B o th a lia 19,2: 1 7 9 - 1 8 2 (19 89 )
A re-exam ination o f the gen us A m p h ib o lia (M esem bryanthem aceae)
H .E .K . H A R T M A N N * a n d M . D E H N *
K e y w o r d s : A m p h i b o l i a , le c to ty p ific a tio n , M e s e m b r y a n th e m a c e a e
ABSTRACT
T h e h o lo ty p e s o f all fiv e s p e c ie s p la ce d in the g e n u s A m p h i b o l i a L . B o l. ex T o elk en & J e s so p have b e e n e x a m in e d . O n ly
tw o o f th e m a g re e w ith th e g e n e r ic d ia g n o s is , b ut th ey are n ot c o n g e n e r ic . T h e ty p e s p e c ie s c h o s e n by H err e is sh o w n to
b e in se rio u s c o n flic t w ith the p rotologu e and a n ew lectoty p e (A . m a rir im a L. B o l. ex T oelk en & Jessop ) is therefore d esign ated .
T h e o th e r fou r s p e c ie s are e x c lu d e d from A m p h i b o l i a .
U IT T R E K S E L
D ie h o lo tip e s van al v y f s p e s ie s w at in d ie g e n u s A m p h i b o l i a L . B o l. ex T o elk en & J e s so p g e p la a s w or d , is o n d e r so e k .
S le g s tw e e van h u lle stem o o r e e n m et d ie d ia g n o s e van d ie g e n u s, m aar h u lle is n ie k o n g e n e r ie s n ie. D aar w ord aa n g etoon
dat d ie tip e s p e s ie w at d eu r H erre g e k ie s is , e m s tig m et d ie p r o to lo o g in stryd is , e n d aar om w ord 'n n u w e le k to tip e (A .
m a r i t i m a L . B o l. ex T o e lk en & J e s so p ) aa n g ew y s. D ie a n d er v ie r s p e s ie s w ord u it A m p h i b o l i a u itg e slu it.
(C R ) at th e d is ta l e n d o f th e c o v e r in g m e m b r a n e s (F ig u r e
IN T R O D U C T IO N
IB ). T h e r e m a in in g th r ee s p e c ie s la c k c lo s in g b o d ie s , b u t
In th e c o u r s e o f a c o m p r e h e n s iv e stu d y o f th e su b trib e
tw o o f th em (A . h a llii and A . g y d o u w e n s i s ) p o s s e s s c lo s in g
R u s c h iin a e (M . D e h n ) an d th e p rep a ra tio n o f a su r v e y o f
ro d le ts (C R in F ig u r e I D ). A . s t a y n e r i sh o w s n o c lo s in g
th e g en er a o f th e A iz o a c e a e ( H .E .K . H a rtm a n n ), th e g e n u s
d e v ic e at a ll.
A m p h i b o l i a L . B o l. e x H e rr e (T o elk e n & J e s s o p 1976) h as
b e e n r e -e x a m in e d . T h e r e su lts an d ta x o n o m ic c o n s e
It s e e m s th at in th e la st th ree s p e c ie s m e n tio n e d th e
p r o m in e n t fr e e e n d o f th e p la c e n ta (P L in F ig u r e ID ) h as
q u e n c e s ar e p r e s e n te d .
b e e n u n d er stoo d to re p r esen t a c lo s in g b o d y (c o m p a r e data
A m p h i b o l i a w a s fir st d e s c r ib e d b y B o lu s (1 9 6 5 ). It w a s
fr o m d e sc rip tio n s in T ab le 1). T h is in terp retation , h ow ever,
le c t o ty p if ie d a n d th u s v a lid a ted b y H e rr e (1971). T o e lk en
h a s to b e r e je c te d . T h e rele va n t p la c e n ta l stru ctu re is n ot
& J e s s o p (1976) r e v ie w e d th e n o m e n c la tu r e a n d su p p lie d
in d ir e c t c o n ta c t w ith th e ta n g e n tia l e n d o c a r p , a s w o u ld
c o r r e c t n a m e s for a ll fiv e s p e c ie s in c lu d e d in th e g e n u s
b e n e c e s s a r y fo r an e n d o c a r p a l c lo s in g b o d y (H a rtm a n n
& L ie d e 1 9 8 6 ). F u r th e r m o r e , fu n ic le s o c c u r u p to th e tip
at th at tim e .
o f th e p la c e n ta (F in F ig u r e 2 ) , p r e c lu d in g , fo r la c k o f
T h e h isto r y o f th e g e n u s sta rted e a r lie r , th o u g h , w h e n
B o lu s (19 62) n o tic e d that th e tw o s p e c ie s sh e h ad d e sc rib ed
in th e g e n u s S t o e b e r i a S ch w a n t. (S. h a l l i i , S. l i t t l e w o o d i i )
d id n ot c o n fo r m w ith th e g e n e r ic ch a ra c ters o f that g e n u s ,
‘a n e w g e n u s is th e re fo r e re q u ir ed for th e m . . . ’ (I.e . p. 14).
T h e d ia g n o s is o f th at n e w g e n u s , n a m e ly A m p h i b o l i a
(B o lu s 1 9 6 5 ), g iv e s th r ee ch a rac ters: p e r e n n ia l, fru its w ith
w in g e d v a lv e s (lik e in L a m p r a n th u s ) an d c lo s in g b o d ie s
(lik e in R u s c h ia an d S t o e b e r i a ) . In a d d itio n B o lu s stated
that th e c a p s u le s d iffe r fro m th o s e o f S t o e b e r i a in so fa r a s
th ey o p e n sp o n ta n e o u sly an d d o n ot c lo s e ag ain . N o fu rth er
featu res w er e m e n tio n e d , b u t H e rr e (1971) p ro v id ed a sh o rt
sp a c e , th e fo r m a tio n o f a p la c e n ta l tu b e r c le (a s d e s c r ib e d
for P l e i o s p i l o s su b g e n u s P l e i o s p i l o s by H ar tm an n & L ie d e
1 9 8 6 ).
R e m a r k a b le ,
even
th o u g h
not
m e n tio n e d
in
a ll
d e s c r ip tio n s , is th e e x is te n c e o f c lo s in g r o d le ts in fo u r o f
th e fiv e s p e c ie s (T a b le 1, C R in F ig u r e IB, D ).
E x a m in a tio n o f a d d itio n a l c h a ra c ters sh o w s d istin c t
d iff e r e n c e s b e tw e e n th e tw o s p e c ie s w ith c lo s in g b o d ie s .
C a p s u le s o f A . m a r i t i m a b re a k o f f e a s ily an d r e g u la r ly
after rip en in g , le a v in g th e p ersisten t p e d u n cle o n th e p lan t,
th u s fo r m in g a sp in e ; c a p s u le s an d p e d u n c le s are lig h t-
d e s c r ip tio n .
c o lo u r e d . a lm o s t w h ite , lik e a ll stem s an d b r a n ch es o f th e
p lant; th e m ar gin s o f th e v a lv e s are o n ly m o d e ra tely raised
RESUL TS
a n d , a s a c o n s e q u e n c e , th e v a lv e s o p e n s o c o m p le te ly that
S in c e
th e
d ia g n o s is
of
A m p h ib o lia
s tr e s s e s
fru it
c h a r a c te r s , th e p r e s e n t r e -e x a m in a tio n a ls o c o n c e n tr a te s
o n c a p s u le s . T h e h o lo ty p e s o f a ll fiv e s p e c ie s p la c e d in
th e g e n u s w e r e e x a m in e d . T h r e e d iffe r e n t c o m b in a tio n s
o f the relevan t ch ara cter e x p r e ssio n s c o u ld b e d is tin g u is h ed
(T a b le 1). A . m a r i t i m a an d A . lit tle w 'o o d ii p o s s e s s v a lv e
w in g s an d c lo s in g b o d ie s ( C B ), a s w e ll as c lo s in g r o d le ts
* Institu t fiiir A llg e m e in e B o ta n ik un d B o ta n isc h e r G a rten , U n iv er sitat
H am b u rg. O h n h o r ststra ss e 18, D - 2 0 0 0 H am b u rg 5 2 .
M S . r e ce iv e d : 1989.02.10.
th e tip s to u c h th e b a s e o f th e fru it. O th e r d is tin g u is h in g
featu res o f A . m a r i tim a are sh ort, in flate d , rou n d ish lea ves,
sm a ll flo w e r s an d p e ta ls b a r e ly re a c h in g th e tip s o f th e
c a ly x lo b e s. In co n tra st, c a p su le s o f A . lit t l e w o o d i i rem ain
o n th e p e d u n c le s an d th e y are d ark r e d d is h b ro w n fr o m
n u m e r o u s ta n n in ife r o u s id io b la s ts , lik e a ll ste m s and
b ran ch es. T h e m arg in s o f th e valves are raised to h ig h rim s
p r e v e n tin g th e v a lv e s fro m o p e n in g fu rth er th an 180 .
O th e r c h a r a c te r istic fea tu r es o f A . l i t t l e w o o d i i are lo n g ,
s le n d e r le a v es and larg er flo w er s w ith th e p eta ls e x c e e d in g
th e tip s o f th e c a ly x lo b e s .
B o th a lia 19,2 (19 89 )
180
F I G U R E 1 .— L o n g itu d in a l s e c tio n s o f c a p s u le s . A , B , A m p h i b o l i a m a r i t i m a ( H a ll 2 8 8 5 , B O L ): lo c u le (L O ) b lo c k e d a p ic a lly by c lo s in g b o d y
(C B ) p ro tru d in g fr o m in n er w a ll (W ) o f c a p su le ; p la ce n ta (P L ) to m to its b as e; in d ry state (s e e n h er e) c lo s in g b od y to u c h e s e x p a n d in g
k e e l (E K ) b ut n o t c o v e r in g m em b ra n e (C M ); c lo s in g ro d le t (C R ) an d ad ja c en t b en d o f o u te r part o f c o v e r in g m em b ra n e (O C M ) p u lled
o u t fr om b e tw e e n v a lv e p r op er an d ex p a n d in g k ee l at first o p e n in g o f c a p su le . C , D , A m p h ib o lia h a l l i i ( H a ll 1741 , B O L ): p la cen ta
(P L ) se p a r ates a p ic a lly fr o m en d o c a r p an d b ea rs fu n ic le s (F ) to its v er y tip ; th ic k e n in g o f ou ter fru it w a ll (T W ) ab o ve p la cen ta (sim ila r
to c o n d itio n in L a m p r a n th u s -ty p e fru it; H artm an n 1 9 8 8 ), d ista l en d o f c o v e r in g m em b ra n e (C M ) b en t ag ain st ex p a n d in g k e el (E K ),
r e s e m b lin g p o s itio n in u n o p e n e d fru it. R C M , re flex ed part o f c o v e r in g m em b ra n e; S , s e e d . S c a le in m m .
B o th a lia 19,2 (1 9 89 )
181
T A B L E 1. — C om p a riso n o f fruit characters o f A m p h ib o lia , R u s c h ia and L a m p r a n th u s as o b se r v ed in th e h o lo ty p e s and as g iv e n in or ig in a l d esc rip tio n
T axon
V alve w in g s
(h o lo ty p e )
C lo s in g b o d ie s
C lo s in g r od le ts
h o lo .
d esc r.
h o lo .
d escr .
h o lo .
d esc r.
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
A . m a ritim a , H a ll 2 8 8 5 !
A . little w o o d ii, L ittle w o o d K G 5 2 2 /5 9 !
A . h a l l i i, H a l l 1741 !
A . g y d o u w e n s i s , L e i p o ld t 4 8 0 1 !
A . s ta y n e r i, S ta y n e r K G 2 5 8 /6 5 !
R u s c h ia
+
( + ) ,+
L a m p r a n th u s
(+ ) ,-
( ) = R are e x p r e s s io n o f ch ara cters.
C O N C L U S IO N S
w ith in th e c l o s e r e la tio n s h ip o f th e g e n u s R u s c h i a (s m a ll
c lo s in g b o d y , n ar ro w v a lv e w in g s o p e n in g in to an u p r ig h t
In
th e
lig h t
of
recen t
fin d in g s
in
fru it
t y p o lo g y
(H a rtm a n n 198 8) an d in g e n e r ic d e lim ita tio n s in th e w id e r
p o s it io n , lo n g s le n d e r le a v e s , d a rk r e d d ish ste m s; D e h n
in p r e p .).
r e la tio n s h ip o f th e R u s c h iin a e (D e h n 1989, a n d in p r e p .),
th e g r o u p s d e s c r ib e d a b o v e c a n b e a llie d to d iffe r e n t fr u it
t y p e s a n d , c o n s e q u e n tly , to d iffe r e n t g e n u s g r o u p s.
O f th e
r e m a in in g
th r ee
s p e c ie s ,
A.
h a llii
an d A .
g y d o u w e n s i s la c k c lo s in g b o d ie s b u t p o s s e s s v a lv e w in g s
an d c lo s in g r o d le ts (T ab le 1). T h is c o m b in a tio n p la c e s
D e h n (1 9 89 , an d in p r e p .) h a s b e e n a b le to s h o w that
th e m c l o s e s t to th e L a m p r a n t h u s - ty p e fru it (H a rtm a n n
v a lv e w in g s m a y o c c u r w ith in th e R u s c h i a - t y p e o f fru it.
19 8 8 ) w h ic h is c h a r a c te r iz e d b y v a lv e w in g s a n d c lo s in g
T h e p o s s e s s io n o f th e s e str u c tu r e s c a n th e r e fo r e n o t b e
le d g e s in th e a b s e n c e o f c lo s in g b o d ie s ; c h a r a c te r istic a ls o
u s e d to e x c lu d e taxa fr o m th at g ro u p . E s s e n tia l c h a ra c ter
is a th ic k e n e d t is s u e w ith in th e o u te r w a ll a b o v e th e z o n e
e x p r e s s io n s are th e p o s s e s s io n o f sm a ll en d o ca rp a l c lo s in g
w h e r e th e p la c e n ta e n d s (s im ila r to T W in F ig u r e I D ).
b o d ie s an d o f c lo s in g r o d le ts at th e c o v e r in g m e m b r a n e s.
P r e lim in a r y
in v e s tig a tio n s in th e g e n u s L a m p r a n t h u s
in d ica te that th e tax o n is p r o b a b ly h e te r o g e n e o u s an d w ill
A . m a r i tim a and A . lit t l e w o o d i i a g re e w ith th e se req u ire
m e n ts, b u t th e y d iffe r in o th e r ch a r a c te r s n o t c o n s id e r e d
b y B o lu s (1 9 6 5 ) in th e d e lim ita tio n o f A m p h i b o l i a . S e e in g
that fru it ch ar a cte rs are p r e s e n d y c o n s id e r e d to b e o f g reat
t a x o n o m ic
im p o r ta n c e
in
M e s e m b r y a n th e m a
h a v e to b e d iv id e d u p , A . h a l l i i an d A . g y d o u w e n s i s
a g r e e in g w ith a c e r ta in p art o f th e m a te r ia l.
S u p e r fic ia lly , A . s t a y n e r i m ay b e p la c e d in th e v ic in ity
( e .g .
o f th e L a m p r a n th u s - X y p t fru it a s w e ll , b u t th e c o m p le te
H a r tm a n n 19 8 3; D e h n 198 9; I h le n fe ld t & B ittr ic h 1 9 8 5 ),
a b s e n c e o f a n y c lo s in g d e v ic e an d th e fo r m a tio n o f rath er
th e tw o s p e c ie s h a ve to b e u n d e r sto o d a s b e lo n g in g to tw o
f r a g ile c a p s u le s in d ic a te s a c lo s e r r e la tio n s h ip to th e
d iffe r e n t g e n e r a w ith in th e g r o u p c h a r a c te r iz e d b y th e
D r o s a n t h e m u m - t y p e o f fru it (H a rtm a n n 1 9 8 8 ). W ith in th is
R u s c h i a - t y p e o f fru it. W h e r e a s A . m a r i t i m a sh o w s sev er a l
grou p ,
s im ila r it ie s to th e g e n u s E b e r l a n z i a (v a lv e w in g s , sm a ll
M e s to k le m a (th is h y p o th e sis is su p p o rted b y th e for m a tio n
c lo s in g b od y , w id e ly o p e n in g v a lv e s, in fla te d sh or t le a v es,
o f a re m a rk a b le sto ra g e ro o t in A . s t a y n e r i , a fe a tu re c o m
w h ite s te m s , p e d ic e ls b e c o m in g s p in e s ; a ll d ata a fte r
m o n in M e s t o k l e m a , b u t fu rth er c h a r a c te r s w ill h a v e to
b e e x a m in e d ).
S tiib e r
u n p u b lis h e d ),
A.
little w o o d ii
can
be
p la c e d
th e
n e a r e st
r e la tiv e
seem s
F IG U R E
to
be
th e
2 . — A m p h ib o lia
d istal o p e n in g
genus
h a llii:
o f lo c u le
of
c a p su le sh o w n in F igu re 1C ,D
se e n fro m a b o ve after rem oval
of
v a lv e ,
s h o w in g
en d
of
p la cen ta (P L ) w h ic h can b e
m istak e n for a c lo s in g b ody.
C M , cover in g m em b ran e; E K ,
expan ding k eel; F, fun icle; P L ,
p lacen ta ; R C M , re flexe d part
o f co v er in g
m em b ran e;
w all. S c a le in m m .
W,
B o th a lia 19,2 (1989)
182
2 5 8 /6 5
T h e re su lts sh o w th at o n ly tw o s p e c ie s , A . m a r i t i m a and
(B O L !).
N o m e n c la t u r a l
sy n o n y m :
R u s c h ia
d i s s i m i l i s R o w le y . 6 2 (1978b) n on R u s c h ia s t a y n e r i L . B o l.
A . l i t t l e w o o d i i , c o n f o r m w ith th e g e n e r ic d ia g n o s is . T h is
im p lie s th at th e le c t o t y p e s p e c ie s (A . h a l l i i ) s e le c te d by
H e r r e (1971) is in s e r io u s c o n f lic t w ith th e p r o to lo g u e
ACKNOW LEDGEM ENTS
(G r e u te r e t a l . 19 88, art. 8 . 2 , b ) an d h a s to b e r e p la c e d
b y o n e o f th e tw o s p e c ie s n a m e d a b o v e . A . m a r i t i m a is
s e le c t e d a s th e n e w le c t o ty p e s p e c ie s b e c a u s e its v a lv e
w in g s ar e p a r tic u la r ly b r o a d , th u s c o n f o r m in g b e s t w ith
th e g e n u s S t o e b e r i a c o n s id e r e d to b e c lo s e s t to A m p h i b o l i a
b y B o lu s (1 9 6 5 ) w h e n s h e e s ta b lis h e d th e la tte r g e n u s.
W e are in d e b ted to th e D ir e c to r o f th e B o lu s H erb a riu m
fo r th e lo a n o f a ll ty p e s h e e ts o f a ll s p e c ie s o f A m p h i b o l i a
an d fo r p e r m is s io n to e x a m in e th e m a te r ia l.
O u r w o r k o n M e s e m b r y a n th e m a c e a e h a s b e e n m a d e
T h e ta x o n o m y o f th e g e n u s A m p h i b o l i a , as g iv e n b elo w ,
r e fle c ts th e p r e s e n t sta te o f k n o w le d g e .
p o s s ib le th r o u g h g ra n ts fr o m th e D e u ts c h e F o r sc h u n g sg e m e in s c h a f t an d fin a n c ia l su p p o r t fr o m th e N a tio n a l
B o ta n ic G a r d e n s. F o r b o th w e are m o s t gr a te fu l. W ork o n
fru its h a s b e e n fu n d e d a ls o b y th e C a c tu s an d S u c c u le n t
S o c ie ty o f A m e r ic a th r o u g h a R ese a rc h G ran t. W e sh o u ld
TAXONOM Y
lik e to th a n k a ll d o n o r s.
A m p h i b o li a L . B o l. e x H e r r e , T h e g e n e r a o f M e s e m b r y a n th e m a c e a e : 7 0 (1971); L . B o l.: 169 (1 9 6 5 ); T o e lk e n ,
H .R . & J e s s o p , J.P.: 6 4 (19 76). L e c to ty p e s p e c ie s (h e r e
W e ar e a ls o th a n k fu l to H o r st G o llin g fo r h e lp w ith
S E M -g r a p h s o f fr u its.
d e s ig n a te d ): A . m a r i t i m a L . B o l. e x T o e lk e n & J e s s o p .
T h is le c t o ty p if ic a t io n s u p e r c e d e s th e le c t o ty p if ic a t io n o f
H e r r e (1971), A . h a l l i i , b e c a u s e th e c h a r a c te r e x p r e s s io n s
o f th a t s p e c ie s are in s e r io u s c o n f lic t w ith th e p r o to lo g u e .
REFERENCES
B O L U S , H .M .L . 1960. N o te s o n M e s e m b r y a n th e m u m an d allied gen era.
A . m a r it im a L . B o l. e x T o e lk e n & J e s s o p in B o th a lia
12 : 6 4 (1976); L . B o l.:
169 (1 9 6 5 ). L e c to ty p e s p e c ie s .
H o lo t y p e : H a l l 2 8 8 5 ( B O L !) . N o m e n c la tu r a l sy n o n y m :
R u s c h ia m a r i t i m a R o w le y : 6 2 (19 78b ).
J o u r n a l o f S o u th A f r ic a n B o ta n y 2 6 : 15 5 —164.
B O L U S , H .M .L .
196 2. C o r r e c tio n s an d a d d itio n al n o te s (n o te s on
S to e b e r ia h a llii and S. little w o o d ii) . J o u rn a l o f S o u th A fric a n B o ta
n y 2 8 : 14.
B O L U S . H .M .L . 1963. N o te s on M e s e m b r y a n th e m u m an d allied gen era.
J o u r n a l o f S o u th A f r ic a n B o ta n y 29 : 11-19 .
B O L U S , H .M .L . 1965. N o te s on M e s e m b r y a n th e m u m and allied gen era.
J o u r n a l o f S o u th A f r ic a n B o ta n y 31: 16 9 —174.
B O L U S , H .M .L . 1966. N o te s o n M e s e m b r y a n th e m u m and allied gen era.
S p e c ie s e x c lu d e d
I.
J o u r n a l o f S o u th A f r ic a n B o ta n y 32 : 1 2 5 —128.
S p e c ie s c l o s e to th e g e n u s R u s c h ia S c h w a n t.:
B O L U S , H .M .L . 1967. N o te s on M e s e m b r y a n th e m u m an d allied gen era.
A . lit t le w o o d ii (L . B o l . ) L . B o l. e x T o e lk e n & J e s s o p
in B o th a lia 12 : 6 4 (1976); L . B o l.: 170 (1 9 6 5 ). B a s io n y m :
S to e b e r ia
little w o o d ii L .
B o l.:
162 (1 9 6 0 ). H o lo ty p e :
L i t t l e w o o d K G 5 2 2 / 5 9 ( B O L !). N o m e n c la tu r a l sy n o n y m :
R u s c h ia m u ta ta R o w le y : 7 (1978a) n o n R u s c h ia l i t t l e w o o d i i
J o u r n a l o f S o u th A f r ic a n B o ta n y 3 3: 3 0 5 - 3 0 9 .
D E H N , M . 1989. U n te r su c h u n g e n zu m M er k m a lsb es ta n d un d zu r
S te llu n g d er G a ttu n g A n tim im a N .E . Br. (M e se m b r y a n th e m a
c e a e ) . M itte ilu n g e n a u s d e m
I n s titu t f u r A l lg e m e in e B o ta n ik
H a m b u r g 22 : 1 9 5 - 2 3 2 .
D E H N , M . in p rep . U n ter su ch u n g en zu m V e rw a n d tsch aftsk re is d er
S u b trib u s R u sc h iin a e (M e se m b r y a n th e m a c e a e ).
L . B o l.
GREUTER.
W.
et
a l.
1988.
In tern a tion a l
C ode
of
B o ta n ic a l
N o m e n c la tu r e . R e g n u m V e g e ta b ile 118.
H A R T M A N N , H .E .K . 1983. U n te r su c h u n g e n zu m M er k m a lsb es ta n d
2 . S p e c ie s c l o s e to th e g e n u s L a m p r a n t h u s N .E . Br.:
u n d zu r T a x on om ie d^*r S u b trib u s L e ip o ld tiin a e (M e s e m b r y a n
A . h a llii (L . B o l . ) L . B o l. e x T o e lk e n & J e s s o p in
B o th a lia 12: 6 4 (1976); L . B o l .: 161 (1 9 6 0 ). B a s io n y m :
S t o e b e r i a h a l l i i L . B o l.: 161 (1 9 6 0 ). H o lo ty p e : H a l l 1741
( B O L !) . N o m e n c la tu r a l sy n o n y m : R u s c h ia a m p h i b o l i a
R o w le y : 7 (1978a) n o n R u s c h ia h a l l i i L . B o l.
A . g y d o u w e n s is
(L.
B o l.) L .
B o l.
ex
T o e lk e n
&
t h e m a c e a e ). B ib l i o t h e c a B o ta n ic a 136: 1 - 6 7 .
H A R T M A N N , H .E .K . 1988. Fruit typ e s in M e se m b ry an th em a. B e itr d g e
z u r B i o lo g ie d e r P fl a n ze n 6 3 : 31 3 —34 9.
H A R T M A N N , H .E .K . & L I E D E , S. 1986. D ie G attu n g P l e io s p i l o s s.
la t.
(M e s e m b r y a n th e m a c e a e ).
4 3 3 -4 8 5 .
H E R R E , H . 1971. T h e
gen era o f
B o ta n is c h e
J a h rb u ch er
106:
M e s e m b r y a n th e m a c e a e . Tafelberg, C ape
B a s io n y m :
T ow n .
IH L E N F E L D T , H .-D . & B IT T R IC H , V. 1985. M o r p h o lo g ie , G lied eru n g
L a m p r a n t h u s g y d o u w e n s i s L . B o l.: 13 (1 9 6 3 ). H o lo ty p e :
un d A b g ren zu n g d er G attu n g P s ilo c a u lo n N .E . Br. s .l. (M e s e m
J essop:
64
(1 9 7 6 ),
L.
B o l .:
306
(1 9 6 7 ).
L e i p o l d t 4 8 0 1 ( B O L !). N o m e n c la tu r a l sy n o n y m : R u s c h ia
g y d o u w e n s i s R o w le y : 7 (19 78a ).
b r y a n th e m a c e a e ). B o ta n is c h e J a h r b u c h e r 105: 2 8 9 - 3 2 2 .
R O W L E Y , G .D . 1978a. R eu n io n o f s o m e gen er a o f M e s e m b r y a n
th e m a c e a e . N a t i o n a l C a c tu s a n d S u c c u le n t J o u r n a l 3 3 . 6 —9.
R O W L E Y , G .D .
3. S p e c ie s p e r h a p s c l o s e to M e s t o k l e m a N .E . B r.:
1978b. R e u n io n o f s o m e g en er a o f M e s e m b r y a n
th e m a c e a e
—
a p o stsc r ip t. N a t i o n a l C a c tu s a n d S u c c u le n t
J o u r n a l 3 3: 6 2 .
A . s t a y n e r i L . B o l. e x T o e lk e n & J e s s o p in B o th a lia
12: 6 4 (1976); L . B o l.: 126 (1 9 6 6 ). H o lo ty p e : S t a y n e r K G
TOELKEN,
H R.
&
JESSO R
J.P.
1976.
M es e m b r y a n th e m a c e a e .
N o m e n c la tu r e o f th e g e n u s A m p h ib o lia . B o th a lia 12: 6 4 .
Wilfried Hartmann