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MEDEDELINGEN LANDBOUWHOGESCHOOL
WAGENINGEN • N E D E R L A N D • 77-20 (1977)
SEEDLING M O R P H O L O G Y OF
SOME A F R I C A N SAPOTACEAE
AND ITS TAXONOMICAL
SIGNIFICANCE
J. B O K D A M
Laboratory of Plant Taxonomy and Plant Geography,
Agricultural University, Wageningen, The Netherlands
(Received26-VII-1977)
7
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CONTENTS
page fig.
1. I N T R O D U C T I O N , 5
2. SEEDLING CHARACTERS 7
2.1. Germination 7
2.2. Root system 7
2.3. Hypocotyl 7
2.4. Cotyledons 8
2.5. Endosperm 9
2.6. Epicotyl 9
2.7. Eophylls 9
3. SEEDLING TYPES 11
SUMMARY 23
RESUME 24
ACKNOWLEDGEMENTS . .25
A P P E N D I X : DESCRIPTIONS A N D D R A W I N G S O FTHE F O L L O W I N G
SPECIES O FA F R I C A N SAPOTACEAE 27
1. Afrosersalisia afzelii (ENGL.) AuBRÉv 29 1
2. - cerasifera ( W E L W . ) AUBRÉV 29 1
3. Aningeria adolfi-friederici ( E N G L . ) ROBYNS et GILBERT 31 2
4. - altissima (A. CHEV.) AUBRÉV. e t PELLEGR 33 2
5. - robusta (A. CHEV.) AUBRÉV. et PELLEGR 33
6. Argania spinosa (L.)SKEELS 34 3
7. Aubregrinia taiensis (AUBRÉV. et PELLEGR.) HEINE 34
8. Autranella congolensis ( D E W I L D . ) A. CHEV 36 4
9. Baillonella toxisperma PIERRE 38 5
10. Breviea leptosperma (BAEHNI) HEINE 38
11. Butyrospermumparadoxum(GAEKTN.F.)HEPPER 40
12. Chrysophyllum azaguieanum MIÈGE 42 6
13. - giganteum A. CHEV 42 7
14. - muerense ENGL 44 8
15. Donella ogowensis (A. CHEV.) AUBRÉV. et PELLEGR 44 9
16. - pruniformis (PIERRE ex E N G L . ) AUBRÉV. et PELLEGR 46 9
17. - ubanguiensis ( D E W I L D . ) AUBRÉV 48 10
18. - welwitschii ( E N G L . ) PIERRE ex AUBRÉV. et PELLEGR 50 10
19. Gambeya beguei (AUBRÉV. et PELLEGR.) AUBRÉV. et PELLEGR 50 11
20. - boukokoensis AUBRÉV. et PELLEGR 52 12
21. - lacourtiana ( D E W I L D . ) AUBRÉV. et PELLEGR 54 13
22. - perpulchra ( M I L D B R . ) AUBRÉV. et PELLEGR 56 14
3
page fig.
23. Gambeya subnuda (BAK.) PIERRE 56 14
24. Gluema ivorensis AUBRÉV. et PELLEGR 58
25. Ituridendron bequaertii D E W I L D 58 15
26. Kanton guereensis AUBRÉV. et PELLEGR 60 16
27. - sp 62 16
28. Malacantha alnifolia (BAK.) PIERRE 62 15
29. Manilkara obovata (SABINE et G. D O N ) J. H. HEMSLEY ( = M. lacera (BAK.)
DUBARD) 63
30. Mimusops sp 63 17
31. Omphalocarpum elatum MIERS 65
32..- lecomteanum PIERRE ex E N G L 66 18
33. - procerum P. BEAUV 68
34. - sp 68
35. Pachystela bequaertii D E W I L D 69 19
36. - brevipes (BAK.) E N G L 71 19
37. Pseudoboivinella oblanceolata (S. MOORE) AUBRÉV. et PELLEGR 71
38. Synsepalum dulcificum (SCHUM.) BÂILLON 72 20
39. - stipulatum ( R A D L K . ) E N G L 72 20
40. - subcordatum D E W I L D 74 20
41. Tieghemella africana PIERRE 74 21
42. - heckelii PIERRE 76 21
43. Tridesmostemon claessensii D E W I L D 77 22
44. - omphalocarpoides ENGL 77 22
45. Tulestea seretii ( D E W I L D . ) AUBRÉV. et PELLEGR 79
46. Wildemaniodoxa laurentii ( D E W I L D . ) AUBRÉV. et PELLEGR 79 23
BIBLIOGRAPHY 83
1. I N T R O D U C T I O N
Seedlings are studied for two reasons mainly. First of all there is an urgent
need amongst plant ecologists t o identify seedlings from a certain region or
habitat. To enable this, seedlings have been described and figured, and identifi-
cation keys have been composed, in temperate regions (LUBBOCK, 1892;
K I N G , 1966; CSAPODY, 1968; M U L L E R , in prep.) as well as in the tropics
( T R O U P , 1921; D U K E , 1965; D E L A MENSBRUGE, 1966; B U R G E R , 1972; D E
K O N I N G , in prep. ; D EVOGEL, in prep.). Moreover, morphological characters
may have ecological significance. T R O U P (1921), R I Z Z I N I (1965) and JACKSON
(1968)emphasized that seedling morphology should bethoroughly investigated
for a better comprehension of germination, establishment andjuvenile growth
during thenatural regeneration of vegetation.
In the second place seedlings are studied because they provide additional
information toourknowledge ofthe life cycle ofplants. Itisgenerally accepted
today, that taxonomical research should not be based exclusively on mor-
phological characters of the mature specimens, but also on characters ofthe
juvenile stages of the plant. Critical examination of correlations between both
groups of characters has resulted repeatedly in a better understanding of
taxonomically difficult taxa. D ECANDOLLE (1825) wasthe first botanist touse
seedling characters for the delimitation of tribes and genera in Leguminosae.
Unfortunately, very few botanists followed his example (e.g. GUILLAUMIN,
1910). Inthepast twodecades however, theinterest inusing seedling characters
for taxonomical purposes has revived (VASSILCZENKO, 1936; D EFERRÉ, 1952;
LÉONARD, 1957; CERCEAU-LARRIVAL 1962; JACOBS, 1966; WEBERLING and
LEENHOUTS, 1966; BRETELER, 1973; BAUDET, 1974). M o r e details a b o u t the
history of blastogeny (study of the development of the seedling) are given
by JACOBS (1966) a n d BAUDET (1974).
In Sapotaceae, as in Leguminosae, the delimitation of subfamilies, tribes
and genera israther complicated. Recent monographs andrevisions of African
Sapotaceae show wide differences in opinion: MEEUSE, 1960, 1963; HEINE,
1963; AuBRÉviLLE, 1964, 1974; BAEHNI, 1965; HEMSLEY, 1968. A c o m m o n
concept seems to be absent and confusion about the proper classification and
nomenclature reigns. F o r this reason an investigation of the seedling mor-
phology ofthe African Sapotaceae seems opportune.
In this paper, 46species outof 25 genera have been studied. Most species are
from West and Central Africa. The seedlings were either directly collected in
the field or obtained after sowing in a nursery in Kisangani (Zaire) or in the
conservatory at Wageningen (Netherlands). Specimens have been conserved as
herbarium orinspirit attheHerbarium Vadense (WAG). Other seedlings have
been studied at the Herbarium of the National Institute for Agricultural
Sciences at Yangambi, Zaire (YBI), or have been kindly made available bythe
Herbarium of the National Botanical Gardens of Belgium (BR). My observa-
2.1. GERMINATION
2.3. HYPOCOTYL
The rootcrown, i.e. the transition between the taproot and the hypocotyl,
isnot always clearly recognizable, but normally it iswellmarked by changes in
* For authors names the reader is referred to tabel 2which ismainly based on the classi-
fication ofAuBRÉviLLE, 1964.In some aberrant casesthese namesare mentioned inthe text.
2.4. COTYLEDONS
The main criteria used for thedescription ofthecotyledons are their position
inrelation to thetesta and their texture.
Seedlings with free cotyledons are called phanerocotylar. The position of
their cotyledons may be horizontal or slightly erect. In cryptocotylar seedlings
thecotyledons remain enclosed in the testa (DUKE, 1965).
According totheir texture thecotyledons of thestudied taxa can be classified
as follows:
A. Cotyledons foliaceous, nervation acro-brochidodromous.
1. Cotyledons papyraceous, dark green, much enlarging during and after
liberation from the testa and the endosperm, long persistent, asymmet-
rical, base abruptly narrowed into a petiole; venation system con-
spicuous.
2. Cotyledons coriaceous, dark to light green, enlarging during and after
liberation from the testa and the endosperm, less persistent, slightly
or not asymmetrical, base abruptly narrowed into a petiole; venation
system inconspicuous.
B. Cotyledons fleshy, nervation indistinct, hyphodromous.
Cotyledons thick, plano-convex, light to dark green in phanerocotylar
seedlings, pale yellowish green to brown in cryptocotylar seedlings,
symmetrical or asymmetrical, caducous to rather persistent (in cryp-
tocotylar seedlings).
The asymmetrical shape of the cotyledons iscaused by the shape of the seed
and it isexpressed in the following features (seefig.4: 3,4):
- the blade halves on either side of the midrib are unequal;
- the margin of the broadest blade half is either concave or straight, in-
stead ofconvexbetween themiddle and the apex of thecotyledon;
8 Meded.Landbouwhogeschool Wageningen 77-20(1977)
- themidrib terminates inthemargin besidetheapex ofthe cotyledon.
The surface is glabrous but on both faces of foliaceous cotyledons minute,
colourless papillae can be observed with strong magnification. In some species
lactiferous ducts are visible in transmitted light. Stipules are never present.
2.5. ENDOSPERM
2.6. EPICOTYL
In nearly all species the epicotyl is distinct, straight, slender and either
cylindrical or compressed. It may be long ( > 4 cm) or short ( < 4 cm). This
limit at 4 cm was chosen after epicotyl length of seedlings with foliaceous
cotyledons was compared with that of seedlings with fleshy ones. Usually an
indumentum of variably developed medifixed hairs covers the epicotyl.
Lateral buds may be observed in the axils of the cotyledons and also in the
axils of cataphylls, brown scale-like rudimentary leaves which are present in
some species. From both types of buds new shoots may develop after destruc-
tion of theapical bud.
2.7. EOPHYLLS
The first well developed green leaves are called eophylls (DUKE, 1965). They
may be opposite or alternate, provided with stipules or not, while their texture
is either papyraceous or coriaceous. The petioles are short to rather long, e.g.
up to 1.5 cm in Tieghemella africana. They are variably shaped and rarely
glabrous. Translucent dots have been observed in Aningeria, Malacantha,
Chrysophyllum giganteum and Chrysophyllum muerense. Lactiferous ducts are
visible in transmitted light in many species. The nervation may be eucampto-
In the available seedling classifications the major criteria are the position of
the cotyledons in relation to the soil surface after germination and their
liberation from the testa. The corresponding terminology, derived from agri-
cultural practices, distinguishes epigeal and hypogeal germination, resulting in
seedlings with the cotyledons above and below the soil surface respectively.
Among the taxa studied I never found seedlings with the cotyledons below
soil surface in the field. The germinating seeds were invariably lying on top
of the soil, often still enclosed in the mouldering fruit or in dung of monkeys
and elephants. However, when seeds of taxa with a short or undeveloped
hypocotyl, are sown at considerable depth, the cotyledons may remain buried
irrespective of their liberation from the testa. Examples of the latter are
Pachystela, Wildemaniodoxa and Baillonella: their seedlings are epigeal in the
field, but may be hypogeal when sown at considerable depth. Because of this,
I prefer the criterion and the terminology of DUKE (1965), who distinguishes
crypto- and phanerocotylar seedlings (see 2.4), but this criterion should be
applied with care as well. Taxa, which have phanerocotylar seedlings with a
short or undeveloped hypocotyl in the field may produce seedlings with
slightly expanded cotyledons iftheir seedsare sown ina solid heavy soil, which
hamperstheunfolding ofthecotyledons.LÉONARD(1957)arrangedtheseedlings
of sometribesof Caesalpiniaceaeintofour types,which maybereduced to two,
following the considerations above. His type B (cryptocotylar, hypogeal) is
identical to type D (cryptocotylar, epigeal), but the seeds of Bhave been sown
at considerable depth. His type C (phanerocotylar, epigeal, hypocotyl not
developed)isbutavariant ofA(phanerocotylar, epigeal,hypocotyl developed).
The taxonomical value of the length of the hypocotyl may be very limited in
certain groups. It has been demonstrated by LAMPRECHT (1945, 1948, quoted
by BAUDET, 1974)that the difference between the well developed hypocotyl of
the phanerocotylar seedling of Phaseolus vulgaris L. and the undeveloped
hypocotyl of the cryptocotylar seedling of Phaseolus coccineus L. was based
on one allel only. Hybridization of the two species was easy and it produced
heterozygotic phanerocotylar Fi seedlings with short hypocotyls, i.e. inter-
mediate between the parents.
This shows that the difference between the phanerocotylar and the crypto-
cotylar seedlingaswellasitstaxonomical significance maybeless fundamental
than is pointed out by LÉONARD, I.e., when he concludes: 'Les plantules de
toutes lesespècesd'un même"bon" genreprésentent lemêmetypede structure,
ou, en d'autres termes, au sein de chaque "bon" genre domine un seul type de
plantule'. The validity of this rule has been criticized already by JACOBS(1966)
on the assumption that characters may have different taxonomical weight in
different taxa.
The delimitation of major seedling types should be based preferably on
Meded. Landbouwhogeschool Wageningen 77-20 (1977) 11
characters with fylogenetic significance. It appears from this study (4), thatin
Sapotaceae the texture of the cotyledons is such a character. It is closely
related tothereduction ofendosperm, a reputed phenomenon of evolutionary
advancement in Dicotyledons (TAKHTAJAN, 1969; CRONQUIST, 1968). The
seedlingsofSapotaceae can beclassified inthis way into two basictypes:
In table 1 the seedling types mentioned here are summarized; the figures
are schematic.
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ad 5. Unisexual flowers are uncommon inAfrican Sapotaceae.
ad 6. All the three subfamilies, distinguished by means of the number of
calyxwhorls,include A- aswellasB-type seedlings.
ad 10. Dorsal appendages are rare in Sideroxyloideae (Kantou, B-type) and
absent in Omphalocarpoideae (predominantly A-type) but common in
Mimusopoideae (A-and B-type).
ad 13. Staminodes occur in taxa of each seedling type. In some taxa sta-
minodes are strongly reduced or lack completely: Pachystela, Zeyhe-
rella, Wildemaniodoxa, all with B-type seedlings. This might indicate a
correlation with the advanced seedling type in Sideroxyloideae only,
but Gambeya subnuda contradicts this view (reduced staminodes and
A-type seedling). Moreover several taxa with B-type seedlings have
well developed staminodes (Synsepalum, Kantou, Gluema, Butyrosper-
mum). Hence AUBRÉVILLE'S opinion that reduction of staminodes is an
advanced character in Sapotaceae isnot supported bythis evidence.
On the other hand more or less well marked correlations seem to exist
between seedlingtype and thefollowing characters:
14. level of insertion of the stamen on the corolla tube;
15. number of locules in the ovary;
16. number of developed seeds in the ripe fruit;
17. scar position on the seed;
18. shape and dimension of the scar;
19. endosperm development in ripe seeds;
20. cotyledon development in ripe seeds.
Evolutionary advancement
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As this investigation was initiated during the period I was working in the
Faculty of Agronomy of the National University of Zaire (1970-1974), my
thanks areduetotheBoard ofthe University andtotheDean ofthe Faculty of
Agronomy for their permission to work on this subject. Cits. YOSWA BOHELE
and MASSUDI M A N D A K E N D A were valuable assistants in the field, Cit. SALUMU
MAJANGA BIN ATIBU was very helpful inthe Herbarium of the Faculty of Agron-
omy.
F o r hospitality and loans, my thanks are due to the Director of the
Herbarium of the National Institute for Agricultural Study and Research
(I.N.E.R.A.) atYangambi, Zaire, andtotheDirector of the National Botanical
Gardens of Belgium at Brussels, Dr.J.J. DEMARET. Dr.ir.J. J. F .E. DE W I L D E
kindly sent me living seeds from Cameroon.
At the Laboratory of Plant T a x o n o m y and Plant Geography of the Agri-
cultural University ofWageningen, I wish to express myacknowledgements to
Prof. dr. H . C. D . DE W I T , Ir. J. C. ARENDS, M . S C , Ir. J. J. Bos, Dr. ir. F . J.
BRETELER and Dr. A. J. M . LEEUWENBERG for their interest in my work and
their critical remarks. M r s .F . N . BATA-GILLOT made thedrawings, which are
a very essential part of this paper. Miss D . M . WASSINK (typescript), M r .
G. BOELEMA (polishing and correcting manuscript and proofs), M r . J.DE
BRUIJN (Herbarium), M r . C. T. DE G R O O T (Library), M r . W. H . GROTENBREG
and the late Miss M . TEULINGS (handling seedlings in the greenhouse) con-
tributed inavery friendly and valuable way tothis paper.
Specimens examined:
IVORY COAST: Cremers 580(BR), 3seedlings; Cremers440(BR), 1seedling; De Koning
2522,2945,3190,3363,3617 (WAG),ca.30seedlings,voucher: DeKoning2249(WAG).
Specimens examined:
IVORY COAST: Cremers 618 (BR), 1 seedling.
ZAIRE: Bokdam 4376 (WAG), 2 seedlings, voucher: Bokdam 4375 (WAG); Dubois 526bis
(BR), 4seedlings, voucher: Dubois 526(BR).
Specimens examined:
ZAIRE: Bokdam 4401 (WAG), 3seedlings, voucher: Bokdam4400(WAG); Bokdam4490
(WAG), 3seedlings, voucher: Bokdam 4489(WAG); Bokdam 4579 (WAG), 2 seedlings;
Bokdam 4619 (WAG), 2 seedlings, voucher: Bokdam 4656 (WAG); Hombert 246 (BR),
1 seedling; Hombert 295(BR), 1seedling, voucher: Mahieu 156(BR); HPLY 3311 (YBI),
voucher: O728 = B210(YBI);Maudoux 109(BR), 2seedlings.
Specimens examined:
ZAIRE: Bokdam4379 (WAG), 1 seedling, voucher: Bokdam4377(WAG); Bokdam 4399B
(WAG),2seedlings, voucher: Bokdam 4399A (WAG).
*In the specimens examined no real eophylls werepresent. Therefore, the description has
beenbased onthefirstleavesofthelateralshoots,replacingthedamagedeophylls.
References ( + figure):
JACKSON, 1968.
dots,sericeouswithgreyishtosilveryhairsbelow.Nervationeucamptodromous;
primary vein straight, weakly developed, impressed above, prominent below;
6-9(11) pairs of secundary veins,divergent at widely acute anglesat the base
and at moderately acute angles near the apex of the leaf, uniformly curved;
many intersecondary veins, parallel to the secondary veins; tertiary veins
originating exmedially at moderate to narrowly acute angles, admedially at
obtuseangles,confluent withtheintersecondary veins,endinginthemidribor
intheeucamptodromous loop;quaternary veinsreticulating together withthe
Meded.Landbouwhogeschool Wageningen 77-20(1977) 43
tertiary veins; aréoles well developed, large, elongate and parallel to the sec-
ondary veins, hairlike veinlets visible in transmitted light.
Specimens examined:
IVORY COAST: / . J. de Wilde 258 (WAG), 1seedling.
References:
D E LA MENSBRUGE, 1966.
Specimens examined:
ZAIRE: Bokdam 4374(WAG), 3seedlings, voucher: Bokdam 4362(WAG); HPLY 2874
(YBI), 1seedling; Teulings 15(WAG), 1seedling,voucher: Bokdam4402(WAG).
TB
FIG. 13. Gambeya lacourtiana ( D E WILD.) AUBRÉV. et PEULEGR. - 1. seed, lateral, l x
2. seed, ventral, 1x ; 3.habit, 1/i x .
After: Bokdam 4549 (1,2), 4608 (3).
Specimens examined:
IVORY COAST: Tolliez 262,273 (BR), 8seedlings.
ZAKE: Bokdam 4297(WAG), 1seedling, voucher: Bokdam 4296(WAG); Bokdam4398
(WAG), 5seedlings, voucher: Bokdam 4397 (WAG); Bokdam 4554, 4569, 4610 (WAG),
10seedlings,voucher: Bokdam4553 (WAG).
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Specimens examined:
ZAIRE: Bokdam4481 (WAG), 2seedlings, voucher: Bokdam4480(WAG); Bokdam 4582
(WAG), 1 seedling,voucher: Bokdam4480(WAG).
Specimens examined:
GABON: Breteler 7189 (WAG), 6 seedlings.
IVORY COAST: Cremers 407 (BR), 2 seedlings.
Hypocotylstraight,cylindrical, 10-20cmlong,brown,glabrous;rootcrown
straight;apicalgreenherbaceouspart4-15mmlong.
Cotyledonsfree, papery, persistent; petiole flattened, 0.5—1.2 mm long;
lamina elliptic, asymmetrical, 7-12 cm x 5-8 cm, base truncate to cordate,
apexrounded;both faceswithminutecolourlesspapillae.Nervation brochido-
dromous;primary veinslightlyzig-zag,weaklydeveloped, prominent onboth
faces;6-10pairsofsecondaryveinsweaklydeveloped,prominentbelow,angle
ofdivergence narrowly to moderately acute, straight, abruptly looped tojoin
the superadjacent secondary vein at a wide angle; few intersecondary veins;
tertiary veins originating exmedially at moderate to widely acute angles,
admedially at right angles,percurrent, sinuous; quaternary veins reticulating;
aréoles imperfect to well developed, at random arranged; veinlets mostly
present.
Epicotylstraight, cylindrical, 1-2.5 (4.5)cmlong,pubescent to glabrescent,
1-2 pubescentcataphyllspresent.
Eophyllsalternate,papery,exstipulate;petiolesemi-terete,canaliculate, 4-7
mm long; lamina narrowly obovate to elliptic, 6-22 cm x 2.8-8 cm, base
cuneate, apex acuminate, acumen rounded, 0.4-2.5cmlong,glabrous above,
glabrescent below, mainly on the midrib. Nervation eucampto-brochido-
dromous;primary veinweakly developed, straight, prominent on both faces;
6-10 pairs of secondary veins, prominent below, angle of divergence moder-
ately acute; tertiary veins originating ex- and admedially at right angles,
sinuously percurrent; quaternary veins reticulating; aréoles imperfect to well
developed, triangular to polygonal, at random arranged;veinlets simple to
repeatedly forked.
Specimens examined:
ZAIRE: Bokdam4390(WAG), 3seedlings, voucher: Bokdam4389(WAG); Bokdam4590
(WAG), 1seedling, voucher: Bokdam 4567(WAG); HPLY 7850(YBI), voucher: O2741
(YBI); HPLY 689 (YBI), voucher: O 373 (YBI); Bokdam 4311, Teulings 8, 12 (WAG),
5seedlings, voucher: Bokdam 4312 (WAG).
FIG. 18. Omphalocarpum lecomteanum PIERRE ex ENGL. - 1. seed, lateral, ilix- ; 2. seed,
ventral, V2x ; 3. habit, V2x ; 4. detail nervation cotyledon, 1 x ; 5. detail nervation
eophyll, 1 x .
After: Bokdam 4389 (1-3), 4590 (4, 5).
Specimens examined:
IVORY COAST: Tolliez 236 (BR), 6 seedlings.
ZAIRE: Bokdam 4630 (WAG), 1seedling, voucher: Bokdam 4507 (WAG).
Specimens examined:
Bokdam 4595(WAG), voucher: greenhouse nr. 713(WAG) ( = Bokdam 4586(WAG));
Bokdam4623(WAG),voucher:greenhousenr.713(= Bokdam4621 (WAG));both voucher
plantsarefloweringand fruiting inthegreenhouseat Wageningen.
Specimens examined:
ZAIRE: Bokdam 4217,4255(WAG), 8seedlings, voucher: Bokdam 4069(WAG); HPLY
6986(YBI), voucher: O 161 (YBI); Teulings 16(WAG),4seedlings, voucher: Bokdam 4550
(WAG).
Hypocotylstraight,cylindricaltoangular,compressedbelowthecotyledons,
ca.6-10cmlong,ca. 3mmindiam.,glabrous,dark green;rootcrown straight.
Cotyledons free, fleshy, light to dark green, rather caducous, sessile,plano-
convex,elliptic-ovate or narrowly so,asymmetrical, 4-5 cm x 1.5-2.2cm x
0.6-0.7cm,glabrous.Nervation hyphodromous.
Epicotyl straight, strongly compressed, (3)5-6.5 cm long, 2.5-3 mm in
diam., glabrous; cataphylls absent.
Eophylls alternate, papery, exstipulate; petiole slightly grooved, glabrous,
5-8 mmlong;laminaelliptictoobovate,ca.7.5cm x 3.5cm,basedecurrent,
apex acuminate-mucronate. Nervationbrochidodromous, with 7-8 pairs of
secondary veins;a pattern of sinuous lactiferous ducts, from baseand midrib
to the margin and parallel to the secondary veins, is conspicuous in young
eophylls.
Specimens examined:
IVORY COAST: De Bruijn 1943(WAG), 2seedlings,voucher: Versteegh 782(WAG).
References:
D E LA MENSBRUGE, 1966.