Preliminary revision of the genus Sopubia (Scrophulariaceae) in Madagascar Hans-Peter HOFMANN Botanisches Institut und Botanischer Garten, Rheinische Friedrich-Wilhelms-Universität Bonn, Meckenheimer Allee 170, D-53115 Bonn, Germany. Eberhard FISCHER Institut für Biologie, Universität Koblenz-Landau, Rheinau 1, D-56075 Koblenz, Germany. efischer@uni-koblenz.de KEYWORDS Sopubia, Scrophulariaceae, Madagascar. ABSTRACT T h e Malagasy taxa of the genus Sopubia Buch.-Ham. ex D . Don are revised, including a discussion of the type species, S. trifida, a variety of which inhabits Madagascar. Two new species are described, S. lemuriana and S. gracilis, and one new synonym is mentioned. A key to the species is provided, along with descriptions, illustrations, and data on their distribution. M O T S CLÉS Sopubia, Scroph ulariaceae, Madagascar. RESUME Les espèces malgaches du genre Sopubia Buch.-Ham. ex D . D o n sont revisées, avec une discussion du type du genre, 5. trifida, dont une variété se trouve à Madagascar. Deux espèces sont décrites comme nouvelles, S. lemuriana et S. gracilis, et un synonyme nouveau est mentionné. Une clé des espèces ainsi que des descriptions, des illustrations et des données sur leur répartition sont présentées. ADANSONIA, sér. 3 • 1998 • 20 (2) 299 Hofmann H.-P. & Fischer E. INTRODUCTION T h e g e n u s Sopubia B u c h . - H a m . ex D . D o n comprises a b o u t 2 5 t o 3 0 species, m o s t o f t h e m in t h e O l d W o r l d tropics. T h e g e n u s is based o n a s o m e w h a t p r o b l e m a t i c t y p e species, S. trífida B u c h . - H a m . from N e p a l , first described in 1 8 2 5 . T h e c o n c e p t o f t h e g e n u s rests o n t h e s t r u c t u r e of t h e a n d r o e c i u m consisting of four stamina, each w i t h t w o different thecae, o n e normal-sized a n d fertile, t h e o t h e r sterile a n d r e d u c e d in size. B E N T H A M ( 1 8 4 6 ) already p u t s t h e g e n u s i n t o t h e t r i b e G e r a r d i e a e , in w h i c h ir is still p l a c e d today. However, because o f t h e u n e q u a l thecae, h e distinguishes w i t h i n t h e tribe a g r o u p o f several g e n e r a (besides Sopubia Graderia, Harveya, Harveya, Aulaya, w h i c h w a s later m e r g e d w i t h a n d Centranthera) as " S o p u b i e a e " a n d o p p o s e s t h e m to the "Gerardieae verae". Neither V O N W E T T S T E I N (1891) in his t r e a t m e n t of t h e Scrophulariaceae in E N G L E R & P R A N T L ' S " N a t ü r l i c h e Pflanzenfamilien" n o r later a u t h o r s subscribe t o this formal d i s t i n c t i o n as a s u b t r i b e . Nevertheless, t h e m o r p h o l o g y o f t h e a n t h e r a e is still t h e t a x o n o m i c c r i t e r i o n o n t h e b a s i s o f w h i c h Sopubia is singled o u t i n m o s t t a x o n o m i c keys. fore already seems t o indicate its t r u e i d e n t i t y as Micrargeria. Since B o N A T l ' s c a t a l o g u e n o f u r t h e r s t u d y o n Sopubia in M a d a g a s c a r h a s b e e n p u b l i s h e d . W h e n r e v i e w i n g h e r b a r i u m s p e c i m e n s for t h e treatment of the genus within the "Flore de M a d a g a s c a r e t d e s C o m o r e s " w e realized t h a t s o m e o f t h e s p e c i m e n s c o u l d n o t b e p l a c e d in either o n e o f t h e three (rsp. t w o ) k n o w n species. O n t h e other h a n d , these specimens d i d n o t seem to relate to t h e African specimens of Sopubia seen e a r l i e r for a p l a n n e d w o r l d w i d e revision o f t h e g e n u s . T h i s s t u d y c o n s e q u e n t l y from i d e n t i f i e s t w o n e w s p e c i e s o f Sopubia M a d a g a s c a r a n d investigates t h e t a x o n o m i c relations b e t w e e n t h e Malagasy species a n d species o f Sopubia from n e i g h b o u r i n g floral regions. MATERIAL A N D M E T H O D S This study is based on the investigation of dried specimens mainly from the Paris herbarium (P). In a d d i t i o n , the following herbaria were c o n s u l t e d (abbreviations according to H O L M G R E N et al. 1990): B O N N , BR, FR, K, M . Material collecred by E. F I S C H E R during a field trip to Madagascar in 1993 and fixed in Ethanol was also studied. All quoted specimens have been seen unless indicated otherwise. T h e species o f Sopubia k n o w n t o d a y are s u b s h r u b s o r herbs w i t h s u b r o t a t e corollae a n d elegant, linear to lanceolate, simple or dissected GENERAL CONSIDERATIONS leaves. T h e y live as h e m i p a r a s i t e s in t h e d r y o r m o i s t grasslands at higher altitudes o r in o r near W e set o u t from B E N T H A M S d e s c r i p t i o n o f S. s w a m p y areas i n c l u d i n g d i s t u r b e d o r cultivated trífida var. madagascariensis w h i c h is t h e o n l y g r o u n d such as rice paddies in t h e lowlands. d e s c r i p t i on o f this species from Madagascar available today. A c c o r d i n g t o B E N T H A M , S. trífiB E N T H A M ( 1 8 4 6 ) was t h e last ( a n d , a p a r t from is a p e r e n n i a l ( a s G e o r g e D O N 1 8 3 8 , t h e only) a u t h o r t o revise t h e da v a r . madagascariensis g e n u s w o r l d w i d e . H e lists six s p e c i e s , a m o n g H A M I L T O N describes t h e type from Nepal) h e r b , t h e m S. trífida var. madagascariensis as t h e o n l y b r a n c h e d , w i t h simple o r rarely trifid leaves a n d species then k n o w n from M a d a g a s c a r . p u r p l e flowers. W h i l e his description is so genSubsequently, B A K E R ( 1 8 8 2 a n d 1 8 8 6 ) described eral t h a t it m a t c h e s m o s t o f t h e material seen, it t w o n e w species, o n e o f w h i c h (S. stricta Baker) does n o t fit i n particular a n u m b e r o f s p e c i m e n filiformis w i t h w h i t e flowers. W e n o t e d f u r t h e r m o r e that, n o w t u r n s o u t t o b e Micrargeria ( S c h u m . & T h o n n . ) H u t c h . & Dalziel. B O N A T I w h i l e t h e r e is little v a r i a t i o n i n leaf c h a r a c t e r s ( 1 9 2 7 ) still lists t h e three species in his catalogue a n d flower m o r p h o l o g y , t h e size a n d n u m b e r o f o f Scrophulariaceae collected in M a d a g a s c a r for flowers, the length of their pedicels a n d rhe t h e M u s e u m d ' H i s t o i r e N a t u r e l l e a t P a r i s . general habit o f t h e p l a n t s vary considerably. However, his descriptio n o f S. stricta relates t o a H A N S E N ( 1 9 7 5 ) describes t h e type s p e c i m e n o f p l a n t w i t h a m o r e z y g o m o r p h i c flower a n d there- S. trífida in his revision o f East African Sopubia, 300 ADANSONIA, ser. 3 • 1998 • 20(2) Sopubia (Scrophulariaceae) as a s l e n d e r h e r b w i t h t w o o r t h r e e s t e m s e x t e n d i n g from t h e thickened lower part of the s t e m . F r o m this h e c o n c l u d e s t h a t t h e plants are likely t o be perennials. Based o n this character h e distinguishes t h e t y p e a n d o t h e r Asian m a t e r i a l from t h e — a n n u a l a n d yellow-flowered—S. eminii E n g l . A c c o r d i n g t o HANSEN, all A f r i c a n m a t e r i a l n a m e d S. trífida falls u n d e r S. eminii, a purely Asian species. m a k i n g S. trífida MlELCAREK ( 1 9 9 6 ) s u b s e q u e n t l y treated S. eminii as a subspecies o f S. parviflora Engl. W e have little d o u b t t h a t these t w o taxa are at least very closely related. B E N T H A M , a n d earlier G e o r g e D O N ( 1 8 3 8 ) , describe a s e c o n d possible c o l o u r for t h e flowers of S. trífida, yellow w i t h a p u r p l e centre, based o n collections b y E D G E W O R T H a n d R O Y L E ( n o t yet seen b y u s ) . B O N A T I ( 1 9 2 7 ) still m e n t i o n s b o t h colours, b u t m o s t m o d e r n Asian Florae d e s c r i b e S. trífida s o l e l y as a y e l l o w - f l o w e r e d b r a n c h e d a n n u a l h e r b (for e x a m p l e Y A M A Z A K I 1985, 1990). B E N T H A M already suspected a c o n f u s i o n o f t w o s p e c i e s , S. trífida sensu H A M I L T O N a n d t h e yellow-flowered species, for w h i c h a n e w n a m e will m o s t p r o b a b l y be required. T h e M a l a g a s y s p e c i m e n o f S. trífida m a t c h B E N T H A M ' S description o f t h e variety madagascariensis well, w h e r e a s Sopubia o f t h e yellow type ( m u c h b r a n c h e d a n d w i t h a n a n n u a l appearance) seem t o be missing. Sopubia trífida var. madagascariensis B e n t h . is therefore retained as t h e a p p r o priate n a m e . B A K E R ( 1 8 8 2 , 1 8 8 6 ) described t w o n e w species of Sopubia from Madagascar, based o n acquisitions b y R . B A R O N . T h e first, S. stricta Baker, t h e t y p e o f w h i c h (Baron 2709) is represented b y a duplicate sheet in P, along w i t h o t h e r specimen s of S. stricta from Madagascar, p r o v e d t o be i d e n t i c a l w i t h Micrargeria filiformis (Schum. & T h o n n . ) H u t c h . & Dalziel. T h e n a m e " stricta " w o u l d have been illegitimate anyway, since G e o r g e D O N already used t h e sam e t o describe a collection b y W A L L I C H from B u r m a , w h i c h differs in s o m e characters from B A K E R ' S description. Sopubia triphylla Baker as t h e s e c o n d o f B A K E R ' S n e w species has simple, u n b r a n c h e d stems, u p r i g h t , s i m p l e , a n d c o m p a r a t i v e l y s h o r t leaves in w h o r l s o f three, a n d flowers i n t h e same disp o s i t i o n . N e i t h e r B A R O N n o r B A K E R specifies t h e ADANSONIA, ser. 3 • 1998 • 20(2) c o l o u r o f t h e flowers. N e i t h e r d o t h e y m e n t i o n t h e p l a n t ' s life c y c l e ( a n n u a l v s . p e r e n n i a l ) . However, t h e n a m e is valid a n d t h e descriptio n fits a n u m b e r o f Malagasy perennial, pale p u r p l e flowered specimen s well. A m o n g the remaining material, two groups c o u l d b e i d e n t i f i e d w h o s e set o f c h a r a c t e r s d o n o t m a t c h either t h e descriptio n o f S. trífida o r t h a t o f S. triphylla. O n e g r o u p h a s w h i t e flowers in a lax i n f l o r e s c e n c e , few m o s t l y trifid leaves w i t h large i n t e r n o d e s , distinctively reflexed bracts, a n d a n a n n u a l r o o t system. T h e s e c o n d g r o u p consists o f smaller plants w i t h small pale purple flowers, m u c h b r a n c h e d stems, a n d s i m p l e leaves. B o t h g r o u p s are in t h e following described as n e w species. S o m e t w o dozen specimens, mostly older or w i t h o u t t h e decisive parts for a p r o p e r d e t e r m i n a t i o n , could n o t b e assigned to o n e of those four species. It c a n n o t b e excluded, in t h e light of t h e material seen so far, t h a t m o r e t h a n four species exist o n M a d a g a s c a r . F u r t h e r s t u d i e s m i g h t therefore p r o d u c e o n e o r t w o a d d i t i o n a l n e w descriptions. IMPORTANT GENERIC CHARACTERS W e find t h a t , i n a c c o r d a n c e w i t h p r e v i o u s a u t h o r s , t h e g e n u s Sopubia is well defined b y its anther morphology. T h e existence of o n e red u c e d , sterile theca a n d t h e shape, size, a n d t h e l o n g - h a i r e d m a r g i n s o f t h e s t o m i u m o f t h e fertile theca s e e m t o b e very c o n s t a n t t h r o u g h o u t t h e g e n u s . All four a n t h e r s are c o h e r e n t at anthesis. T h e a n d r o e c i u m o f S. gracilis (Fig. 1) is given as an e x a m p l e . W e d i d n o t n o t i c e a n y r e m a r k a b l e variations between t h e African, a n d Malagasy m a t e r i a l a n d A s i a n m a t e r i a l o f S. trífida s e e n . W h e r e t h e a n t h e r s t r u c t u r e differs from this patt e r n , as i n S. delphinifolia G . D o n ( t r e a t e d as s y n o n y m o u s w i t h S. fastigiata Bonati by Y A M A Z A K I 1 9 8 5 , 1990), further studies m i g h t lead t o t h e c o n c l u s i o n t h a t this species s h o u l d n o t be i n c l u d e d in t h e genus. A n o t h e r typical character seems t o b e t h e pattern o f pilose a n d glabrous l o n g i t u d i n a l rows a l o n g t h e stems a n d b r a n c h e s . A p a r t o f t h e stem o f S. trífida (Fig. 2) is given as a n e x a m p l e . A t 301 Hofmann H.-P. & Fischer E. lobes are c o v e r e d w i t h l o n g w o o l l y - t o m e n t o s e hairs. T h e inside o f t h e calyx t u b e is glabrous. S o m e of these characters have b e e n m e n t i o n e d for o n e species or another. T h e y have n o t , h o w ever, b e e n recognized as d i s t i n g u i s h i n g characters for t h e g e n u s (with t h e exception o f t h e a n t h e r s c o n s i s t i n g o f o n e fertile a n d o n e sterile t h e c a ) . W e feel t h a t a r e v i s e d c o n c e p t o f t h e g e n u s s h o u l d p u t s o m e e m p h a s i s o n these characters as well. PHYTOGEOGRAPHY T h e four Malagasy species d o n o t m a t c h any of t h e A f r i c a n s p e c i e s s e e n b y us so far. T h e Malagasy S. triphylla bears s o m e resemblance in its g e n e r a l h a b i t t o t h e p a n - A f r i c a n S. mannii Skan var. tenuifolia (Engl. & Gilg) H e p p e r , b u t differs in s o m e leaf a n d flower characters. A closer s t u d y of t h e Asian material still p e n d i n g , t h e Malagasy S. trifida as a purple-flowered p e r e n n i a l seems t o be very similar to s o m e s p e c i m e n s labeled as S. trifida seen from Sri L a n k a a n d I n d i a itself. Sopubia triphylla, S. lemuriana, a n d S. gracilis are m o s t p r o b a b l y Malagasy e n d e m i c s . Fig. 1.—A, a r r a n g e m e n t of s t a m e n s in Sopubia: B, frontal view; C, s t a m e n , s h o w i n g hairy m a r g i n of t h e s t o m i u m . All from S . gracilis, Keraudren 165). t h e n o d e s , pilose rows split in t h e m i d d l e , each h a l f b r a n c h i n g off t o , a n d c o n t i n u i n g o n t h e m a r g i n o f t h e n e i g h b o u r i n g c a u l i n e leaf. Glabrous strips end immediately below the insertion p o i n t o f t h e leaves at t h e s t e m . Above t h e leaves, n e w pilose strips c o n t i n u e a l o n g t h e s t e m . If a leaf axil carries a b r a n c h or pedicel, t h e ad- a n d t h e abaxial sides of t h e b r a n c h or pedicel are pilose, t o o . T h i s p u b e s c e n c e c a n b e r a t h e r dense t o sparse. In d r y i n g , t h e rows of hairs t e n d t o t h i n o u t , s o m e t i m e s giving t h e p l a n t a w h o l l y glabrous a p p e a r a n c e . Lastly, t h e hair p a t t e r n of t h e calyx is distinctive. As in c a u l i n e leaves a n d bracts, t h e o u t s i d e m a r g i n s of t h e calyx lobes a n d , t o a lesser extent, t h e m i d r i b are pilose w i t h s h o r t , recurved hairs o n w h i t i s h - h y a l i n e c u s h i o n s (in all four species these cushions m e r g e i n t o a w h i t i s h crust w h e n d r y i n g ) . T h e i n n e r tips a n d m a r g i n s o f t h e calyx 302 Sopubia is well r e p r e s e n t e d in t r o p i c a l Africa w i t h a p p r o x i m a t e l y 15 t o 2 0 species, a n d p r o b a bly has its centre o f diversity in t h e h i g h l a n d s to t h e East of t h e c o n t i n e n t . T h e species r e p o r t e d from t h e S o u t h w e s t , in p a r t i c u l a r from A n g o l a a n d N a m i b i a , seem t o deserve a closer look; see for e x a m p l e C U C C U I N I ( 1 9 9 1 ) . I n t h e N o r t h , Sopubia reaches u p t o t h e borders of t h e S a h a r o Sindic region ( T c h a d , S u d a n , E t h i o p i a ) , w i t h o u t e x t e n d i n g i n t o it. It seems to be missing from t h e m o u n t a i n s o f Yemen a n d O m a n , as well as from t h e a r c h i p e l a g o s o f t h e I n d i a n O c e a n . Several species are r e p o r t e d from t h e I n d i a n s u b c o n t i n e n t , as well as f r o m B u r m a , T h a i l a n d , L a o s , C a m b o d i a , V i e t n a m and C h i n a . To the south, t h e a n n u a l yellow-flowered species figures as S. trifida in B A C K E R & B A K H U I Z E N ' S Flora of Java ( 1 9 6 5 ) . G o i n g o n t h e Pacific R i m , w e have seen s p e c i m e n s of Sopubia from t h e Phillipines u n d e r t h e n a m e of S. trifida w h i c h d o n o t m a t c h either H A M I L T O N ' S or Y A M A Z A K I ' S description , as well as L E I C H H A R D T ' S collection from Q u e e n s l a n d q u o ted in B E N T H A M ' S Flora Australiensis ( 1 8 6 9 ) , ADANSONIA, ser. 3 • 1998 • 20(2) Sopubia (Scrophulariaceae) d e f i n i t e l y a Sopubia b u t p r o b a b l y also n o t b e l o n g i n g to S. trifida B u c h . - H a m . T h e r e are n o reports of Sopubia from t h e N e w W o r l d . A p p a r e n t l y , Sopubia occurs in t w o d i s t i n c t areas, t h e first c o m p r i s i n g all o f tropical Africa (incl. S o u t h Africa), t h e s e c o n d s t r e t c h i n g from I n d i a to S o u t h East Asia a n d s o u t h to Australia, w i t h t h e Sahara-Sindic desert belt separating t h e t w o ranges. O n M a d a g a s c a r , Sopubia trifida is t h e m o s t w i d e s p r e a d species, i n h a b i t i n g m o s t of t h e c e n tral h i g h l a n d b e t w e e n A m b a t o n d r a z a k a a n d Ihosy. Sopubia triphylla has so far been collected m a i n l y from t h e A n k a r a t r a m o u n t a i n s b e t w e e n Antananarivo and Antsirabe, with one certain find from t h e I t r e m o area. Sopubia gracilis seems to be confined to t h e I t r e m o m o u n t a i n s . Sopubia lemuriana covers a wider range with finds s t r e t c h i n g from t h e A n t a n a n a r i v o area to t h e I t r e m o region. K e y t o t h e species o f Sopubia 1. in Madagascar Flowers white, corolla 12 to 15 m m in diameter. Plant up to 60 cm high. Cauline leaves few S. lemuriana 1'. Flowers dark or pale purple 2 2. Flowers large, corolla 15 to 18 m m in diameter, calyx up to 6 m m long. Stems simple or only lightly branched 3 2'. Flowers smaller, corolla 8 to 10 mm in diameter, calyx 3-4 m m long, stems normally much-branched S. gracilis 3. Cauline leaves mostly in whorls of three, simple, 20 to 25 m m long, 1.5 to 3 m m wide, upright. Flowers pale purple, mostly in whorls of three S. triphylla 3'. Cauline leaves alternate, opposite (decussate) or in whorls of three or four, lower ones (rarely some bracts as well) dissected, with three (sometimes two) linear segments, or simple, 25 to 40 mm long, 1.5 to 1.8 mm wide, spreading. Flowers more or less darkpurple, alternate, opposite or in whorls of three or four S. trifida S o p u b i a trifida B u c h . - H a m . var. madagascariensis B e n t h . I n D C . P r o d r . 10: 522 (1846). T Y P E . — L y a l l 2 3 1 , Madagascar (holo-, K). Perennial, 4 5 to 6 0 c m h i g h . Stems a s c e n d i n g to erect, o n e or m o r e from t h e s a m e r o o t stock. T h e axils o f t h e cauline leaves carry shoots t h a t n o r m a l l y stay s h o r t . O n l y if t h e m a i n axis is disturbed do the shoots immediately below the p o i n t of d i s t u r b a n c e grow to normal-sized b r a n c h e s a n d f o r m inflorescences. If t h e m a i n axis r e m a i n s u n d i s t u r b e d a n d able t o t e r m i n a t e its g r o w t h w i t h an inflorescence, it stays w i t h o u t b r a n c h e s or develops o n l y a few (two to three) branches i m m e d i a t e l y below t h e inflorescence. S t e m 2.5 t o 3 m m in d i a m e t e r , w o o d y at t h e base, t h e l o w e r p a r t c y l i n d i c a l , a n g u l a r a b o v e , w i t h the characteristic p a t t e r n of l o n g i t u d i n a l rows of hairs alternating w i t h glabrous strips. ADANSONIA, ser. 3 • 1998 • 20(2) Leaves few to m a n y , a l t e r n a t e , o p p o s i t e (decussate) o r in w h o r l s of three or four, o n b r a n c h e s m o s t l y o p p o s i t e , fewer t h a n o n t h e m a i n axis. Internodes 10-15 m m long (sometimes longer int t h e u p p e r p a r t o f t h e s t e m ) . C a u l i n e leaves b e l o w m o s t l y three-fid ( s o m e t i m e s o n l y t w o segments) or s i m p l e , above a n d in t h e inflorescence m o s t l y simple ( a l t h o u g h two-fid or even three-fid bracts o c c u r ) , linear or w i t h linear s e g m e n t s , 2 5 t o 4 0 m m long, 1.5 t o 1.8 m m w i d e , m a r g i n s slightly revolute, m i d r i b a n d m a r g i n s pilose. Inflorescence a lax r a c e m e , i n t e r n o d e s m o s t l y longer t h a n t h e flowers o n their pedicels. Bracts r e s e m b l e s m a l l e r c a u l i n e leaves, s p r e a d i n g t o reflexed. P e d i c e l s 12 t o 14 m m l o n g , s h o r t e r t h a n t h e b r a c t s (in d r y i n g , b r a c t s a n d c a u l i n e leaves b e c o m e rather brittle a n d t e n d to break off easily). Flowers alternate, o p p o s i t e (decussate) o r in w h o r l s o f three or four. F l o w e r s m o r e o r less d a r k p u r p l e . B r a c t e o l e s linear t o n a r r o w l y lanceolate, 2 t o 3 m m l o n g , 303 Hofmann H.-P. & Fischer E. Fig. 2 . — S o p u b i a trifida. A, habit; B, d i s s e c t e d corolla; C, d i s s e c t e d calyx s h o w i n g ovary a n d style; D, part of s t e m with fruit; E, s t e m leaf; F, bract; G, b r a c t e o l e . A, D from Viguier & Humbert 1213; B-C from Waterlot 716; E-G from Lowry4515. 304 ADANSONIA, ser. 3 • 1998 • 20(2) Sopubia (Scrophulariaceae) m a r g i n s a n d m i d r i b densely pilose, n o t p r o t r u d ing over t h e calyx t u b e . Calyx u p to 6 m m l o n g , its teeth o b t u s e , their l e n g t h equalling t h e t u b e . I n n e r tips a n d m a r g i n s of t h e teeth densely w o o l ly, m i d r i b a n d o u t e r m a r g i n s of t h e teeth sparsely pilose. C o r o l l a 1 5 to 1 6 m m in diameter, its segments subequal. C a p s u l e 6 to 7 m m l o n g , glabrous. Seeds t r u n cate, their surface reticulate. W e t a n d d r y grasslands u p to 2 3 0 0 m , borders of s w a m p s , rice paddies, a n d o t h e r m o i s t places.—Figs. 2 , 6 . M A T E R I A L S T U D I E D . — A c a d e m i e Malgacke s.n., Ambatondrazaka, Mar. 1905 (P); Academie Malgache s.n., Imerina, Dec. 1908 (P); Baron 142, Madagascar (P); Benoist s.n., Tananarive, Antsirabe, Apr. 1951 (P); Benoist 646, Antsirabe, 22 Dec. 1950 (P); Boissieu s.n., s . l o c , s.d. ( P ) ; Bosser 18856, A n k a r a t r a , R t e . d'Ambatolampy a Faratsiko, 2200-2300 m, Feb. 1964 (P); Bosser 19419, A n d r i n g i t r a , A p r . 1 9 6 4 ( P ) ; Campenon s.n., I m e r i n a , 1 8 8 7 ( P ) ; Cours 130, A n k a r o k a ( A m b a t o n d r a z a k a ) , 3 0 J a n . 1 9 3 8 (P); d'Alleizette 70, Ivohibe, Nov. 1924 (P); d'Alleizette 832M, Tananarive, lieux h u m i d e s , O c t . 1906 (P); Deans Cowan s.n., Ankafana, 1880 (P); Decary s.n., A m b a t o l a o n a , M a r . 1 9 1 9 ( P ) ; Decary 643, A m b o h i m a n g a , 27 Mar. 1921 (P); Decary 13411, T s i a f a j a r o n a , S e p . 1 9 3 8 ( P ) ; Decary 17208, Manankazo, Lampokotsa d'Ankazobe, 3 Jan. 1942 (P); Geneaud 8, environs de Tananarive, Apr. 1892 (P); Guillaumet 2021, Anjozorobe, 28 Mar. 1968 (P); Hildebrandt 3713, Andrangoloaka ( O s t - I m e r i n a ) , Nov. 1880 (K, M ) ; Homolle 1200, Mt. Boby, 1946 (P); Humbert 1324, Tananarive, Ambrahi, Nov. 1935 (P); Humbert 28218, montagnes a l'Ouest d'ltremo (West Betsileo), 1500-1700 m, Jan. to Apr. 1955 (P); Humbert 29985, m o n t a g n e s a l ' O u e s t d ' l t r e m o (West-Betsileo), 1500-1700 m, Jan. to Apr. 1955 (P); Humbert 30016, m o n t a g n e s a l ' O u e s t d ' l t r e m o (West-Betsileo), 1500-1700 m, Jan. to Apr. 1955 (P); Humblot 623, Madagascar, s.d. (P, K); Jard. Bot. Tananarive 3190, s . l o c , J a n . 1 9 3 8 ; Jard. Bot. Tananarive 4749, M t . Tsitondroina, 15 Apr. 1941 (P); Keraudren 40, Angavokely (Tananarive), Feb. 1960 (P); Keraudren 260, Ambatofiterahana, entre Ambositra et Fianarantsoa, 1500 m, Mar. 1960 (P); Lowry 4515, A n d r i n g i t r a R e s . , A n t a n i f o t s y (Ambalavao), 1400 m, 5 Mar. 1989 (P); Morat 1289, Andringitra, J u n e 1965 (K, P); Reserves Naturelles 3038-RN, Radafindrakolo, Sendrisoa (Fianarantsoa), 6 M a y 1 9 5 1 ( P ) ; Reserves Naturelles 3990-RN, Radafindrakolo, Vohitsaoka (Ambalavao), 30 Mar. 1934 (P); Reserves Naturelles 5587-RN, Randriamiera, Sendrisoa (Ambalavao), 15 June 1953 (P); Reserves ADANSONIA, ser. 3 • 1998 • 20 (2) Naturelles 9627-RN, Rakotovao, Ambatosoratra ( A m b a t o n d r a z a k a ) , 16 D e c . 1 9 5 8 ( P ) ; Reserves Naturelles 11298-RN, Rakotovao, Momaka (Ambatondrazaka), 19 Nov. 1960 (P); Peltier 1060, Tananarive, July 1959 (P); Peltier 1182, Amboasary ( A m b a t o l a m p y ) , 22 O c t . 1959 (P); Peltier 1558, Sabotsy, Massif de 1'Antongona, 4 Dec. 1959 (P); Peltier 1612, Ambatofotsy (Tananarive), 18 D e c . 1959 (P); Peltier 3444, s.loc, s.d. (P); Perrier de la B&thie 11405, Ambositra, prairies, Sep. 1911 (P); Perrier de la Bathie 14995, Andilamena, marais, Nov. 1922 (P); Viguier & Humbert 1213, Ambatolaona ( M a n j a k a n d r i a n a ) , 1 4 0 0 m , 11 N o v . 1 9 1 2 ( P ) ; Viguier & Humbert 1490, Ambohiponana (Antsirabe), 1400 m, 20 Nov. 1912 (P); Viguier & Humbert 1641, Itasy, Massif d'Ankaratra, 2600 m, 27 Nov. 1912 (P); Waterlot s.n., Tananarive, Dec. 1917 (P); Waterlot 716, Angavo (Manjakandriana), Mar. 1923 (P). S o p u b i a triphylla Baker J. B o t . 20: 220 (1882). TYPE.—Baron 141, C e n t r a l M a d a g a s c a r (holo-, K; iso-, P) Perennial, b u t s o m e w h a t delicate, stems erect, 3 5 t o 4 0 c m h i g h , o n e o r m o r e from t h e s a m e r o o t stock. T h e axils o f t h e cauline leaves carry shoots t h a t n o r m a l l y stay s h o r t . O n l y if t h e m a i n axi s is d i s t u r b e d d o t h e s h o o t s i m m e d i a t e l y below t h e p o i n t of d i s t u r b a n c e g r o w to n o r m a l sized b r a n c h e s a n d f o r m i n f l o r e s c e n c e s . If t h e m a i n axis r e m a i n s u n d i s t u r b e d a n d able to t e r m i n a t e its g r o w t h w i t h a n inflorescence, it n o r m a l l y stays w i t h o u t b r a n c h e s . S t e m 2 to 2 . 5 m m in diameter, slightly w o o d y at t h e b a s e , t h e l o w e r p a r t c y l i n d i c a l , a n g u l a r a b o v e , m o r e d e n s e l y leafed t h a n t h e p r e c e d i n g species. Leaves m o s t l y in w h o r l s of three, at times alternate. C a u l i n e leaves rather erect, s i m p l e , narrowly triangular to linear, 2 0 to 2 5 m m long, 1 . 5 to 3 m m wide, margins slightly revolute, m i d r i b a n d m a r g i n s pilose. Inflorescence a lax to dense r a c e m e , i n t e r n o d e s m o s t l y shorter t h a n t h e flowers o n their pedicels. Bracts resemble smaller c a u l i n e leaves, u p r i g h t . Pedicels 1 2 t o 1 8 m m l o n g , m o s t l y longer t h a n the b r a c t s . Flowers in w h o r l s of t h r e e , rarely alternate. 305 Hofmann H.-P. & Fischer E. Flowers pale p u r p l e . Bracteoles linear t o n a r rowly lanceolate, 2 to 3 m m long, m a r g i n s a n d m i d r i b densely pilose, n o t p r o t r u d i n g over t h e calyx. Calyx u p to 6 m m long, its teeth t r i a n g u lar, m o r e a c u t e t h a n in t h e p r e c e e d i n g species, t h e i r l e n g t h e q u a l l i n g t h e t u b e . I n n e r tips a n d m a r g i n s of t h e teeth densely woolly, m i d r i b a n d outer margins of the teeth sparsely pilose. C o r o l l a 15 t o 18 m m in diameter, its s e g m e n t s subequal. C a p s u l e 6 to 7 m m long, glabrous. Seeds t r u n cate, their surface reticulate. M o u n t a i n o u s regions u p to 2 2 0 0 m . — F i g s . 3 , 6. Fig. 3 . — S o p u b i a triphylla. 11496. 306 MATERIAL STUDIED.—Baron 141, Central M a d a g a s c a r , 1 8 8 9 , t y p e (K, P ) ; Baron 242, Madagascar, s.d. (P); Baron 5242, N W Madagascar, 1887 (K); Bosser 18349, Tananarive, Iarinandriana (P); Catat287, Ankaratra, 2200 m, 2 May 1889 (P); Catat 299, Ankararra, 2 May 1889 (P); Catat 300, Ankaratra, 2 0 0 0 m, 2 May 1889 (P); Catat 349, A n k a r a t r a (versant E s t ) , 1 8 8 9 (P); Catat 1264, M i a n t s o a r i v o , 13 M a y 1889 (P); Decary 17433, Valozoro, Ambohimahasoa, 6 Feb. 1942 (P); Decary 17655, A n t s a m p a n d r a n o (Ankaratra), 1700 m, 10 F e b . 1 9 4 2 ( P ) ; Jard. Bot. Tananarive 2362, Votovorona (Antsirabe), 4 July 1935 (P); Jard. Bot. Tananarive3173, Behenjy, 12 Feb. 1938 (P); Le Myre de Viliers s.n., s.loc. (P); Terrier de la Bâthie 11496, Mt. Ambona jusqu'ŕ 2200 m (Antsirabe), May 1912 (P). A, habit; B, d i s s e c t e d corolla; C, d i s s e c t e d calyx s h o w i n g ovary a n d style. All from Perrier de la Bathie ADANSONIA, sér. 3 • 1998 • 20(2) Sopubia (Scrophulariaceae) Fig. A—Sopubia E from Fischer lemuriana 197. A, habit; B, i n f l o r e s c e n c e ; C, d i s s e c t e d corolla; D, calyx; E, calyx d i s s e c t e d . A from Schlieben ADANSONIA, sér. 3 • 1998 • 20(2) 8192; B- 307 Hofmann H.-P. & Fischer E. S o p u b i a l e m u r i a n a H.-P. H o f m a n n & E b . Fisch., sp. nov. Herba annua, caulis erectus. Folia pauca, inferiores trífida laciniis linearibus, superiores integra, lineariangustissima. Inflorescentia racemosa, laxa. Bracteae distincte reflectae. Flores albi. Corollae 12 usque ad 15 mm diámetro. Ab S. trífida differt floribus albis internodiisque distincte longioribus. T Y P E . — S c h l i e b e n 8192, Madagascar, Ambalamanaka, 18 Dec. 1959 (holo-, M ; iso-, BR, K). A n n u a l p l a n t , 5 0 to 6 0 c m h i g h . Stems erect, simple to strongly b r a n c h e d . S t e m 2.5 t o 3 m m in diameter, slighty w o o d y at t h e base, t h e lower p a r t cylindrical, a n g u l a r above, w i t h t h e characteristic pattern of longitudinal rows of hairs. C a u l i n e leaves few, s o m e t i m e s c r o w d e d at t h e base o f t h e s t e m , w i d e l y s p a c e d i n t h e m i d d l e section of t h e s t e m w i t h i n t e r n o d e s a p p r o a c h i n g 3 0 m m , o p p o s i t e (decussate) or alternate, s o m e times in w h o r l s o f three or four. C a u l i n e leaves b e l o w m o s t l y three-fid ( s o m e t i m e s w i t h o n l y t w o s e g m e n t s ) , d e l i c a t e , segm e n t s linear, 2 5 t o 3 5 m m l o n g , 1.5 m m w i d e , a b o v e u n d i v i d e d a n d s h o r t e r . M a r g i n s slightly revolute, m i d r i b a n d m a r g i n s pilose. Inflorescence a lax raceme, i n t e r n o d e s m o r e or less equalling t h e flowers o n their pedicels. Bracts u n d i v i d e d , linear t o n a r r o w l y triangular, distinctively reflexed. Pedicels u p to 15 m m long, m o s t ly l o n g e r t h a n t h e b r a c t s . Flowers d e c u s s a t e or rarely alternate. Flowers w h i t e . Bracteoles linear to narrowly lanceolate, 2 t o 3 m m long, m a r g i n s a n d m i d r i b densely pilose, rarely p r o t r u d i n g over t h e calyx. Calyx 5 to 6 m m long, its teeth o b t u s e , shorter t h a n t h e t u b e . I n n e r tips a n d m a r g i n s of the teeth densly woolly, m i d r i b a n d o u t e r m a r g i n s of t h e teeth sparsely pilose. C o r o l l a 12 t o 15 m m in diameter, its s e g m e n t s s u b e q u a l . C a p s u l e 4 to 5 m m long, glabrous. Seeds t r u n cate, their surface reticulate. M o s t l y d r y grasslands u p t o 1 9 0 0 m , p r o b a b l y h i g h e r . — F i g . 4 , 7. PARATYPES.—MADAGASCAR: Benoist 459, M a n j a k a r o m p o ( A n k a r a r r a ) , 19 D e c . 1 9 5 0 ( P ) ; Benoist 1643, Manjakatompo (Ankaratra), 20 Dec. 308 1951 (P); Bosser 7912, Ambatomenaloha (Itremo), Sep. 1956 (P); Cam 1117, Managary, 9 June 1889 (P); Croat 29776, Fianarantsoa, Itremo, Jan. 1975 (P); Decary 13105, environs d'Ambarofinandrahana, 1 6 0 0 - 1 8 0 0 m, 20 Feb. 1938 (P); Decary 17205, Manankiazo, s.d. (P); Decary 17610, Antsampandrano (Ankaratra), 1900 m, 9 Feb. 1942 (P); Fischer 197, Ambalamanaka, secondary grassland in m o u n t a i n forest, 2 9 M a r . 1 9 9 3 ( B O N N ) ; Guillaumet 2042, Andringirra, along R N 7, 8 Mar. 1968 (P); Humbert & Capuron 28052, Betsileo, Ambatofinandrahana, bois des pentes occidentales sur gneiss, 16 Jan. 1955 (P); Jacquemin 1303, Tananarive, Mt. Iharanandriana, Mar. 1973 (P); Leeuwenberg & Rafamantanantsoa 14435, Fianaranrsoa, Ambalamana, Feb. 1994 (K, P); Phillipson et al. 3875a, 50 km W of A m b a r o f i n a n d r a h a n a (Irremo), 1670 m, 13 Mar. 1 9 9 2 ( M O , n o t s e e n , P ) ; Schlieben 8192, Ambalamanaka, 18 Dec. 1959, type (BR, K, M). S o p u b i a gracilis H.-P. H o f m a n n & E b . Fisch., sp. nov. Suffrutex perennis, caules plures ex eadem radice, erecti, ramosi. Rami caulim principalum non superandi. Folia integra, lineari-angustissima. Inflorescentia racemosa, laxa usque ad densa. Ab aliis speciebus madagascariensibus generis Sopubia flores minores, purpurei pallidi. Corollae 8 usque ad 10 mm diametro. Ab S. triphylla differt floribus minorìbus, caule valde ramificato et foliis rare tri-verticillatis. Ab S. trifida differt foHis integris. TYPE.—Keraudren 165, Madagascar, Rre. d'Ambositra à Ambatofinandrahana, Feb. 1960 (holo-, P). P e r e n n i a l little s u b s h r u b , 3 5 t o 4 5 c m h i g h . Stems erect, o n e or m o r e from the same root s t o c k , m u c h - b r a n c h e d in a p y r a m i d a l f a s h i o n , t h e b r a n c h e s n o t p r o t r u d i n g over t h e ( u n d i s t u r b ed) m a i n axis. Stems 2 to 2.5 m m in diameter, w o o d y at t h e base, t h e l o w e r p a r t c y l i n d i c a l , a n g u l a r a b o v e , w i t h the characteristic p a t t e r n of l o n g i t u d i n a l rows of hairs, m o r e or less densely leafed. Leaves s u b o p p o s i t e (decussate), s o m e t i m e s in w h o r l s of three or four, rarely alternate. Leaves simple, linear, 15 to 2 0 m m long, 1 m m w i d e , m a r g i n s slightly t h i c k e n e d o r revolute (in d r y i n g , m a r g i n s curl u p w a r d s , giving t h e leaves a grooved a p p e a r a n c e ) , m i d r i b a n d m a r g i n s pilose. Inflorescence a lax t o dense raceme, i n t e r n o d e s e q u a l l i n g t h e flowers o n t h e i r p e d i c e l s . Bracts ADANSONIA, sér. 3 • 1998 • 20(2) Sopubia (Scrophulariaceae) Fig. 5 . — S o p u b i a gracilis: A, habit; B, d i s s e c t e d corolla; C, d i s s e c t e d calyx s h o w i n g ovary a n d style. All from Keraudren ADANSONIA, ser. 3 • 1998 • 20 (2) 165. 3Q9 Hofmann H.-P. & Fischer E. r e s e m b l e smaller c a u l i n e leaves, u p r i g h t to s p r e a d i n g , appressed to t h e pedicels at least for 1 m m o f their l e n g t h . Pedicels u p t o 8 m m long, m o s t l y longer t h a n t h e bracts. Flowers o p p o s i t e (decussate), in w h o r l s of three, rarely alternate. Flowers pale p u r p l e ( p r o b a b l y s o m e t i m e s even whitish). Bracteoles resembling small bracts, linear, 2 m m l o n g , n o t p r o t r u d i n g over t h e calyx t u b e . Calyx 3 to 4 m m l o n g , its teeth o b t u s e to r o u n d e d , s h o r t e r t h a n t h e t u b e . I n n e r tips a n d m a r g i n s of t h e teeth densely woolly, m i d r i b a n d outer margins of the teeth sparsely pilose. C o r o l l a 8 t o 10 m m in d i a m e t e r , its s e g m e n t s subequal. C a p s u l e 3 to 4 m m long, glabrous, at m a t u r i t y n o t p r o t r u d i n g over t h e calyx. Seeds t r u n c a t e , their surface reticulate. Mostly dry places on o p e n g r o u n d , up to 1 8 0 0 m , p r o b a b l y h i g h e r . — F i g . 5, 7 . P A R A T Y P E S . — M A D A G A S C A R : Catat 1150, Ambohip r o n a n a , 20 M a y 1889 (P); Decary 12918, env. d'Ambatofinandrahana, 1600-1800 m, 15 Feb. 1938 (P); Decary 12966, env. d ' A m b a t o f i n a n d r a h a n a , Fig. 6 . — G e o g r a p h i c distribution of Sopubia: 310 left S. trifida; right S . triphylla. ADANSONIA, ser. 3 • 1998 • 20(2) Sopubia (Scrophulariaceae) A(DS) A(DS) -15°S -15°S h-25° 00 50"E 45 Fig. 7 . — G e o g r a p h i c distribution of Sopubia: 1600-1800 m, 16 Feb. 1938 ( P ) ; Decary 13075, env. d'Ambatofinandrahana, 1600-1800 m, 19 Feb. 1938 ( P ) ; Decary 13127, env. d ' A m b a t o f i n a n d r a h a n a , 1600-1800 m, 20 Feb. 1938 ( P ) ; Dorr 3871, M t . Ibity (south), Fianarantsoa, 6 Feb. 1985 ( M O , not seen, BR, K); Keraudren 165, Rte. d'Ambositra a Ambatofinandrahana, March 1960, type ( P ) ; Perrier de la B&tbie 12366, entre l'lvato et la Mania (Itremo), 1500 m, Feb. 1919 ( P ) . 200 km left S. gracilis; right S. lemuriana. Acknowledgements W e would like to thank the keepers of the following herbaria for the loan of specimens: B R , F R , K, M, P , and express our gratitude to Drs. P . B A M P S ( B R ) and F . B A D R F . ( P ) for additional information and practical advice. REFERENCES E x c l u d e d species: Sopubia stricta B a k e r = Micrargeria ( S c h u m . & T h o n n . ) H u t c h . & Dalziel. ADANSONIA, sér. 3 • 1998 • 20 (2) B A C K E R C.A. filiformis & BAKHUIZEN VAN D E N BRINCK R.C. 1965.—Flora of Java 2: 514. B A K E R J.G. 1 8 8 2 . — C o n t r i b u t i o n s to the Flora of Central Madagascar./. Bot. 20: 218-222. 311 Hofmann H.-P. & Fischer E. 1 8 8 6 . — F u r t h e r C o n t r i b u t i o n s to the Flora of Central Madagascar: Sopubia strida, n. sp. /. Linn. Soc, Bot. 2 1 : 407-455. B E N T H A M G . 1846.—Scrophulariaceae: 521-523, in D C , Prodromus Systematis Naturalis Regni Vegetabilis 10. B E N T H A M G . 1869.—Flora Australiensis 4: 512-513. B O N A T I G . 1927.—Scrofulariacées de Madagascar. Bull. Soc. Bot. France 74: 98-99. C U C C U I N I P. 1991.—Sopubia patris (Scrophulariaceae), a new species from Central Angola. Webbia 45:309-315. D O N G . 1838.—A General System of Gardening and Botany 4: 559-560. H A M I L T O N F. 1 8 2 5 . — S o p u b i a : 8 8 , in D O N D., Prodromus Flora Nepalensis. H A N S E N O . J . 1 9 7 5 . — T h e East African Species of Sopubia (Scrophulariaceae). Kew Bull. 30: 543-558. BAKER J.G. F.N. 1 9 6 3 . — 1 5 3 . Scrophulariaceae : 3 5 2 374, in H E P P E R F.N, Flora of West Tropical Africa, 2, second edition. HEPPER H O L M G R E N P.K., HOLMGREN N.H. & BARNETT L.C. 1990.—Index Herbariorum. Part L: The Herbaria of the World, eighth edition. Utrecht. M l E L C A R E K R. 1996.—Les Scrophulariaceae dans la Flore d Afrique Centrale (ex Lindernieae). Fragm. Flor. GeobotAl: 199-220. W E T T S T E I N R. von 1891.—Scrophulariaceae: 39-107, natürlichen in E N G L E R A. & P R A N T L K., Die Pflanzenfamilien 4, 3b. Leipzig. Y A M A Z A K I T . 1985.—Scrophulariaceae: 1-217, in L E R O Y J.-F., Flore du Cambodge, du Laos et du ViêtNam 2 1 . Paris. Y A M A Z A K I T . 1990.—Scrophulariaceae: 223-225, in S M I T I N A N D T . & L A R S E N K., Flora of Thailand 5. Bangkok. Manuscript received 26 February 1998; revised version accepted 12 june 1998. 312 ADANSONIA, ser. 3 • 1998 • 20 (2)