BLUMEA 47 (2002) 363-383
Seed morphology of some tribes of Brassicaceae
(implications for taxonomy and species
identification for the flora of Egypt)
K.+Abdel Khalik & L.J.G. van der Maesen
Summary
of 23 of Brassicaceae examined Seed morphology 45 taxa belonging to genera were using light and scanning electron microscopy. The taxa included representatives of the tribes Arabideae,Euclidi-
eae, Hesperideae,Lunarieae,Matthioleae, and Sisymbrieae, which all occur in Egypt. Macro- and
micromorphological characters, including seed shape, colour, size, the position of the radicle relative
to the cotyledons, epidermal cell shape, anticlinal boundaries, outer periclinal cell wall and relief
of outer cell walls, are presented. Three types ofbasic anticlinal cell wall boundaries are recognised
and six different shapes of the outer periclinal cell wall are described. A key for the identification
of the investigated taxa based on seed characters is provided.
Key words '. Brassicaceae, SEM, seed coat.
Introduction
The Brassicaceae (Cruciferae) are one ofthe largest angiosperm families, comprising
approximately 340 genera and more than 3350 species in 10 poorly defined tribes,
distributed throughout the world, chiefly in temperate regions of the northern hemi-
sphere (Al-Shehbaz, 1984). The major centres of distributionofthe family are in the
Irano-Turanian, Mediterranean, and Saharo-Sindianregions (Hedge, 1976).
In the flora ofEgypt, Brassicaceae are one of the four largest plant families, repre-
sented by about 102 species belonging to 55 genera,assigned to 9tribes (Schulz, 1936).
Species of the tribes Lepideae, Brassiceae, and Alysseae (63 species belonging to 32
genera) have been the subject of an earlier study (El Naggar, 1987; El Naggar & El
Hadidi, 1998). The remaining Brassicaceae, belonging to the tribesArabideae, Eucli-
deae,Hesperideae, Lunarieae,Matthioleae,and Sisymbrieae (Schulz, 1936) (39 species
in 23 genera) exhibit great diversity, as shown in this revision. The membersof these
tribes distributed wide of habitats of farmland and are over a range as weeds waste
places, desert land, Mediterraneancoastal land, oases; mountains of Sinai and Elba,
as well as inland flats and canal banks.
that data the and microstructure of Most systematists agree concerning macro-
seeds are very significant for the classification of angiosperm taxa. Heywood (1971)
drewattention to the importance and impact of scanning electron microscopy (SEM)
in the study of systematic problems, as very valuable informationhas been provided
1) South Valley University, Faculty of Science, Botany Department, Sohag, Egypt.
2) Nationaal Herbarium Nederland, Wageningen University branch, Biosystematics Group,
Generaal Foulkesweg 37, 6703 BL Wageningen,The Netherlands. 364 BLUMEA Vol. 47, No. 2, 2002
this the last decades, scholars have SEM to by using technique. During many applied morphological studies of seeds and small fruits. Micromorphology and ultrastructural
data have contributed useful informationfor evolutionand classification ofseed plants
and play an important role in the modern synthetic systems of angiosperms (Dahlgren,
1979-1980).
Most studies focused on intrageneric seed coat variation(Chuang & Heckard, 1972;
Hill, 1976; Heyn & Herrnstadt, 1977; Clark & Jernstedt, 1978;Newell & Hymowitz,
1978; Canne, 1979; Crow, 1979; Wofford, 1981; Juan et al., 2000; Segarra & Mateu,
2001) or on variation among several closely related genera(Musselman & Mann, 1976;
Seaveyetal., 1977;Canne, 1980; Chance & Bacon, 1984; Matthews& Levins, 1986;
Fayed & El Naggar, 1988,1996;Haridasan & Mukherjee, 1993; Shanmukha & Leela,
1993; Kanak Sahai et al., 1997; Karam, 1997; Koul et al., 2000). Less commonly,
SEM level variation was used to place taxa into tribes (Whiffin & Tomb, 1972).
For furtherinformationand references concerning seed forms and internal structures
see Corner (1976). Seed appendages were studied by Kapil et al. (1980); for seed size
and number see Aniszewski et al. (2001); for the colours of seeds refer to Berggren,
1962; Huber, 1969; Dahlgren & Clifford, 1982; Barthlott, 1984; while epidermal cell
patterns and distributionofelements such as trichomes, glands, and stomata are given
by Stace (1965). The distributionofepidermal cells is of importance between species
and genus level (Barthlott, 1981, 1984; Haridasan & Mukherjee, 1993); while outer
periclinal walls are also a good diagnostic character for the lowesttaxonomic categories
(Barthlott, 1981, 1984; Fayed & El Naggar, 1988, 1996).
In Brassicaceae, the external features of the seeds colour and shape are studied by
Kondo (1917), McGugan (1948), Musil (1948), and Murley (1951). Berggren (1962)
investigated the external morphology and anatomy ofseeds testa ofBrassica. Vaughan
& Whitehouse (1971) studied the external morphology and anatomy of seeds of 90
generaand 200 species of the Cruciferae and paid special attentionto the relationships between seed structure and the existing taxonomy. Prasad (1976) investigated the
seed coat structure and development in certain species of Brassicaceae. Mulligan &
Baily (1976) and Stork et al. (1980) studied some taxa of Brassica seeds with SEM
reclassification of and they suggested other data that help in the the genus. Fayed &
El Naggar (1988) investigated the seed coat sculpture in species of the tribe Brassiceae
in Egypt by using SEM; several details offered taxonomic value at the generic and
subtribal such the seed and cells anticlinal levels, as shape size, epidermal pattern,
wall boundaries, and outer periclinal cell walls. Jonsell (1975) studied the seed coat of Lepidium in East tropical Africa; Fayed & El Naggar (1996) investigated the seed
morphology and taxonomy with SEM in the tribe Lepidieae in Egypt.
Jonsell (1986) studied the seed coat of genusFarsetia by using SEM to distinguish between species. Stork (1971, 1972) and Stork & Wiiest (1978) investigated the seed
coats and development of epidermal slime bodies ofMalcolmiaand foundlarge differ-
ences between the species. Jonsell(1979) used SEM on the seed surface to distinguish
Matthiola, Morettia, and Diceratella. Stork & Wiiest (1980) studied the seed coat
morphology, particularly the epidermal slime structure ofMorettia.Koul et al. (2000)
investigated the seed coat microsculpturing in Brassica and alliedgeneraand provided evidence for the close relationships among various generawithin subtribe Brassicinae and of subtribes Raphaninae and Moricandiinaewith the Brassicinae. K. Abdel Khalik & L.J.G. van der Maesen: Seed morphology of Brassicaceae tribes 365
The deal with the and & present investigations macro- micromorphological (LM
SEM) characters ofseeds of23 genera(39 species) of6 tribes from thefamily Brassica-
for ceae in Egypt, to decideon the importance of seed characters as a criterion sepa-
in this The used here follows authors rating genera and species family. terminology
such as Barthlott (1981, 1984), Fayed & El Naggar (1988, 1996), Karam (1997), and
Koul et al. (2000) with some modifications by the author.
MATERIAL AND METHODS
Some of the investigated seeds were collected from mature plants in Egypt during
1998-2000.The otherswere supplied by INIA, Madrid, Spain or taken from herbarium
specimens (see Table 1).
Only mature seeds were taken for investigation. The dried seeds were first examined
by light microscope (Carl Zeiss 475002), and 7-15 seeds for each taxon were chosen
stubs with double adhesive to cover the range of variation. Seeds were mounted on
with for 2-3 min. in Polaron tape. The stubs were sputter-coated gold-palladium a
Equipment Ltd. SEM coating unit E 5100.After coating, the specimens were examined
•with a Jeol JSM 5200 scanning electron microscope, using accelerating voltages at
of Plant and 15-20 KV. All photomicrographs were taken at the department Cytology
Morphology, SEM laboratory, Wageningen University, The Netherlands.
RESULTS
Seed characters are very important to separate among genera and species levels in
Brassicaceae. Seed shape varies fromorbicular, suborbicular, oblong, ovoid, to oblong-
ovoid. Seed size varies from large 3-8 by 2.5-7 mm to small 0.5-1 by 0.3-0.9 mm
and helps to separate between species. Seed colour varies from brown, yellow-brownish
to red-brownish. Radicle position relative to cotyledons varies from accumbent to
incumbent. Epidermal cell shape varies from isodiametric, 4-5-6-polygonal to elongate
in one direction. Anticlinal boundaries vary from raised to channelledand anticlinal
sinuous and the surface varies from smooth walls from straight to or slightly sinuous;
fine folds. Periclinal cell walls showed variation to or coarse a large among genera
and species level, and can either be flat, concave, convex, or domate; smooth, micro-
or macroreticulate, striate, and possess fine to coarse folds and papillae.
of radicle relative The seed shape, absence/presence a wing, size, colour, position
to the cotyledons, epidermal cell shape, and the characters of anticlinal boundaries
and periclinal cell walls of each taxon (39 species belong to 23 genera) are given in
Table2.
DISCUSSION OF SEED CHARACTERS
1. SEED SHAPE
variation. of The shape of seeds among the investigated taxa showed a large Most
the seeds vary from oblong or ovoid to oblong-ovoid or oblong-ellipsoidal (Table 2),
however, they are orbicular in Ricotia lunaria (Fig. 11); sub-orbicularto sub-orbicular-
ovoid in Matthiolalongipetala subsp. hirta, Arabis alpina subsp. caucasica, Neslia 366 BLUMEA Vol. 47, No. 2, 2002
Table 1. List of Brassicaceae seed specimens used in Scanning Electron Microscope (SEM) studies.
taxon collection no.n 1.1. Tribe Arabideae Arabis alpina L. subsp.subsp. 1313 Morocco, Taza,Taza, around gebel Tazzeka (NMGM) caucasica (Willd.)Briq. Switzerland,Valois, Val de Arabis nova Vill. 20273 Switzerland,Valois, Val de Bagnes (BR) Nasturtium officinaleofficinale R.Br.R. Br. 415 Rorippa indica (L.) Hiern s.n. Egypt, Qena, El Mahrosa islandisland (SHG) Rorippa integrifolia Boulos 9 Egypt, Middle of Sinai, aboutabout 5 km beforebefore Ras Sedr (SHG) Rorippa palustris (L.)(L.) Besser 20 Egypt, along the canal of Ismailiya (SHG) 2. Tribe Euclidieae Anastatica hierochunticahierochuntica L. 327 Egypt, Cairo-Suez road, 95 kmkm fromfrom Cairo (CAI) Neslia paniculatapaniculata (L.) Desv. 64 Iran,Iran,nearnear Gere between Abushir andand Shiraz (WAG) Ochthodium aegyptiacum (L.)(L.) DC. 129 Palestine, Jerusalem: Mt. Scopus (BM) Schimpera arabica Hochst. & Steud.Steud . 3747-75 Spain, Madrid (INIA) 3. Tribe Hesperideae Eremobium aegyptiacum (Spreng.) 29 Egypt, El Arish areaarea beside thethe airport, N Sinai Asch. & Schweinf. subsp. (SHG) aegyptiacumaegyptiacum Eremobium aegyptiacum subsp. s.n. Egypt, El Tor desert, SinaiSinai (L)(L) linearelineare (Delile) Abdel Khalik in near (LY) Eremobium aegyptiacum subsp. s.n. Egypt, in sandssands Asswan longisiliquumlongisiliquum (Coss.)(Coss.) Maire Erysimum repandum L. 1163-67 Spain, Madrid (INIA) LeptaleumLeptaleumfllifoliumfilifolium (Willd.) DC. s.n. Egypt, Wadi Chafura, north Galala (BR) Malcolmia africanaafricana (L.) R.Br. 2 Egypt, Deir el Rahba, Wadi el Arbain, SS Sinai (SHG) Malcolmia Boiss. 28 El Arish-Rafah road, 23 km fromfrom El Arish Malcolmia nana (DC.) Boiss. 28 Egypt, El road, 23 km (SHG) Malcolmia Boiss. 22 Alexandria-Matruh coastal 26 pygmaea (Delile) Egypt, road, km, Sidi Krer, (SHG) 4. TribeTribe Lunarieae Ricotia lunaria (L.) DC. 45,00445,004 Syria, Golan, Yahudiya forest (B) 5. TribeTribe Matthioleae Diceratella elliptica (DC.)(DC.) JonsellJonsell 17 Egypt, Gebel Elba, Wadi el ShallalShallal (SHG)(SHG) Matthiola arabica Boiss.Boiss. 12 Egypt, Wadi el Arbaien, S. Sinai (SHG) Matthiola fruticulosa (L.) Maire 1221-66 Spain, Madrid (INIA) Matthiola longipetala (Vent.) DC. 43794379 Turkey, Tassia, inin sandssands near Saban Dagi (BR)(BR) subsp. bicornisbicomis (Sibth.)(Sibth.) Ball Matthiola longipetala subsp. 2323 Egypt, Alexandria-Matruh coastal road, near Burg hirtahirta (Conti) Greuter & Burdet elel Arab (SHG) Matthiola longipetala subsp. 77 Egypt, Cairo-Alexandria desertroad,road, 100 km fromfrom kralikiikralikii (Pomel) Maire Alexandria (SHG) Matthiola longipetala subsp. 27 Egypt, Cairo-Ismailiya desert road,road, 23 km before lividalivida (Delile) Maire IsmailiyaIsmailiya (SHG)(SHG) K. Abdel Khalik & L.J.G. van der Maesen: Seed morphology of Brassicaceae tribes 367 Table 1 (continued) taxon collection no. place of collections Matthiola longipetala subsp. 25 Egypt, 2828 km before Matruh, Alexandria-Matruh longipetala(Vent.)(Vent.) Boulos road (SHG) Matthiola parviflora (Schousb.) 0907-66 Spain, Madrid (INIA) R.Br. Morettia canescensBoiss. 1098-67 Spain, Madrid (INIA) Morettia parviflora Boiss. 16984/216984/2 SaudiSaudi Arabia, Wadi Harjab, rocky hillsideshillsides (BR) 3 Morettia philaeana (Delile)(Delile) DC. 3 Egypt, 35 km from Qenaonon the road Qena-Safaga (SHG) Notoceras bicorne (Aiton) CaruelCamel 1052 Egypt, Gebel Elba, Saddle between Gebel Asotriba and Makin (CAI) 6. Tribe Sisymbrieae Crucihimalaya kneuckeri (Bornm.) 49785 Egypt, Sinai, E slopesslopes of Gebel Catherine (CAI) Al-Shehbaz, O'Kane & R.A. Price Descurainia Webb & 1475 sophiaSophia (L.)(L.) Jordan, El-Jubeiha, University campus (B) Berthold Nasturtiopsis coronopifoliacoronopifolia (Desf.) 3663-76 Spain, Madrid (INIA)(INIA) Boiss.Boiss. Neotorularia aculeolata (Boiss.) 5801 Iran, West of Jaz Murian (BR) Hedge & J. Leonard Neotorularia torulosa (Desf.) 1461-681461-68 Spain, Madrid (INIA) Hedge & J. Leonard Olimarabidopsis pumilapumila (Stephan) 779 Syria, nearDamascus, roadroad of Zahroub toMerdj, Al-Shehbaz,O'Kane & R.A. PricePrice Baustam el Mahar (L) Robeschia schimperi (Boiss.)(Boiss.) 170170 Egypt, Sinai Mts, between graniticgranitic rocks (B)(B) O.E. Schulz Sisymbrium erysimoides Desf. 1919 Egypt, Gebel Elba, Wadi Drawina (SHG) Sisymbrium irio L.L. 31 Egypt, S Sinai, Deir el-Rahebat, Wadi Feiran, as weed in garden (SHG) Sisymbrium orientale L.L. II11 Egypt, S Sinai, Deir Feiran, Wadi Feiran (SHG) Sisymbrium polyceratum L.L. 6627-846627-84 Spain, Madrid (INIA) Sisymbrium runcinatum Lag. ex DC. s.n.s.n. Algeria, El Kantara, near Constantine (BR) Sisymbrium septulatum DC. 3719-753719-75 Spain, Madrid (INIA) paniculata, Eremobium aegyptiacum subsp. lineareand Diceratella elliptica (Fig. 1, 7, Seed was found useful the allied Matthiola 12). shape to separate closely genera with oblong seeds; Morettia with ovoid seeds and Diceratella with suborbicular to elliptical and these results congruence with the results of Jonsell (1979). The shape of seeds showed a significant difference between the subspecies of Eremobium in and aegyptiacum: they are oblong subsp. aegyptiacum subsp. longisiliquum and sub-orbicular to ovoid in subsp. lineare (Fig. 7, 8). Most of the seeds examined have no wings or only narrow wings, but in Arabis alpina subsp. caucasica, Eremobiumaegyptiacum subsp. lineare, Matthiolafruticulosa and Matthiolalongipetala subsp. hirta seeds have a broad wing (Fig. 1, 7). continued (text on page 376) 368 BLUMEA Vol. 47, No. 2, 2002 fine to toto striatestriate loto micro- micro-micro- to central toto loto concaveconcave central slightly mediummedium folds papilla; folds toto smooth oror smoothsmooth convex; smooth concave; convex; smooth smoothsmooth wall globularglobular with concave; flat radiate-striate fine cell position; central smoothsmooth with withwith macro-reticulate concave; slightlyslightly concave;concave; convex to slightly slightly convex;convex; convex; concave;concave; macro-reticulate micro-reticulate micro-reticulatemicro-reticulate convex micro-papillate folds micro-reticulate or to to to smooth toto to to toto smooth to reticulatereticulate reticulate to periclinal central smooth papilla; sunken smooth foldsfolds folds fine periclinal domate domatedomate flat; flatflat flat flat flat concave; flat flatflat flat flat flat fine fine to fine fine to smooth sinuous; straightstraight sinuous; slightlyslightly to slightlyslightly to slightly to toto slightly to folded to to to folds sinuous; foldsfolds straight; straight;straight; toto to to to smoothsmooth to straightstraight straight; fine finefine slightly boundaries sinuous; smooth folds smooth smooth raised-channelled, smooth smooth straightstraight folds straight; straightstraight straightstraight straight to raised, toto to raised, raised,raised, straight folds foldsfolds sinuous;sinuous; folds slightly coarse sinuous;sinuous; medium sinuous;sinuous; sinuous; smooth raised-channelled,raised-channelled, smoothsmooth straight sinuous; smooth smooth sinuous; coarse fine finefine fine foldsfolds foldsfolds anticlinal channelled, raised,raised, raised,raised, raised,raised, raised,raised, raised, channelled, to slightly slightlyslightly slightly slightlyslightly raised, in shape elongateelongate direc- polygonal 5-6-poly-5-6-poly- elongate polygonalpolygonal 4-5-gonal direc- 4-5-6- 5-6-gonal,5-6-gonal, polygonalpolygonal elongate one cell one directiondirection directiondirection in in polygonalpolygonal direction one one elongate Brassicaceae. epidermal gonal inin tion, one gonal tion epidermal tion, tion in 4-5-polygonal, 4-5-6-gonal, 4-5-polygonal, 5-6-gonal, one isodiametric, isodiametric,isodiametric, in isodiametric,isodiametric, isodiametric, isodiametric 4-polygonal4-polygonal elongateelongate 5-6-gonal, isodiametric, isodiametric,isodiametric, isodiametric, of accumbent accumbentaccumbent accumbent accumbentaccumbent accumbentaccumbent accumbent accumbent incumbentincumbent incumbent incumbent accumbent accumbentaccumbent accumbentaccumbent descriptions radicle Seed colour brown brownbrown 2. orange-brownorange-brown yellow-brownish red-brownishred-brownish seed light lightlight brown red-brownivcl-hnmn brown brown brownbrown brownbrown brown brownbrown Table (mm)(mm) 1.2-1.3 0.3-0.6 0.3-0.4 0.75-0.9 0.7-0.80.7-0.8 0.5-0.750.5-0.75 1-1.7 1.5-1.7 size xx x x x 1.3 1.2 1.6 x 1 seed 1.3-1.5x1.3-1.5x1.2-1.3 0.7-1x0.3-0.4 l-1.9x1-1.9x1-1.7 1.5x1.3x 2.5-3x1.5-1.7 1.2-2.7x0.5-1.2 xx 2.1x1.6x x 0.7-1 0.9-1 0.8-10.8-1 0.8-10.8-1 0.5-0.9 1.5 2.5-3x 2.2 2.1 2.12.1 wing wide none narrow narrow none nonenone nonenone none none none narrownarrow widewide narrow seed toto ellipsoidellipsoid loto sub-sub- shape tolo toto sub-orbicular oblong-ovoidoblong-ovoid cordiform ovoidovoid orbicularorbicular ovoid-oblongovoid-oblong oblong-ovoid sub-orbicularsub-orbicular ovoid seed ovoid ovoid ovoidovoid ovoid ovoidovoid oblongoblong oblongoblong aegyptia-aegyptia- officinaleofficinale arabica Hesperideae aegyptia- aegyptia- lineare aegyptia- longi-longi- ArabideaeArabideae caucasicacaucasica indica integrifoliaintegrifolia palustrispalustris Euclidieae paniculatapaniculata alpina novanova hierochuntica subsp.subsp. subsp.subsp. subsp.subsp. taxontaxon TribeTribe subsp. Tribe hierochuntica aegyptiacumaegyptiacum Tribe siliquum Arabis subsp. Arabis NasturtiumNasturtium Rorippa RorippaRorippa RorippaRorippa Anastatica NesliaNeslia OchthodiumOchthodium Schimpera Eremobium cum cum EremobiumEremobium cum EremobiumEremobium cum 1. 2. 3. K. Abdel Khalik & L.J.G. van der Maesen: Seed morphology of Brassicaceae tribes 369 1) i to3 X X rt E — p Z 3 3 y -3 5 z a to2 3 c folds micro- c fine coarse a 3 central 1 folds •x smoothE smoothE 0 --J u Sxj medium w central * 4, u fi E « to c o u 4i ■g concavec 3 k > > « -3 sunken3 0 3 s ,zfolds striateE flat to 0 id fine a 3 3 3 coarseC 3 smooth 'J > u u 3 •J I or ■5 Si 3 3 3 3 3 toQ 3 3 -O 3 striate striate concave;O c concave;0 concave;O V convex; convex; fine wall C smooth fine p X C '4= O 0 O 5 with1 to 1 with X globular flat XJ 00. ■ u Q tog xT* C radiate-striate 3 — "C S 'a. fi -3 ■a Z X x 4) cell5 3 fine u X position; 43 papilla; X xs > > X "a, > smooth — folds"e foldsc folds smooth£ a "J 50 51j 5/J eo a "a with with 3 3 C ifi z 3 z 3 P P convex;Z slightly1_ slightly slightly slightly z slightly convex convex - convex; convex; c a convex;C _3 folds ij *" *3 7^ Z o 8*S u g 1« u u 'J 3 p « --> — M smoothE V 3 fine fine _ o 1 u C- 3 '& a 0 a micropapillate jf3 3 tog to to2 Sto tog to3 s to to or or to 3 - papillate papilla; 0) portion;p 0 3 E § central central striate coarse — fine 3 to to folds E 3 o 3 u 3 3 periclinal 5 — 3 3 n —■ convex;c convex;3 domate domate - flat flat flat flat flat- flat flat flat flat flat flat; flat a "3 -3 zz 53 c C3 o € c c 53 € C u c X to BS >> u >. u toO 0 >. 5J 3 3 — a c t3 c c X X O X X 3 fine fine fine fine 5fl xa u= -5 J= '4= X3 (3 xt« X 3 c 3 s tog o tog to o c c 5 smooth bJj-o H ~ 00 50 sinuous;3 sinuous; 0 to c c c slightly slightly S slightly to O to to '5 '3 c J= £ C3 s s 3 folds _e folds smooth 3 smoothE smooth smoothE E X X straight; fine smooth| JH straight; Si 1 straight X o a o c £ to to to— to to B 3 O u oj *- 0 0 c z 3 — 3 — 0 z c 3 '5 E sinuous straight — r. X -3 J3 ■d fine X! W3 x w fine X J= ■d X X 4= x c boundaries '1 E E £ ■q £ 5 S smooth smooth Q smooth so-a -a smooth u smooth 60~0 5JJ straight; 5lj 50 50 60 9 51j op C '" 2 to "O . 'S3 '3 c '3 c '3 to '3 a p r1 '§ raised, » folds folds raised, raised, folds ■3 rt straight; c« straight; straight; straight; d straight; — 2 straight;jj — straight straight J3 x £ s _c jg 3 3 foldsc o 4> _= >> 0 0 >^ 0 -^ . . . 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"u e E E E E E E E E E E E 3 3 E £ 3 3 3 3 3 3 3 3 '■3 3 3 3 3 u 0 0 0 O O 0 incumbent incumbent incumbent incumbent incumbent accumbentO accumbent accumbentu accumbent accumbent accumbent accumbent accumbent accumbent accumbent accumbent•j radiclew O u O 0 ■-< o u u u Q 3 3 3 3 3 u « 3 x XI -3 X3 X X X 3 3 3 3 3 3 3 S s * =: 3 3 " 3 3 3 C C p P O is -C X X X colour"o x X 0 "T brown brownP X - i is 3 X £ s 3 S s 3 3 3 3 s p _c _c _C _c _c j= yellow-brownish yellow-brownish yellow-brownish X yellow-brownish yellow-brownish yellow-brownish yellow-brownish 0 z c p p p seed r;fj Ti ti 5ij u brown u flj brown light brown brown brown brown brown light >-, >* js >. s J5 X X X >> >. 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'a. to to to to to2 -a 2 2 2 > > reticulate 5 reticulate o c 3 c a *T a portion;0 g papilla;« 5 portion; S papilla; papilla; "3 papillae; striate central central striate *- Sea folds folds li B u folds 1 S a a u w 0or CI- a •- «*•- fl periclinal domateO convex;o convex;o flat flat domate convex; domateo domateo domate0 flat flat flat domate domate TS flat T3 T3 T3 C TS -0 a. -_i o c EC c ■0 5 d S3 ■5 x >* to2 U ~ M> O g 3 c c illfine 1 -S deep fine x tSfine to b0 O to smooth straight .5? c 00 slightly 3=2to sinuous; 2 8 to to2 to « g 2 8 >i slightly slightly slightly ° slightly — - jr 45 '3 smooth smooth folds 3 smooth 'g folds "*- — straight; -5 straight; s § 00 Si O G O T3 X -5 o£ iU o foldso 3 c 0 Ss to to to U to to js B ,,«Q to— 3 - 0 c — 4- a -8 8 ,SP si a « o — O c G ° c sinuous;g c slightly 0 fine sinuous; _' straight1 sinuous fine S ■ -o -3 ■/ -3 T3 x 3 2 ILS-M - - boundaries smooth| -G J= smooth « E E ■a fl '9 u "3 "3 ' "3 "3 "3 >. c5 "3 "3 >-. 75 c 1 i 3 in.S 0 vO c C >. 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