A synoptical revision of Operculicarya (Anacardiaceae)

10 Pages·1995·5.3 MB·English

by  U Eggli

A
(Anacardiaceae)
Operculicarya
synoptical revision of
genus
Summary Based on herbarium material, supplemented by living material of 2 taxa, the
:
recognized species of Operculicarya are described in
Operculicarya synoptically revised. All 5
is
included, as well
are
avesonly), and notes on d ition
new from extreme N. of Madagascar,
as a key to species. 3 species are described as O. borealis the
:
O and pachypus from near Tulear in S.-W.
from inland region of Ihosy, O.
hirsutissima the
unnamed mate-
Madagascar. An additional possible species of the genus is left because insufficient
decani
Only O. an\
with Operculicarya uncertain. is -
nibble, and generic affiliation is
un its
ria! is
be narrowly endemic, and their ranges to not over-
appear
the others to
all
The of Operculicarya with
pachypus with O. decaryi. relationships
except those of O.
lap
Poupartia and Lannea are shortly discussed.
specimens
de
du genre Operculicarya a partir d
Revision synoptique
:
-ie."
Les 5 especes •
r<
:.
leur distribution,
lement) des notes sur leurs affinites et
;
de
O. borealis
nees Trois especes nouvelles sont decrites
:
pachypus au S.-W. de Madagascar, pres
O.
dans region d'lhosy, et (
tissima de l'interieur, la
Une espece additionnelle
possible e
.
d'une parte
irya
Mythenquai CH-8002 Zurich, Switzerland.
Sukkulenten-Sammlung, 88,
Urs Stadtische
Eggli,
INTRODUCTION
among of succulents and other xero-
collectors
caudiciform plants
With
the recent interest in
now
and
have been introduced into cultivation are
members genus Operculicarya
phytes, of the
Attempts to identify the material in
and private collections.
encountered public
not infrequently in
(Perrier, 1944) or the treatment in the Flora
descriptions
with help of the original
cultivation the
was
unsurmountable obstacles, and this the reason
have met with
Madagascar 1946)
of (Perrier,
which revealed the presence of additional taxa
of genus,
revision the
for undertaking a synoptical
subsumed under wide-spread
and which were partly the
recognized
have been previously
that not
O. decaryi.
SUMMARY
HISTORICAL
The
genus Operculicarya was described by Perrier 1944 on
in the basis of rather scanty
material, none of the three species initially described for the new genus was completely known
at
The
that time. subsequent treatment of the group for the Flora of Madagascar (Perrier, 1946) did
not provide significant additional information, apart from the fact that three taxa were
all illus-
by
trated line drawings.
According
to the information available Perrier, genus had
to the a restricted distribution in
and
the arid semi-arid regions of S.-W. Madagascar.
Capuron
removed On
In 1962, O. monstruosa Perrier from Operculicarya. the basis of addi-
tional material, especially fruits and seeds, became apparent that not even a member of the
it is
it
Anacardiaceae, but belongs Commiphora monstruosa Capuron
to (as C. (Perrier) Burseraceae).
;
The
seeds leave no doubt Capuron's was
arillate that decision well
justified.
GENERIC
RELATIONSHIPS
Capuron
(1975) proposed an addition to Operculicarya when he published the new combi-
nation O. gummifera for Poupartia gummifera Sprague (Capuron, 1975 mentioned
casually
first
;
Capuron,
in 1962 271). There can be no doubt that the genera Poupartia and Operculicarya are
:
closely related most notably, they both have operculate Of between two,
the differences the
fruits.
;
as indicated in the key of Perrier 946), the number of stamens does not seem to have any signif-
1
(
and
icance, the isostemonous androecium
reported for Operculicarya seems be an error of
to
observation.
The main
difference between the two genera according to Capuron (1975) appears to be the
number
of opercula of
the fruit Operculicarya has a single operculum, while species of Poupartia
:
have
2-5 opercula. Moreover, anatomy
the of the fruit also appears different, and these character-
istics provided the base for Capuron's transfer of Poupartia gummifera. Because of very
the dif-
ferent appearance of this species, which has much larger leaves without a winged rachis and
female
flowers arranged in spicate to racemose inflorescences, further evaluation of affinities
its
necessary must
but be postponed
is
until full material available for anatomical studies.
is
Consequently, gummifera
O. excluded from
is the present concept of Operculicarya.
The
consistency of
the alleged difference in operculum number between Operculicarya and
&
Poupartia must
be questioned however, Teichman Hardy
at least in part, as (1992) report the
%
occasional occurence of more than one operculum Operculicarya examined
in of 110 stones
(8
what
in they refer to as O. decaryi). Moreover, they found that Operculicarya very similar to
is
Lannea,
specifically to L. discolor and argue that the current status of these two genera needs
reconsideration. Lannea,
also in the tribe Spondiadeae, a genus of some 40 species mainly from
is
tropical Africa and
is not currently reported from Madagascar. seems from
to differ
It
Operculicarya mainly
in its spicate to paniculate female inflorescences (vs. solitary in
Operculicarya), and
in the shoot system, which does not exhibit the contracted short shoots of
Operculicarya and more
overall appears
similar Poupartia.
to
JCO"
p^v" .00
If
y
fi Dr,
pcnhxpus Y.
F,
<7.
; :
OPERCULICARYA H
Perrier
Madag.
Mem. 248 (1944) 14 18 (1946).
Mus. Natl. Hist. Nat., ser. nov., 18 (7) Fl. 1 16,
: ; :
Capuron
Lectotype (designated by 1962 271).
O. decaryi H. Perrier
:
:
known
trunk
Gnarled compact or open shrubs or small trees, completely dioecious as far as
;
somewhat
always thickened
and in rela-
pyramidally swollen base or bole-like parallel-sided,
at
bumpy-warty or smooth with superficial fissures ultimate pri-
tion to plant-size bark irregularly
;
;
mary glabrous or tomentose, straight or bent ± zig-zag-wise
branches (long shoots) slender,
;
with condensed internodes, producing a
leaves internodes long short shoots spur-like
alternate,
;
imparipinnate with 3 to 20 pairs of leaflets, rachis slightly to pro-
of leaves each season leaves
set
;
coriaceous or with
nouncedly winged, glabrous and glossy or dull, or velvety to hairy, thin,
leaflets
many
few
to veins.
mm
many-flowered
few- spicate
Flowers 3-4 diam. male flowers solitary or in to
in
stellate,
;
from shoots sepals petals
the short 5,
inflorescences arising 5,
to paniculate solitary or fascicled
;
spreading, stamens filaments broadened and
10, thickened at base, centre of the flower with a
conspicuous,
thickish, discoid irregularly lobed nectary and a 3- to 5-cleft indistinct or distinct pis-
tillodium female flowers male
solitary, similar to flowers but with anthers ovary globu-
sterile
;
;
dome-shaped
lar to with 5 subsimilar short, stumpy styles originating the periphery stigma
at
;
small, appearing
shortly glandular.
Fruits solitary, sessile or shortly stalked with the calyx persistent, elongate-roundish ovoid,
to
mm
ca. 7 in diam., fleshy, normally with a unilocular stone which normally has one elongate-
ellipsoidal operculum.
Known
Distribution only from Madagascar
(excluding the moist forests of the E. slopes).
:
Key
to
species
Leaflets glabrous or almost so, dark brown with paler underside when dry venation darker brown on
;
underside, lateral veins 3-6 on each side of the main
axis.
Shrub
m,
2. to trunk an irregularly pyramidal caudex with apically spreading crown ultimate twigs
1
;
glabrous, zig-zag-wise between
the nodes, frequently with pungent flowers yellowish-
sterile tips
;
green
pachypus
O.
m m
Shrub
2'. to 3 or tree to 6(-15?) with ± bole-like or conical trunk ultimate twigs minute-
straight,
;
ly white-tomentose, without pungent tips flowers purplish-red O. decaryi
;
±
Leaflets densely hairy to tomentose, colour various veins >
lateral
7.
;
3. Leaflets all equal, (ll-)14-18(-20) pairs roundish-rectangular with revolute margins and raised
;
venation on lower
side hyphaenoides
o.
3'. Leaflets not all equal, in less than 14 pairs, never roundish-rectangular, without revolute margins
4. Leaflets not emarginate.
5. Leaflets in (4-)5-7 pairs, olive-brownish when dry margins ± veins 8-
ciliate lateral
;
;
cm
12, distinctly visible on lower side leaves 2.8-5 long fruits sessile. O. borealis
; ; .
5'. Leaflets in (9-)ll-13(-14) pairs, dark dull brownish when margins not
dry, ciliate
;
mm
cm
lateral veins 5-10, hardly visible leaves 5.5-7 long 8-12
fruits stalked, stalk
;
;
lor
4\
Leaflets
Operculicarya
borealis
Eggli,
sp. nov.
(8-12 utroque latere
ur sables au pied sud-
Small tree trunk and bark not known more
ultimate twigs or less straight, glabrous or very
;
;
insignificantly tomentulose, when
longitudinally striate dry, dark brown older twigs sometimes
;
with
scattered, small, transverse concolorous
lenticels short shoots often with or 2 elon-
starting
1
;
gated
internodes before producing
the typical congested internodes and thus appearing stalked
;
—
—
153
cm
narrowly
leaves 2.8-5 long, with (4-)5-7 pairs of leaflets overlapping or just separate rachis
;
winged, with scattered shaggy hairs or short bristles lowest leaflets almost orbicular and smaller,
;
mm mm
roundish-ovate
the other leaflets in upper 2/3 of leaf 4-8(-13) long, 3-4(-7) broad, to ellip-
brownish and
apex rounded, never emarginate, upper side of dry leaves pale slightly
soidal,
glossy, glabrous or ± tomentose, under side dull pale olive, ± to densely white-tomentose, veins
somewhat
8-12 on each of midvein margins often
darker, conspicuous, side the ciliate.
;
unknown.
Flowers
some-
Fruit almost globular, the calyx lobes appressed stone subglabrous, laterally
sessile,
;
what compressed, operculum narrowly elongate, always close to the margin.
Diego deciduous
Distribution Extreme northern Madagascar in the region of Suarez, in partly forests,
:
Notes This taxon easily recognized by the medium brownish dry leaves with ciliate margins and
is
:
the numerous closely set side-veins readily discernible on the underside of the leaflets. In addition, the short-
ly to conspicuously stalked short shoots provide an character.
SF
SF 22967 SF 23097
Material examined Capuron 22726 22954bis (type).
(P), (P), (P),
:
Operculicarya decaryi H. Perrier
249 Madag. 114 19-20,//g. 1-6 21
18 (1944) Fl. 5,
(7)
: :
: ;
Madag.
Type Decary 9305 H. Perrier, 14 20.
(holo-, P!) cf. Fl.
1
:
: ;
m m
m
Shrub 3 or more frequently tree-like to 6 (occasionally to 15 ?) trunk bole-like,
to
;
m
dark
up diam., bark wrinkled to irregularly tuberculate,
bottle-shaped or conical, to in
1
soon
medium brownish, glabrous or minutely whitish-tomentose but then
greyish ultimate twigs
;
cm
dark green,
more between the nodes leaves 2.5-6 long, fresh
glabrescent, or less straight
;
when brown brown above, with
glossy above, paler below, dry typically dark to blackish
slightly
and below medium brown with slightly darker venation completely
or without a silvery sheen,
;
some also the
glabrous or especially the rachis with greyish to white hairs or bristles (rarely
somewhat conspicuously
somewhat predominantly on the lower side) rachis to
leaflets hairy,
;
lowest
suborbicular (especially the pair)
broadly with (4-)5-7(-8-9) pairs of leaflets leaflets
alate
;
;
mm
mm
from
to ovate, largest in upper 2/3 of leaf, 4-7 long, 3-4 broad, normally well separated
even very emarginate venation indistinctly
each apex round, rarely truncate or slightly
other,
;
main
on of
on lower with 3-4 veins either side the vein.
visible the side, lateral
and tomentose
minutely bracteose inflo-
Male paniculate,
flowers solitary or in short,
margin spreading-ascending,
with minutely erose petals
rescences sepals spreading-ascending,
;
;
filaments thick base, round in cross section, nec-
dark red stamens ascending-spreading, at
10,
;
mm,
Female flowers pedicel 2-3
2- solitary,
pistillodium to 5-cleft.
tary discoid, irregularly lobed,
male
caducous of the pedicel similar to male flowers, filaments as in
easily the articulation
at
;
and apparently nectary discoid, thickish,
male flowers sterile,
flowers, anthers smaller than in
around periphery
originating the
with thickish styles
ovary depressed-globose 5
irregularly lobed,
and thickened and shortly papillate.
deflexed, stigma slightly
mm
Fruit globose-discoid or droplet-shaped, with a slender stalk 5-6 long, with appressed
remnants
of
the calyx, reported as green tinged with red stone pale ochre honey-brown, almost
to
;
smooth
or slightly verrucose, with broadly trapezoid concolorous operculum.
deciduous
forests.
1
M
Vernacular
names "
Jiabiha (Sussmann " "
170, reports fruit as edible) Saby (Bosser 3735)
:
;
;
" "
Tabily {Humbert 20202) " Zabihy " (Decary " "
4383) Zabily (Service des Eaux Forets 1635 SF)
et
; ;
;
" Zaby " (Humbert 14158bis, " Zaby " name
etc.). is the vernacular most frequently cited on herbarium
labels.
Notes
This taxon
frequently encountered
is in cultivation, but material here newly described as
:
pachypus
O. is equally frequent in collections and is invariably labelled as O. decani (Rowley, 1987 197,
:
&
Teichman
Hardy, The two
ill. 1992). taxa are easily separable, however, despite the similarity in leaf
;
characters O. decaryi has dirty pinkish to dark purple flowers and is a much larger plant with a bole-like,
:
more
or
less parallel-sided trunk with
smooth
(at least in cultivation) quite longitudinally structured bark.
pachypus
O. has pale greenish-yellow and
flowers, an
altogether smaller bonsai-like shrub
is with a pyra-
midal
basal stem-thickening
the bark is irregularly bumpy-warty and conspicuously silvery-grey old
in
;
The
specimens.
ultimate branches
are distinctly zig-zag, and their tips frequently become and hardened.
sterile
Material examined
Bosser 3735 10185 10528 SF SF
(P), (P), (P) Capuron 344 9470 8523
(P), (P),
:
;
SF
SF
(P), 18678 (P), 20460 SF (P), 27955 SF (P), 28303 SF (P) Croat 31412 (MO) Decary 3294
(P),
;
;
4383 (P), 9305 (type) Fosberg 52472 (MO) Humbert
12456ter (P), 12798bis (P), 14158bis (P), 20202 (P)
; ;
;
&
n 1502 (P) Leandri 4468 (P) McWhirter Capuron 163 MO), 2525 (MO,
(P) Phillipsnn 2 ?/ -"
; ;
:
i
&
P) Roosli Rechberger
s.n. (cult.) Service des Eaux Forets 1635 SF 865 Sussimum
et (P) Sexria P)
; ;
; < :
170 (MO) '
va
;
Operculicarya
hirsutissima
Eggli,
sp.
i
Type
Leandri "
3447, Madagascar,
restes de forets de vallee de Menarahaka
la la
:
t
Am,
Ivohibe,
vers
ition
29.X. 1960
(nolo-,
P!).
m
Small 5-6 m,
tree to trunk to 0.4 diameter bark unknown
in ultimate twigs glabrous or
;
;
tomentose,
slightly inconspicuously when
longitudinally greyish-brown
striate dry, straight, to
medium
brown
with inconspicuous
small
transverse and
lenticels short shoots spur-like sessile,
;
but able to produce long shoots leaves 5.5-7 cm brown when
long, dark lower
dull side
dry,
;
slightly paler, with (9-)l 1-13(14) pairs of leaflets rachis inconspicuously broadly winged
to
;
;
leaflets often asymmetrically
arranged,
slightly overlapping or separate, lowest almost
leaflets
mm mm
orbicular and
smaller, other leaflets all about equal, 6-9 long, 3-4.5 broad, broadly ovate
to rounded-oblong to elliptic, apex rounded or subtruncate, rarely very emarginate both
slightly
;
sides conspicuously and
densely
greyish hairy venation hardly with 5-10
visible, lateral veins.
;
—
1-2. Leaf characters of Operculicarya A, O.
hirsutiss,
:
Capuron 22726 SF) C, O. pachvpus (drawn from the holotype)
;
.Al-El:wh<
(scale bar = 2 mm). E3 = from above
leaflet (scale as
/
Male
flowers
in fascicled spicate to paniculate inflorescences epipetalous filaments
slightly
;
united with
petals at base nectary discoid, conspicuous, ca. 10-lobed, pistillode 3-cleft almost to
;
the base. Female flowers unknown.
mm
Fruit pendent, with a slender stalk 8-12 long, broadly ovoid, no remains of calyx
observed somewhat more
stone laterally compressed, or less smooth, operculum near margin,
;
irregularly broadly ellipsoid to trapezoid, concolorous or paler than the rest of the stone.
deciduous ami
Ihosy, forest
I
Notes
Material of species has
this
r
:
nizable because of the densely hairy leav
—
sessile
ers in its fruits.
i
Operculicarya hyphaenoides
H. Perrier
249 (1944) Madag. 114 18-19, 7" /7
3 (7) Fl. 5,
fig- :
: ; :
Type 19169
Perrier
(nolo-, P!).
:
common
Tree to 1.5 m, with several thick gnarled trunks from base, each with a spreading
m
much-branched crown, trunks 0.4 diameter with irregularly bumpy-warty bark ultimate
to in
;
cm medium
more dark
twigs glabrous, brownish-grey, or less straight leaves 2.5-6 long, to olive-
;
when
green olive-brown with (ll-)14-18(-20) pairs of parts densely hairy
to dry, leaflets, all
;
mm
mm
winged 4 long and 2
minutely conspicuously
rachis regularly to leaflets all similar, ca.
;
with
broad, roundish-rectangular, apex truncate to slightly emarginate, margins revolute, underside
5-7 conspicuously raised lateral veins on either side of the main vein.
mm
Male flowers with a densely tomentose peduncle 4-7 long with 2 small bracte-
solitary,
probably dark
oles, sepals ascending-spreading, probably dark red, petals ascending-spreading,
red with pink margins filaments broad and thickish, inserted below the thickish, 10-lobed, dis-
;
coid nectary, anthers yellow. Female flowers reputedly with yellow-green ovary.
Fruit pendent, discoid-subglobose, reported as green with grey bloom, with a slender minute-
mm
ly tomentose stalk to 8 long which carries several vestigious bracts stone roundish, flat-
1
;
±
tened, slightly rugulose with droplet-shaped, angular, paler operculum.
on
Distribution S.-W. Madagascar, region of Lac Tsimanampetsotsa, xerophytic scrub limestone.
in
:
named The
species
of the nature reserve for the lake. is
which have revolute margins and conspic-
leaflets
Operculicarya pachypus
Eggli, nov.
sp.
&
Hardy
Teichman
i
Perrier
(1944).
Operculicaryae decarvi aspeciu tolioruni simiiis seel
ti
;
m
Compact bonsai-like shrub to tall trunk conical or irregularly pyramidal, tapering
1
;
crown
bumpy-warty grey bark spreading
towards the apex, with irregularly silvery-grey to dull
; ;
conspicuously zig-zag between the nodes, glabrous, pale grey to
ultimate twigs indistinctly to
cm
becoming and pungent leaves 1.5-3.6 long, glabrous,
greyish-brown, the tips often sterile
;
dark green and glossy above, slightly paler below, or when dry dark olive-brown to brownish-
conspicuously
black above and medium dark brown below, appearing coriaceous rachis
to
;
winged, with 3-4(-5) pairs of leaflets leaflets almost orbicular to roundish droplet-shaped, just
;
mm
mm
touching each other or well separated, largest in the middle of the leaf, 7-8 long, ca. 4
on with
broad, apex rounded or rarely shallowly emarginate, venation indistinct the underside,
main
3-4 veins on each side of the vein.
lateral
Male flowers almost sessile and easily falling at an articulation of the short pedicel
solitary,
somewhat mar-
medium spreading with revolute
greenish-yellow,
sepals spreading, green, petals
almost
thickened base nectary thickish, discoid,
gins stamens ascending, with filaments at
10,
;
;
unknown.
somewhat Female flowers
unlobed pistillodium small, indistinctly chanelled or cleft.
;
somewhat compressed, with a slender stalk
probably pendent, subglobular, laterally
Fruits
mm
operculum broadly ellipsoid.
3-4 long stone slightly rugulose, indistinct,
;
Distribution S.-W. Madagascar, region of La Table near Tulear, on limestone outcrops in low xero-
:
decani. See
as O.
i-identit'ied
Encountered frequently
quite
i
:
which have appeared especially on the European market (the
The numbers of field-collected plants
large
conservation
elsewhere unknown) are reason for concern as to its status.
situation
is
10158 Capuron & Chauvet 20807 SF (P) Chauvet 167 (P), 203 (P),
Material examined Bosser (P)
; ;
:
&
Rechberger
224 Humbert 2614 mpe) Razafmdnitsiro s.n. (cult.) Roiisli s.n. (cult.).
(P)
;
:
;
A
Operculicarya
sp.
m smooth young twigs
probably
greyish-brown,
bark greyish to
Small 3
tree to tall ;
;
cm on
when dry leaves 3-4 long, congested in rosettes
glabrous and longitudinally striate
slightly
;
some
whitish
7- rachis only slightly alate, with bristly stiff
spur-like short shoots to 9-foliolate
;
;
when
largest
and pale olive-green dry, dull,
coriaceous
hairs especially towards the base leaflets
;
mm
mm
conspicuously emarginate, with fine
broad,
near of 7-10 long, 4-6(- 7)
the leaves,
tip
dense venation visible on the lower side.
unknown.
and flowers
Inflorescences
mm
tomen-
some minute minutely
4-5 with bracts,
with persistent calyx, stalk
Fruits solitary
becoming then
" yellow purple,
described as red above, pale at first,
tose elongate-ovoid,
fruit
;
operculum (immature material studied
" without
smooth, probably
black 2890) stone
{Phillipson
;
only).
23400 Sambirano
known from two one (Capuron SF, region,
This taxon only collections,
is
with
from extreme N. of Madagascar, the other (Phillipson 2890, Ejeda-Betioky, fruits)
the
sterile)
from the S.-W. While the general facies of the plant is more than suggestive of Operculicarya. the
examined have and very smooth texture and appear to lack an operculum.
unripe stones a different
two cannot be resolved and for the present
Without more of these collections
material the status
they limbo as a possible additional species of Operculicarya.
are left in
SF 2890 (MO,
23400
examined Capuron (P) Phillipson P).
: ;
EXCLUDED TAXA
gum-
= Poupartia
Operculicarya gummifera (Sprague) Capuron, Adansonia, ser. 2, 14 (4) 571 (1975)
:
408
mifera Sprague, Bull. Herb. Boissier, ser. 2, 5 (1905).
:
572
Capuron Capuron, Adansonia, 14 (1975).
Operculicarya gummifera (Sprague) var. seyrigii ser. 2, (4)
:
Mem. Mus. 18 249-250 (1944) type
Operculicarya monstruosa Perrier, Natl. Hist. Nat., ser. nov., (7) :
: ;
270
= Commiphora monstruosa Capuron, Adansonia, 2 (1962).
Perrier 12783, P! (Perrier) ser. 2, (2)
:
MO
Acknowledgments Material for this study was kindly made available by the herbaria and P, and
:
ZSS,
their curators are thanked for their help. In addition to living material cultivated in the collection at
&
some were made by W. Roosli B. Rechberger, Zurich, and their help has been inva-
collections available
K. U. Kramer very kindly assisted with the Latin diagnoses. Words of thank go to the edito-
luable. (Z,
t)
Wov who made
who French summary, and U. (Erlenbach, Switzerland) the
staff (P) contributed the to
rial
drawings.
REFERENCES
—
Capuron, R., 1962. Contributions a l'etude de la flore forestiere
i
Ada
Commiphora nouveaux du sud de Madagascar.
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