r e d the s e c o n d species of Capirona
(Ducke, I 9 1 3 ) . U n f o r t u n a t e l y he p r e s e n t e d the n e w species as a nomen n u d u m , C . ducke Huber i n Ducke ( 1 9 1 3 ) . A f t e r m a k i n g two additional c o l l e c t i o n s of the species in 191 a n d I 9 2 2 , he validly p u b l i s h e d it as C . h u b e r i a n a (Ducke, 1 9 2 2 ) .
In 1 9 1* 1 , Benoist collected the same species in French G u i a n a , but his col lectio w a s s t e r i l e . In 1920, M. W a c h e n h e i m , s u p e r v i s o r of the penal a d m i n i s t r a t i o n in Frenc G u i a n a , c o l l e c t e d the s p e c i e s w i t h f l o w e r s in the vicinity of the penal e n c a m p m e n t a G o d e b e r t . Benoist (1921) then used both c o l l e c t i o n s as the basis of his n e w specie C . l e i o p h l o e a w h i c h has p r i o r i t y o v e r that of D u c k e .
In the p r e p a r a t i o n of the R u b i a c e a e for the F l o r a o f S u r i n a m e ( B r e m e k a m p , 133^a) Bremekamp (l93^b) p r e s e n t e d the n e c e s s a r y t a x o n o m i c c h a n g e s a n d n o v e l t i e s in an extensi\e prior p u b l i c a t i o n . A m o n g the n o v e l t i e s is a new s p e c i e s of C a p i r o n a , C . surinamensis He felt that it showed a strong r e s e m b l a n c e to C . l e i o p h l o e a , but h e w a s u n a b l e t o b t a i n the original d e s c r i p t i o n of C . l e i o p h l o e a . If h e had been a b l e to see it, I am c o n f i d e n t that h e w o u l d n o t h a v e p u b l i s h e d his s p e c i e s w h i c h i s , in my o p i n i o n , a taxo o m i c s y n o n y m of C . l e i o p h l o e a .
In I 9 3 6 , Standley (1936a) p r e s e n t e d the n e w P e r u v i a n genus L o r e t o a and its single new s p e c i e s , L . p e r u v i a n a . It is p r o p h e t i c that S t a n d l e y s t a t e s , "The w r i t e r takes rw p a r t i c u l a r p r i d e in d e s c r i b i n g this tree as a n e w g e n u s . . . " This is w i t h o u t d o u b t the third n a m i n g of the g e n u s . He u n f o r t u n a t e l y had a s p e c i m e n w i t h o n l y a s i n g l e very small foliar calyx lobe w h i c h he a p p a r e n t l y o v e r l o o k e d .
In 1 9 ^ 0 , S t a n d l e y p u b l i s h e d a n e w species of C a p i r o n a , C . e r y t h r o x y l o n , based on a c o l l e c t i o n m a d e by R u i z and P a v o n in Peru in 1800. He w a s o v e r impressed by t h e foliar calyx lobe a n d failed to e x a m i n e t h e f l o w e r s in d e t a i l . If he h a d , he w o u l d h a v e disco vered that their a e s t i v a t i o n is v a l v a t e a n d the locules c o n t a i n m a n y s m a l 1 . h o r i z o n t a l ly a t t a c h e d o v u l e s . T h i s s p e c i e s is a m e m b e r of the g e n u s P o g o n o p u s K l o t z s c h .
T h e last s p e c i e s of C a p i r o n a , C . w u r d a c k i i , w a s p r e s e n t e d by S t e y e r m a r k ( 1 9 6 * 0 in the " B o t a n y of the G u a y a n a H i g h l a n d s , " H e d i s t i n g u i s h e d h i s s p e c i e s f r o m C . decorticans by its s h o r t e r , p i n k - c o l o r e d c o r o l l a s , m o r e broadly i n f u n d i b u 1 i f o r m limb of t h e c a r o l l a longer foliar calyx lobe, shorter s t y l e , rounded calyx lobes a n d s o m e w h a t s h o r t e r stip-u l e s . W h e n C . w stip-u r d a c k i i is c o m p a r e d w i t h s p e c i m e n s from the e n t i r e known range of C. d e c o r t i c a n s , it is o b v i o u s that these " d i f f e r e n t i a l " c h a r a c t e r i s t i c s o c c u r sporadically t h r o u g h o u t the latter s p e c i e s . C a p i r o n a w u r d a c k i i is a local v a r i a t i o n w h i c h hasbrought
these c h a r a c t e r i s t i c s t o g e t h e r , but it does not m e r i t , in m y o p i n i o n , formal taxonomic r e c o g n i t i o n as a s p e c i e s . C a p i r o n a is p r e s e n t e d h e r e as c o m p r i s i n g t w o s p e c i e s , C . d e c o r t i c a n s , t h e type, a n d C . l e i o p h l o e a . C a p i r o n a S p r u c e , J. P r o c . L i n n . S o c . , B o t . 3: 200. 1 8 5 9 . T y p e : C . d e c o r t i c a n s Spruce. (Figs. 2 , 3, k). M o n a d e l p h a n t h u s K a r s t e n , F l . c o l u m b . I: 6 7 , t . 3 3 - i860. T y p e : M . f l o r i d u s K a r s t e n .
L o r e t o a Standley, F i e l d M u s . N a t . H i s t . , Bot.Ser . 1 1 :221. 1836.Type: L . p e r u v i a n a
Trees without r a p h i d e s , g l a b r o u s or w i t h v a r i o u s o r g a n s s e r i c e o u s , the branches rete; stipules i ntrapet'iolar, p e r s i s t e n t ; leaves o p p o s i t e , p e t i o l a t e , the petiole t e -te and abaxially c a n a l i c u l a t e to w e a k l y s o , w i t h the blade varyng from m o d e r a t e to large in size, w i t h those s u b t e n d i n g inflorescences s i g n i f i c a n t l y smaller than those not subtending inflorescences; inflorescences terminal and a x i l l a r y reduced t h y r s e s , s e s s i l e or pedunculate, with the p e d u n c l e t e r e t e , w i t h o n e bract subtending e a c h b r a n c h ; f l o w e r s puHicellate to subpedicei 1 a t e , the pedicel t e r e t e , w i t h 1 or 2 bracts subtending each flower, the calyx tubular w i t h the tube c a m p a n u l a t e , w i t h 5 l o b e s , sometimes w i t h one lobe expanded into a large b r i g h t l y c o l o r e d foliar o r g a n w i t h a p s e u d o p e t i o l e and a p5eudoblade, with the p s e u d o b l a d e e l l i p t i c , a t t e n u a t e at the b a s e , o b t u s e at the a p e x , with k or 5 acrodromus p r i m a r y v e i n s , the corolla g l a b r o u s w i t h a sericeous band inter nally at the level of stamen s e p a r a t i o n , w i t h the a e s t i v a t i o n d e x t r o s e c o n t o r t e d , w i t h the tube plicate in bud w i t h 15 f o l d s , after e x p a n s i o n strongly infundibular, w i t h 5 lobes, the stamens w i t h the f i l a m e n t s c o n n a t e near their b a s e and separating from the corolla tube near its b a s e , w i t h the included a n t h e r s b a s i f i x e d , n a r r o w l y oblong, thedisk cup-shaped or rarely c i r c u l a r , the s t y l e t e r e t e , in the bottom of the disk or its cen ter, the stigma included b i l o b a t e , the 2 locules t u r b i n a t e , the p l a c e n t a narrowly t u r b i -nate, afixed to the c e n t e r of the s e p t u m , w i t h n u m e r o u s v e r t i c a l l y imbricate p e l t a t e ovules, the ovules e l l i p t i c or o b o v a t e ; fruits c a p s u l a r , t u r b i n a t e , s p e t i c i d a l , c r o w n e d with the persistent c a l y x , the n u m e r o u s seeds b i w i n g e d , n a r r o w l y e l l i p t i c , w i t h the apices of the w i n g s o b t u s e or i r r e g u l a r , the testa c e l l s rectangular to rarely e l l i p -tic, with the w a l l s thick, w i t h large ci rcular pits in the internal w a l1 .
Distribution
The genus C a p i r o n a is known to o c c u r in the A m a z o n v a l l e y , the llanos of Colombia -and the eastern G u i a n a s (Fig. 1 ) . It is a typical element of the flora of the Amazon
basin and defines its limits as does the genus H e v e a (Ducke & B l a c k , 1 9 5 3 , 195*0. C a p i -rona decorticans is found in the w e s t e r n end of the A m a z o n v a l l e y , both north and south of the Amazon R i v e r , reaching as far as the gallery forests in the llanos of C o l o m b i a where they meet the A n d e s . To the n o r t h , it has been c o l l e c t e d along the C a s i q u i a r e ca rial and once in the very uppermost reaches of the Rio O r i n o c o h e a w a t e r s . In the eastern end of the A m a z o n in the state of P a r a , B r a z i l , it is a found only south of the Amazon River.
The other species of C a p i r o n a , C . l e i o p h l o e a , is found in the e a s t e r n G u i a n a s , that is S u r i n a m e a n d F r e n c h G u i a n a , and e x t e n d s s o u t h w a r d into the state of P a r á , B r a -zil. It occurs b o t h north a n d south of the A m a z o n R i v e r , and south of the river,it has been more f r e q u e n t l y c o l l e c t e d in the lower d r a i n a g e of the Rio T a p a j o z .
T a x o n o m ϊ c Pos i t i o n
W h e n Spruce (1857) p r e s e n t e d the g e n u s C a p i r o n a , he p l a c e d it in f a m i l y Rubiaceae tribe C i n c h o n e a e , and it has remained there e v e r since ( H o o k e r , 1 8 7 3 ; B a i1 Ion,1881;Schu m a m m , 1 8 8 9 , 1 8 9 1 ; W e r n h a m , 1916; S t a n d l e y , 1930 , 1 9 3 6 b : Bremekamp, 1 9 3 4 a ; S t e y e r m a r k , 1971*).
It has b e e n m a i n t a i n e d in the t r i b e b a s e d o n its lack of r a p h i d e s , c o n t o r t e d corolla a e s t i v a t i o n , b i l o c u l a t e o v a r y , p l a c e n t a a f f i x e d to the c e n t e r of the s e p t u m , numerous v e r t i c a l l y imbricate o v u l e s , c a p s u l a r f r u i t , n u m e r o u s b i w i n g e d s e e d s , a n d testa cells w i t h n u m e r o u s large pits in the internal wall (Fig. 2 ) . Even in the n e w e s t c l a s s i f i c a
-tions (Verdcourt, 1 9 5 8 ; B r e m e k a m p , 1966) of the R u b i a c e a e , it will c o n t i n u e in tribe C i n c h o n e a e .
W i t h i n C i n c h o n e a e , C a p i r o n a has a l w a y s been a s s o c i a t e d w i t h t h o s e g e n e r a that have c o n t o r t e d a e s t i v a t i o n of the c o r o l l a and o n e c a l y x lobe s o m e t i m e s e x p a n d e d into a large c o l o r e d foliar o r g a n , S c h i z o c a l y x W e d d . and Ca 1 y c o p h y 1 1 u m D C . ( H o o k e r , 1 8 7 3 ; S c h u m a n n , I 8 9 I ; W e r n h a m , 1916; S t a n d l e y , 1 9 3 0 ) . It is easily d i s t i n g u i s h e d from these g e n e r a by its intrapetiolar s t i p u l e s , lobate c a l y x and c o n n a t e s t a m e n s .
Key to the S p e c i e s o f C a p i r o n a
1 . Trees w i t h all o r g a n s g l a b r o u s ; A m a z o n i a n C o l o m b i a , E c u a d o r , and P e r u , the uppermost h e a d w a t e r of the Rio O r i n o c o , V e n e z u e l a , and A m a z o n i a s , A c r e , R o n d ô n i a and s o u t h e r n P a r á , B r a z i l .
C . d e c o r t i c a n s 1 . Trees w i t h various o r g a n s densely to s p a r s e l y s e r i c e o u s or g l a b r a t e ;
S u r i n a m e , French G u i a n a and P a r á , B r a z i l .
C . leiophloea
C a p i r o n a d e c o r t i c a n s S p r u c e , J . P r o c . L i n n . S o c . , B o t . 3 : 2 0 0 . 1859. T y p e : S p r u c e ^ 2 0 2 , Oct I 8 5 5 , T a r a p o t o , San M a r t i n , Peru (holotype: Κ; isotypes; F, K, N Y , P ) . (Figs. 2 , 3 ) . M o n a d e l p h a n t h u s f l o r i d u s K a r s t e n . F l . c o l u m b . 1 : 6 7 , t. 3 3 - i860. T y p e ; Karsten s n , J i r a m e n e C a n o G i r a m e n a , L l a n o de San M a r t í n , M e t a , C o l o m b i a ( l e c t o t y p e : W ; isotype: L E ) .
L o r e t o a p e r u v i a n a S t a n d i . , F i e l d M u s , N a t . H i s t . , B o t . S e r , 1 1 : 2 2 2 . 1 9 3 6 , Type: Klug 2 0 2 2 , M a r - A p r 1 9 3 1 , F l o r i d a , Rio P u t u m a y o , at the m o u t h of the Rio Z u b í n e t a (holo-type: F) .
C a p i r o n a wurdackii S t e y e r m . , M e m . New Y o r k B o t . C a r d . 1 0 : 1 9 0 , 1964. T y p e : Wurdack & A d d e r l e y '•3383, C a n o D u q u i a p o , Rio C a s i q u i a r e , 2 km a b o v e S o l a n o , A m a z o n a s , V e n e z u e l a
(holotype: N Y ; isotypes: F, Κ, N Y , P, S P , U, V E N ) .
T r e e s 7 - 3 3 m tall w i t h a trunk d i a m . of 1 0 - 3 5 ( - 1 0 0 ) c m , g l a b r o u s ; stipules ovate or rarely e l l i p t i c , a c u t e t o n a r r o w l y so or rarely n a r r o w l y a c u m i n a t e at the a p e x ,
1 . 5 . 1 4 . 5 ( - 8 ) χ 0 . 6 - - 2 . 2 ( - 2 . 5 ) c m ; leaves subtending the inflorescences slightly smaller than those w h i c h do n o t , w i t h the p e t i o l e 1 - 3 cm long, w i t h the b l a d e e l l i p t i c , cuneate or a t t e n u a t e o r o b t u s e at the b a s e , a c u m i n a t e o r a c u t e a t the a p e x , 11 -^+1 x 6 - 2 ^ c m , 1 . 5 - 2
times longer than w i d e , w i t h ( 6 - ) 7 - 1 0 ( - 1 7 ) a r c u a t e s e c o n d a r y nerves on e a c h side of the midrib, the midrib and nerves s u b p r o m i n e n t a b o v e a n d p r o m i n e n t b e n e a t h ; inflorescences 7-20 χ 10-30 cm, w i t h 2 5 - 1 5 0 f l o w e r s , w i t h 1 - 6 flowers or rarely none w i t h an e n l a r g e d foliar calyx lobe, w i t h the p e d u n c l e 5 - 8 c m long, w i t h 2 - 3 pairs of lateral branches
3-10 cm long, w i t h o n e bract s u b t e n d i n g e a c h b r a n c h , the b r a c t s o v a t e or el 1 i p t i c , a c u t e or acuminate at the a p e x , 0. 6 - 3 χ 0 Λ - 1 c m ; flowers w i t h the pedicel 2 - 7 m m long,with I or 2 bracts subtending e a c h f l o w e r , the b r a c t s o v a t e , a c u t e at the apex,3-4.5x2-3mm,with the hypanthium t u r b i n a t e , 5 - 6 m m t a l l , 3 - 4 ( - 5 ) Bid in d i a m . , the calyx w i t h the interior sericeous, with the tube ( 1 - ) 3 - 5 m m t a l l , ( 5 - ) 7 - 1 1 m m in d i a m . at the o r i f i c e , w i t h the lobes very broadly a c u t e or rarely very broadly o b t u s e , 0. 5 - 3 x 3-5 m m , w i t h the foliar lobe red, with the p s e u d o p e t i o l e 2- 4 cm long, w i t h the p s e u d o b l a d e 4 - 8 . 5 x 2 . 5 - 5 c m , the corolla tube w h i t e to p i n k , 2 0- 3 5 m m long, w i t h the lobes w h i t e to pink,ovate to broadly so, obtuse at the a p e x , 11-15 x 8 - 1 5 m m , the stamens separating from the corolla tube 2-3 mm from the b a s e , c o n n a t e for 4 - 6 m m from their b a s e , w i t h the free portion 9-14 mm long, with the a n t h e r s 6 - 9 m m long, the d i s k circular or c u p - s h a p e d up to 2 mm in diam. and depth, the s t y l e 5 - 8 m m long, the stigma lobes o v a t e , o b t u s e at the a p e x , 2 χ 0 . 5 m m , the locules 3 . 5 - 4 . 5 mm t a l l , 1 - 1 . 5 in d i a m . , the placenta 3 - 4 m m t a l l , ca 0.5mm in diam., with the s u b - b a s a l l y p e l t a t e o v u l e s e l l i p t i c or o b o v a t e w i t h the apex truncate to subtruncate, 0. 6 - 0 . 8 χ 0 . 5 m m ; fruits 2 - 3 . 5 cm tall including the c a l y x , 0 , 8 - l c m in diam., the seeds 6 - 8 χ 1 - 2 m m , the testa c e l l s rectangular to e l l i p t i c , 1 0 4 - 3 0 1 ( x = l 8 6 ) μιη long, 3 6 - 7 5 ( x = 6 l ) urn w i d e , w i t h the w a l l s 6 - 7 ]im thick, w i t h 1 4 - 3 4 large pits in the
internal w a l l , the pits ( 5 - ) 1 2 - 3 2 ( x = 1 7 ) lim in d i a m .
Specimens e x a m i n e d . C O L O M B I A . M e t a : La S e r r a n i a , L l a n o G r a n d e , 26 Nov 1939, Cuatrecasas 7 8 9 0 (COL, F ) ; Llano de San M a r t i n , T r i a n a 1791 ( 3 2 6 4 . 1 ) (COL, F, P ) . Vicha da: ca 10 km W of Las G a v i o t a s along road to Puerto G a i t a n , 180 m , 30 Dec 1 9 7 3 , Davidse & Llanos 5 3 3 7 (COL, M O ) . V E N E Z U E L A . A m a z o n a s : C e r r o S i p a p o (Paráque) , 2 5 Jan 1 9 4 9 , M a gulre & Politi 28613 ( N Y ) . FRENCH G U I A N A . V a l l é e e n c a i s s é e d e la c r í q u e P a r é p o u , sur des col lines ã e n v i r o n 18 km de son c o n f l u e n t , 2 3 Sep 1968, O í d e m a n 2 8 5 2 ( N Y ) ; pied du saut Couéki, riv. Y a r o u p i , a f f l u e n t de 1 ' O y a p o c k , 10 A p r 1970, O l d e m a n 3 0 6 5 (NY)ECUADOR. Pastaza: Tzapino, 2 3 May 1976, O l d e m a n & A r e v a l o 4 7 ( U S ) . P E R U . L o r e t o : V a r a d e r o - M a z ã n , distrito Mazán, p r o v í n c i a M a y n a s , 1 Feb 1 9 6 3 , Arostegui 90 ( F ) ; V a r a d e r o A m a z o n a s , M a z ã n distrito Mazán, p r o v í n c i a M a y n a s , 2 2 Jun 1 9 6 3 , A r o s t e g u i 1 0 7 ( F ) ; trail from c a s e r i o of San Andrés, Rio M o m o n , t r i b u t a r y of Río N a n a y , d i s t r i t o Iquitos, província M a y n a s , 2 9 Mar 1 9 7 7 , Rimachi
2936
( F ) ; c a m p a m e n t o dei B o s q u e Nacional A l e x a n d e r von H u m b o l d t , d i s -trito Callería, p r o v í n c i a Coronel P o r t i l l o , 23 Feb 1 9 7 2 , S c h u n k e 5 2 7 5 (F). San M a r t i n : Tarapoto, Dec 1 9 2 9 , W i l l i a m s s n ( F ) . B R A Z I L . A m a z o n a s : M a r a l , Rio J a p u r á , 18 Apt 1970 Albuquerque & Lima 3 3 5 ( I N P A ) ; M a n a u s - C a r a c a r a Γ R o a d , krn 148, 2 7 Sep 1 9 7 3 , B e r g , B i s b y , Steward & Ramos Ρ18-Ϊ34 (INPA, K, M O , NY , U) ; south A l a l a u r i v e r , entre o A m a z o n a s e Ter_ rítório Roraima, 26 Feb 1 9 6 8 , B o y a n 2 6 9 ( I N P A ) ; e s t r a d a M a n a u s -1 t a c o a t i a r a , km 1 1 8 , Jul 1976, Coelho 7 9 8 (INPA); T a b a t i n g a , 8 Mar 1 9 4 4 , D u c k e 1 5 5 6 (F, Κ , N Y ) ; C u c u h y , Rio Ne -gro, 26 Sep 1935, Ducke 3 5 0 6 0 ( F ) ; M a r a r á , P a d a u i r y , Rio N e g r o , 4 Oct 1947, Froes 22511INSTITUTO NACIONAL DE
PESQUISAS DA AMAZÓNIA
; IΛΝ, U) ; Reserva Florestal Ducke, M a n a u s , 5 Jun 1964, Rodrigues S Loureiro 5838 (I Ν PA, RB); Sao Gabriel , R i o Negro, 8 Mar 1975, Ν . T. Silva 3 7 5 5 (IAN). Pará: San tarem,Curuã-Jna ( B a r r e i r i n h a ) , 17 Oct 196¾, Campbell S Osmarino sn (I Ν PA) ; Boa Vista ,8 Mar 1 933,Capu cho 550 ( F ) ; Juruty V e l h o , 28 May 1 9 2 7 , D u c k e 2 1 6 8 5 (Κ, P, U) ; Rio V e r m e l h o , 23 A p r 1961.
Froes 2 6 9 8 2 (IAN, U, U B ) ; s e r r a r i a , Serra dos C a r a j á s , M a r a b á , 1 Apr 1 9 7 7 , S i l v a 4 Bahia 299D (UB) ; entrada da e s t r a d a para a s e r r a r i a , Serra dos C a r a j á s , M a r a b á , 5 Apr 1 9 7 7 , Ξ Γ1 να c, Bahia 3D42 (UB). A c r e : Colônia Dias M a r t i n s , Rio B r a n c o , 12 Feb 1962, Vasconcelos y. Coelho sn (ΙΝΡΛ) . Rondônia: Porto Velho (Rio M a d e i r a ) , 9 Jan 1 9 3 0 , Ducke 22844 (Κ, P); C a c h o e i r a de Santa C r u z , Rio Jarnart, 28 Jun 1965, Pires í. Martin 9950 (iNY, RB, U B ) ; 1 Ια-s o u t h of Riberao, road Abunã-Guaj a ra-M i r i rn, 27 Jul 1968, Prance, Forero, Wig ley , RamoΙα-s S Farias 6564 ( F , I M P A ) . Mato Grosso: Dardanelos, estrada Santa Elena, Rio A r i p u a n ã , 16] Jun 1974, C o r d e i r o 122 ( I AN , N Y ) ; A r i p u a n ã , 20 Aug 1976, Gomes S Miranda 216 (INPA), t\ Mar I 9 7 7 , G o m e s , Miranda í Oliveira 946 INPA); A r i p u a n a , próximo ao Aeroporto, 24 May
I 9 7 6 , M o n t e i r o , Leite & A n d r a d e 1043 (INPA); rnarqern direita do Rio Juruena ,a r redores do tampo de a v i a ç ã o , 28 Jun 1 9 7 7 , Rosa & Santos 2 1 9 0 (INPA, M O , NY, RB) ; e s t r a d a para Ari-puanã, P o n t a n i l h a , A r i p u a n ã , Rio J u r u e n a , 30 Jun 1 9 7 7 , S i l v a & M a r i a 3 2 3 4 (MO, NY , UR). Common n a m e s , COLOMBIA: Palo de indio. E C U A D O R : O y u w ã e . PERU: C a p i r o n a , C a p i r o n a de a l t u r a , Capirona n e g r o , Meta q u a i s , Meto h u a y o . B R A Z I L : K u l a t e i r o , Pau m u l a t o .
Discussion. O l d e m a n 2 8 5 2 and 3 0 6 5 , both s t e r i l e , have been identified as C . decorticans on the basis of their totally g l a b r o u s c o n d i t i o n . T h e y have the largest stipules known
in the g e n u s , up to 8 cm long arid 2 . 5 cm w i d e . Flowering c o l l e c t i o n s are needed to c o n f i r m wi thout d o u b t , the presence of C . d e c o r t i c a n s in French G u i a n a .
C a p i r o n a leiophloea Beno i st , Bu 1 I, M u s . M i s t . N a t . (Paris) 2 7 : Jdj. 1 9 2 1 . T y p e s : Wachen-heim 2 1 5 , 20 Dec 1920, e n v i r e n s du camp de Godebert , French Guiana (lectotype: Ρ; isole ctc-types: F, K, P ) ; Benoist 1 1 9 2 , 8 May 1914, Saint Jean du M a r o n i , French Guiana (para
types: P ) . (Figs. 2 , 4 ) .
C a p i r o n a Duckey Huber in D u c k e , B o i . M u s . P a r a e n s e H i s t . N a t . 3 : 1 8 5 . 1 9 1 3 · Nom. nud .
C a p i r o n a h u b e r i a n a D u c k e , A r c h . J a r d . B o t . Rio de J a n e i r o 3:257- 1922, T y p e s : Ducke I I 8 6 5 , 2 9 Jun 1912, entre os Rios Curni nã -πι 7 r i m e A r i r a m b a , Rio Trombetas , Pa rã ,Bra zil (lectotype: RB; iso 1 e c t o t y p e s : F, P) ; Ducke 1 0 4 6 8 , 12 Oct 1 9 2 2 , Bella V i s t a , R i o Ta • a j o z , P a r a , Brazil ( p a r a t y p e s : R B , U ) ; Ducke 17126, 18 Jul 1918, perto de Cachoeira T e r m i n a l , A l t o A r i r a m b a , Rio B r a n c o - Ó b i d o s , P a r a , Brazil (paratypes : F, P. R B ) .
C a p i r o n a s u r i n a m e n s i s B r e m . , Recuei 1 T r a v . Bot. N e e r l . 3 i :2 β 1 . 1 9 3 4 . T y p e s : Forest Bureau 1829 (tree 110. 7 1 0 ) 9 May 1916, Bosch forest reserve, sectie 0, Suriname (lecto-type: U, 'solectotypes: K ) ; Forest Bureau 11¾] (tree no. 710) 8 Nov 19)6 (para(lecto-type: U ) , 2)00 (tree no. )115) 18 Jul 1916 (paratype: M 0 , U ) , 2 4 0 7 (tree no. 106)13 Nov I 9 1 6 (para type: U ) , 2416 (tree no. 7 1 0 ) 6 Sep 1916 (paratype: U ) , 4246 (tree n o . I O 6 ) 13 Feb 1919 (paratype: υ) , 4 3 9 7 (tree no. 106) 3 Nov 1919 (paratypes: Κ, NY, RB, υ) , Bosch forest reserve, sectie 0, S u r i n a m e .
Trees 11-32 m tall with a trunk d i a m . of 20-45 c m , the branches sparsely sericeous • ι very young and then glabrate to g l a b r o u s ; stipules ovate or el 1iptic to narrowly so, acute at the apex, 2.5-5 x 1-1.6 c m , w i t h the abaxial surface sparsely s e r i c e o u s , with lhe adaxial surface with a few a p p r e s s e d h a i r s ; leaves of v e g e t a t i v e b r a n c h e s w i t h the pfiiole 2-2,5 cm long, sparsely s e r i c e o u s , w i t h the blade el Iiptic or obovate to n a r r o w l y so, obtuse or cuneate at the b a s e , broadly acute or a c u m i n a t e or o b t u s e at the a p e x , 2 2 -40 χ 8-22 cm, 1 . 5- 3 . 3 times longer than w i d e , g l a b r o u s or g l a b r a t e a b o v e and sparsely
riceous or with a few a p p r e s s e d h a i r s b e l o w , w i t h (10-)12-24 a r c u a t e secondary nerves on each side of the m i d r i b , the m i d r i b and nerves plane above and prominent beneath,the leaves of fertile branches w i t h the petiole t e r e t e , 0.5-1.5 cm long, sparse 1 y ser i'ceous with the blade narrowly to broadly e l l i p t i c , o b t u s e or rarely cordate at the b a s e , b r o -adly acute or obtuse at the a p e x , 5-15 x 2 - 8 . 5 c m , 1.2-2 times longer than w i d e , glabro US or glabrate above and sparsely s e r i c e o u s b e n e a t h , w i t h 5-9 a r c u a t e secondary nerves on each side of the m i d r i b , the m i d r i b and n e r v e s plane and subprominent to prominent beneath; inflorescences 9-25 χ 12-30 c m , w i t h ( 1 0 - ) 2 5 - 1 0 0 f l o w e r s , w i t h 1 or 2 flowers or rarely none w i t h an e n l a r g e d foliar calyx lobe, w i t h the peduncle 68 cm long, s p a r -sely sericeous, with 2-5 pairs of lateral b r a n c h e s 2.5-15 cm long, spar-sely s e r i c e o u s , With the one bract subtending each b r a n c h o v a t e , a c u t e at the a p e x , 1.2-2 χ 0. 5 - 0 . 8 cm, with the abaxial surface g l a b r a t e to sparsely s e r i c e o u s , w i t h the adaxial surface g l a b -rous to glabrate; flowers w i t h the pedicel 2-7 m m long, g l a b r a t e to sparsely s e r i c e o u s , with one bract subtending e a c h f l o w e r , the bract broadly o v a t e , acute at the a p e x , 4 . 5 -6.5 χ 3-5 m m , with the abaxial surface glabrate to sparsely s e r i c e o u s , with the adaxial surface glabrous to g l a b r a t e , w i t h the h y p a n t h i u m f u s i f o r m , 4-7 m m t a l l , 3-4 m m indiam., sericeous, the calyx w i t h the e x t e r i o r s e r i c e o u s to w e a k l y s o , w i t h the interior densely sericeous, with the tube 4- 6 m m t a l l , 7-10 mm in d i a m . at the o r i f i c e , w i t h the lobes very broadly acute or very broadly o b t u s e , 1-2 χ (3-)4-7 m m , w i t h the foliar lobe with
the pseudopetiole 1-4.5 cm long, sparsely s e r i c e o u s to g l a b r a t e , w i t h the p s e u d o b l a d e 2.5-7 Χ 1-4 cm, glabrate a b o v e , sparsely sericeous b e l o w , the corolla tube 2 5 - 3 0 m m l o n g , with the lobes c i r c u l a r , o b t u s e at the a p e x , 10-13 χ 10-13 m m , the stamens separating
from the corolla tube 3 mm from the b a s e , connate for 5- 6 m m from their b a s e , with the free portion of the f i l a m e n t s 10-12 m m long, w i t h the a n t h e r s ca 8 m m long, the disk cup-shaped, 1.5 m m in d i a m . and d e p t h , the style 9 m m long, the stigma lobes e l l i p t i c , obtuse at the a p e x , 2 χ 1 m m , the locules 5- 6 m m t a l l , 1-1.5 m m in d i a m . , the placenta ca 4 mm tal 1, ca 0.5 m m in d i a m . , w i t h the c e n t r a l l y peltate ovules e l l i p t i c , 0 . 6 - 0 , 9 χ 0.4 mm; fruits 2 . 5 - 3 . 5 cm tall including the c a l y x , 1-1.3 cm in d i a m . , glabrate or spar sely sericeous, the seeds 6- 8 χ 1.5^2 m m , the testa cells r e c t a n g u l a r , 156-333(x = 232) \im long, 35-67(x = 54) y m w i d e , w i t h the w a l l s 4- 8 y m thick, w i t h 1 4 - 2 7 large pits in the internal w a l l , the pits 12-33(x = 21) y m in d i a m .
Specimens e x a m i n e d . S U R I N A M E . H o o g b o s a r b o r e t u m Kamp 8 M a p a n e g e b i e d , 3 Sep 1970, Elburg & Roberts 1 2 6 3 4 ( U ) ; Bosch forest r e s e r v e , sectie 0, F o r e s t B u r e a u 1 7 2 , Dec 1942 (K, NY, U ) , 1 7 2 a , May 1945 (COL, Κ, M O , N Y , S P , U ) ; from c o n f l u e n c e of Lucie and Oost
Rivers to 5 km e a s t , a l t . 2 7 5 - 3 2 5 m, 8 Sep 1 9 6 3 , Irwin, P r a n c e , S o d e r s t r o m & Holmgren 55528 (COL, F, Κ , M O , N Y , S P , U ) ; J o d e n s a v a n n e - M a p a n e k r e e k area (Suriname R . ) , 2 Apr 1 9 5 3 , L i n d e m a n 3681 ( U ) , 2 Mar 1 9 6 1 , Schulz 8583 (F, U ) ; Brownsberg N a t u r e Park near m a i n park h e a d q u a r t e r s , 90 km S of P a r a m a r i b o , Mazaroni P l a t e a u , 26 Sep 1976, Mori & B o l t e n 8 M 3 (Κ, M O , N Y , U ) ; B r o w n s b e r g , 2k Jun 1 3 7 0 , T a w j o e r a n 12851 (U) .FRENCH GUIANA. Route de C a y e n n e , km 7,980, 2 3 Mar 1956, c o l l e c t o r unknown 7 4 0 8 (P, U ). B R A Z I L . Pari: Igarapé do L i m a , p l a n a l t o de S a n t a r é m , 2 5 J u n 1951», F r o e s 3096O ( I A N , Κ , N Y , U) ; Itaitu b a , Rio T a p a j o z , k Feb 1 9 5 2 , P i r e s ^ 0 5 ^ ( I A N ) ; e s t r a d a e n t r e P i l ã o e R e p a r t i m e n t o , re -gião d o J a r i , 6 May 1 9 7 0 , N . T . S i l v a 3 1 1 3 (IAN); e s t r a d a do M u n g u b a km 10, re-gião do J a r i , 22 Oct 1970, Ν . Τ . Silva 3388 (IAN). Common n a m e s . S U R I N A M E : A k a t o m b e , A k a t o m n o , A k e g o e m i o , A k e p e m i o , A k h o r o k , «.a li d a n , K a l i d a n , K a n t a s i e h o e d o e , M o e n t e n e , M o e n t e n e h e , T o e l a l a h o e d o e . FRENCH G U I A N A : Bouchi B a n d a .
D i s c u s s i o n . T h e p u b e s c e n c e d e n s i t y of C . l e i o p h l o e a forms a c l i n e from n o r t h to south. The most d e n s e l y s e r i c e o u s m e m b e r s of the s p e c i e s are found in the n o r t h e r n part of its range, S u r i n a m e a n d French G u i a n a . T h e m o s t sparsely s e r i c e o u s forms a r e found at the southern limits of its r a n g e , south o f the A m a z o n River along the lower d r a i n a g e of the Rio T a p a j o z . T h o s e c o l l e c t i o n s from n o r t h o f the A m a z o n River a n d s o u t h o f the Guianas exhibit a n intermediate p u b e s c e n c e d e n s i t y . T h e area in w h i c h the s p a r s e s t pubescence
is f o u n d , is also the z o n e of c o n t a c t b e t w e e n C . l e i o p h l o e a a n d C . d e c o r t i c a n s . Perhaps the sparser p u b e s c e n c e of C . l e i o p h l o e a in this region indicates a p o s s i b l e e x c h a n g e of genes sometime in the p a s t . Our p r e s e n t k n o w l e d g e is insufficient to e v a l u a t e this s i t u a t i o n . Intensive c o l l e c t i n g in the z o n e of c o n t a c t a n d h y b r i d i z a t i o n studies are needed to c l a r i f y this s i t u a t i o n .
E x c l u d e d S p e c i e s
C a p i r o n a b o i v i n i a n a Ba i11 o n , B u l l . M e n s . S o c . L i n n . Paris 1: 2 7 0 . 1 8 8 O , is a typo graphic error for P l e u r o c o f f e a b o i v i n i a n a Bail Ion.
Bail Ion (1880) p r e s e n t e d the n e w g e n u s P l e u r o c o f f e a w i t h a s i n g l e s p e c i e s , P. boi v i n i a n a , a n d v a l i d a t e d both j o i n t l y . He c o m p a r e d his n e w g e n u s to P l a t y c a r p u m H u m b . & B o n p l . , C a p i r o n a , C o f f e a L., P o s o q u e r i a A u b l . a n d C a n t h i u m L a m . P l e u f o c o f f e a boiviniana w a s t r a n s f e r r e d to the g e n u s C o f f e a a n d w a s remained there (de W Í1 d e m a n , 1 9 ^ 1 ; C h e v a 1 i e r ,
1 9 ^ 7 ) . In c o m p a r i n g his g e n u s to C o f f e a , Bail Ion (188 0) m e n t i o n e d " C . b o i v i n i a n a " . Un doubtedly " c " w a s s u b s t i t u t e d for " p " by m i s t a k e . T h e first genus beginning w i t h " c "
to appear before " C . b o i v i n i a n a " w a s C a p i r o n a , so in v o l u m e I o f Index K e w e n s i s (Jack -son, I 8 9 3 ) b o i v i n i a n a w a s cited a s a n e w species of both P l e u r o c o f f e a a n d C a p i r o n a .
C a p i r o n a e r y t h r o x y l o n S t a n d l e y , P u b l . Field M u s . N a t . H i s t . , B o t . S e r . 2 2 : 4 8 . 19Ί0. = Pogonopus e r y t h r o x y l o n (Standi.) K i r k b r i d e , c o m b . n o v . T y p e : R u i z 8 Pavon s n , Peru (holotype: MA (specimen w i t h a u t o g r a p h i c label, " P e n t a n d r i a M o n o g y n i a , Genus novum
[crossed o u t . ] . Vulgo Palo C o l o r a d o . F. H. D. 1 3 9 . L 4 9 5 . A n o de I 8 O O . Ia S p . (Macro c n e m u m ) . " ) ; isotype: MA (duplicate specimen w i t h o u t a u t o g r a p h i c l a b e l ) .
Stand 1ey was a p p a r e n t l y o v e r i m p r e s s e d w i t h the calyx structure o f C . e r y t h r o x y l o n , If he had looked at the s p e c i m e n s in m o r e d e t a i l , he w o u l d h a v e d i s c o v e r e d that they have
vdlvate aestivation and many s m a l l , horizontal o v u l e s in each locule. The only genus combining these characters w i t h o n e calyx lobe sometimes e x p a n d e d into a large c o l o r e d
foliar organ, is Pogonopus K l o t z s c h . T h e g e n u s w a s reported to have two s p e c i e s , P. speciosus (Jacq.) Schum. and P. t u b u l o s u s (DC.) S c h u m . ( O e r s t e d , 1 8 5 2 ; S c h u m a n n , 1889,
(891;
Standley,)938;
S a n d w i t h ,19^9;
S t e y e r m a r k ,) 9 7 ^ ;
D w y e r ,1 9 8 0 ) ,
w h i c h d i f f e r mark edly from P. e r y t h r o x y l o n by p e r s i s t e n t s t i p u l e s , g e n e r a l l y s m a l l e r leaves and slightlylonqer, slenderer c o r o l l a s .
ACKNOWLEDGMENTS
I thank the following institutions for loaning or a l l o w i n g access to c o l l e c t i o n s for this study: C O L , F, IAN, INPA, L E , M A , NY, P, R B , S P , U, and W, and I thank Κ for supplying a list of their c o l l e c t i o n s a n d p h o t o g r a p h s of their t y p e s .
Apn.ZAzn.ta-* ζ um hÍÁtÕAÍ.co taxonÕmico do gznzho Capinona, *inonÍmia completa, dz*-CKÍÇOZA, noma* va-tgaAZA ζ di*cu**õz* AZ^ΖΑζηΧ.ζΑ, tanto paAa o gznzAo, como poJia ai> duo* thpicizh, C. deconZicam ζ C. Iziophloza. A di.*tAÍbiução gzogfia^iza do gznzAo abAangz at> bacia* do* Rio A Amazona* ζ So limo ζ*, e coincide com a di*th,Vouicão do gZnzno H e v e a . CapiAona dzcoAticxm* ζ conhzcida do oz*t.z ζ do *udz*tz do Amazona* ζ C. Iziophloza do ÍUtz da* Guiana* ζ do noAdzAtz do Amazona*. A zona dz contacto zntAZ a* duo* ZApzeiz* é a bzixa *ul do Rio Amazona* ζ a paxtz baixa da bacia do Rio Japa jo* no E*tado do Ροκά, BIOAÍÍ.
FIG. 2. Floral a n a l y s i s of C a p i r o n a d e c o r t i c a n s : a) the c o r o l l a , b) the c o r o l l a spread o p e n , c) s e c t i o n of the b a s e of the c o r o l l a w i t h a filament,d) the h y p a n t h i u m , e) transverse section of the h y p a n t h i u m and locules w i t h the p l a c e n t a s and ovules r e m o v e d , f) c r o s s - s e c t i o n of the h y p a n t h i u m and locules w i t h the p l a centas a n d o v u l e s r e m o v e d , g) a n o v u l e , h) the style a n d s t i g m a , a n d i) the h y p a n t h i u m w i t h one calyx lobe a large foliar o r g a n (Boyan 269 INPA).
2mm 4mm 5 mm J 8mm o.b.i 14 mm
F I G ; 3. F l o r a l a n a l y s i s of C a p i r o n a l e i o p h l o e a : a) the c o r o l l a , b) the c o r o l l a spread o p e n , c) s e c t i o n of the b a s e of the c o r o l l a w i t h a filament, d).the h y p a n t h i u m , e) t r a n s v e r s e s e c t i o n of h y p a n t h i u m a n d locules with the p l a c e n t a s a n d ovules r e m o v e d , f) c r o s s - s e c t i o n of the h y p a n t h i u m w i t h the p l a c e n t a s and ovules r e m o v e d , g) a n o v u l e , h) the s t y l e and s t i g m a , i) the h y p a n t h i u m w i t h one c a l y x lobe a large f o l i a r o r g a n , a n d j) c r o s s - s e c t i o n of the c o r o l l a wall s h o w i n g the p l i c a t e f o l d s , e x t e r i o r to the left a n d i n t e r i o r to the right (Silva 3113 I A N ) .
Referências b i b l i o g r á f i c a s
Hkillon, M. H. - 1 8 8 0 . Sur le P l e u r o c o f f e a . B u l l . M e n s . S o e . L i o n . P a r i s , 1 (34): f-270.
1881. T h e Natural H i s t o r y of P l a n t s . L o n d o n . L o v e l l R e e v e & C o . , v. 7.
Sftjoist, Μ. R. - 1 9 2 1 . L i s t e de p l a n t e s r é c o l t é e s par M . W a c h e n h e í m en Guyane F r a n ç a i s e . Buli. M u s . H i s t . N a t . , P a r i s , 2 7 : 3 6 6 - 3 6 8 .
Bremekamp, C. Ε. Β. 1 9 3 4 a . R u b i a c e a e . I n : P u l l e , A. F l o r a of S u r i n a m e , 4 ( 1 ) : 1 1 3 -' 298.
~ 1934b. N o t e s on the R u b i a c e a e of S u r i n a m , R e c . T r a v . B o t . N é e r l . , S o e . B o t . Néerl., 31 ( 1 - 2 ) : 2 4 8 - 3 0 8 .
1966. R e m a r k s o n the p o s i t i o n , the d e l i m i t a t i o n a n d the s u b d i v i s i o n of the Rubiaceae. A c t a B o t . N é e r l . , 1 5 : 1-33.
Chevalier, A. - 1 9 4 7 . L e s C a f é i e r s d u G l o b e , fascicule I I I . P a r i s , L e c h e v a l i e r .
Ducko, A, - 1913. E x p l o r a ç õ e s c i e n t í f i c a s no E s t a d o do P a r á . B o i . M u s . Goeldi (Mus. Paraense) H i s t . N a t . E t h n o g r a f i a , 3: 1 0 0 - 1 9 9 .
- 1922. Plantes n o u v e l l a s o u peu c o n n u e s de la region a m a z o n i e n n e (II e P a r t i e ) . Arch. Jard. B o t . Rio de J a n e i r o , 3: 3-282.
Ducke, A.& B l a c k , G . Α . - 1 9 5 3 . P h y t o g e o g r a p h i c a l n o t e s o n the B r a z i l i a n A m a z o n . A n a i s Acad. Bras. C i e n . , 25 : 1-46 .
1954. N o t a s sobre a f i t o g e o g r a f i a da A m a z o n i a b r a s i l e i r a . B o i . T e c . I n s t . A g r o n . Norte, 29: 1-62.
Dwyer, J. D. - 1 9 8 0 . 6 5 . P o g o n o p u s . I n : W o o d s o n J r . , R. Ε. & S c h e r y , R. W. - F l o r a of Panama. Part I X , F a m i l y 1 7 9 . R u b i a c e a e - Part II. A n n . M i s s o u r i B o t . G a r d . , 62
(2): 329-333. Hooker, J. D. - 1 8 7 3 . R u b i a c e a e . I n : B e n t h a m , G . & H o o k e r , J . D . - G e n e r a p l a n t a r u m 2(1): 7-152, L o n d o n . L o v e l l R e e v e & C o . .Jackson, B. D . - 1 8 9 3 . I n d e x K e w . O x f o r d , C l a r e d o n . v. 1. Oersted, A. S . - 1 8 5 2 . C e n t r a l a m e r i k a s R u b i a c e e r . V i d e n s k . M e d d e l . D a n s k N a t h u r i s t . Foren. K j o b e n h a v n , 1852 ( 2 - 4 ) : 2 3 - 6 1 . Schumann, C. - 1889. R u b i a c e a e . I n : v o n M a r t i n s , C . F. P. e d . F l o r a b r a s i l i e n s i s , 6 ( 6 ) : 1-466.
Schumann, C. - 1 9 8 1 . R u b i a c e a e . I n : E n g l e r , A. & P r a n t l , K. e d s . , D i e natu'rlichen Pflanzenfamilien, 4 ( 4 ) : 1-156.
Spruce, R. - 1 8 5 9 . On five n e w p l a n t s from E a s t e r n P e r u . J . P r o c . L i n n . S o c . , B o t . , 3: 191-204.
1970. N o t e s of a b o t a n i s t on the A m a z o n & A n d e s . R e p r i n t of 1 9 0 8 . New Y o r k , Johnson R e p r i n t C o r p .
Standley, P. C. - 1 9 3 0 . The R u b i a c e a e of C o l o m b i a . P u b l , F i e l d . C o l u m b i a n M u s ., B o t . Ser., 7: 3-175.
1936a. S t u d i e s of A m e r i c a n P l a n t s . - V I . F i e l d M u s . N a t . H i s t . , B o t . S e r . , 11 (5): 1 4 5 - 2 7 6 .
1936b. Rubiaceae. I n : MacBride, J. F. e d . -Flora of P e r u . Field. M u s . N a t . H i s t . , B o t . Ser. , 13: 3-261.
- 1938. Pogonopus Klotzsch. In: Flora of Costa Rica (part IV). P u b l . Field M u s . N a t . H i s t . , B o t . S e r . , 18(4):1339-1340.
- 1940. Studies of American Plants - IX. P u b l . F i e l d . M u s . N a t . H i s t . , B o t . S e r .
22 (1): 3-62.
Steyermark, J . A. - 1964. Rubiaceae. I n : B. Maguirre & colaboradores, Botany of the
Guyana Highland - Part V. M e m . N e w Y o r k B o t . G a r d . , 10 (5): 186-278.
- 1974. Rubiaceae, primeira parte. I n : Lasser, Ύ. - F l o r a de V e n e z u e l a , 10 (1):
1-593.
Verdecourt, B. - 1958. Remarks on the classification of the Rubiaceae.Bull . J a r d . B o t .
E t a t . Brxixelles, 28 (3): 209-281.
Wernham, H. F. - 1916. Tropical American Rubiaceae. - VII. - The Genera. J . B o t . , 54:
322-334.
Wildeman, E. de - 1941. Etudes sur le genre C o f f e a L. Bruxelles. Académie Royale de
Belgique.
