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B o th a lia 3 7 ,2 : 1 7 7 - 1 8 2 (2 0 0 7 ) N ew species and notes on H e s p e r a n th a (Iridaceae) in sou th ern A frica p. G O L D L A T T * an d J.C . M A N N IN G * * K e y w o r d s : H e s p e r a n t h a K er G a w l., Ir id a cea e, so u th ern A fr ic a , ta x o n o m y A BSTRACT F ie ld stu d ie s o f th e su b -S a h a ra n A fric a n an d la r g e ly so u th ern A fr ic a n H e s p e r a n t h a c o n d u c te d sin c e 2 0 0 3 h a v e re su lte d in th e d is c o v e r y o f th ree n e w sp e c ie s in th is g e n u s, b r in g in g th e to ta l to 8 2 . H e s p e r a n t h a lo n g is t y la J .C .M a n n in g & G o ld b la tt. k n o w n fro m o n e c o lle c t io n fro m th e m o u n ta in s o f S W N a m ib ia , is a d w a r f p la n t w ith m o d e r a te ly lo n g -tu b e d , p u rp le flo w e r s an d u n u su a lly lo n g sty le b ra n ch es, p o s s ib ly a llie d to th e K a m ie sb e r g s p e c ie s , H . l a t i f o l i a . A se c o n d s p e c ie s , H . h e lm e i G o ld b la tt & J .C .M a n n in g , a lso k n o w n fro m a s in g le c o lle c t io n fro m th e in te rio r m o u n ta in s o f E a ste rn C a p e n ea r G ra a ffR e in e t, h a s terete le a v e s and sm a ll flo w e r s w ith te p a ls ± 8 x 2 .5 m m , ab o u t a s lo n g a s th e perianth tu b e. It is a p p a ren tly m o st c lo s e ly a llie d to th e R o g g e v e ld s p e c ie s , H . d l i o l a t a . A third n o v e lty , H . lit h ic o la J .C .M a n n in g & G o ld b la tt. re str ic te d to th e S w a rtru g g e n s ra n ge in th e ea ste r n C o ld B o k k e v e ld , h a s b e ll-s h a p e d c o r m s w ith to o th ed m a rg in s, le a v e s 1 - 2 m m w id e , and w h ite f lo w e r s w ith a tu b e 1 0 - 1 2 m m lo n g , an d a p p ea rs m o st c lo s e ly a llie d to th e w id esp r e a d H . f a l c a t a . A n e w c o lle c t io n o f H k a r o o i c a fro m n o rth e a st o f th e H a n ta m sb er g rep r ese n ts a sm a ll bu t sig n ific a n t ra n ge e x te n s io n fo r th is lo c a l en d e m ic p r e v io u s ly k n o w n fro m ju s t tw o c o lle c t io n s n ea r C a lv in ia , so u th o f th e se m o u n ta in s. T h e flo w e r s iz e , e s p e c ia lly d im e n s io n s o f th e te p a ls, c o n firm s its statu s as a sep a r a te s p e c ie s a llie d to H . v a g i n a t a . L a stly , n e w c o lle c t io n s o f th e r e la tiv e ly u n c o m m o n , y e llo w -flo w e r e d va ria n t o f H . a c u t a s h o w that th is p la n t, c o n fin e d to th e e a s te m p o rtio n o f th e ra n g e o f the s p e c ie s , d iffe r s c o n sis te n tly fro m th e w h ite -flo w e r e d fo rm in se v e r a l flo ra l fea tu res, an d it is ra ised to s u b s p e c ie s rank a s H . a c u t a su b sp . tu g H e llia e . IN T R O D U C T IO N TAXONOM Y H e s p e r a n t h a K e r G aw l. (Irid ace ae: C ro c o id ea e) is d is ­ trib u ted acro ss su b -S ah a ra n A fr ic a b u t is m o st d iv e rs e an d sp ec ies-ric h in so u th ern A frica . T h e g en u s h as tw o c e n ­ tre s o f d iv e r s ity h e re , th e w in te r ra in fa ll w e s t (G o ld b la tt 1 9 8 4 ), a n d th e c o a s t a n d a d ja c e n t m o u n ta in s o f th e e a s t­ e m , s u m m e r ra in fa ll h a l f o f th e s u b c o n tin e n t (H illia rd & B u rtt 1 9 8 6 ). In th e m o s t re c e n t re v is io n o f th e g e n u s , 79 s p e c ie s w e re re c o g n iz e d (G o ld b la tt 2 0 0 3 ). A lth o u g h H e s p e r a n t h a is n o w w e ll u n d e r s to o d ta x o n o m ic a lly , th re e c o lle c tio n s m a d e s in c e th e p u b lic a tio n o f th is re v is io n re p re s e n t n o v e ltie s , a ll fr o m a re a s o f s o u th e rn A fr ic a th a t are p o o r ly c o lle c te d . W e d e s c rib e th e m h er e. In a d d itio n , a n e w c o lle c tio n a n d im p o rta n t ra n g e e x te n ­ sio n o f th e p o o rly k n o w n H . k a r o o i c a G o ld b la tt c o n fir m th at flo w e r s ize , e s p e c ia lly d im e n s io n s o f th e te p a ls , c o n ­ s is te n tly d iffe rs fro m th a t in th e c lo s e ly a llie d H . v a g i ­ n a t a (S w e e t) G o ld b la tt, r e m o v in g d o u b ts a b o u t its s ta tu s as m e r e ly a d e p a u p e ra te fo rm o f th e latter. In H . a c u t a n e w c o lle c tio n s h a v e p ro v id e d c o n v in c in g e v id e n c e th a t th e e a s te m , y e llo w -f lo w e re d p o p u la tio n s o f th e s p e c ie s c o n s is te n tly h a v e la rg e r flo w e rs th a t d iffe r in s e v e ra l a d d itio n a l fe a tu re s fro m th e w e s te m , w h ite -f lo w e re d p o p u la tio n s . T h e y e llo w -f lo w e re d p o p u la tio n s c le a r ly c o n s titu te a s e p a ra te ra c e o f th e s p e c ie s , w h ic h w e r e c o g ­ n iz e as H . a c u t a su b s p . t u g w e l l i a e . T h e to ta l n u m b e r o f s p e c ie s in H e s p e r a n t h a is th u s in c re a s e d to 8 2: 4 3 o f th e s e o c c u r in th e s o u th e m A fr ic a n w in te r ra in f a ll z o n e ; 3 9 in th e s o u th e m A fr ic a n s u m m e r ra in fa ll z o n e s o u th o f th e L im p o p o ; a n d fo u r in tro p ic a l A fr ic a. H e s p e r a n t h a \ o n g h X y \ 2L J . C . M a n n i n g & G o l d b l a t t , sp. no v . * B .A . K rukofT C u rator o f A fr ic a n B o ta n y , M isso u r i B o ta n ic a l G a rd en , R O . B o x 2 9 9 , St. L o u is , M isso u r i 6 3 1 6 6 , U S A . E -m a il: p e te r .g o ld b la tt@ m o b o t.o r g . * * C o m p to n H erb a riu m . S o u th A fr ic a n N a tio n a l B io d iv e r s ity In stitu te , P la n ta e ± 3 0 m m alta e , c o rm o c a m p a n u la to ± 10 m m d ia m . m a rg in e s p in is ra d ia tis a r m a to , c a u le ad 8 m m lo n g o , fo liis 3 fa lc a tis 3 0 - 5 0 x 2 . 5 - 3 .0 m m , s p ic a 1- v el 2 -f lo ra , b ra c te is v irid ib u s 7 - 1 0 m m lo n g is , flo rib u s m a n ife ste p u rp u r e is a d fa u c e m tu b i fla v is , tu b o p e ria n th ii 2 0 - 2 5 m m lo n g o , te p a lis s u b a e q u a lib u s a n g u s te o v a tis p a te n tib u s , s ta m in ib u s a d s c e n d e n tib u s , a n th e r is ± 5 m m lo n g is , ra m is sty li ± 10 m m lo n g is la x e p a te n tib u s . T Y P E .— N a m ib ia , 2 7 1 6 (W itp u tz ): s o u th o f S e b ra fo n te in , 1 4 0 0 m , ( - D D ) , 2 8 A u g u s t 2 0 0 3 , P. B r u y n s 9 4 8 4 A (N B G , h o lo .). P la n ts ± 3 0 m m h ig h ; co rm b e ll- s h a p e d w ith flat b a s e , ± 10 m m d ia m . a t b a s e , m a rg in w ith r a d ia tin g s p in es . S t e m re a c h in g u p to 8 m m a b o v e g ro u n d , u n b ra n c h e d . L e a v e s 3 , fa lc a te , 3 0 - 5 0 x 2 .5 - 3 .0 m m ; c a ta p h y lls m e m ­ b ra n o u s . S p i k e 1- o r 2 -flo w e re d ; b ra c ts g re e n , 7 - 1 0 m m lo n g , o u te r a b o u t as lo n g as in n e r o r s lig h tly lo n g e r, in n e r fo rk e d a t a p e x . F l o w e r s e v id e n tly p u rp le , y e llo w in m o u th o f tu b e ; p e r ia n th tu b e 2 0 - 2 5 m m lo n g , s le n d er , w id e n in g n e a r a p e x ; te p a ls s u b e q u a l, n a r ro w ly o v a te , ± 10 X 6 m m , o u te r s lig h tly w id e r th a n in n er, s p re a d in g h o riz o n ta lly . S t a m e n s s y m m e tric a lly d is p o s e d , s lig h tly s p re a d in g ; fila m e n ts ± 4 m m lo n g , e x s e r te d ± 3 m m fr o m tu b e ; a n th e r s ± 5 m m lo n g . O v a r y o v o id , ± 1 . 5 m m lo n g ; sty le d iv id in g at m o u th o f p e r ia n th tu b e , b ra n c h e s ± 10 m m lo n g , s p re a d in g laxly . C a p s u l e s a n d s e e d s u n k n o w n . F l o w e r i n g t i m e : m id -A u g u s t to e a rly S e p te m b e r. F ig u re lA . P riv a te B a g X 7 , 7 7 3 5 C la re m o n t, C a p e T o w n . E -m a il: m a n n in g @ s a n b i.o r g . M S . re c e iv e d : 2 0 0 7 - 0 1 - 0 9 . D i s t r i b u t i o n a n d e c o l o g y : f o u n d in ro c k c re v ic e s at h ig h e r e le v a tio n s in th e m o u n ta in s o f s o u th w e s te r n B o th a lia 3 7 ,2 (2 0 0 7 ) 178 FIG U R E 1.— A , H e s p e r a n th a lo n g i­ s ty la , B r u y n s 9 4 8 4 A \ B , H . h e lm e i, H e lm e 3 1 4 4 . S ca le bars: 10 m m . Artist; J.C. M a nning. N a m ib ia n e a r R o sh P in ah , in th e w in te r ra in fa ll p a rt o f th a t c o u n try (F ig u re 2). P la n ts w e re g ro w in g a m o n g d o lo m ite ro c k s a n d in c re v ic e s o n th e s u m m it o f a h ig h rid g e , w h e re th e y w o u ld h a v e b e n e fite d fro m a n y a d d i­ tio n a l m o is tu r e th at c o n d e n s e d fro m s e a fo g. It is lik e ly th a t th e s p e c ie s o c c u rs e ls e w h e re in s u ita b le p la c e s in th is ru g g e d , la rg e ly in a c c e s s ib le , a n d b o ta n ic a lly p o o rly e x p lo re d re g io n . K n o w n o n ly fr o m th e ty p e c o lle c tio n , m a d e b y C a p e T o w n su c c u le n t s p e c ia lis t P e te r B ru y n s in 2 0 0 3 , H e s p e r a n t h a l o n g i s t y l a is a s u rp ris in g d isco v er y , r e p re s e n tin g th e o n ly s p ec ie s o f th e g e n u s k n o w n fr o m N a m ib ia . D i a g n o s i s a n d r e l a t i o n s h i p s ', d is tin c tiv e in its s h o rt s ta tu re , fa lc a te lea v e s, re la tiv e ly lo n g p e r ia n th tu b e , 2 0 25 m m lo n g , a n d d is p ro p o rtio n a te ly lo n g sty le b ra n c h e s , H e s p e r a n t h a l o n g i s t y l a h a s th e b e ll-s h a p e d c o rm w ith a flat b a s e th a t c h a ra c te riz e s a h a n d fu l o f o th e r s p e c ie s in th e g e n u s fro m th e so u th e rn A fric a n w in te r ra in fa ll z o n e to th e so u th , in c lu d in g w h ite -flo w e re d H . f a l c a t a ( L . f ) K e r G a w l. as w e ll as th e tw o p u rp le -flo w e re d sp e c ie s, H . l a t i f o l i a (K la tt) M .P .d e V os a n d H . p a n c i f l o r a (B a k e r) G .J .L e w is. T h e last tw o sp e c ie s o c c u r in N a m a q u a la n d , th e fo rn ie r re s tric te d to th e K a m ie s b e rg a n d th e la tte r re a c h in g its n o rth e rn lim it th e re , w e ll to th e so u th o f B o th a lia 3 7 ,2 (2 0 0 7 ) 17 9 F IG U R E 2 .— K n o w n d istrib u tio n o f H e s p e r a n t h a l o n g i s t y l a . A ; H . h e lm e i, tu g w e U ia e , • . N a m ib ia . N e v e rth e le s s , th e p ro x im ity o f th e ir ra n g e s , an d th e ir s im ila r flo w e r c o lo u r s u g g e s t th a t H . l o n g i s t y l a is a llie d to th e s e tw o s p e c ie s . H . l o n g i s t y l a is m o s t s im i­ lar to th e lo n g -tu b e d H . l a t i f o l i a b u t d iffe rs in its s m a lle r flo w e rs , w ith te p a ls ± 10 m m lo n g , a n d in th e m a rk e d d iffe re n c e in th e re la tiv e p r o p o r tio n s o f s ta m e n s a n d sty le. T h e fila m e n ts in H . l o n g i s t y l a are re la tiv e ly sh o rt, 3 - 4 m m lo n g a n d th e s ty le b ra n c h e s p r o je c t ± 3 m m b e y o n d th e tip s o f th e a n th e r s , th u s a p p e a rin g u n u s u a lly lo ng . T h e flo w e rs o f H . l a t i f o l i a , in c o n tra s t, a r e g e n e r­ a lly m u c h la rg e r, w ith te p a ls 1 5 -2 3 m m lo n g , th e fila ­ m en ts ( 4 - ) 7 - 1 0 m m lo n g , a n d th e sty le b r a n c h e s d o n o t e x te n d m u c h b e y o n d th e tip s o f th e an th e rs . H e s p e r a n t h a h e lm e i G o l d b l a t t & J . C . M a n n i n g , sp. n ov . P la n ta e 1 8 0 - 2 5 0 m m a lta e , c o r m o g lo b o s o a s y m m e trico ± 9 m m d ia m ., c a u le e r a m o s o in q u a r ta s u p e rio re fo liu m s q u a m ifo rm e m g e r e n te , fo liis 3 , in fe r io r ib u s d u a te re tib u s c o s ta tis , s u p e r io r e v a g in a n ti, s p ic a 1- v el 2 -flo ra, b ra c te is v irid ib u s , ± 9 m m lo n g is , flo rib u s p allid e m a lv in is , tu b o p e r ia n th ii 6 - 8 m m lo n g o , te p a lis su b e q u a lib u s a n g u s te o v a tis ± 8 x 2 .5 m m p a te n tib u s , filam en tis ex tu b o ± 2 m m e x s e rtis , a n th e r is ± 4 m m lo n g is , ra m is sty li ± 3 . 5 m m lo n g is. T Y P E .— S o u th A fr ic a , E a s te r n C a p e , 3 2 2 4 (G ra a ffR e in et): ± 5 0 k m E o f G ra a ff -R e in e t, u p p e r n o rth e r n slo p e s o f N a rd o u w s b e r g , 1 k m N o f m a in p e a k , A n n e x S tijlfo n te in 2 5 0 , 2 3 0 0 ' [7 0 0 m ], ( - B B ) , 19 N o v e m b e r 2 0 0 4 , N . A . H e l m e 3 1 4 4 (N B G , h o lo .). P la n ts 1 8 0 - 2 5 0 m m h ig h ; c o rm g lo b o s e , ± 9 m m d ia m ., a s y m m e tric , w ith c o n c e n tric tu n ic s . S t e m s le n ­ d er, u n b ra n c h e d , b e a r in g sh o rt, s c a le -lik e l e a f in u p p e r fo u rth . L e a v e s 3 , lo w e r 2 te re te , c o n s p ic u o u s ly g ro o v e d , b la d e s 8 - 1 5 x 1.2 m m , u p p e r m o s t le a f s h e a th in g lo w e r h a l f to tw o th ird s o f s te m ; c a ta p h y lls m e m b ra n o u s . S p i k e 1 - o r 2 -flo w e re d ; b ra c ts g re e n , ± 9 m m lo n g , in n e r a b o u t as lo n g as o u te r a n d fo r k e d at a p e x . F l o w e r s p a le m a u v e . k; H . U th ic o la , ■ ; H . a c u t a su b sp . a c u t a , O ; an d H . a c u t a su b sp . d a r k e r at m o u th o f tu b e , o u te r te p a ls w ith b ro a d p u rp le s tre a k o n re v e rs e ; p e ria n th tu b e 6 - 8 m m lo n g , s le n d e r, w id e n in g n e a r a p e x ; te p a ls s u b e q u a l, n a r r o w ly o v a te , ± 8 .0 X 2 .5 m m , o u te r s lig h tly la rg e r th a n in n er, s p r e a d ­ in g s lig h tly a b o v e h o riz o n ta l. S t a m e n s s y m m e tric a lly d is p o s e d , s lig h tly s p re a d in g ; fila m e n ts e x s e rte d ± 2 m m fro m tu b e ; a n th e r s ± 4 m m lo n g . O v a i y o v o id , ± 2 m m lo n g ; s ty le d iv id in g at m o u th o f p e r ia n th tu b e , b ra n c h e s ± 3 .5 m m lo n g , a s c e n d in g , u ltim a te ly re a c h in g to a b o u t m id d le o f an th e rs . C a p s u l e s a n d s e e d s u n k n o w n . F l o w e r i n g t i m e : N o v e m b e r a n d p r o b a b ly D e ce m b er . F ig u re IB . D i s t r i b u t i o n a n d e c o l o g y , a p p a r e n tly re s tr ic te d to th e s o u th e rn e d g e o f th e in te rio r e s c a rp m e n t, w h e re it is k n o w n fro m a s in g le c o lle c tio n m a d e at h ig h e le v a tio n s in r o c k y g ra s s la n d in th e S n e e u b e rg e , e a s t o f G ra a ffR e in e t (F ig u re 2 ). T h e p la n ts w e re f o u n d in o p e n a re a s in M e r x m u e l l e r a g ra s s la n d a m o n g d o le rite b o u ld e rs . T h e s e g ra s s la n d s b u m r e g u la rly e v e ry fe w y e a rs d u rin g th e d ry s e a s o n a n d a re re g u la rly c o v e re d w ith s n o w in w in te r (N .A . H e lm e p e rs . c o m m .) T h e tip s o f th e lo n g , b a s a l le a v e s in a ll th e s p e c im e n s h a v e b e e n b u rn e d o r fr o s te d o ff, in d ic a tin g th a t th e s p e c ie s b e g in s le a fin g e a rly in th e g ro w in g s e a s o n w h e n th e d a n g e r o f fire o r fro st is n o t y e t o v er. D i a g n o s i s a n d r e l a t i o n s h i p s : H e s p e r a n t h a h e l m e i is n a m e d fo r its d is c o v e re r, th e C a p e T o w n e c o lo g is t a n d e n e rg e tic a n d w id e - ra n g in g p la n t c o lle c to r , N .A . H e lm e . T h e s p e c ie s is d is tin c tiv e in its r e la tiv e ly sm a ll, p a le m a u v e flo w e r w ith a p e r ia n th tu b e 6 - 8 m m lo n g , n a r ro w te p a ls ± 8 .0 x 2 .5 m m , a n d te re te , fin e ly rib b e d lea v e s. A n o th e r u n u s u a l a ttrib u te o f th e s p e c ie s is th e s m a ll, s h e a th in g , s c a le -lik e le a f in th e u p p e r p a r t o f th e stem . T h is d is tin c tiv e fe a tu re is c h a ra c te ris tic o f a s m a ll g ro u p o f s p e c ie s a llie d to H . p i l o s a (L .f.) K e r G a w l., all n a tiv e to th e w in te r ra in f a ll z o n e o f s o u th e r n A fric a . A m o n g th e s e s p e c ie s , H . p i l o s a is la rg e ly a s p e c ie s o f th e C a p e flo ris tic re g io n , w h e re a s H . c i l i o l a t a G o ld b la tt, H . t e r e t i f o l i a G o ld b la tt a n d th e h a ir y - le a v e d H . p s e u d o p i l o s a B o th a lia 3 7 ,2 (2 0 0 7 ) 18 0 G o ld b la tt a re c e n tr e d o n th e R o g g e v e ld E s c a rp m e n t o f th e W e ste rn K a ro o (G o ld b la tt 1 98 7). A m o n g th e s p e ­ c ie s in th is g ro u p , H . h e l m e i is p ro b a b ly m o s t c lo s e ly r e la te d to H . c i l i o l a t a a n d H . t e r e t i f o l i a , w h ic h a lso h a v e te re te , p ro m in e n tly rid g e d lea v e s, th e g ro o v e s lin e d w ith fin e c ilia te h a irs th a t ar e ab s e n t in H . h e l m e i . T h e flo w ­ e rs o f H . c i l i o l a t a a n d H . t e r e t i f o l i a are s lig h tly la rg e r th a n th o s e o f H . h e l m e i , w ith a p e ria n th tu b e ± 8 m m lo n g a n d te p a ls ± 10 m m lo ng . H e s p e r a n t h a t e r e t i f o l i a in p a r tic u la r is b ro a d ly s im ila r to H . h e l m e i in g e n e ra l a p p e a ra n c e , b u t h a s m in u te ly c ilio la te le a f rib s, a sm all, s c a le -lik e lea f, 2 - 4 m m lo n g , a n d la rg e r a n th e rs , ± 8 m m lo n g . H . h e l m e i , in co n tra s t, h as g la b ro u s le a v e s, a la rg e r s c a le -lik e lea f, 8 - 1 0 m m lo n g , a n d s m a lle r a n th e rs , ± 4 m m lo n g . W h ile th e p re s e n c e o f an e n d e m ic sp e c ie s o f H e s p e ­ r a n t h a in th e S n e e u b e rg e is su rp ris in g , it is n o t w ith o u t p re c e d e n t in th e Irid a c e a e . A h a n d fu l o f sp e c ie s o f th e fa m ily o c c u r th e re , in c lu d in g ta x a fr o m b o th th e w in te r ra in fa ll z o n e to th e w e st a n d th e s u m m e r ra in fa ll z o n e to th e ea st. T h o s e fr o m th e w in te r ra in fa ll z o n e in c lu d e M o r a e a c ilia ta , M . c r is p a and M im g u ic u la ta . T he Irid a c e a e fr o m th e s u m m e r ra in fa ll z o n e in c lu d e o n e D i e r a m a s p ec ie s, R o m u l e a m a c o w a n i i , th re e sp e c ie s o f S y r i n g o d e a , in c lu d in g th e n e a r e n d e m ic S . p u l c h e l l a , a n d a d is tin c tiv e s h o rt, b ro a d -le a v e d fo rm o f B a b i a n a b a i n e s i i . T h is m ix o f s u m m e r a n d w in te r ra in fa ll s p ec ie s re fle cts th e tra n s itio n a l p o s itio n o f th e s e m o u n ta in s , th a t h av e a g r a s s la n d -d o m in a te d flo ra c h a ra c te ris tic o f th e s u m m e r ra in f a ll re g io n at h ig h e r e le v a tio n s b u t w h ic h re c e iv e p r e d ic ta b le a m o u n ts o f w in te r ra in fa ll th a t p e rm it th e p e r s is te n c e o f a h a n d fu l o f sp e c ie s fr o m th e w in te r ra in fa ll z o n e to th e w e st. H e s p e r a n t h a lith ic o la J . C . M a n n i n g & G o l d b l a t t , sp. nov. P lan tae ( 4 0 - ) 8 0 - 2 0 0 ( - 2 9 0 ) m m altae, co rm o ca m p an u lato , 7 - 1 0 m m d iam ., ad b ase m d en tato , ca u le era m o so , foliis 4 , in fe rio rib u s 2 su b erectis v el falc atis ( 3 0 - ) 5 0 1 0 0 (- 1 6 0 ) X 1 -2 m m , sp ic a 1 ^ - f l o r a , flo rib u s alb is, te p a lis e x te r io rib u s e x tra ru b ris, tu b o p e ria n th ii 1 0 -1 2 m m lo n g is, te p a ls s u b a e q u a lib u s, 1 0 -1 5 x 4 .0 - 6 .5 m m , fila m e n tis 3 - 5 m m lo n g is, a n th e ris ± 6 m m lo n g is, ra m is sty li 4 .0 - 5 .5 m m lo n g is. T Y P E .— S o u th A fric a , W e ste rn C a p e , 3 2 1 9 (W u p p e r­ tal): S w a rtru g g e n s , F arm K n o lfo n te in , b e w e e n K a tb a k k ies P a ss a n d S k itte ry k lo o f, in lo am a n d l e a f litte r o n s a n d s to n e p a v e m e n t, ( - D C ) , 12 S e p te m b e r 2 0 0 6 , P. G o l d b l a t t cfe J . C M a n n i n g 1 2 8 1 7 A (N B G , h o lo .; M O , P R E , iso .). P lan ts ( 4 0 - ) 8 0 - 2 0 0 ( - 2 9 0 ) m m h ig h ; c o rm b e ll-s h a p e d w ith flat b a s e , 7 - 1 0 m m d ia m ., b re a k in g in to s e g m e n ts b elo w , m a rg in s fr in g e d a n d o ften w ith ra d ia tin g sp in es. S t e m u n b ra n c h e d o r ra re ly w ith o n e b ra n c h fr o m n e a r b as e . L e a v e s 4 , lo w e r 2 b as a l, s u b e re c t o r fa lc a te , o ften u n d u la te o r lo o s e ly c o ile d , m id rib p ro m in e n tly ra is e d an d m a rg in s s q u a re d a n d m in u te ly s c a b rid a lo n g b o th e d g e s, b la d e s ( 3 0 - ) 5 0 - 1 0 0 ( - 1 6 0 ) x 1 -2 m m , u p p e r­ m o s t le a f e n tire ly sh e a th in g ; c a ta p h y lls m e m b ra n o u s. S p i k e 1^ - f l o w e r e d ; b ra c ts g re en , flu sh e d re d d is h a n d b e c o m in g d ry ap ic a lly , 1 1 -2 0 m m lo n g , in n e r a b o u t as lo n g as o u te r a n d fo rk e d at ap e x . F l o w e r s w h ite , o u te r te p a ls re d o u tsid e , o p e n in g in e v e n in g a n d p ro d u c in g r o s y c itru s fr a g ra n c e ; p e ria n th tu b e 1 0 -1 2 m m lo n g , n o t o r s lig h tly e x s e rte d b e y o n d b ra c ts , sle n d e r, w id e n ­ in g n e a r ap ex ; te p a ls s u b e q u a l, e llip tic , 1 0 -1 5 x 4 .0 - 6 .5 m m , o u te r s lig h tly la rg e r th a n in n er, s p re a d in g h o riz o n ­ ta lly w h e n fu lly o p en . S t a m e n s s y m m e tric a lly d is p o s e d , s lig h tly s p re a d in g ; filam en ts 3 - 5 m m , e x s e rte d 2 - 3 m m fr o m tu b e ; a n th e rs h o riz o n ta l, p a le y e llo w , ± 6 m m lo n g . O v a r y ' o v o id , ± 3 m m lo n g ; sty le d iv id in g at m o u th o f p e r ia n th tu b e, b ra n c h e s 4 .0 - 5 .5 m m lo n g . C a p s u l e s e llip s o id , 9 - 1 0 x 4 m m . S e e d s s u b g lo b o s e w ith fla tte n e d c h a la z a a n d p e rs is te n t ftin icle , g o ld e n b ro w n , 1 .0 -1 .5 m m d ia m ., ru g o s e , te s ta ce lls d o m e d . F l o w e r i n g t i m e : S e p te m b e r. F ig u re 3. D i s t r i b u t i o n a n d e c o l o g y ’: k n o w n fro m s a n d s to n e ro c k p a v e m e n t in th e S w a rtru g g e n s M o u n ta in s in th e e a s te rn C o ld B o k k e v e ld , w h e re th e p la n ts g ro w in ro c k c re v ic e s o r s h a llo w h u m u s , o fte n in th e s h e lte r o f sm all s h ru b s (F ig u re 2). D ia g n o s is a n d r e l a t i o n s h i p s '. H e s p e r a n th a lith ic o la is d is tin c tiv e in its s y m m e tric a l, b e ll-s h a p e d c o rm a n d n a rro w , o fte n tw is te d o r c o ile d lea v e s. It is p ro b a b ly m o s t c lo s e ly a llie d to H . f a l c a t a , w h ic h it re s e m b le s in g e n e ra l a p p e a ra n c e , b u t fr o m w h ic h it d iffe rs in its c o n ­ s is te n tly n a r ro w e r lea v e s, 1 -2 m m w id e (v s. 4 - 8 m m in H . f a l c a t a ) , an d lo n g e r p e r ia n th tu b e , 1 0 -1 2 m m v s. 4 - 9 m m . T h e tw o s p ec ie s d iffe r also in th e ir h a b ita ts . H e s p e r a n t h a l i t h i c o l a is r e s tr ic te d to s h a llo w so ils o n s a n d s to n e ro c k p av e m e n t, w h e re a s H . f a l c a t a is a c o m ­ m o n a n d w id e s p re a d s p e c ie s o f s e a s o n a lly m o is t, d e e p e r c la y o r s a n d y so ils. T h is e c o lo g ic a l d iffe re n c e is v e ry e v id e n t at th e ty p e lo cality , w h e re H . f a l c a t a o c c u rs o n s a n d y flats n o t fa r fro m ro c k y o u tc ro p s to w h ic h H . l i t h i ­ c o l a is re s tric te d . H e s p e r a n t h a l i t h i c o l a is re m a rk a b ly s im ila r in g e n ­ e ra l a p p e a ra n c e to H . a c u t a (L ic h t. ex R o em . & S ch u lt.) K e r G a w l. b u t th is sp e c ie s h a s an a s y m m e tric a l co rm w ith o n e sid e flatten e d a n d e x te n d e d d o w n w a rd . W ith o u t c o rm s , h o w e v e r, th e tw o ta x a ar e e s s e n tia lly in d is tin ­ g u is h a b le , a lth o u g h H . a c u t a d o e s n o t o c c u r o n th e C o ld B o k k e v e ld a n d is ty p ic a lly fo u n d o n c la y s o ils, ra re ly lim e s to n e o r in p o c k e ts o f lo am o n s a n d sto n e o u tc ro p s . O th e r m a te r ia l e x a m in e d W ESTERN C A P E .— 3 2 1 9 (W up p ertal): K a tb a k k ies e x p o s e d sa n d sto n e slo p e , ( - D C ) , 2 0 S ep te m b er 1 9 9 1 , C 139 F a m i. R e id 1 3 8 9 ( N B G , P R E ); S w a rtru g g en s, 6 0 k m N E o f C er e s, K n o lfo n te in , 1 2 0 5 m , ( - D C ) , 14 S e p te m b er 2 0 0 5 , /. & C . J a r d i n e 1 2 6 (N B G ); G r o en fo n tein , Z e e k o e g a t, w e s t o f R iet R iv er, 9 0 0 m , ( - D C ) , 15 S ep te m b e r 2 0 0 1 , M .S . S t o b i e 7 ( N B G ). T A X O N O M IC N O T E S H e s p e r a n t h a k a r o o i c a G o ld b la tt K n o w n fro m ju s t a h an d fu l o f p la n ts c o lle c te d at tw o lo c a tio n s a ro u n d C alv in ia in N o rth e rn C ap e, H e s p e r a n t h a k a r o o i c a is re c o g n iz e d a m o n g th e sp e c ie s o f th e g e n u s w ith larg e , a s y m m e tric c o rm s w ith o v e rla p p in g tu n ic s b y its lo w s ta tu re , 3 0 - 5 0 m m h ig h , a n d sp ik e s o f 1 o r 2 y e llo w flo w e rs w ith a tu b e ± 5 m m lo n g a n d te p a ls ± 2 0 m m lo n g (G o ld b la tt 1984, 2 0 0 3 ). In g e n e ra l a s p e c t B o th a lia 3 7 ,2 ( 2 0 0 7 ) 181 2 0 0 3 ), th e re h a s r e m a in e d s o m e d o u b t as to w h e th e r th e y a re r e a lly d is tin c t. N e w c o lle c tio n s o f H e s p e r a n t h a v a g i n a t a fro m n e a r L o e r ie s fo n te in a n d o f y e llo w -f lo w e re d p la n ts th a t a c c o rd w ith H . k a r o o i c a fro m w e ll to th e e a s t o f C a lv in ia , c o n ­ trib u te to o u r u n d e r s ta n d in g o f th e tw o ta x a . T h e la tte r p o p u la tio n , f o u n d s o m e 50 k m N N E o f C a lv in ia o n th e ro a d to K lip w e r f in th e a re a k n o w n a s th e A g te r H a n ta m { G o l d b l a t t & P o r t e r 1 2 7 4 7 ) , c o n s is ts o f p la n ts w ith b rig h t y e llo w flo w e rs , th e o u te r te p a ls s h a d e d b r o w n o u t­ s id e, a n d r a n g in g in h e ig h t fr o m 50 to 100 m m , w ith o n e p la n t f o u n d in th e s h a d e o f a lo w s h ru b a tta in in g a h e ig h t o f 160 m m . T h e flo w e rs h a v e a p e r ia n th tu b e 6 - 7 m m lo n g , th u s lo n g e r th a n u n til n o w r e p o r te d fo r H . k a r o o ­ i c a a n d m a tc h in g H . v a g i n a t a , b u t th e te p a ls ar e ± 20 X 9 - 1 0 m m lo n g a n d th e a n th e r s ± 10 m m lo n g , th u s ± as re c o rd e d fo r th e tw o o th e r c o lle c tio n s o f H . k a r o o i c a ( G o ld b la tt 1984). T h e c o lle c tio n o f H e s p e r a n t h a v a g i n a t a fr o m n e a r L o e r ie s fo n te in { G o l d b l a t t & P o r t e r 1 2 7 7 5 ) , fr o m th e F a rm R ie tfo n te in , a n d w h ic h c o m p r is e s a m o d e s t ra n g e e x te n s io n (G o ld b la tt 1984), c o n s is ts o f p la n ts 5 0 -1 2 0 m m h ig h , th e s m a lle s t o f th e s e u n u s u a l fo r th e sp e c ie s . A ll in d iv id u a ls h a v e flo w e rs w ith th e b r o w n m a rk in g s o n th e in s id e o f th e te p a ls th a t a re ty p ic a l o f th e s p e c ie s a n d a n th e r s th a t a re ± 12 m m lo n g a n d te p a ls 3 0 x ± 12 m m . E v e n th e s m a lle s t p la n ts h a v e flo w e rs ty p ic a l o f th e s p e c ie s . W e c o n c lu d e th a t p la n t h e ig h t a n d p e r ia n th tu b e le n g th a re n o t fa c to rs th a t c o n s is te n tly d is tin g u is h th e tw o s p e c ie s , b u t th a t a n th e r le n g th a n d te p a l s iz e a lw a y s d iffe r. T h e te p a l s iz e e s p e c ia lly is s u b s ta n tia lly s m a lle r in H e s p e r a n t h a k a r o o i c a . B o th th is s p e c ie s a n d H . v a g i n a t a ty p ic a lly g ro w o n h e a v y re d c la y d e r iv e d fr o m d o le rite , a lth o u g h p o p u la tio n s o f H . v a g i n a t a fr o m th e w e s t o f its ra n g e a ls o o c c u r o n tillite -d e riv e d c la y o r a m ix o f c la y a n d s a n d d e r iv e d fr o m T a b le M o u n ta in S a n d s to n e . R a n g e e x te n s io n s H e s p e r a n th a k a r o o ic a NORTHERN C A P E .— 3 1 1 9 (C a lv in ia ): A g te r H antam . NNE of C a lv in ia on road b e tw e e n M o o rd en a a rsp o o rt an d K lip w e r f, sto n y d o l­ er ite c la y fla ts . ( - B B ) , 1 S e p te m b er 2 0 0 6 , G o l d b l a t t & P o r t e r 1 2 7 4 7 (M O , N B G , P R E ). H e s p e r a n th a v a g in a ta N O R T H E R N C A P E .— 3 1 1 9 (C a lv in ia ): Farm R ie tfo n te in , ± 4 8 km F IG U R E 3 .— H e s p e r a n t h a l i t h ic o la , G o l d b l a t t & M a n n i n g 1 2 8 1 7 A . A , w h o le p la n t; B , flo w e r; C , c a p su le; D , s e e d . S c a le bars: A - C , 10 S E o f L o e r ie sfo n te in , lo w d o ler ite c la y h ill. ( - B C ) , 5 S e p te m b e r 2 0 0 6 . G o l d b l a t t & P o r t e r 1 2 7 7 5 (M O . N B G , P R E ). m m ; D , 1 m m . A rtist: J.C . M a n n in g . H e s p e r a n t h a a c u t a (L icht. ex R oem . & S ch u lt.) K e r G aw l. th e s p e c ie s is v e r y lik e th e m o re w id e s p r e a d B o k k e v e ld P la te a u s p e c ie s , H . v a g i n a t a (S w e e t) G o ld b la tt, w h ic h is n o rm a lly ta lle r, 1 2 0 - 1 8 0 m m h ig h , a n d h a s s p ik e s o f ( 1 ) 2 - 4 y e llo w flo w e rs , u s u a lly m a rk e d w ith c o n tra s tin g b ro w n m a rk in g s in th e c e n tr e a n d o n th e tip s o f th e o u te r tep als. P la in y e llo w -f lo w e re d p la n ts a re ^ o w n fro m a few site s b u t b o th th e s e a n d p la n ts w ith m a r k e d te p a ls h a v e a p e r ia n th tu b e 5 - 8 m m lo n g a n d te p a ls 3 0 - 3 5 x 1 5 -1 7 m m . A lth o u g h th e tw o s p e c ie s h a v e b e e n r e c o g ­ n iz e d in re c e n t r e v is io n s o f th e g e n u s (G o ld b la tt 1 9 8 4, N e w a n d w e ll-p r e s e r v e d c o lle c tio n s o f th e y e llo w flo w e re d v a r ia n t o f H e s p e r a n t h a a c u t a ( G o ld b la tt 1984), u n til n o w b e lie v e d to b e ra re , h a v e p r o v id e d u s w ith a d d itio n a l in fo rm a tio n a b o u t th e s p e c ie s . T y p ic a l w h ite flo w e re d H . a c u t a is c o m m o n in th e w e s te rn K a ro o , a n d b lo o m s p ro lific a lly in y e a rs o f a d e q u a te ra in fa ll. T h e flo w e rs , in c lu d in g th e a n th e r s , a r e w h ite , a lth o u g h th e o u te r te p a ls a re b ro w n to re d o n th e o u ts id e . C a re fu l m e a s u r e m e n ts o f th e s ta m e n s a n d s ty le s s h o w th a t th e B o th a lia 3 7 ,2 (2 0 0 7 ) 182 T A B L E 1.— C o m p a riso n o f y e llo w - and w h ite -flo w e re d variants o f H e s p e r a n t h a a c u ta . O n ly w e ll-p r e ss e d , fu lly o p e n flo w e r s w e r e m e a ­ su re d C a p e ], P rin c e A lb e rt, T u g w e l l s . n . { N a t . B o t . G a r d . i l 5 1 / 2 9 in K , h o lo .!, B O L , M O , iso .!). S e le c te d s p e c im e n s P erian th c o lo u r Y ello w W h ite Perian th tu b e le n g th (m m ) 1 2 .5 -1 5 .0 8 .5 - 1 2 .0 T epal le n g th (m m ) 1 2 .5 -1 6 .0 ( 9 .5 - ) 1 1 .0 -1 6 .0 W e e k s P oort a lo n g L a d ism ith -L a in g b u r g road , ( - A C ) , 9 S ep te m b e r A n th er le n g th (m m ) ^8 5 .3 - 7 .0 F ila m en t e x s e r tio n (m m ) ±4 1 .5 -2 .5 1 9 7 1 , T h o m a s s .n { N B G 9 3 8 2 I ) \ S w artb erg, b e tw e e n M a tje sv le i an d G a m k a sp o o rt, ( - B C ) , O ct. 1 9 8 6 , V lo k s .n . (M O ); S w artb erg S ty le tip le v e l ± m id d le o f anthers 2 m m b e lo w to 1 m m a b o v e anther tips W E S T E R N C A P E .— 3 3 2 1 (L a d ism ith ): 3 0 m ile s b e y o n d S e v e n M tn s. road to G a m k a sk lo o f, ( - B D ) , 9 S ep te m b e r 2 0 0 1 , G o l d b l a t t & P o r t e r 1 1 8 5 9 (M O , N B G ). 3 3 2 2 (O u d tsh o o m ): P rin ce A lb ert, Farm K le in sle u te lfo n te in , ( - A B ) , 1 A u g u st 1 9 9 5 , M a r i n c o w i t z s u b S n ijm a n 1 4 9 2 (N B G ); K a reed o u w P a ss b e tw e e n K la arstro om an d P rin c e A lb ert, fila m e n ts a re e x s e rte d 1 .5 -2 .5 m m fro m th e tu b e a n d th a t th e sty le b ra n c h e s re a c h fro m ± 2 m m b e lo w th e a n th e r tip s to 1 m m a b o v e th em . T h e tu b e its e lf is 8 .5 - 1 2 .0 m m lo n g a n d th e te p a ls 9 .5 - 1 6 .0 m m lo n g (T a b le 1). P la n ts w ith flo w e rs fa llin g in th e u p p e r ra n g e o f th e s e d im e n ­ s io n s ar e m o st c o m m o n a lo n g th e C e d e rb e rg b u t a ls o o c c u r o v e r th e e n tire ra n g e o f th e w h ite -flo w e re d v a ria n t o f th e sp e c ie s (F ig u re 2 ). S m a lle r-flo w e re d p la n ts o c c u r m a in ly in th e n o rth w e s t o f th e ra n g e , a lo n g th e fo o th ills o f th e B o k k e v e ld a n d M a ts ik a m m a -G if l^ e rg M o u n ta in s. Y e llo w -flo w e re d p la n ts , re fe rre d b y R .C . F o s te r (1 9 4 8 ) to H . t u g w e l l i a e , o c c u r in th e ea s t o f th e ra n g e o f H . a c u t a , in th e n o rth e r n fo o th ills o f th e S w a rtb e rg M o u n ta in s , alo n g th e K a m m a n a s s ie M o u n ta in s , a n d in th e L o n g K lo o f to th e so u th in th e v ic in ity o f J o u b e r tin a (F ig u re 2). N e w c o lle c tio n s m a d e sin ce 2 0 0 2 (lis te d b e lo w ) sh o w th a t p la n ts h a v e a p e ria n th tu b e 1 2 .5 -1 5 .0 m m lo n g a n d te p a ls 1 2 .5 - 1 6 .0 m m lo n g , th u s c lo s e ly m a tc h in g th e w h ite -f lo w e re d p la n ts , b u t th e fila m e n ts a re e x s e rte d ± 4 m m a n d th e a n th e rs are 6 - 8 m m lo n g , w h e re a s th e sty le b ra n c h e s re a c h o n ly to a b o u t th e m id d le o f th e a n th e rs (T a b le 1). T h e s e c o n s is te n t d iffe re n c e s in th e d im e n s io n s o f a n th e rs a n d s ty le p ro v id e p ro o f, in a d d itio n to flo w e r co lo u r, th a t th e e a s te rn p o p u la tio n s c o n s titu te a s e p a ­ ra te , la rg e r-flo w e re d ra ce , a n d w e b e lie v e it is u s e fu l to re c o g n iz e th e y e llo w -flo w e re d p o p u la tio n s as a se p a ra te ta x o n . B e c a u s e o f th e sm all d iffe re n c e s b e tw e e n th e tw o , a n d c o m p le m e n ta r y g e o g ra p h ic ra n g e s , w e re c o m m e n d s u b s p e c ie s ra n k fo r th e ta x o n , a n d th u s m a k e th e c o m b i­ n a tio n H . a c u t a su b sp . t u g w e l l i a e . H e s p e r a n t h a a c u t a su b sp . tu g w e llia e ( R . C . F o s t e r ) G o l d b l a t t & J . C . M a n n i n g , c o m b , et sta t nov. H e s p e r a n ­ t h a t u g w e l l i a e R .C .F o s te r, C o n trib u tio n s fr o m th e G ra y H e rb a riu m 166: 2 6 (1 9 4 8 ). T ype: S o u th A fric a , [W e stern ( - A D ) , 10 S e p te m b er 2 0 0 2 , G o l d b l a t t & P o r t e r 1 2 1 9 1 (M O , N B G ); G ro o t D o o m R iv er, ( - C D ) , A u g u st 1 9 3 1 , T h o r n e s .n . (S A M S 1 7 1 0 ) . 3 3 2 3 (W illo w m o r e ): ea stern slo p e s o f K a m m a n a ssie M tn s, Farm W ild ep a a r d efo n tein , ( - C A ) , 18 A u g u st 1 9 8 6 , V iv ie r s & V lo k s.n . (M O ); K a m m a n a ssie M tn s, lo w e r ste e p slo p e s a b o v e B uflfelsd rif, ( - D B ) , 8 A u g u st 1 9 8 3 , M a t t h e w s 1 1 2 8 (N B G ). E A S T E R N C A P E .— 3 3 2 3 (W illo w m o r e ): 8 km w e s t o f J o u b ertina , ( - D D ) , 2 4 A u g u st 1 9 8 1 , S n ijm a n 4 5 0 (N B G ); h ill north o f J o u b ertina , ( - D D ) , A u g u st 1 9 2 3 , F o u r c a d e 2 6 9 0 (B O L , K , N B G ). ACKNOW LEDGEM ENTS S u p p o rt fo r th is w o rk b y g ra n ts 7 3 1 6 -0 2 a n d 7 7 9 9 0 5 fro m th e N a tio n a l G e o g ra p h ic S o c ie ty is g ra te fu lly a c k n o w le d g e d . W e th a n k P e te r B ru y n s, N ic h o la s H e lm e , a n d Iv o r a n d C o ra J a rd in e fo r d ra w in g o u r a tte n tio n to th e n o v e ltie s d e s c rib e d h ere. W e a ls o a c k n o w le d g e o u r d e b t to In g rid N a n n i a n d L e n d o n P o rte r fo r th e ir a s s is ­ ta n c e a n d c o m p a n io n s h ip in th e fie ld a n d to R o y G e re a u fo r c h e c k in g o u r L a tin d e s c rip tio n s . C o lle c tin g p e r m its w e re p ro v id e d b y th e N a tu re C o n s e rv a tio n a u th o ritie s o f W e ste rn C ap e P ro v in c e , S o u th A fr ic a . REFERENCES F O S T E R , R .C . 19 48. S tu d ie s in th e Irid a cea e V. S o m e n e w o r n o te ­ w o rth y s p e c ie s o f H e s p e r a n t h a . C o n t r i b u t i o n s f r o m th e G ra y' H e r b a r i u m 166: 3 - 2 7 . G O L D B L A T T , P. 1 9 8 4 . A r e v isio n o f H e s p e r a n t h a (Irid a ce a e ) in the w in te r ra in fa ll area o f sou th ern A frica . J o u r n a l o f S o u th A f r i c a n B o ta n y 50: 1 5 -1 4 1 . G O L D B L A T T , P. 198 7. N e w sp e c ie s an d n o te s o n so u th ern A fric a n H e s p e r a n t h a (Irid a cea e). S o u th A f r i c a n J o u r n a l o f B o t a n y 53: 459^63. G O L D B L A T T , P. 2 0 0 3 . A sy n o p tic r e v ie w o f th e A fric a n g e n u s H e s p e r a n t h a (Irid aceae: C r o c o id e a e ). A n n a l s o f th e M i s s o u r i B o t a n i c a l G a r d e n 90: 3 9 0 - 4 4 3 . H IL L IA R D , O .M . & B U R T T , B .L . 1 9 8 6 . H e s p e r a n t h a (Irid a cea e ) in N atal and nearby. N o t e s f r o m E d in b u r g h 43 : 4 0 7 ^ 3 8 . th e R o y a l B o t a n i c G a r d e n