TAXONOMIC STUDIES OF NORTH AFRICAN
CARYOPHYLLACEAE WITH SPECIAL
REFERENCE TO THE FLORA OF
LIBYA
BY
MOHAMED NURI ABUHADRA
THESIS PRESENTED FOR THE DEGREE
OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF GLASGOW
1996
ProQuest N um ber: 11007819
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2
ABSTRACT
The thesis deals with the Caryophyllaceae, a family well represented in the
countries of North Africa including Libya which has about 22 genera and 80
species within the three subfamilies as currently recognised. The principal
part of the thesis concerns the description of the seeds of the Libyan
species primarily with the use of scanning electron microscope (SEM).
SEM produces very revealing images of the testa cells and their often
striking ornamentaion. These seed characters as well as seed colour,
shape and size, hilum position and radicle shape have been applied to
particular problems at several taxonomic levels up to that of the subfamilies,
the
limits
of which
have
long
been controversial.
Within
the
Caryophyllaceae, whatever its scope, seed characters have always been
given importance but that importance is often under-rated.
A survey of the calcium oxalate crystals in the genera Arenaria,
Minuartia and Moehringia has been undertaken to an extent and in a detail
not previously carried out within any large genus of the family. The size,
type and distribution of the foliar crystals has taxonomic significance
particularly at the sectional and series. There are especially large crystals
in Minuartia subgenus Minuartia which appear to be of taxonomic
importance.
The micromorphology of the capsular walls of Silene and a few
other genera has been investigated and has taxonomic importance but the
results are not discussed in detail. Presented as appendices, there are keys
for the identification of the seeds of Libyan Caryophyllaceae, for fruiting
material of Libyan Silene as well as for the infrageneric taxa of Arenaria,
Minuartia and Moehringia based on crystals.
CONTENTS
ACKNOW LEDGEMENTS
LIST OF FIGURES
LIST OF PLATES
LIST OF TABLES
CHAPTER
1. GENERAL INTRODUCTION
13
CHAPTER
2. MATERIALS AND METHODS
20
2.1
PLANT MATERIAL
20
2.2
SEEDS
20
2.3
CRYSTALS
21
2.4
CAPSULES
22
CHAPTER 3. SEEDS
3.1
INTRODUCTION
24
3.1.1 HISTORY OF THE USE OF SEED
MORPHOLOGY
24
3.1.2 SCANNING REFLECTION ELECTRON
MICROSCOPY (SEM) AND SEED
MORPHOLOGY
28
3.2 SEED MORPHOLOGY OF THE LIBYAN
3.3
SPECIES
29
3.2.1 LAYOUT
29
3.2.2 GLOSSARY
32
3.2.3
33
DESCRIPTIONS
DISCUSSION
94
CHAPTER 4. CRYSTALS
4.1
INTRODUCTION.
4.2
MORPHOLOGY AND DISTRIBUTION OF FOLIAR
154
4
CRYSTALS IN THE GENERA ARENARIA,
MOHERINGIA AND MINUARTIA
157
4.3
181
DISCUSSION
CHAPTER 5. CAPSULES AND NUTLETS
5.1 INTRODUCTION
198
5.2 SILENE
198
5.3 OTHER GENERA
202
CHAPTER 6. GENERAL DISCUSSION AND
C O N C LU S IO N S
215
6.1 CHAPTER 2 METHODS
215
6.2 CHAPTER 3
215
6.2.1 TESTA MICROMORPHOLOGY: SPECIES AND
INFRASPECIFIC TAXA
215
6.2.2 SEED SHAPE AND TESTA MICROMORPHOLOGY
GENERA/SUBGENERA
217
6.2.3 SEED SHAPE AND TESTA MICROMORPHOLOGY:
TELEPHIUM
218
6.2.4 SEED SHAPE: THE DORSIVENTRAL GENERA 219
6.2.5 SEED CHARACTERS: THE TRIBES
220
6.2.6 SEED SHAPES AND TESTA MICROMORPHOLOGY:
SUBFAMILIES AND ILLECEBRACEAE
225
6.3 CHAPTER 4
232
6.4 CHAPTER 5
234
6.5 SOME GENERAL CONCLUDING POINTS.
235
R EFEREN CES
237
APPENDIX I. LIST OF SPECIES WITH AUTHORITIES
252
APPENDIX II. LIST OF STUDIED SPECIMENS
258
5
APPENDIX III.
AN ARTIFICIAL KEY TO THE LIBYAN
SPECIES BASED ON SEED CHARACTERS
271
APPENDIX IV. AN ARTIFICIAL KEY TO THE LIBYAN
SPECIES OF S IL E N E USING FRUITING
PLANTS
281
APPENDIX V. AN ARTIFICIAL KEY FOR THE
IDENTIFICATION OF THE SUBGENERAf
SECTIONS AND SERIES OF THE GENERA
ARENARIA, MOEHRINGIA AND MINUARTIA
USING CRYSTALS
283
6
LIST OF FIGURES
page
Fig 1. Typical seed measurements.
120
Fig 2.
121
Range of outline shapes in the subfamilies.
Fig 2A. Subfamily Paronychioideae.
„
Fig 2B. Subfamily Alsinoideae.
Fig 2C. Subfamily Caryophylloideae.
Fig 3.
Range of radicle outlines in the subfamilies.
Fig 3A. Subfamily Paronychioideae.
122
„
Fig 3B. Subfamily Alsinoideae.
Fig 3C. Subfamily Caryophllyoideae.
Fig 4. Shapes and distribution of calcium oxalate in mature leaves of
Arenaria, Minuartia and Moehringia.
191
L I S T OF P L A T E S
The plates from no. 1 to 31 are SEMs of the seeds showing details of the
testa ornamentation located near the midzone of the lateral face,
the
marginal face, the radicle and the hilar notch as following.
1. Spergula fallax (1,2,3), Spergularia bocconii (4,5,6), Spergularia
diandra (7,8,9).
123
2. Sperguiaria maritima (1,2,3,), Spergularia rubra (4,5), Spergularia salina
(6,7,8).
124
3. Spergularia salina (1,2), Polycarpaea carnosa (3,4,5,6), Polycarpaea
divaricata (7,8).
125
4. Polycarpaea divarcata (1), Polycarpaea repens ( 2,3,4,5,6), Polycarpaea
robbariea (7,8).
126
7
5. Polycarpaea smithii (1,2,3,4), Polycarpaea tenuis (5,6,7,8).
127
6. Polycarpon prostratum (1,2), Polycarpon tetraphyllum (3,4), Loeflingia
hispanica (5,6,7,8).
128
7. Pteranthus dichotomus (1,2), Sclerocephalus arabicus (3,4,5),
Gymnocarpos decander (6,7,8).
129
8. Paronychia arabica (1,2,3), Paronychia argentea (4,5,6), Paronychia
capitata (7,8).
130
9. Paronychia capitata (1), Paronychia chlorothyrsa (2,3,4,5), Paronychia
kapela (6,7,8).
131
10. Herniaria cinerea (1,2), Herniaria cyrenaica (3,4), Herniaria ericifolia
(5.6.7.8).
132
11. Herniaria fontanesii (1,2,3), Herniaria glabra (4,5,6,7), Herniaria
hemistemon (8).
133
12. Herniaria hemistemon (1,2), Telephium sphaerospermum (3,4,5),
Arenaria serpyllifolia (6,7,8).
134
13. Arenaria serpyllifolia (1,2,3,4), Arenaria leptoclados (5,6), Arenaria
leptoclados Davis 50344 (7,8,9).
135
14. Cerastium dichotomum (1,2), Cerastium glomeratum (3,4), Cerastium
comatum (5,6), Cerastium ligusticum (7,8).
136
15. Cerastium pumilum (1,2), Cerastium semidecandrum (3,4), Cerastium
siculum (5,6),Stellaria media (7,8,9).
137
16. Stellaria media subsp. cupanina (1,2,3,4), Stellaria media
(5.6.7.8).
138
17. Stellaria pallida (1,2,3,4), Minuartia campestris (5,6), Minuartia
geniculata (7,8).
18. Minuartia geniculata (1,2,3,4,5,6,7,8,9).
19. Minuartia geniculata (1,2), Minuartia hybrida (3,4)
139
140
8
Minuartia mediterranea (5,6,7), Minuartia montana. (8,9).
141
20. Sagina apetala (1,2,3,4), Sagina maritima (5,6) , Gypsophila elegans
(7,8).
142
21. Gypsophila pilosa (1,2,3,4,5), Vaccaria pyramidata (6,7,8).
143
22. Vaccaria pyramidata (1,2), Dianthus crinitus (3,4,5,6,7,8).
144
23. Petrorhagia illyrica (1,2), Petrorhagia velutina (3,4,5,6) Silene apetala
morphotype A (7,8).
145
24. Silene apetala morphotype B (1,2,3), Silene apetala morphotype C
(4.5.6), Silene articulata (7,8).
146
25. Silene articulata (1),S/7e/?e behen (2,3), Silene cerastoides (4,5)
Silene colorata subsp. colorata (6,7,8).
147
26. Silene colorata subsp. lasiocalyx (1,2) Silene conoidea (3,4), Silene
cyrenaica (5,6,7), Silene fruticosa.(8).
148
27. Silene fruticosa (1,2), Silene fuscata (3,4), Silene gallica (5,6),Silene
italica (7,8).
149
28. Silene longipetala (1,2,3,4), Silene marmarica (5,6,7), Silene
muscipula (8).
150
29. Silene muscipula (1), Silene nocturna (2,3,4), Silene rubella
(5.6), Silene sedoides (7,8).
151
30. Silene sedoides (1,2,3) Silene succulenta (4,5), Silene tridentata (6,7),
Silene villosa (8).
152
31. Silene villosa (1,2), Silene vivianii (3,4) Silene vulgaris (5,6),
Agrostemma githago (7,8).
153
The plates from no. 32 to 37 are photomicrographs and SEMs of crystals
in tissues of mature leaves.
32. Moehringia stellarioides (1), Arenaria alsinoides (2,3), A. cucabaloides
(4), A. grandiflora (5,6), A. montana (7,8).
192
33. Arenaria kingii (1,2), A. paludicola (3,4,5), A. pungens (6), Minuartia
rimarum (7), M. bulgarica (8).
193
34. Minuartia campestris( 1,2), M. geniculata (3), M. globulosa (4), M.
hybrida (5,6), M. mediterranea (7,8).
194
35. Minuartia mesogitana (1), M. hamata (2), M. montana (3), M. saxifraga
(4), M. stricta (5), M. dichotoma (6), M. douglasii (7), M. pestalozzae
(8).
195
36. Arenaria al$inoides( 1), Moehringia trinervia (2), Arenaria steveniana
(3,4), Minuartia cerastifolia (5), M. bulgarica (6,7,8).
196
37. Minuartia mediterranea (1), M. tenella (2), M. mediterranea (3,4),
M. montana (5,6).
197
The plates from no. 38 to 44 are SEMs of capsule wall (midzone) and LMs
of impressions of capsule wall of Silene.
38. Silene acaulis (1,2), S. fruticosa (3), S. nicaeensis (4), S. succulents
(5), S. villosa (6), S. armeria (7), S. atlantica (8).
204
39. Silene dioica (1,2), S. alba (3), S. dioica (4), S. nutans (5,6),
S.noctiflora (7),S. nutans (8).
205
40. S. conoidea (1,2), S. colorata (3,4), S. apetala (5), S. colorata (6),
S. cyrenaica (7), S. rubella (8).
41. S. longicaulis (1,2,3), S. neglecta (4),
206
S. longipetala (5,6,7), S. calaryi
(8).
42. S.
italica (1,2), S. boryi (3),
207
S. italica (4), S. mollissima (5), S.
protensis (6), S. pomelii (7,8).
208
43. S. tridentata (1,2,3), S. gallica (4),S. cerastoides (5), S.conica
(6,7,8).
44. S. colirosa (1,2,3,4), S. laeta (5,6,7), Lychnis flos-cuculi (8)
209
210
10
The plates no 45 and 46 are LMs of capsule walls showing cell shape and
ornamentation in the midzone areas of Polycarpon, Polycarpaea, Spergula
and Spergularia.
45. Polycarpon arabicum (1), P. bivonae (2), P. depressum (3),
P. eploides (4), P. polycarpoides (5), P. tetraphyllum (6),
Polycarpaea repens (7), P. robbairea (8)
211
46. Spergula arvensis( 1), S. fallax (2), S. morisonii (3),
S. pentandra (4), S. viscosa (5), Spergularia salina (6), S. maritima
(7), S. rubra (8).
212
The plates no. 47 and 48 are SEMs of capsule wall showing surface
ornamentation of Arenaria, Paronychia and Herniaria
47. Arenaria serpyllifolia (1), A. leptoclados (2), Paronychia arabica (3), P.
argentea (4), P. capitata (5), P. chlorothyrsa (6),
P. kapela(7,8). 213
48. Herniaria cinera (1), H. cyrenaica (2), H. glabra (3,4),
H.fontanesii
(5,6), H. hemistemon (7,8).
214
LIST OF TABLES
page
Table 1. Classification of Caryophyllaceae in different Floras.
18
Table 2. Number of genera and species of Caryophyllaceae family in
North Africa.
19
Table 3. Measurements of British in (GL) and Mediterranean specimens of
Arenaria leptoclados.
98
Table 4. Measurements of British in (GL) and North African specimens of
Arenaria serpyllifolia
99
Table 5. Comparison between Polycarpaea, Polycarpon and Robbairea
seeds.
109
11
Table 6. Seed characteristics of the subfamily Alsinoideae (genera found
in Libya).
228
Table 7. Seed characteristics of the subfamily Paronychioideae (genera
found in Libya).
229
Table 8. Seed characteristics of the subfamiy Paronychioideae (genera
found in Libya).
230
Table 9. Seed characteristics of the subfamily Caryophylloideae (genera
found in Libya).
231
12
ACKNOLEDGEMENTS
I am indebted to the Nasser University and Ministry of Higher Education for
financial support.
I wish to thank my superviser, Dr. J. Dickson for his advice, help and
encouragement throughout the period of reserch.
/
I am grateful to the Department of Botany, Glasgow University for allowing
me to carry out this work and to the Regius Keeper, Curator and staff of the
Royal Botanic Gardens, Edinburgh, for allowing me to work in the
herbarium during this study. Dr. R. R. Mill, Prof. J. Morton of the University of
Waterloo Canada and Dr. D. Hind of Kew made helpful comments and
advice. I would like to thank all members of the Botany Department, also P.
Ainsworth in the electron microscopy laboratory and photographer D.
Maclean in the Geology Department who have helped me.
I wish to thank my colleague M. Magrabi in the Tripoli National Herbarium
for helping me to obtain recent collections of plants and seeds from Libya
material.
I would also like thank my parents, my wife and my children for their support
through some very difficult periods.
13
Chapter 1
General In tro d u c tio n
The title of this thesis is taxonomic studies of North African Caryophyllaceae
with particualr reference to the flora of Libya. The thesis has three main
aims, the first being the taxonomic revision of all the Libyan
Caryophyllaceae sensu lato with reference to seed morphology. Scanning
electron microscopy (SEM) in particular has been used to investigate seed
morphology in relation to taxonomic value at various levels from
infraspecific to subfamilial. The second aim was to investigate the
taxonomic significance of calcium oxalate crystals in the genera Arenaria,
Moehringia and Minuartia of subfamily Alsinoideae. The third aim was the
study of capsular micromorphology of the genus Silene, subfamily
Silenoideae particularly species of North African origin; a few other genera
were also examined in this way.
In the family Caryophyllaceae as now recognised, Linnaeus (1753)
recorded 24 genera and about 132 species. Genera Piantarum (1789) of
de Jussieu contained only 31 genera of the family. By 1904 Willis in the
second edition of his well known dictionary listed 60 genera and 1300
species, but in the (1966) seventh edition the figures had become 70 and
1750. Heywood (1978) and Abdul Gafoor (1987) gave totals of 80 genera
and 2000 species, and more recently Mabberley (1987) recorded 89
genera and 2070 species. According to Bittrich (1993) the family consists of
about 86 genera and about 2200 species. More than half of the
Caryophyllaceae belong to only 6 genera, Silene has 400 species,
Dianthus 250, Arenaria 250, Gypsophila 125, Stellaria 100 and Cerastium
100 according to Cronquist (1981). According to Bittrich (1993), however,
the genus Silene has nearly 700 species, Dianthus about 300, Arenaria
150, Gypsophilla 150, Stellaria about 150-200, and Cerastium about 100.
14
The following diagnosis of the family is taken from Bittrich (1993), but
with additions and alterations from various other sources.
Caryophyllaceae A. L. de Jussieu, Gen. Plant. : 299 (1789), nom. cons.
Annual or perennial herbs, rarely shrubs or small trees, usually
hermaphrodite, rarely gynodioecious or dioecious. Stems often swollen at
the nodes, usually with anomalous secondary growth in older stems, often
woody at base, erect to prostrate, branching often dichotomous, rarely
irregular; leaves opposite, occasionally alternate or spirally arranged,
sometimes whorled, entire, often petiolate, occasionally connate at base, or
rarely with completely adnate margins, exstipulate or with usually
membranous stipules. Inflorescence dichasial cymose or monochasial,
flowers terminal or in axils of leaves solitary or in clusters, sessile or on long
peduncles. Flowers actinomorphic, 5- rarely 4- merous hypogynous or
perigynous. Sepals (4-)5 or rarely more or fewer, free or connate into a
tube, or only slightly adnate at base, green or with membranous margins,
glabrous or hairy; petals (4-)5 or rarely more, usually conspicuous,
occasionally small, inconspicuous (subfamily Paronychioideae) stamens
up to 10, rarely fewer by abortion, usually in 2 whorls of 5, the second whorl
often absent or reduced into staminodes, free from one another, inserted
with the petals in the rim of the cup-shaped receptacle or in a perigynous
disc; gynoecium of 3-5 carpeled, rarely more or only 2; ovary superior, 1loculed or 3-5 imperfectly celled at base, 1- numerous ovules in commonly
free central placentation; styles 2-5, free or united, ending with simple
capitate or lobed stigmas. Petals stamens and ovary sometimes borne on
an elongate internode (asithophore). Fruit usually a dehiscent capsule, with
more or less numerous seeds, opening by as many or twice as many valves
or teeth as the number of styles; rarely an indehiscent achene, enclosed in
15
a persistent calyx or a berry or pseudo-berry. Seed small, reniform or
globose to pyriform, rarely peltate, usually with variously sculptured testa,
rarely smooth, the back sometimes caniculate or winged, mostly exarillate
or rarely arillate; embryo peripherally curved around the starchy perisperm,
sometimes ± straight, rarely spiral, endosperm little or absent, x = 5-19,
sometimes polyploid series occur.
This large family of relatively uniform characters is usually conceived
in modern treatments such as Bittrich (1993) as comprising three
subfamilies: Alsinoideae, Paronychioideae and Caryophylloideae. The
Caryophylloideae are sometimes referred to as Silenoideae, as in Flora
Europaea. The subdivision of the family has long been controversial ,
particularly the status of Subfamily Paronychioideae. The lllecebraceae is
classified within the Caryophyllaceae by some e. g. Tutin et al. (1964-93),
Zohary (1966), and Stace (1991), but by others is classified as a separate
family e.g. Davis (1967), Meikle (1977) and Abdul Gafoor (1977). The
anatomical characters of the genera included by Bentham and Hooker in
family lllecebraceae are so similar to those of the Caryophyllaceae that
both groups have been described together as in the system of Engler and
Prantl (Metcalfe & Chalk, 1950). Boulos (1979) listed Paronychiaceae
separately from Caryophyllaceae.
The arrangement of the genera into subfamilies and tribes varies
with the author. See Table 1. In Flora Europaea and Flora Palestlna the
genera are separated into three subfamilies Alsinoideae, Paronychioideae,
Silenoideae (Tutin et al. 1964-1993, Zohary 1966). In Flora of Turkey the
genera are split into five groups and three subfamilies (Davis, 1966). Flora
o f Cyprus
divided the genera into tribes Caryophylleae, Alsineae,
Polycarpeae (Meikle, 1977). The New Flora of the British Isles arranged the
16
genera in to three subfamilies Alsinoideae, Paronychioideae and
Caryophylloideae (Stace, 1991).
The detailed account by Bittrich (1993) is the most up-todate of the family and he adopted this classification.
I. Subfam. Paronychioideae (A. L. Juss.) Meisn. (1838)
1. Tribe Polycarpeae DC. (1828)
2. Tribe Paronychieae (A. L. Juss.) Dumort. (1827)
3. Tribe Corrigioleae Dumort. (1827)
II. Subfam. Alsinoideae (DC.) Fenzl (1840)
1. Tribe Alsineae DC. (1824)
2. Tribe Pycnophylleae Mallfeld (1922)
3. Tribe Geocarpeae Palmer & Steyermark (1950)
4. Tribe Habrosieae (Fenizl) Pax (1927)
5. Tribe Sclerantheae (A. L. Juss.) DC. (1828)
III. Subfam. Caryophylloideae
1. Tribe Caryophylleae
2. Tribe Drypideae Fenzl (1840)
3. Tribe Sileneae DC. (1824)
According
to
Cronquist
(1981)
the
relationship
of
the
Caryophyllaceae to the other families of the order Caryophyllales was
confirmed by embryology, pollen morphology, the frequent occurrence of
anomalous secondary growth and a special type of sieve-tube plastid. The
book Caryophyllales Evolution and Systematics by Behnke and Mabry
(1994) deals with many modern aspects such as DNA, gene sequences,
chemotaxonomy and cladistics. There is, however, little or no treatment of
seeds.
17
The Caryophyllaceae is mainly distributed in the temperate regions
of the northern hemisphere with a centre in the Mediterranean and IranTuranean region ( Bittrich, 1993) (Table 2). There are 10 species belonging
to 7 genera (Arenaria, Cerastium, Melandrium, Minuartia, Sagina, Silene,
Stellaria), growing in Greenland between Victoria Fjord, and Danmark
Fjord, the northernmost land in the world, (Holmen, 1957) and the
species,Colobanthus quitensis, is one of only two species of Angiospermae
growing on the Mainland of Antarctica (Green, 1970).
18
NAME OF WORK
SYSTEM OF CLASSIFICATION
Flora Europaea 1962,93
Subfamilies: Alsinoideae, Paronychioideae,
(following Pax and
Silenoideae/Caryophylloideae
Hoffman)
Flora Palestina 1966
New Flora of the British
Isles 1991
Flora of Turkey 1966
Group I Subfam. Alsinoideae, Group II Sub
fam.Paronychioideae, Group III Subfam.
Paronychioideae, Group IV one genus
Thurya, Group V Subfam. Silenoideae.
Flora of Cyprus 1977
Tribe 1. Caryophylleae, Tribe 2 Alsineae,
Tribe 3 Polycarpeae.
Table 1. Classification of Caryophyllaceae in different Floras.
COUNTRY
NO OF GENERA
NO. OFSPP.
Britain
19
91
Egypt
20
57
Libya
22
80
Tunisia
25
115
Algeria
27
175
Morocco
26
229
Table 2. Approximate numbers of genera and species of the
Caryophyllaceae in North Africa, and in Britain.
20
Chapter 2.
MATERIALS AND METHODS
2.1
The
Plant
M aterial
plant
m aterial
used
in
this
study
was
entirely
dried
specimens obtained from many herbaria (BM, E, GL, K, L, M, ULT,
T).
Full details are given in Appendix 1, which is a list of all the
taxa studied during this research.
2.2 Seeds
In obtaining material from herbarium specimens great care was
taken to select only fully ripe, undamaged seeds from mature
capsules. In the early stages of the research the seeds were
exam ined
using
a
Philips
500
electron
m icroscope
in
the
Biological Science Electron Microscopy Laboratory. Seeds were
mounted
on stubs with double sided sticky tape
(Sellotape),
surrounded by conductive silver paint, and vapor coated with gold
(200-400 A thickness) while being rotated at an angle of 45.
Later it was found that the use of the Leo Microscopy Stereoscan
360 and the Emscope sputter coater with a gold target, in the
Geology Department and a different technique in mounting the
seeds produced results of better quality. With few exceptions,
the
photographs
presented
here
were
microscope and the following technique.
obtained
with
that
21
The heated stubs were evenly rubbed with wax sticks to leave a
smooth, thin layer which hardened after a few second's. Using a
dissecting microscope, the seeds were carefully placed on the
wax layer. Material was then kept in a dry and dust-free place to
avoid dust contamination and to stop hydration. Measurements of
the seed size were made using a micrometer eyepiece.
2.3
C rystals
The m orphology and distribution of calcium oxalate crystals in
the leaves of the genera A re n a ria , M oehringia and Minuartia w as
studied. The technique devised was modified from the method
used by Bokhari (1970) in order to reduce the time needed for
clearing. Two or many mature leaves from
herbarium
sheets
were placed in a small tube, a few drops of 10% KOH were added,
and boiled in a water bath for five minutes. After washing with
distilled water, the material was spread on a slide, blotted dry
with a clean tissue and mounted in New Aquamount.
By far the most common component of crystals in plants is
calcium oxalate; see Section 4.1 No elaborate procedure was
undertaken
to
prove that the
crystals
studied
were
calcium
oxalate. However, following Dormer (1961), it was shown that
the crystals were soluble in HCI but not in acetic acid.
2 2
SEM
species
difference
of
photographs were taken of crystals from
Arenaria,
in
M oehringia and M inuartia
to
different
show
the
shape and fine structure between the druses and
the sandy crystals, particularly the elongate ones in M in u a rtia .
Mature leaves were kept over night in 10% KOH containing a few
drops of H 2 O 2 . They were then boiled in distilled water until the
tissues macerated. Crystals and macerated cells were selected
by using a needle and forceps under the light microscope and
placed on the stubs, following the same technique used for the
seeds.
2.4
Capsules
The methods used were are those of Payne (1970) and a new
technique
detailed
below.
A complete
capsule
of
S i i e n e (or
fragment from the middle) was laid on a watch glass, flooded
with acetone, and a small square of cellulose acetate film placed
over the capsule before flooding again with acetone. The acetone
dissolves the acetate, which settles against the surface of the
specimen and in a few moments the acetone evaporates and the
rehardened acetate film is peeled away (Payne 1970). It was
found that the peels obtained by this technique were not very
satisfactory because a dense cover of air bubbles often obscured
the preparation. In trying to make better preparations, it was
23
decided to use superglue. In testing a few brands, Loctite was
found to be the most satisfactory. Impressions of fully ripe
capsules were made as follows. One or two drops of superglue
were put on a slide and the middle of the capsule rather than the
upper part was pressed into the superglue and left for one to two
minutes to harden. Then the capsule was removed. No coverslip
was placed over the sample. Photographs can be taken by using a
light microscope. Only one capsule was, in general, examined of
each
specim en.
Capsule
epiderm al
cells
of
Polycarpon,
Polycarpaea, Spergula and S p e rg u la ria were studied by clearing
the whole capsule using the same method used for studying the
crystals. A few SEMs were taken of the S ilene capsules and the
nutlets of Herniaria and Paronychia.
24
Chapter 3.
Seeds
3.1 Introduction
3.1.1 History of the Use of Seed Morphology
The taxonomic value of seed morphology had been realised in the
eighteenth century by Linnaeus in his book Genera Plantarum (1737-1767)
The examples quoted below are taken from genera No 567, 569 and 586
567- Silene — plurima, reniformia . 569 - Arenaria
— Plurima, reniformia
586 - Spergula— plurima, depresso-globosa, margine emarginato cineta
However in his book Species Plantarum he makes no mention of seeds .
A.De. Jussieu (1791) in his book Genera Plantarum, used different sexual
organs in his descriptions and ignored seed characters. De Candolle in
1828 published the first of the many volumes of Prodromus systematis
naturalis regni vegetabliis. For the Caryohyllaceae he used various
taxonomic characters but made almost no use of
seed morphology.
Endlicher however in his Genera Plantarum (1836-1840) treated 6235
species of vascular plants. He used detailed seed characters as part of the
diagnoses of genera. In the Caryophyllaceae, for instance, he described
seed shape and colour as well as ornamentation of the testa. The examples
quoted below are taken from page 959 and onwards.
TRIBUS II. PTERANTHEAE R. Brown in Wallich Plant. As. rar. I. 17
Seminum chalaza lateralis, ab umbilico distincta. Embryo planus, rectus,
albuminis farinacei lateri adplicitus .
Pteranthus Forsk.
Semen erectum, oblongum, compressum, testa tenui, laevissima, chalaza
infra medium laterali. Embryo semini conformis, rectus, albuminis lateri hinc
applicitus, radicula tereti, prominula infera .
25
TRIBUS IV. Telephieae DC. Prodr. Ill . 366 .
Telephium Tournef.
Semina plurima, columellae centrali liberae affixa, globuloso-reniformia.
Embryo fere annularis, albumen farinaceum cingens.
TRIBUS V. Polycarpeae DC. Prodr. III. 373.
S p e rg u la ria Pers.
Semina plurima, pyrifromia, lenticulari-compressa, saepissime margine
scarioso cincta, laevia, granulata v. muricata. Embryo uncinatus v.
annularis, albumen farinaceum cingens. Cotyledonibus incumbentibus.
Within the Tribe Sileneae De Candolle described the seeds of
Dianthus as follows (p. 355). “Semina compressa hinc convexa inde
concava, peltata”. However, Endlicher used seed shape as parts of tribal
diagnoses. Wihin Subordo IV Sileneae DC., his Tribus I Diantheae Kunth
Flor. berol. 1.106 has the following.
“ Semina oblonga v. ovalia, compressa, dorso convexa, facie plana v.
convexiuscula, medio longitudinaliter carinata. v. marginibus involutis
canaliculata, placentae centrali columellari peltatim affixa.”
And his Tribus II Lychnideae Fenzl msc.
“Semina globosa, reniformia v. lenticularia, umbilico marginali funiculus
distinctis affixa.”
Bentham and Hooker (1862-67) in their Genera Plantarum used seed
morphological characters for most genera under ordo Caryophyllaceae but
do not give detailed descriptions. Within their Tribus I. Sileneae they
separated the genera Velezia, Dianthus and Tunica because of “ Semina
peltata, hilo faciali”. In his Flora of Syria, Palestine, and Sinai, Post (1883)
had two tribes within the order Sileneae as follows.
“Tribe I. Diantheae. Seeds shield-shaped, with hilum on face. Styles 2.
Embryo straight.”
26
“Tribe II. Lychnideae. Seeds kidney-shaped or nearly globular, with lateral
hilum. Embryo peripheral or spiral.”
In his monograph of the genus Silene published in 1868, Rohrbach
used seed characters in a key to genera. For instance he separated
Petrocoptis from Lychnis by “semina barbata” and “semina ebarbata”. He
included two plates of drawings showing seed shape, ornamentation and
sections and he gave detailed seed descriptions for each species of Silene.
For S. cucubalus (p.85, = S. vulgaris) he stated “semina magnetudine
variantia, rotundata-reniformia, dorso plano-convexiuscula, faciebus
levisime concava vel plano-concaviuscula rarove plana, seriatim
tuberculata.” For S. maritima With, (p.84, = S. uniflora) "semine dorso leviter
canaliculata, faciebus plana tuberculata "These are two highly polymorphic
species which since Rohrbach's day have received much study. In their
well-known book The Bladder Campions, Marsden-Jones and Turrill (1957)
illustrated a series of seeds of S.uniflora ssp. uniflora that varied from
markedly papillate to non-papillate and used the terms armadillo and
tubercled. These terms have been adopted by Aeschimann in his
numerous studies of S. vulgaris as for instance in his 1985 paper.
In 1882 Luerssen in his book Handbuch der systematischen Botanik gave
very brief information about the seeds of some genera of Caryophyllaceae.
Because of these taxonomic uses of seed morphology, it is somewhat
surprising that Pax (1894), and Pax and Hoffman (1934) in the highly
significant work Die naturlichen Planzenfamilien make little mention of seed
characters within the Caryophyllaceae.
Various seed atlases and books of drawings of plants have depicted
seeds of Caryophyllaceae; only four examples are mentioned here.
Bertsch’s Fruchte und Samen (1941) has four plates of seeds and
Beijerinck’s Zadenatlas Der Nederlandsche Flora (1947) has six. The well
27
known Drawings of British Plants by Ross-Graig include drawings of seeds
and fruits; part 5 devoted to Caryophyllaceae has seeds drawn with the
attention to detail typical of the artist. Much more important to this thesis is
the work of Berggren (1981) whose work covers seeds and small fruits of
northwest European plants. Part 3 deals with the Caryophyllaceae.
In the first and second editions of Flora Europaea, seed shape,
colour, size and ornamentation have been used with the other
morphological characters to describe most genera, sections and species
eg. Arenaria, Petrorhagia and Silene. On the other hand for some genera
and species, the seeds have been ignored or largely so eg. in the genera of
subfamily Paronychioideae.
Variation in seed coat morphology between different populations of
the same species has been noted in Spergula arvensis (New 1958) and he
showed that the smooth and papillate seed coat forms of Spergula arvensis
responded differently to temperature and humidity. According to Ball and
Heywood (1964) , the only reliable morphological character which can be
used to distinguish between Petrorhagia velutina, P. prolifera and P.
nanteuilii is the structure of the seed coat or testa. Seed morphology and
anatomy of 42 species and 3 varieties of the genus Dianthus have been
described by Kowal & Wojterska (1966). Wojterska (1969) gives results for
the genus Cerastium. Because seed characters are only very slightly
influenced by environmental factors, these features are very important
criteria for the classification of species and genera. The genera and species
of Dianthus , Petrorhagia, Silene, and Spergula show this very well. In
many cases seed morphologly varies at infraspecific level e. g. Spergula
arvensis and Spergularia media with or without wings (Stace 1989).
In his recent survey of the family Bittrich, (1993, p. 214) stated “Seed
morphology, especially the form of testa cells was variously found to be of
28
great diagnostic value mainly for separating taxa at the species level”. He
gave no importance to seed characters at the subfamily and tribal levels but
often listed seed size, colour and ornamentation as significant at the
generic level.
3.1.2 Scanning Reflection Electron Microscopy (SEM) and seed
morphology
Since the late 1960s the scanning reflection electron microscope has been
used
to provide much taxonomic information about
plant surfaces,
particularly those of pollen and seeds. As early as 1969 Heywood (p.7)
stated “ The Caryophyllaceae in fact shows a very wide range of external
features of the seed, including elaiosomes, and these are often related to
reproductive capacity, germination differences and other biological
problems as many studies have already shown. A detailed survey of the
seeds in this family is needed and scanning electron microscopy now
available as a rapid technique it is likely that such a survey will be
underaken”.
The taxonomic significance of seed morphology of 15 species of
Sagina from North America, Europe and eastern Asia was studied utilizing
SEM (Crow 1979). Seeds of 13 southeastern United States taxa of
Arenaria were examined with SEM. With a few exceptions, the external
morphology was distinctive and valuable for separation at the species level
(Wofford 1981). The seed-coat of twenty nine taxa belonging to the section
Minuartia of the genus Minuartia, has been investigated by SEM. Almost
all species, and sometimes several varieties of the same species could be
identified by the details of their fine structure (Celebiogiu etal. 1983). With
SEM the seed-coat was described in detail as one of the major
morphological characters in Minuartia glaucina, Minuartia smejkalii and
29
M. orthophylla by Dvorakova (1985,1988,1991). A seed-coat study by SEM
was made concerning Arenaria tetraquetra populations in southern Spain
by Fararger et al. (1988). The papillae of the seed-coat cells possess a
very distinct morphology, which enables the four subspecies of Moehringia
intricata to be clearly distinguished (Guardia etal. 1991).
Melzheimer (1980) revised some Balkan species of Silene sec. inflatae
using several taxonomic features including SEM micrographs of the seed
coat as a good diagnostic character. In Flora Iranica Rechinger etal. (1988)
dealt with about 140 species of the large genus Silene . For many of these
species they provided high quality SEM micrographs of the mid-zone testa
cells; this revealed striking differences between the species. Since this
important investigation there have been at least two more SEM studies of
Silene. Testa cells were used with other characters to diagnose a new
subsp.,S//ene bupleuroides L. ssp. ganiatsasiana, in the Greek flora
(Voliotis 1991). The seed colour and testa ornamentation were used to
separate Silene haussknechtii from S. laconica and the petal, capsule and
the ridge on the seed back were used to distinguish
Silene. aegaca from S. pentelica ( Oxelman 1995).
3.2 Seed Morphology of the Libyan Species.
3.2.1
Layout
Subfamilies, tribes and genera are laid out in the order of
B ittrich
(1993).
W ithin
each
genus
the
species
are
in
alphabetical order. Nomenclature follows Greuter et al. (1984).
The
description for each
species
begins with
a direct
quotation from the Flora of Libya (FI. Lib.). Abdul Ghafoor's
30
statem ents
on
seed
necessarily
based
shape,
solely
size
on
and
light
colour are terse
microscopy.
He
and
provided
drawings of the seeds of most species at x10, 15, 25, 30 or even
50.
These
draw ings
by
Mohamed
Rafiq
satisfactory but SEM inevitably reveals
are
m ostly
very
important details very
difficult or impossible to observe by light microscopy.
Abdul Ghafoor considered 59 species of Caryophyllaceae
and 14 species of lllecebraceae in FL. Lib. He gave descriptions
of the seeds of 56 and 13 of these species respectively and he
illustrated
38
species
in Caryophyllaceae
lllecebraceae. The species
described
S ilene
the
are
D ianthus
listed
3 species
by Abdul Ghafoor and
se rru la tu s , P etrorhagia
in
not
illy ric a
and
biappendiculata. The following descriptions total 57 of
Caryophyllaceae and all
relevant,
and
descriptions
have
14 of the
been
lllecebraceae.
given
of
the
W here
seeds
of
subspecies.
The following species are listed from Libya by Greuter e t
al. (1984) but have not been studied for this thesis:
s e rra tifo liu s
Sm.,
P e tro rh a g ia
ru p e s tris
S p e rg u la ria
m unbyana Pomel,
A d d itio n a lly,
within
d ifficu lt
M in u a rtia
s a n d w ith ii Maire
Brullo & Furnari, Silene
the
Dianthus
& Sim pson,
uniflora
T ele ph iu m b a rb e a n u m
taxo no m ically
and
Roth,
Born.?
no m e n cla tu ra lly
genus Cerastium under " semidecandrum aggr." Greuter
e ta l. (1984,p. 183) list C, balearicum F. Hermann and C. diffusum
31
Pers. Neither has been studied. Monnier (1975) provided fine line
drawings of the seeds of S p e rg u la ria
m unbyana. M in u a rtia
sa n d w ith ii from one locality in cyrenaica has been discussed by
McNeill (1963) who thought that it may be conspecific with M .
meryeri (Boiss.) Bornm.
The new descriptions in this thesis are based primarily on
Libyan material but often supplemented by material from other
Mediterranean countries, mostly North African, but also where
appropriate
from
countries
such
as
Britain
and the
Arabian
states.
With regard to size, the seeds have been placed into three
cateogries of length with few exceptions: < 1mm, 1-2 mm and > 2
mm. The length measurement was taken as shown in (Figure. 1).
The few exceptions are as follows.
In the case of dorsiventrally flattened seeds, the length
measuremnts was taken from the tip of radicle to the opposite
end of the seed. In the case of more or less globular seeds the
diameter was measured. Each species description ends with a
statem ent
on
geographical
distribution
Floras and Greuter et al. (1984).
derived
from
various
32
3.2.2
Glossary
Capitate papillae or tubercles : Long papillae or tubercles ending
with structure like a head.
Callus : A brittle, membranous ring around the hilum (cf. Berggren)
Caruncule : An outgrowth near the micropyle and the seed (Usher, 1966).
Strophiole = Caruncle.
Cogwheel : Testa cells with an outline resembling a cogwheel.
Collar cells : Cells often distinctive, around the hilum and the radicle
(Chuang & Ornduff,1992).
Discoid : Referring to a protuberance ending with a more or less flat,
circular head.
Granule : Minute or very minute protuberance on testa cell; granules may
densely cover the cells, (adj. granular).
Hilar notch : Indentation in which the hilum is situated.
Keeled : With a ridge along the periphery.
Midzone : Middle area of the lateral face of a seed (Figure. 1).
Pad : Padlike structure on each side of the hilar notch in Silene (cf.
Berggren).
Papilla : Single pimplelike protuberance, only one from the middle of a
lateral face of a testa cell, and large in comparison with overall size
of the cell (adj. papillate).
Prickle : Small, sharp protuberance from testa cell; cf. Spergularia.
Revolute : With the margin rolled so that the upper side is expand and
concealed the lower side .
Spurs : Arms of a testa cell, short to long, unbranched or branching a few
times, with sharp or blunt ends.
Stelliform : Testa cell with radiating branches so as to appear starlike.
33
Tubercle : Single large, conspicuous, pimplelike protuberance from a
testa cell
of the marginal face (adj. tuberculate).
W art: Small, pimplelike protuberance from testa cell; usually there are
more than one or several or more per cell and small in comparison
with overall size of the cell.
3.2.3
D escriptions
I. Subfamily Paronychioideae (A. L. Juss. ) Meisn. (1838)
1. Tribe Polycarpeae DC. (1828)
S p e rg u la L.
Spergula L.f Sp. Pl.:440 (1753).
S Ja lla x (Lowe) E. H. L. Krause
(Plate 1 )
F I.L ib . Seeds winged, lenticular, c. 1 mm in diam. (Excld.wing),
black, wing nearly as broad as seed, tubercular or smooth.
Shape, size and colour: Circular, 1-2 mm, dark brown.
T esta
surface:
Lateral face convex, cells stelliform , fla t or
slightly convex; spurs regular, pinnate, each spur ending in one
sm all
but
stelliform ,
d istin ct
raised
po re-like
into
conical
d e p re ssio n ;
papillae;
m arg in al
surface
very
cells,
fine
granular; wing membranous, pale, up to half or more the width of
the seed, with a deep V-shaped notch opposite the hilum, edge
|
m ostly entire; small prickles merging
into papillae where the
f
wing joins the seed; surface mostly smooth or slightly wrinkled .
M easu rem en ts: Mid zone cells 45-50 pm long, 20-25 pm wide;
(
34
spurs 12-16 pm long.
S pecim ens exam ined: Appendix 2 No 628, 630.
D is tr ib u tio n : N. Africa, Saharo Arabian and Sudanian territories
S p e rg u la ria (Pers.) J. Presl and C. Presl, nom. cons.
Spergularia (Pers.)J.Presl and C. Presl, FI. Cech.:94 (1819).
S .bocconei (Scheele) Graebner in Ascherson & Graebner
(Plate 1)
FI.Lib.S eeds triangular-ovate, c.0.5 mm long, all wingless,
greyish-brown, finely tubercled.
Shape, size and colour: obovate-obtriangular, < 1 mm, brown.
T esta
s u rfa c e :
Lateral faces slightly convex, cells irregularly
and deeply lobed, papillae sparse, scattered irregularly, spurs
irregular with very faint margins; marginal face surrounded with
a distinct
raised
rim
leaving
a groove
between
lateral
and
marginal faces, cells faint, covered with conical papillae, whole
papillae crowned with a flat-topped head-like structure; radicle
compressed laterally, slightly tapering; surface granular.
M e a su re m e n ts:
Lateral face cells 40-60 pm long,
10-20 pm
wide; marginal face papillae c. 6 pm long, radicle 30 pm wide
near apex.
S pecim ens exam ined: Appendix 2 No 638, 639.
D is tr ib u tio n : S.W. Europe, North Africa, S. Britain, eastwards to
Iran, Atlantic Islands .
35
S p e rg u la ria d ian dra
(Guss.) Boiss.
(Plate 1 )
F I.L ib . Seeds triangular-obovate, c. 0.5 mm long, all wingless,
dark
brown
to
black,
beset
with
rigid
bristly
papillae
or
rugulose.
Shape, size and colour : Obovate, < 1mm, dark brown .
Testa
surface:
Lateral faces
have the same shape
slightly
biconvex,
cells
(i.e. Jigsaw like), protrusive,
mostly
irregular,
deeply lobed, spurs broad, few in number, with blunt apices,
papillae very small and indistinct, marginal surface broad with a
distinct
raised
rim,
shallow ly
grooved
between
marginal faces, tip of radicle slightly curved;
lateral
and
Surface, finely
granular.
M easurem ent:
Testa surface cells
18-48 pm long, c.10 pm
wide; spurs 13, c.7 pm long; radicle extended, c. 20 pm long.
Specimens examined: Appendix 2 No 643, 644.
D is tr ib u tio n :
M e d iterra ne an
region
and
eastw ard
to
(P la te
2)
Afghanistan, Iran, Pakistan and India .
S. m aritim a (All.) Chiov.
in FI.Lib. [S .m e dia
(L.) C. Presl]
FI.Lib.Seeds brown, compressed, rounded, smooth or tuberculed,
c. 0.8-1 mm in diameter, mostly all winged, wavy membranous,
entire or laciniate.
Shape, size and colour: Obovate, < 1 mm,
red-brown.
36
Testa surface: Lateral faces slightly biconvex, cells irregular,
jigsaw like, raised into blunt papillae; spurs lobed with blunt
apices; marginal face with slightly raised rim, cells raised into
distinct conical papillae, with a very shallow groove between
the
lateral and
marginal face;
wing
margin
slightly sinuate,
making a V shape towards the hilum, covered with small prickles
along the marginal face; surface minutely granular.
Measurements:
wing c. 312 pm wide; radicle extended up to c.
63 pm; papillae 6-14 pm long; spurs 6-9 pm long.
Specimens examined: Appendix 2 No 648, 649.
Distribution: Europe, Russia, Mediterranean region, China and S.
W.Asia.
S.rubra (L.) J. Presl & C. Presl
(Plate
2)
F I.Lib.Seeds obovate triangular or ± trigonous, c. 0.5mm long, all
wingless, dark brown or black, finely tuberculate.
Shape, size and colour: Obovate, <1 mm, red brown.
Testa
surface:
Lateral faces biconvex, cells mostly elongate,
distinct or faint, some cells raised into globular papillae; spurs
deeply lobed irregular; radicle mostly straight, tip broad, thick
and round; marginal face with thick rim, most cells raised into
distinct
conical
papillae
with
discoid
heads,
with
a
narrow
groove between lateral face and marginal face; surface covered
with
dense granules.
37
M easurem ents:
Lateral face cells c. 65 pm long, c. 10 pm wide
; spurs number 9-11, c. 10 pm long; papillae on lateral surface
10-14 pm long, papillae on marginal surface
10-20 pm long;
radicle up to c. 50 pm long, c. 70 pm wide near apex.
Specimen examined: Appendix 2 No 653, 654.
Distribution: Widespread in N. hemisphere, Mediterranean and
Euro-siberian, in some parts of Asia .
S.salina J. Presl & C. Presl
(Plate
2)
[in FI.Lib.S. marina (L.) Gariseb.]
FI.Lib.
Seeds light brown, compressed,
rounded,
smooth
or
tubercled, c.0.6-0.8 mm in diameter, wingless or winged, both in
the same fruit, wings scarious, erose to laciniate .
Shape, size and colour: Circular-obovate, < 1mm, red brown.
Testa surface: Lateral faces slightly biconvex, cells elongate
or ovate, with sparse spherical or elongate papillae; spurs lobed,
irregular and faint ; marginal face rounded and thick, papillae
club
shape
between
dense
dispersed
lateral face
m inute
on
and
granules
or
m arginal
side,
shallow ly
grooved
marginal face, wings covered with
prikcles,
margin irregular,
deeply
sinuate; surface minutely granular.
Note : Winged and
Measurement:
mm .
unwinged seeds have the same features.
papillae 6-30 pm long ; radicle extended up 1
38
Specim ens exam ined: Appendix 2 No 655, 656.
D is trib u tio n :
W idespread
in tem perate,
costal areas of the
northern hemisphere in Europe, North Africa, India, Pakistan,
China .
P olycarpaea Lam.
Polycarpaea Lam., J. Hist. Nat.2:3, t.25 ("Polycarpea”) (1792)
P. repens (Forsk.) Asch. & Schweinf.
F I.L ib .
Seeds
obovoid-oblong,
(Plate 4)
less than
1mm
long,
slightly
compressed.
Shape, size and co lo u r: Oblong-curved (crescent-shaped)
tapering to radicle, <1 mm, creamy.
Testa s u rfa c e :
with
Lateral faces slightly biconvex, cells indistinct
no distinct spurs;
d istin ct,
shallow
dorsal
furrow ,
surface
cells
strongly
in d istin ct;
convex,
ve n tra l
with
surface
concave; radicle terminal, slightly curved, surrounded by distinct
cells; hilum subterminal; surface mostly smooth .
M e a s u re m e n ts : Hilar cells c.13 pm long; radicle c. 78 pm long.
S pecim ens exam ined: Appendix 2 No 434, 435.
D is trib u tio n :
Saharo Arabian region, North Africa
P. robbairea (O. Kuntze) Greuter & Burdet
[In FI. Lib. R obbairea delileana M ilne-R edhead]
FI. Lib. Seeds minute, smooth, shiny.
(P la te
4)
39
Shape, size and colour: Broadly elongate tapering to radicle,
<1 mm, colour creamy.
Testa surface: Lateral faces biconvex, cells indistinct, dorsal
face convex with distinct furrow; ventral face concave; radicle
terminal, slightly curved, surrounded by a collar of cells; hilum
subterminal; short, smooth line connecting the radicle and the
hilum; surface generally glossy, smooth .
Measurements:
Radicle cells
8-10 jam long,
6-8 ^m wide;
short smooth line c. 50 ^im long.
Specimens examined: Appendix 2 No 436, 437, 438.
D istribution:
N. Africa, Egypt, Palestine, Iraq and Saudi Arabia
and Sudan.
P o lyca rp o n L.
Polycarpon L. syst. Nat., ed. 10:881, 1360 (1759)
P. p ro s tra tu m ( Forsk. ) Aschers & Schweinf.
(Plate
6)
F I.L ib . Seeds ± triangular - reniform , light brown , nearly 0.5
mm long.
Shape, size and colour:
Cuneate, tapering to the radicle, <1
mm, creamy.
Testa
papillae
surface:
Lateral faces biconvex, cells raised into big
(dome-shaped),
spurs
indistinct;
dorsal face
strongly
convex, cells similar to those of the lateral face; ventral face
concave, covered with very dense papillaeas compared with the
40
lateral
and
surrounded
dorsal
faces;
radical
term inal,
sligh tly
curved,
by elongate cells, dark brown; hilum subterm inal;
surface finely granular.
M e a s u r e m e n ts :
Lateral face cells 15*30 pm diam , radicle
cells 16-27 pm long, c. 8 pm wide .
Specimens examined: Appendix 2 No 455, 457.
Distribution:
P.
Tropical and subtropical countries .
te tra p h y llu m ( L . ) L.
FI.Lib.
(Plate
6)
Seeds ± triangular-ovoid, brown, usually c.0.5 mm long,
punctulate.
Shape, size and colour: Cuneate, tapering to the radicle, <1
mm, creamy.
Testa
su rface:
globular,
domed
structures
at the
Lateral
papillae
base,
cells ovate or elongate
cells
faces
biconvex,
surrounded
spurs
indistinct;
by
cells
raised
irre g u la r,
into
th ick
dorsal face convex,
arranged in rows; ventral face concave,
irregular ; radicle terminal, slightly curved, surrounded by
mostly elongate cells; hilum subterminal; surface smooth .
Measurements:
Lateral face papillae 10-20 u diam., dorsal face
cells 30-54 pm long, c. 18 pm wide, radicle cells 12-36 pm
long, c. 12 pm wide.
Specimens examined: Appendix 2 No 463, 465, 466.
Distribution: central Europe, Euro-Siberian region, Arabia and
41
Sudan; now cosmopolitan as a weed
L o e flin g ia
L.
Loeflingia L.f Sp. PI.: 35 (1753).
L o e flin g ia
h is p a n ic a
L.
(Plate
6)
FI. Lib. Seeds ± compressed, obliquely obovate, c. 0.5 mm long,
finely papillose, white.
Shape,
size
and
colour:
Broadly elongate, tapering to the
radicle, <1 mm, white-gray .
Testa
surface: Lateral faces biconvex, cells indistinct or faint,
spurs
faint,
compressed
blunt
with
laterally,
thick
covered
walls;
with
dorsal
surface
keeled,
distinct
papillae;
ventral
surface slightly curved, papillae concentrated at the top and the
base;
radicle
term inal,
sligh tly
curved,
m icropyle
discoid,
ornamentation indistinct surrounded by collar of papillae; hilum
raised into an elongate ridge; surface smooth.
M easurements:
wide;
spurs
Lateral surface cells 39-65 pm long, 10-26 pm
5-10,
3-13
pm
long;
radicle
c.
83
pm
long,
micropyle c. 42 pm diam., papillae c. 10 pm long; dorsal face
keel 17 pm high.
Specimen examined: Appendix 2 No 248.
D is trib u tio n : Mediterranean area, South Iran.
42
2. Tribe Paronychieae (A. L. Juss.) Dumort. (1827)
P te ra n th u s
Forssk.
Pteranthus Forssk., FI. Aegypt.-Arab.: 36 (1775).
P. d ic h o to m u s
Forsk.
(Plate
7)
FI.Lib. Seed erect, compressed, endospermous.
Seed
shape,
size
and
colour:
N a rro w -o b o v a te ,
s lig h tly
dorsiventrally flattened, >2 mm, faint yellow.
Testa
surface:
Dorsal face convex,
cells
indistinct,
surface
wrinkled; dark brown circle near the middle of the dorsal face;
no spurs; ventral face
sim ilar to the
dorsal face; thick rim
between the dorsal and ventral faces; radicle flattened, broad
and round at apex, cell ornamentation is more distinct than on
the other parts of the seed testa; hilum in the ventral surface
below radicle; surface smooth not granular.
M easurem ents: Radicle c. 0.5 mm long.
Specimens examined: Appendix 2 No 468, 469.
D is trib u tio n :
Mediterranean,
Saharo-Arabian
and
Sudanian
te r r ito r ie s .
S c le ro c e p h a lu s Boiss.
Sclerocephalus Boiss., Diagn. PI. Orient. I, 1(3): 12 (1843);
Chaudhri, Meded. Bot. Mus. Herb. Rijksuniv. Utrecht 285:1-64 (1968), rev.
43
S .a ra b ic u s
Boiss.
(Plate
7)
F I.L ib .S e e d s obovate-reniform 2-5-3 X 1.5-2 mm, compressed,
glabrous.
Seed shape, size and co lo u r: Obovate, >2 mm, light-brown.
T esta
s u rfa c e : Lateral face flat, compressed laterally, slightly
slanted towards the hilum, cells indistinct, raised into wrinkled
ridges; no spurs; marginal face flat round with a thick rim and
with a distinct groove separating the lateral and marginal faces;
radicle
dark
brown,
extended
straight to the front,
tapering
gradually from the base to the apex, cells mixed, mostly elongate
or polygonal;
between
hilum subterminal, with deep groove separating
the radicle and the hilum;
Seed m easurem ents:
surface wrinkled, smooth.
Radicle c. 0.75 mm long, elongate, cells
24-96 pm long, c. 12 pm wide.
Specim en exam ined: Appendix 2 No 480.
D is tr ib u tio n :
N. Africa, lower Jordan Valley, Dead Sea, Arabia
and Iran .
G y m n o c a rp o s Forskal
Gymnocarpos Forskal, FI. Aegypt.: 65(1775).
G .decander
Forsskal
(P la te
FI.Lib.Seed oblong-reniform, compressed.
Seed shape, size and colou r: Obovate, 1-2 mm, brown.
Testa
s u rfa c e : Lateral face flat, compressed laterally, cells
7)
44
faint, elongate; no spurs; marginal face flat, cells similar to the
lateral face, slightly concave, groove separating the lateral and
marginal faces; radicle tapering gradually from the base to the
top, surrounded by elongate or polygonal cells, ending in a flat
apex turned over the hilum; a deep groove separates the radicle
and the hilum; surface smooth.
M easurem ents:
Radicle c. 403 pm long, elongate cells 24-56
pm long.
Specimens examined: Appendix 2 No 218, 220.
Distribution:
N. Africa, Iran, Syria, Jordan, Palestine, Arabia,
Afghanistan, W. Pakistan, N. W. China.
P a ro n y c h ia Miller
Paronychia Miller, Gard. Diet. abr. ed. 4:3 (1754); Chaudhri
P .arabica
(L.) DC.
(Plate
8)
FI.Lib. Seeds lenticular, ± compressed, c. 0.75-1 mm, glabrous.
Seed shape, size and colour:
Circular, <1 mm, light-brown,
glossy.
T esta
s u rfa ce :
marginal
rim thick,
Lateral
laterally
faces
biconvex,
compressed;
cells
radicle
in d istin ct;
dark-brown,
thick, slightly curved, with circular rim around the apex filled
with
shrunken,
surrounded
by
faint,
polygonal
cells;
hilum
subterminal, depression surrounded by a collar of rectangular,
square or polygonal cells between the hilum and the radicle;
45
surface smooth, glossy.
Measurements:
Cells around the hilum
rectanglar
12-18
jam
long, 6-14 pm wide; square c. 11 x 11 pm; radicle apex c. 54 pm
diam.
Specimens examined: Appendix 2 No 404, 406.
Distribution:
N. Africa, Syria, Lebanon, Palestine, Turkey and
Saudi Arabia.
P .a rg e n te a
FI.Lib.
Lam.
Seed
(Plate
ovoid-round,
c.
1 mm
in
diam.,
8)
brownish-red,
glabrous.
Seed
shape,
size
and
colour:
C ircular-obovate,
1-2
mm,
brown.
Testa
surface:
Lateral
faces
biconvex,
cells
indistinct
or
faint, polygonal; marginal face round with thick rim, cells faint,
with a shallow groove separating the lateral and marginal faces;
radicle curved, tapering gradually from the base to the top, its
apex with a circular rim filled with shrunken cells surrounded by
distinct, elongate cells; hilum subterminal, raised, surrounded by
a collar of protrusive, ovate or elongate cells; surface smooth.
Measurements:
Radicle c.1.58 mm long, radicle apex c. 315 pm
diam .,radicle cells 315-756 pm long, 216-315 pm wide; hilum
378 pm diam, hilum cells 63-189 pm long, c. 63 pm wide.
Specimens examined: Appendix 2 No 410, 412, 413.
46
D is trib u tio n :
N.
Africa,
Europe,
Jordan,
Syria,
Palestine,
Sahara-Arabian and Sudanian territories.
P .c a p ita ta (L.) Lam.
(Plate
8,9)
FI.Lib. Seeds oblong-lenticular, c. 1.2-1.5 mm long.
Seed
shape,
size
and
colour:
Broadly
elliptic,
faces
biconvex,
cells
1-2
mm,
brown.
Testa
surface:
Lateral
in d istin ct;
marginal rim thick, laterally compressed and shallow-grooved;
radicle curved with tapering apex surrounded by mostly narrow,
elongate and faint cells;
hilum subterminal, surrounded
by a
conspicuous collar, cells round; surface smooth.
M ea su re m e n ts : Hilum c. 94 pm diam., cells around hilum c. 12
pm diam.; radicle 250 pm long, cells surrounding radicle 31-38
pm long, c. 6 pm wide.
Specimens examined: Appendix 2 No 415, 416.
Distribution: N. Africa, Sahara and south Europe.
P. c h lo ro th y rs a M urb.
(Plate
9)
FI. Lib. Seeds c. 1 mm long , yellow-brown.
Seed shape, size and colour: Broad
elliptic, 1-2 mm,
light
brown .
Testa
surface:
Lateral
surfaces
biconvex,
cells
in distinct;
marginal face keeled compressed laterally, cells faint, wrinkled
47
on one marginal side, otherwise smooth; radicle flattened,
compressed
laterally from
both
sides,
surrounded
by distinct
cells which are mostly elongate; hilum subterminal,
protrusing, surrounded
strongly
by conspicuous cells; surface smooth,
glossy .
M e a s u re m e n ts : Keel c. 65 pm wide;
hilum c.75 pm diam.;
radicle c. 120 pm long.
Specimens examined: Appendix 2 No 418, 419.
Distribution: South Libya, Egypt to Morocco and Eritrea.
P. kapela (Hacq.) A. Kenner
(Plate 9)
FI. Lib. Seeds c. 1.25-1.5 mm long, brown.
Seed shape, size and colour: Broadly elliptic, 1-2 mm, brown.
Testa
surface:
m arginal
face
Lateral
strongly
faces
biconvex,
compressed
cells
laterally,
in d istin ct;
surrounded
by
narrow, elongate cells; radicle tapering gradually, ending in a
pointed
apex,
surrounded
by
narrow,
elongate
cells;
hilum
subterminal, surrounded by an inconspicuous collar of polygonal
cells; surface smooth.
Measurments:
Radicle c. 330 pm long, surrounding cells 42-90
pm long, c. 12 pm wide; hilum 113 pm diam., surrounding cells
6-13 pm long.
Specimens examined: Appendix 2 No 421, 422.
Distribution: Morocco, S. E. Spain, France, Libya.
4 8
H e rn ia ria L.
Herniaria L., Sp. PI.: 218 (1753); Chaudhri, Meded. Bot. Mus.
Herb. Rijksuniv. Utrecht 285: 297-398 (1968), rev.
H. cinerea DC.
(Plate
10)
FI.Lib. Seeds shining black, lenticulate, c. 0.5 mm long, erect,
with
brittle
testa.
Seed shape, size and co lo u r: Broadly elliptic , <1 mm, brown.
T e s ta
s u rfa c e : Lateral
faces
biconvex,
ce lls
in d istin ct;
marginal face keeled, compressed laterally, dark brown, around
the
circum ference;
radicle
thick
slightly
curved
toward
the
hilum, apex circular, filled with shrunken cells, surrounded by a
collar of square and rectangular cells; hilum subterminal, with a
plug of many, compressed, small cells, ± 3 cells high, and with a
collar
of square
or
polygonal
cells;
surface
smooth,
highly
glossy.
M e a s u re m e n ts : Radicle apex 60 pm diam., square cells 18x18
pm, rectangular cells 9-24 pm long ,12-15 pm wide; hilum c.30
pm.diam., ring cells 6-18 pm high, collar cells 15-21 pm long,
12-21 pm wide.
Specim ens exam ined: Appendix 2 No 224, 226.
D is tr ib u tio n :
Atlantic
N. Africa, S. E. Europe, Turkey, Pakistan and
Islands.
49
H .cyre n a ica
F. Hermann
(Plate
10)
FI. Lib. Seeds lenticular, c. 0.5 mm, shining brown.
Seed shape, size and colour: Circular, <1 mm, glossy brown.
Testa
surface:
L a te ra l
faces
biconvex,
cells
in d istin ct;
marginal face keeled, strongly compressed laterally; the radicle
and the hilum in prominent, terminal positions where the cells
have rectangular and polygonal are with deep cavity separating
the hilum and the radicle; surface smooth, glossy .
M e a s u re m e n ts : Radicle tip
30 pm wide, hilum 30 pm diam.,
groove 36 pm deep; cells surrounding hilum and radicle 9-18 pm
long, 6-15 pm wide.
Specimen examined: Appendix 2 No 227.
D istribution: Endemic to Libya.
H .e r ic ifo lia
Townsend
(Plate
10)
FI. Lib. No description.
Seed shape, size and colour: Circular, <1 mm, brown .
Testa
surface:
La te ra l
face
biconvex,
cells
in d is tin c t;
marginal face keeled around a part of the circumference; radicle
slightly curved, surrounded by a collar of conspicuous, narrow,
elongate or polygonal cells; hilum subterminal surrounded by a
collar of polygonal and rectangular cells, ending
in a dome
shaped structure; a shallow groove separates the hilum and the
radicle; surface smooth, glossy .
50
Measurments:
Radicle c. 40 pm diam., cells 10-40 pm long, 4-
12 pm wide; hilum c. 30 pm diam., cells 6-12 pm long, 4-8 pm
wide.
Specimens examined: Appendix 2 No 228, 229.
D istrib u tio n : Endemic to Libya along sea shore west of Tripoli .
H. fo n ta n e s ii Gay
(Plate
11)
FI.Lib. Seeds compressed, ovoid, c. 0.75 mm, light brown .
Seed shape, size and colour: Broadly elliptic, <1 mm, brown.
Lateral
surface:
Lateral faces
biconvex,
indistinct;
m arginal
face is strongly compressed laterally; radicle surrounded by a
double
c o lla r
of
elongate
and
recta ng ula r
cells;
hilum
subterminal, surrounded by a collar of rectangular or polygonal
cells; a deep cavity separates the
hilum and the radicle; surface
smooth, glossy.
Measurements:
Radicle apex c. 30 pm diam.; hilum apex c. 36
pm diam.; cavity c. 40 pm long, c. 10 pm wide; rectangular cells
6-18 pm long, 2-6 pm wide.
Specimens examined: Appendix 2 No 231, 234.
Distribution: Spain, Sicily, Canary Is. , N. Africa .
H .g la b ra
L.
(Plate
11)
F I.L ib . Seeds lenticular, c. .5 mm in diam., glabrous, shining
brown.
51
Seed shape, size and co lo u r: Circular-obovate , <1 mm, dark
brown.
T e s ta
s u rfa c e .
Lateral
fa ce s
biconvex,
cells
in distinct;
marginal face keeled around the circumference; radicle slightly
curved, surrounded
walls; hilum
by a conspicuous ring of cells with faint
subterminal, surrounded by a collar of polygonal
cells; surface smooth and glossy.
M easurem ents:
Micropyle c. 40 pm diam., hilum apex c. 48 pm
diam., space between radicle and hilum c. 60 pm long.
S pecim ens exam ined: Appendix 2 No 237, 238.
D is trib u tio n : Europe, N. Africa, Iran, Iraq, Turkey and Russia.
H .h e m is te m o n Gay
(P la te
11,12)
FI.Lib. Seeds ovoid-suborbicular, c. 0.5-0.7 mm long.
Seed shape, size and colou r: Circular-obovate, <1 mm, brown.
T e sta
s u rfa c e :
Lateral
faces
biconvex,
cells
in d istin ct;
marginal face keeled, compressed laterally; radicle thick curved
ending
in
a discoid
apex
filled
with
shrunken
cells;
hilum
compressed laterally with a thin apex surrounded by a collar of
faint, polygonal and elongate cells; a shallow space separates
the hilum and radicle; surface smooth and glossy.
M e a s u re m e n ts : Radicle apex c. 30 pm diam.; hilum apex c. 40
pm diam.; space between hilum and radicle c. 50 pm.
S pecim ens exam ined: Appendix 2 No 240, 241.
52
Distribtion:
Tribe
N. Africa, Iran-Turanian region.
C orrigioleae Dumort. (1827)
T e le p h iu m L.
Telephium L. , Sp. PI. :271 (1753); Wiliam, J. Bot. 44:289-304, rev.
T e le p h iu m
sp h a e ro sp e rm u m
Boiss.
(Plate 12)
F I.L ib .Seeds brown, sphaerical, c. 0.75 mm in diameter, not
com pressed, papillate.
Seed shape, size and colour: Globose, <1 mm, brown .
Testa
surface:
Lateral faces convex,
cells mostly spherical
marginal face convex; radicle slightly curved with discoid apex;
rectangular cells in about 10 parallel rows in a band extending
about half way round the seed; hilum in marginal notch; a marked
feature of this seed is a very distinct small caruncle
consisting
of a mass of small globular cells; surface rough.
Measurem ents:
Lateral surface
cells c.13 pm diam.,
apex c. 50 pm diam. ; caruncle cells c. 7 pm diam.
Specimens examined: Appendix 2 No 667.
D istrib u tio n: N. Africa, Egypt, Saharo-Arabia region.
II. Subfam. Alsinoideae (DC.) Fenzl (1824)
1.
Tribe
A re n a ria
Alsinoideae
L.
Arenaria L., Sp. PI.: 423 (1753); McNeil, Notes R. Bot. Gard.
radicle
53
Edinb.24: 102-129 (1962),245-309 (1963), part, rev., reg. rev.
A.
s e rp y llifo lia
L.
(Plate
F I.L ib .Seeds circular-reniform ,
12)
c. 0.5-0.7 mm long, blackish-
brown, with papillate tubercles.
Seed shape, size and colour: Subreniform
mm,
Testa
to
circular,
<1
brown.
surface:
Lateral faces plane or slightly convex, midzone
cells mostly ovate, round or elliptic, protrusive; spurs regular
cogwheel-like;
cells
marginal
in regular rows,
face
raised
plane
to
slightly
grooved
with
into
blunt, conical tubercules;
radicle curved, with distinct cells; hilum terminal in hilar notch;
surface granular.
Measurements:
Lateral face elliptic cells 80-156 pm long , and
c. 45 pm wide, ovate cells 45-78 pm long and 35-45 pm wide;
marginal face cells c. 43
pm diam. , spurs 14 - 16 ,
c. 6 pm
long.
Specimens examined: Appendix 2 No 157, 161, 164.
Distribution: N. Africa, Europe, Asia and N. America.
A .le p to c la d o s (Reichenb.) Guss.
(Plate
13)
FI.Lib. Not included in the Flora of Libya.
Shape, size, and colour: Circular-reniform, <1 mm, brown.
Testa
surface: Lateral faces slightly biconvex, midzone cells
mostly elongate or elliptic in regular rows; spurs regularly
54
spaced,
mostly short and with single blunt or sharp apices;
marginal face mostly convex cells round raised into blunt conical
tubercles,
and
in regular rows; radicle curved with elongate cells
sinuate
margins;
hilum
term inal
in
hilar
notch;
surface
minutely granular.
M easurem ents:
Lateral surface elliptic cells c. 42 pm long and,
c. 21pm wide, elongate cells 42-84 pm long and, c. 14 pm wide;
radicle cells 21-56 pm long, c. 14 pm wide; spurs 10-14 , 4-10
pm long.
Specim ens exam ined: Appendix 2 No 98, 100, 111.
A. le p to cla d o s
(Davis
50344)
(Plate
13)
FI. Lib. Not included in the Flora of Libya.
Seed shape, size and co lo u r. Circular-reniform, < 1 mm., dark
brown.
Testa surface: Lateral faces biconvex, cells mostly elongate or
elliptic, rarely ovate, mostly raised into distinct, round papillae;
spurs irregular in length, with sharp or blunt apices; marginal
face convex,
cells raised
into very distinct conical tubercles
with regular spaced spurs; radicle cells elongate, covered with
2-5 warts in each cell; hilum
mostly term inal, in hilar notch;
surface granular.
M easurem ents:
wide;
Lateral face cells 48-80 pm long and 16-25 pm
radicle cells 18-70 pm long 9-18 pm wide; spurs
55
5-18, 3-9pm long; papillae c. 6 pm long; tubercles c.18 pm long.
Specimens examined: Appendix 2 No 101.
C e ra stiu m
L.
Cerastium L. Sp. PI.: 437 (1753).
C. com atum Desv.
(Plate
14)
in FI. Lib. [C. illy ric u m D e sv.]
F I.L ib .Seeds obovate, c. 0.5 mm long, light brown, m inutely
tub ercu late.
Seed
shape,
size
and
colour:
Broadly-cuneate,
<1
mm,
orange-brown.
Testa
surface:
Lateral faces, flat, slightly sloping to notch,
cells elongate in regular rows, slightly curved, long and short,
radially arranged; spurs short, regularly spaced; marginal face
grooved,
cells
conically
tuberculate;
radicle
slightly
curved;
hilum in hilar notch; surface granular.
M easurem ent:
Lateral face cells 60-103 pm long, and 9 pm
wide; spurs number ± 17, c. 10 pm long .
Specimens examined: Appendix 2 No 192, 193.
Distribution: N. Africa, S. Europe, Turkey and Palestine .
C .d ic h o to m u m
L.
F I.L ib . Seeds
±
tu b e rcu la te .
(Plate
obovate,
c.
1
mm
long,
14)
reddish-brow n,
56
Seed shape, size and colour. Broadly cuneate, <1 mm, light
orange.
Testa
s u rfa c e .Lateral faces
circular,
raised
into
blunt
biconvex,
or
sharp
cells mostly ovate to
papillae,
dom e-shaped,
smooth at the apices, arranged in regular rows; spurs irregularly
branched, covered with small granules, arranged like necklaces;
marginal face convex, cells raised into conical tubercles with
tapering
apices; radicle curved;
hilum
in hilar notch; surface
minutely granular except the upper apices of the cells.
M e a s u re m e n ts : Lateral face cells
pm
wide;
spurs
8-18,
25-100
pm
45-120 pm long and c. 45
long;
m arginal
surface
tubercles 28-70 pm long.
Specimen examined: Appendix 2 No195, 197.
D istribution: Mediterranean, Western Asia .
C .g lo m e ra tu m
Thu ill.
(Plate
F I.L ib .Seeds subreniform,
pale-brown,c.
0.5
mm
14)
long, finely
tuberculate.
Seed shape, size and colour. Broadly cuneate, <1 mm, orange
but marginal face darker.
Testa
cells
su rfa ce .Lateral faces, flat, slightly sloping to the notch,
elongate,
slightly
curved
and
distinctly
raised,
radially
arranged, mostly regular, spurs long, tapering gradually toward
apices; marginal face grooved, cells raised into long tubercules
;
57
radicle
curved
about equaling the cotyledons, cells with
no,
distinct spurs; hilum in hilar notch; surface finely granular.
M e a s u re m e n ts :L a te ra l face cells 33-100 pm long and 10-23
pm
wide; spurs 10-16, 4-16 pm long; radicle cells 50 - 60 pm
long.
Specimens examined: Appendix 2 No 200, 201.
Distribution: Europe, Mediterranean, Western Asia .
C .lig u s tic u m
V iv.
(Plate
14)
FI.Lib. Seeds ovoid-reniform, c. 0.5-0.9 mm long, brown, sharply
and minutely tuberculate.
Seed shape, size and colour: Broadly cuneate, <1 mm, pale
brown.
Testa
s u rfa c e :L a te ra l
stelliform ,
distinctly
faces
raised,
flat,
cells
irregularly
narrow,
elongate
distributed;
spurs
or
of
differing lengths and irregular arrangement, mostly branching at
apices;
marginal face deeply grooved,
raised
into
short
conical
tu b e rcle s;
cells mostly stelliform
radicle
longer
than
cotyledons; hilum in hilar notch; the space between the radicle
and the hilum wide; surface minutely granular .
Measurements:
Lateral face cells 26-63 pm long and 6-16 pm
wide, spurs 8-11,13-20 pm long, tubercles c. 16 pm length.
Specimens examined: Appendix 2 No 202, 203.
D is trib u tio n ^ . Europe, Asia minor and N. Africa (Libya,Algeria)
I
5 8
C .p u m ilu m Curtis
(Plate
F I . L i b . Seed ± reniform,
15)
pale brown, c. 0.5 mm long, finely
tu b e rcu la te .
Seed
shape,
size
and
colour:
B roa dly-cun ea te ,
<1
mm,
orange-brown.
Testa
surface:
stelliform ,
mostly with
curved; spurs
m arginal
Lateral faces flat, cells narrowly elongate or
face
tuberculate;
distinctly
of differing
mostly
radicle
ridges,
often,
slightly
lengths and irregular arrangement;
rounded,
equals
raised
grooved,
cotyledons,
cells
curved;
stelliform
hilum
and
in hilar
notch; surface minutely granular.
M e a s u re m e n ts : Lateral faces cells stelliform , 3 3 - 50 pm long
and 23-33 pm wide ; lateral face spurs 12-14 , 10-16 pm long;
marginal cells rounded, 33 pm diam, marginal cells spurs 11 12 , 10 - 16 pm long ; cells at junction of lateral and marginal
faces elongate, 60 - 66 pm long and 15 - 18 pm wide , spurs 10
- 13 , c. 8 pm long, tubercles c. 35 pm long.
Specimen examined: Appendix 2 No 206, 207.
Distribution: N. Africa, Iran, Turkey and Caucasia .
C e rastiu m
se m id e c a n d ru m
L.
(Plate
15)
FI.Lib.Seeds somewhat rounded-reniform, pale brown, c. 0.4-0.5
mm long, finely tuberculate.
59
Seed shape, size and colour: Broadly cuneate, <1 mm, light
orange.
Testa
s u rfa c e :
Lateral faces
biconvex,
slightly
sloping
to
notch, cells mostly elongate or elliptic, with distinctly raised,
m ostly
blunt
papillae,
radially
arranged;
spurs
long,
sharp,
sometimes divided at apices; marginal face rounded grooved,
cells
stelliform
and
tuberculate;
radicle
equaling
cotyledons,
slightly curved;
hilum in hilar notch; surface minutely granular.
Measurements:
Lateral face long cells 35-67 pm long, and 14-
21pm wide, stelliform cells 21-35 pm diam.; spurs 10 -12 , 12
- 31 pm long; marginal face tubercles c. 20 pm.
Specimens examined: Appendix 2 NO 209, 210.
Distribution: Irano-Turanian region, Europe, N. Africa .
C .s ic u lu m Guss.
(Plate
15)
F I.L ib .Seeds rounded-reniform , 0.4-0.5 mm long, pale-brown,
finely tubercled.
Seed shape, size and colour : Broadly
cuneate,
<1
mm,
orange-brown.
T esta
surface: Lateral
middle,
cells
d is tin c t
granules
mostly
papillae;
arranged
fla t
stelliform ,
spurs
in
faces
raised
sharp,
regular
slightly
into
narrow
rows
on
concave
round
and
spurs;
in
the
or conical
cogw heel-like;
marginal
rounded, cells stelliform raised into conical papillae, dark in
face
60
colour;
radicle
curved,
equaling
or
slig h tly
longer
than
cotyledons; hilum in hilar notch; surface minutely granular.
M e a s u re m e n ts :
Lateral face cells stelliform,
12-25 pm in
diam.;
spurs
10-3
, 10-25
pm
long;
radicle
cells
16-50 pm
long,
marginal face tubercles c. 18 pm long.
Specimen examined: Appendix 2 No 212, 213.
Distribtion: N. Africa, W. Med. from Portugal to Italy and Sicily.
S t e l I a r ia
L.
Stellaria L., Sp. PI.: 421 (1753).
S. media
(L.) Vill.
(Plate
15)
FI.Lib. Seeds reniform, rounded, up to 1.5 mm long, compressed,
light to dark brown.
Subsp. m edia
Shape, size and colour: Circular-reniform, <1mm, dark-brown.
Testa
surface:
Lateral faces flat, cells stelliform
or elongate
in concentric regular rows, distinctly raised; spurs many, long,
with
sharp,
single
or
divided
ends,
with
2-4
small
warts
regularly arranged on each spur regularly; marginal face rounded,
cells
stelliform ,
raised
into
long
tubercules
(tongue-shaped),
decreasing gradually towards the hilum, covered by dense warts;
radicle equal and very close to the cotyledons, cells elongate
with very small or
no spurs, warts arranged on margins;
61
surface finely granular.
M e a s u r e m e n ts :
Lateral face, stelliform cells 30-80 pm diam,
elongate cells 60-113 urn long and, c. 26 pm wide; radicle cells
c. 7; marginal tubercules 60-120 pm long and, 53-106 pm wide.
Specimens examined: Appendix 2 No 660, 661, 662.
D istribution: Throughout the world.
S. media (L.) Cyrill.
Subsp.cupaniana
(Plate
16)
(Jordan & Fourr.) Nyman
FI. Lib. This taxon is not recorded from Libya.
Seed shape, size and colour:C ircular-obovate, <1 mm, darkbrown.
Testa
surface: Lateral
faces
slightly
convex,
cells
m ostly
stelliform or curved, elongate, distinct by raised, in concentric
rows; spurs irregular, covered with three or more small warts;
marginal face rounded, cells stelliform with very long tubercles,
flat
(tongue-shaped), spurs covered with
small warts;
radicle
equal and close to the cotyledons, cells elongate, lacking spurs,
covered with small warts, arranged on the margins; hilum in
hilar notch; surface finely granular.
M e a s u r e m e n ts :L a te r a l
face
stelliform
cells
67
pm
diam.,
elongate cells 67-133 pm long and 20-33 pm wide; spurs 11-14,
7-33 pm long; radicle cells c. 7 urn long; marginal tubercles 87167 pm
long, 53-113 pm wide .
62
Specimen examined: Appendix 2 No 659.
Distribution: Widespread in the Mediterranean area particularly
in E. Mediterranean.
S. pallida (Dumort)
Pire
(Plate
17)
FI. Lib. Not described.
Seed shape size and colour: Circular-reniform , <1
mm, red
brown.
Testa
cells
surface:
m ostly
short,
Lateral faces
stelliform ,
slightly
arranged
in
sloping to the
concentric
notch,
rows;
spurs
cogwheel-like, with one or two warts on each
spur;
marginal face cells round, tuberculate, spurs many, each covered
with one or two warts; radicle curved equal to the cotyledons,
cells elongate, spurs not distinct or very short, warts dense,
arranged
on the
cell
margins;
hilum
in
hilar
notch;
surface
m inutely granular.
M e a s u re m e n ts :Lateral face stelliform, c. 40 pm diam., elongate
cells c. 60 pm long, c. 27 pm wide; spurs 12-14 , 10-20 pm long;
radicle cells c. 33
pm long; marginal tubercles c. 47
pm long.
Specimens examined: Appendix 2 No. 665, 666.
Distribution: Mediterranean and Euro-Siberian regions.
6 3
M inuartia
L.
Minuartia L. , Sp. PL: 89 (1753), McNeil, Notes R. Bot. Gard.
Edin. 24: 133-150 (1962), 311-401 (1963), veg. rev.
M. ca m p e stris L.
FI.Lib.
(Plate
Seeds rounded-reniform ,0.4-0.7mm
long,
17)
brown, finely
tuberculate.
Shape, size and colour: Retortiform, <1 mm, orange.
Testa
surface: Lateral faces slightly biconvex to plane, cells
mixed, mostly ovate or elongate raised into blunt papillae; spurs
regularly
spaced,
cogwheel-like;
radicle
curved,
cells
lacking
spurs; marginal face convex, cells mostly ovate, rising into blunt
tubercles, spurs regular, cogwheel-like, with a few papillae on
notch cells; hilum in hilar notch; surface minutely granular.
M easurem ents: Lateral face cells elongate, 40-80 pm long and
20-24 pm wide, ovate cells
40-72 pm long and 28-40 pm wide
near the base; spurs ± 12-15, 4-12 pm long; radicle cells 13-54
pm long.
Specimen examined: Appendix 2 No 262.
Distribution: S.W. Europe, N. Africa.
M. geniculata (Poiret) Thell.
(Plate
17)
FI.Lib.S eeds rounded-reniform, c.0.5-0.82 mm long, brown, with
slightly rugose margin.
Seed shape, size and colour: Reniform, <1 mm, orange, tip of
64
radicle dark brown.
T esta
s u rfa c e : Lateral faces plane or slightly slanted, cells in
concentric rows radially arranged, elongate, smooth; spurs short,
blunt and regularly spaced; marginal face slightly concave, cells
covered with dense, small warts in the European specimen, the
Moroccan
and
Libyan
specim ens
lack
such
warts;
radicle
tapering, curved, with dark brown apex; hilum in hilar notch and
the adjacent
cells covered with dense small warts, in European
and Libyan specimens, but few in the Moroccan material; surface
generally smooth .
M e a s u re m e n ts : lateral face cells in European and Moroccean
specimens 40-120 pm long and 18-20 pm wide but in
Libyan
specimen 30-75 pm long and c. 15 pm wide; spurs in European
and Moroccean material spurs 10-22 but in Libyan material 7-16,
spur length in European specimen c. 12 pm but in Libyan and
Moroccan specimens c. 6 pm.
S pecim ens
exam ined: Appendix 2 No 295, 296, 297, 300.
D is trib u tio n :W id e ly distributed in the Mediterranean region.
M . h y b r id a (Vill.) Siskin
F I.L ib .
Seed
reniform ,
(P late
0.3-0.6
mm
long,
brown,
19)
fin e ly
tu b e rcu la te .
Seed shape, size and c o lo u r:
orange but radicle dark brown.
E lliptic-retortiform ,
<
1 mm,
Testa
the
surface:
notch,
elongate;
curved,
Lateral surfaces slightly biconvex, sloping to
cells
radially
arranged
from
the
notch,
m ostly
spurs short, blunt, straight or slightly curved; radicle
narrowing
gradually;
marginal face
slightly
concave,
cells raised into blunt tubercles; hilum in hilar notch, adjacent
cells
raised
into
elevated,
conical
papillae;
surface
m inutely
granular.
M e a s u re m e n ts : Lateral face cells 6-20 pm long and 10-16 pm
wide; spurs 10-24; 2-6 pm long; papillae near hilum c. 18 pm,
radicle cells 18-42 pm long.
Specimens examined: Appendix 2 No 316, 322, 324.
D istrib u tio n: Mediterranean area, S.W. Asia and eastwards to
A fganistan.
M.
m e d ite rra n e a (Link) K. Maly in Glasn.
(Plate 19)
FI.Lb.Seeds reniform, c.0.4-0.6 mm long, smooth.
Seed
shape,
size
and
colour:
C ircular-retortiform ,
<1
mm,
orange.
Testa
surface:
Lateral surfaces slightly
biconvex, depressed
near hilum, cells elongate, arranged radially, a few with blunt,
faint warts; spurs short, blunt and regularly spaced; marginal
face convex, shallowly grooved, cells raised into blunt tubercles;
radicle curved; hilum in hilar notch surrounded by elongate
66
papillae; surface minutely granular.
M e a s u re m e n ts :Lateral face cells 35-85 pm long and 10-15 pm
wide; spurs ± 20-26, c. 4 pm long, papillae near hilum c.14
pm,
marginal tubercles c. 10 pm long.
Specimen examined: Appendix 2 No 331.
Distribution: Throughout the Mediterranean area .
M. montana L.
(Plate
19)
FI.Lib.Seeds rounded-reniform, 0.6-0.8 mm long, dark brown,
obscurely and minutely tuberculate.
Seed
shape,
size
and
colour.
C ircu la r-re n ifo rm ,
<1
mm,
orange .
Testa
surface:
Lateral faces biconvex, cells mostly ovoid or
elliptic; spurs regularly spaced, narrow with sharp apices;
m arginal
face
convex,
cells
elevated
into
dom e-shaped
tubercules arranged in ± 6 rows, spurs regularly cogwheel-like;
radicle curved; hilum in hilar notch, adjacent cells raising into
distinct papillae; surface fine granular .
Measuremnts:
wide; spurs 22,
Lateral face cells 33-83 pm long and c. 26 pm
5-7 pm long; radicle cells 33- 65 pm long;
marginal face cells 28-49 diam. .
Specimens examined: Appendix 2 No 336, 339, 341.
Distribution: N. Africa, S. Europe.
6 7
Sagina L.
Sagina L. Sp.PI.: 128 (1753); Crow, Rhodora 80: 1-91 (1971),
reg. rev.
S.apetala Ard.
(Plate
F I.L ib .Seeds compressed,
reniform,
less than 0.4 mm
20)
long,
brown.
The illustration cannot be a seed of Sagina. It shows a seed
closely resembling those of Polycarpon.
Shape, size and colour: Cuneate to elongate-reniform, <1 mm,
lig h t-b ro w n .
Testa
surface:
Lateral faces
slightly
biconvex,
cells
mostly
elongate or round, large and few in number, ± 24 in each face;
spurs
shortly
pinnate
or cogwheel-like,
marginal face
rounded
equaling
cotyledons;
the
and
grooved,
hilum
near
with
cells
the
pointed
apices;
stelliform ;
radicle
surface;
surface
minutely granular.
Measurements
: Lateral faces,
cells stelliform
25-32
pm
diam. , elliptic, 41 - 50 long , spurs 9 - 15 , 4 - 16 urn long.
Specimens examined: Appendix 2 No 471, 474.
Distribution: N. Africa, Euro-Siberian area.
See 3.3 for a detailed discussion.
in
6 8
S. m aritim a
G. Don
(Plate
20)
F I.L ib .Seeds compressed, reniform to ± triangular, c. 0.4 mm
long, brown papillose.
Shape,
mm,
size
and
colour:
cuneate to triangular-reniform ,
<1
light-brown.
Testa
surface:
Lateral faces slightly biconcave, cells mostly
elongate or stelliform; spurs sharp, pinnnate or cogwheel-like;
marginal face slightly concave, rounded; radicle slightly curved;
hilum near the surface; surface minutely granular.
Measurements:
15-20
Lateral face cells elliptic 62 - 50 pm long and,
pm wide , stelliform cells c. 25 pm in diam. ; spurs 12 -
13 , 4 - 10 pm long.
Crow (1979) published an SEM of a seed of this species from
Sweden.
This
seed
appears
broadly
sim ilar
to
the
Libyan
M editerranean
coast,
m a te ria l.
Specimen examined: Appendix 2 No 476, 477.
D i s t r ib u t io n :
W idespread
along
the
Atlantic Islands, Northern and Western Europe.
III.
Subfam .C aryo p hyllo id eae
I.Tribe
Caryophylleae
G y p s o p h ila L.
Gypsophila L., Sp. PI.: 406 (1753); Barkoudah, Wentia 9:35-157 (1962).
69
G.elegans MB.
(Plate
FI. Lib. Seeds ± rounded-reniform, c.
20)
1.5 mm long, obtusely
tubercled.
Shape, size and colour: Circular-reniform, 1-2
Testa
in
surface:
regular rows,
papillae;
mm, brown.
Lateral faces slightly biconvex,cells arranged
mostly ellipitic
or ovate,
raised
spurs regularly spaced, mostly blunt;
with
blunt
marginal face
convex, cells stelliform, raised into pointed, conical tubercles;
radicle longer than cotyledons, thick and curved, cells mostly
similar to those of the lateral face; hilum in hilar notch; surface
finely granular.
Measurements:
Lateral face elliptic cells 85-187 pm long and
c. 51pm wide, ovate cells 51-85 pm long and c. 45 pm wide;
spurs ±10-15, 9-30 pm long; marginal face tubercles c. 85 pm
long.
Specimen examined: Appendix 2 No 222.
Distribution: Central Europe, Irano-Turanian region.
G. pilosa Hudson
(Plate
21)
FI.Lib. Seeds reniform, c. 1.5 mm long , obtusely tubercled.
Seed shape, size and colour: Circular-reniform,
Testa
surface:
cells elliptic,
1-2,
brown.
Lateral faces concave, slanted to the hilum,
raised into distinct conical papillae; spurs blunt,
curved down; marginal face broad, cells narrowly elliptic,
70
arranged in ± 7 regular rows with low or high tubercules; radicle
curved; hilum in hilar notch; surface rough and wrinkled.
M e a s u re m e n ts :L a te ra l face cells 100-210 pm long and c.40 um
wide; spurs 20, c. 20 pm long ; marginal face cells 120-250 pm
long and c. 40 pm wide .
Specimen examined: Appendix 2 No 223.
Distribution:
N. Africa, W. Asia from Turkey and Palestine into
adjacent regions.
Vaccaria
W o lf
Vaccaria Wolf, Gen. Ill (1776); Gen. Sp.: 234 (1781); Rechinger,
Flora Iranica, Cont. 163: 337-341 (1988), reg. rev.
V. hispanica (Miller)
(In FI. Lib. Vaccaria
Rauschert
p y ra m id a ta M edik.)
(Plate
22)
FI.Lib.Seeds globose.
Seed shape, size and colour: Globose, 1-2 mm, dark brownblack
Testa
surface:
Lateral faces
strongly
convex
with
m ostly
broad, elliptic cells, each cell raised into blunt papillae; spurs
shallow and curved; radicle indistinct; hilum occupies a terminal
depression, surrounded by an inconspicuous collar of polygonal
cells; a band of narrowly rectangular cells extend laterally from
the hilum about half way round the seed; surface finely granular
to smooth.
71
M ea su re m e n ts : Lateral face cells 62-92 pm long and 34-46 pm
wide; spurs 10-15, 2-11 pm long; hilum c. 125 pm diam.; band
cells c. 62 pm long and c. 24 pm wide.
Specimens examined: Appendix 2
No 668, 669.
Distribution: S. Europe, N. Africa, Irano-Turanian region.
Dianthus
L.
Dianthus L., Sp. PI.: 409 (1753).
D. c rin itu s Sm.
(Plate
22)
FI. Lib. Seeds ovate, 2-3 mm long, finely papillate.
Seed
shape,
size
and
colour:
D orsiventrally
com pressed,
broadly elliptic, >2 mm, black.
Testa
surface:
Dorsal faces plane or slightly concave, cells of
two types, one type extended along the radicle to the middle
ridge area with
differeing
shapes,
mostly elongate,
ovate or
polygonal, the other type covers the whole area of the dorsal
face, mostly narrow and elongate; spurs short, blunt or sharp;
ventral face plane, slightly convex near the hilum, cells mostly
narrow elongate
m ostly
round
com pressed;
or with differing
e llip tic
radicle
or
ovate;
stra ig h t
and
shapes, around the
m arginal
extended,
face
hilum
stro n g ly
longer
than
cotyledons with round apex; hilum on the surface, crateriform;
surface minutely granular.
Measurements:
Dorsal face narrow, cells elongate, c. 63-195
72
pm long and c.17 pm wide, spurs ± 10-21, c. 3 pm long; ventral
face narrow, elongate cells c. 60-120 pm long and c.17 pm wide,
spurs c. 3 pm long; radicle c. 264 pm long.
Specimens examined: Appendix 2 No 216, 217.
Distribution:
Europe, Asia, Australia.
It is cultivated as an
ornamental plant.
P e tro rh a g ia
(Ser.) Link
Petrorhagia (Ser.) Link, Handb. 2: 235 (1831); Ball and Heywood, Bull. Brit.
Mus. (Nat. Hist.) Bot. 3: 121-172 (1964).
P M lyrica ( Ard. ) Ball & Heywood
(Plate
23)
FI.Lib. seeds oblong , black , 1.5 - 2.3 x c. 1mm, smooth, margins
thin.
Seed
shape,
size
and
brodly elliptic (leaf-shaped),
Testa
colour:
D orsiventrally
com pressed,
1-2 mm, black.
surface: Dorsal surface plane, slightly
concave near the
centre, cells irregular and narrow or broadly elongate; spurs
broad, with blunt or sharp apices; ventral face slightly concave,
cells mostly elongate; radicle protruding extended with broad,
round apex; hilum near the lower third of the seed; surface
minutely granular.
Measurements
: dorsal face cells16-33 pm long and c. 13
wide, spurs 6-11, 4.2-8.3 pm long; radicle c. 220 pm long .
pm
73
Specimens examined: Appendix 2 No 423, 424.
Distribution: N.
P.velutina
Africa, S. Europe .
(Gauss.) P. W. Ball & Heywood
FI.Lib. Seed
black,
boat-shaped,
tuberculate or cylindrical-papillate
Seed
shape,
size
and
colour:
(Plate
1-1.3x0.7-0.8
mm,
23)
strongly
.
D orsiventrally
com pressed,
(boat-shaped), 1-2 mm, black.
Testa
surface: Dorsal surface extremely convex, cells mostly
stelliform in regular rows, each cell raised into conical papillae
with different heights; spurs deeply lobed, often single or bifid
with sharp or blunt apices; middle ridge area very distinct in its
elongate
cells
which
have
blunt
warts;
radicle
broad
and
conspicious; ventral side deeply concave, hilum near the lower
third, surrounded by prickles or blunt papillae; surface minutely
granular.
Measurements:
wide;
Middle ridge cells 42-70 pm long and c.14 pm
dorsal stelliform cells 28-56 pm diam, spurs 10-12 , 14-
40 pm long; papillae 21-42 pm long; radicle c. 170 pm long .
Specimens examined: Appendix 2 No 427, 428.
D is tr ib u tio n :
N. & S Africa, S. Europe., Cyprus, West Asia,
Hawaii, Western Australia.
SEM
photographs
of seeds of this widespread
species were
produced by Thomas (1980). They show testa features identical
74
to those of the seeds described here. She (p. 152) stated that
"The morphology
between
of the testa was found to be very consistent
different
papillae and spurs
populations".
The
SEM
of the testa
cells
in Cutter (1978) shows strong sim ilarities
with our material.
2.Tribe
Sileneae DC. (1824)
Silene L.
Silene L. Sp. PI.: 416 (1753); Melzheimer, Flora Iranica, Cont.
163: 341-508 (1988), reg. rev.
S .ap etala
Willd. Morphotype A.
F I. L ib . Seeds
rounded-reniform ,
(Plate
c.1
mm
long,
dull
23)
blackish-
brown, faces plane, deeply grooved on back with undulate wings .
Seed
shape,
size
and
colour:
C ircular-reniform ,
1-2
mm,
with undulate wings, orange-brown.
Testa
surface: Lateral faces slightly concave slanted in the
middle; cells narrowly elongate, tapering at the ends,
in the
midzone each cell is raised into one wart, cells of the undulate
wings narrow, elongate, raised into 6-9 warts on each cell; the
spurs undulate or indistinct in midzone but in the undulate wing
the spurs regularly spaced, with sharp or blunt apices; marginal
face with
deep groove; the
radicle equalling
cotyledons; the
hilum sunken in hilar notch, surrounded by long finger-shaped
papillae; pads with distinct cells; surface minutely granular .
75
M e a s u re m e n ts :
Lateral face cells 85-175 pm long, 14-21pm
wide; papillae in the midzone area c. 7 pm
hilar notch 14-35 pm long; spurs on the
long, papillae in the
undulate wing spurs
number ± 13-22, c. 7 pm long.
Specimens examined: Appendix 2 No 484, 487, 488, 489.
Distribution: Mediterranean area, SW. Asia, Tropical Africa .
S. apetala. Morphotype B.
(Plate
24)
FI.Lib. Not included in the Flora of Libya.
Seed shape, size and colour: Circular-reniform, 1-2 mm long,
with undulate wings, brown.
Testa
surface: Lateral face biconcave at the midzone, cells
narrow elliptic or elongately, raised into round or blunt warts;
spurs broad and very short, the cells of the wings undulate
raised into distinct warts; marginal face deeply grooved; pads
with
distinct
cells
raised
into
radicle equalling the cotyledons;
elongate
or
conical
hilum sunken
papillae;
in hilar notch
surrounded by elongate papillae; surface finely granular.
Measurements:
Lateral face cells 40-136 pm long and 10-24
pm wide, warts c. 3.7 long; papillae of the pads c. 23 pm long;
hilum papillae c. 39 pm long,
specimen examined: Appendix 2 No 485.
7 6
S.apetala. Morphotype C.
(Plate
24)
Seed shape, size and colour: C ircular-reniform ,1-2 mm
long,
FI.Lib. Not inculded in the Flora of Libya.
with undulate wings, brown.
Testa surface: Lateral face biconcave near midzone, cells
narrowly elliptic and elongate, raised into large conical papillae
and small warts; spurs lacking; cells of undulate wings narrow,
elongate
and
raised
into
many
warts;
radicle
equalling
the
cotyledons; hilum sunken in the hilar notch, surrounded by dense,
long papillae (finger-shaped); surface finely granular.
Measurements:
Lateral face cells 34-136 pm long, and 7-20
pm wide, lateral face warts 3.4-34 pm long; papillae of the
pads
c. 20 pm long; hilum papillae 14-40 pm long.
Specimen examined: Appendix 2 No 486.
S. articu la ta
V iv .
(Plate
25)
FI. Lib. Seeds Circular-reniform, much compressed, ± 2 mm
long, with 2 undulate wings, finely punctate-papillose.
Seed
shape,
size
and
colour:
C ircular-reniform ,
1-2
mm,
brown, with undulate wings.
Testa
the
surface: Lateral faces slightly concave, slanted toward
m argins,
midzone
cells
narrow ly
elongate, tapering
apices, without spurs and no papillae; cells of the
elongate, with many conspicuous regular warts on
at
undulate wing
77
each cell;marginal face deeply concave; radicle equalling the
cotyledons; hilum sunken in hilar notch; pads lacking; surface
finely
granular.
Measurements:
Lateral face cell 120-160 pm long, and c.14 pm
wide .
Specimens examined: Appendix 2 No 495.
Distribution: Endemic.
S.behen L.
(Plate
25)
FI.Lib. Seeds rounded -reniform , c. 1.5 mm long , faces concaveconvex, back wide, 4-5 furrowed with 4 rows of acute, conical
tubercles, brown.
Seed shape, size and size: Circular-reniform, <1 mm, orangebrown.
Testa
surface:
Lateral
faces
compressed
laterally,
slightly
slanted near hilar notch, cells elongate, arranged in 4 regular
rows,
m ostly
bifurcate
simple
at the
tapering
to
apex,
each
other
one
cell
apex
raised
but
som etim es
into
blunt or
spherical papillae at one end, in regular rows; spurs regularly
spaced,
grooved
short
w ith
and
mostly
d is tin c t
cotyledons; hilum sunken
with
sharp
tu b e rc le s ;
apices;
ra d icle
marginal face
eq ua llin g
the
in hilar notch; pads large, swollen;
surface minutely granular.
M easu re m e n ts : Lateral face cells 110-185 pm long and 26-56
78
pm wide; marginal face tubercles
c.
53 (am long; spurs ± 20-30,
c. 5 jam long; pad cells 26-86 urn long .
Specimen examined: Appendix 2 No 498.
D is trib u tio n :^ Africa, S. Europe, W. Syria, Cyprus.
S . c e r a s to id e s
L.
(Plate
25)
FI.Lib.Seeds rounded-reniform, dark brown, ± 0.6 mm long, faces
deeply
concave,striate,
back
wide,
shallow
obtusely
narrow
grooved.
Seed shape, size and colour: Reniform, < 1 mm, gray.
Seed
testa:
the margin,
Lateral faces deeply biconcave, slanted towards
cells mostly elongate and elliptic, radially arranged,
each cell raised into 4-10 round warts; spurs regularly
mostly with
spaced,
blunt apices; marginal face slightly grooved, with
elongate and ovate cells, each cell raised into blunt tubercles;
radicle equalling the cotyledons; hilum sunken in hilar notch,
surrounded
with
long, finger-shaped
papillae;
pads flat,
with
polygonal cells; surface mostly smooth.
M e a s u r e m e n ts :
Lateral face cells
36-75 jam long and 10-15
pm wide; pad cells 23-32 pm long; papillae around hilum 10-30
pm long .
Specimens examined: Appendix 2 No 503, 504, 505.
Distribution: S.Europe, and N. Africa.
7 9
S . c o lo r a t a
F I. L ib . seeds
Poiret.
rounded-reniform ,
1-1.5
mm
long,
dark
brown,
faces plane and smooth to somewhat tuberculate, deeply grooved
with 2 undulate or wavy wings .
Subsp.
Seed
colorata
shape,
(Plate
size
and
colour:
C ircular-reniform ,
25)
1-2
mm,
with undulate wings, brown.
Testa
surface: Lateral faces slightly concave, slanted towards
the margin, cells narrow, elongate with tapering apices, lacking
papillae
and spurs; slightly undulate wings raised
into many
blunt warts arranged in one line; marginal face deeply concave ;
radicle
equalling
cotyledons;
hilum
sunken
in
hilar
notch,
surrounded by long finger-shaped papillae; pad cells raised into
conical papillae; surface finly granular.
M ea su re m e n ts : Lateral face cells 60-185 pm long and c. 8 pm
wide;
papillae around hilum c. 30 pm long; pad papillae 8-20
pm long.
Specimens examined: Appendix 2 No 509, 511, 512.
D istribution: Mediterranean area, N. Iraq, Syrian Desert, Sinai,
eastwards to Arabia and Pakistan.
Subsp. t r ic h o c a ly c in a Fenzl.
Var. la sio c a ly x S.-W. et Godr.
FI. Lib. Not included in the
Flora of Libya.
(Plate
26)
80
Seed
shape,
size
and
colour:
C ircular-reniform ,
1-2
mm,
with undulate wings, brown .
Testa
surface:
midzone,
cells
Lateral
faces
slightly
narrowly elongate and
biconcave
lanceolate;
near
the
spurs very
distinct with regularly spaced and blunt apices; undulate wing
with narrow, elongate cells, each one raised into many blunt
warts;
m arginal
cotyledons;
face
deeply
concave;
radicle
equalling
the
hilum sunken in hilar notch; distinct pads with cells
raised into round and conical papillae; surface finely granulated .
M e a s u r e m e n ts :
Lateral face cells 54-179 pm long and10-20
pm wide; spurs 4-8 pm long; papillae around the hilum 100-170
pm long.
Specimens examined: Appendix 2 No 510.
Distribution: Mediterranean, Russia, Turkey and Pakistan.
S .c o n o id e a L.
(Plate
26)
F I.L ib . Seeds rounded-reniform, 1.25-1.5 mm long, concave on
one side, bluntly tuberculate, dark brown.
Seed shape,
size and colour:
C ircular-reniform -,
1-2
mm,
dark brown .
Testa
surface:
Lateral faces plane,
slightly slanted towards
the hilur notch, cells elliptic narrowly to broadly ovate shapes
raised into blunt papillae at the midzone; spurs regularly spaced,
with sharp apices; marginal face plane to slightly convex, cells
81
raised into blunt tubercules; radicle equalling cotyledons; hilum
sunken in hilar notch; pads large and globular; surface finely
granular.
Specimens examined: Appendix 2 No 517, 520.
M e a s u re m e n ts :
Lateral face cells 84-217pm long and 34-108
pm wide; spurs ± 20, c. 4-13 pm long; pad cells
Mediterranean
D is tr ib u tio n :
region,
Iraq,
32- 50 pm long.
Iran,
Turcomania,
Afganistan, Pakistan .
The SEM of a seed of this species in Melzheimer (1988) shows
strong similarities with the Libyan material.
S .c y r e n a ic a Maire & Weiller
(Plate
26)
FI. Lib. Seeds rounded reniform, c. 2 mm long, faces plane, back
grooved with 2 undulate wings.
Seed
shape,
size
and
colour:
C ircular-reniform ,
1-2
mm,
with undulate wings, brown.
Testa surface: Lateral faces mostly plane, slanted towards the
wing,
cells
indistinct
narrowly elongate, tapering
or blunt;
undulate
wing
cells
to the
distinct
apices;
spurs
in 3 regular
rows, each cell raised
into many blunt warts, arranged
single
marginal face
regular
line;
deeply grooved;
in a
radicle
equalling the cotyledons; hilum sunken in hilar notch surrounded
by
conical
papillae
and
distinct
callus;
pad
cells
distinct,
irregularly ovate or polygonal, raised into conical or round
82
papillae; surface finely grandular.
Measurements:
Lateral face cells 100-238 pm long and 7-18
pm wide; pad cells
c. 20 pm long,
c.10 pm wide, papillae
c.20
pm high.
Specimens examined: Appendix 2 No 523, 524.
Distribution:
E ndem ic.
S . f r u t ic o s a L.
(Plate
26)
FI.Lib. Seeds rounded-reniform c. 1.5 mm long, faces plane or
sligthly concave, striate, obtusely grooved on the back.
Seed shape, size and colour: Circular-reniform ,
1-2
mm,red-
brown.
Testa
surface: Lateral faces slightly biconcave, cells narrowly
elongate
or elliptic,
many cells
raised
into
conical
papillae;
spurs distinct, of varied irregular sizes; marginal face concave,
cells arranged into ± 6 regular rows, cells raised into pointed
conical tubercles; radicle equalling the cotyledons; hilum sunken
in hilar notch; pads distinct; surface granular.
M easu rem ents: Lateral face cells 84-168 pm long and c. 30pm
wide; spurs 15, 5-28 pm long; tubercles c. 79 pm long.
Specimen examined: Appendix 2 No 530.
D istribution:
N. Africa, Greece and Med. Islands, from Sicily to
Cyprus and, Turkey .
83
S .fu s c a ta
Brot.
(Plate 27)
FI.Lib. Seeds reniform, c. 1mm long , faces concave or
subexcavate and tuberculate,
back tuberculate,
ungrooved
or
shallowly grooved, or ± convex .
Seed shape, size and colour: Reniform, <1 mm, gray .
Testa
surface:
midzone
area,
Lateral face biconvex, slightly concave near
cells
ellip itic
or
elongate,
raised
into
blunt
papillae arranged in ± 4 regular rows; spurs short, distinct, with
regularly spaced and sharp apices; marginal face cells oval or
circular,
arranged
cotyledons;
in
concentric
hilum sunken
in
hilar
rows;
radicle
notch;
pads
equalling
the
distinct, flat
black; surface granular .
M e a s u re m e n ts :
Lateral
wide; spurs 12-22,
c.8.5
face cells 103-170pm and c. 36
pm
pm long; pad cells46-51 pm long.
Specimens examined: Appendix 2 No 531, 532.
Distribution: N. Africa, W. Europe, Lebanon, Palestine.
S. ga llica L.
(Plate 27)
FI.Lib. Seeds rounded - reniform , c. 1 mm long , dark brown to
black, with deeply concave, striate faces and wide, plane or
concave back, slightly tuberculate.
Seed shape, size and colour: Reniform, <1 mm, gray.
Testa
surface:
Lateral faces biconcave, cells broadly elliptic,
each cell raised into 2-3 distinct round warts; spurs lacking or
84
indistinct, 2-3; marginal face slightly convex cells arranged in
dense rows with different shapes, raised into conical tubercles
with
pointed
apices;
radicle
equalling
the
cotyledons;
hilum
sunken in hilar notch, surrounded by figer-shaped papillae; pads
globular, dark brown; surface granular.
M e a s u re m e n ts : Lateral face cells
66-116 pm long , 23-45 pm
wide; papillae c. 9 pm long; pad cells 28-70 pm long, 14 pm
wide.
Specimens examined: Appendix 2 No 534, 536, 539.
Distribution: N. Africa, W. Euro-Siberian, E. Tropical Africa and
Australia .
S.italica (L.) Pers.
FI.Lib.
(Plate
Seeds rounded-reniform,
27)
1-1,5 mm long, faces plane,
obtusely grooved on the back.
Seed shape, size and colour: Reniform, 1-2 mm, brown.
Testa
slightly
surface:
slanted
Lateral
faces
towards
the
com pressed
hilum,
cells
laterally,
ellip tic
or
plane,
ovate,
arranged in ± 5 regular rows, each cell raised into one round or
elongate papilla or into small, elongate warts; spurs regularly
spaced mostly with sharp apices; marginal face grooved; radicle
equalling the cotyledons; hilum sunken in hilar notch; pads large
and globular, cells elongate or polygonal, smooth or raised into
round papillae; surface finely granular.
85
Measurements:
Lateral face cells 95-186 pm long and 27-53
urn wide; spurs number ± 5-20 , 5-15 pm long; pad cells 15-50
pm long, c. 15 pm wide
Specimens examined: Appendix 2 No 546, 548.
Distribution: Mediterranean, Turkey and Russia.
S . lo n g ip e ta la
Vent.
(Plate
28)
FI.Lib. Seeds rounded-reniform , ± 2 mm long, plane or ± concave
on faces, obtusely grooved on the back , finely wrinkled .
Seed shape, size and colour:
Broad-reniform,
1-2 mm, red-
brown.
Testa
surface:
Lateral
faces
compressed
laterally,
slightly
biconcave, cells narrower elongate or elliptic, strongly raised, a
few cells with blunt papillae; spurs regularly spaced with sharp
apices;
m arginal
overlapping,
face
arranged
concave,
ce lls
in ± 4 regular
m ostly
rows;
spurs
elo n g a te ,
indistinct;
radicle equalling the cotyledons; hilum sunken in hilar notch;
pads distinct, cells mostly ovate; surface granular.
M e a s u re m e n ts : Lateral face cells 95-247 pm long and c.
20
pm wide; pad cells 38-76 pm long, c.19 pm wide.
Specimens examined: Appendix 2 No 558, 559.
D is trib u tio n : Widespread in Europe, N. Africa, Central Turkey
and Northern Iran.
86
S .m arm arica B eguinot
(Plate 28)
FI.Lib. There is no seed description .
Seed shape, size and colour: Reniform, <1 mm, red brown.
T esta
surface:
narrow ly
elongate,
Lateral
strongly
face
com pressed
raised;
spurs
laterally,
regularly
cells
spaced,
apices sharp or blunt; margial face concave, cells raised into
blunt conical tubercles; radicle equalling the cotyledons; hilum
sunken in hilar notch; pads large and globular, cells elongate,
curved or ovate; surface minutely granular.
M e a s u r e m e n t s : Lateral face cells 60-160 pm long, 20-40 pm
wide, spurs 6-22, 4-13 pm long ; marginal tubercles 64-84 pm
long.
Specimens examined: Appendix 2 No 560, 561.
Distribution:
Endemic.
S. m u s c ip u la L.
(Plate
28,29)
F I.L ib . Seeds rounded - reniform , c. 1 - 1.2 mm long , faces
plane, finely tuberculate, obtusely grooved on back, dark brown
to blackish - brown.
Seed shape, size and colour: Reniform, <1 mm, red brown.
Testa surface: Lateral faces plane, slanted toward hilum, cells
narrowly elongate or elliptic, arranged
spurs
regular
in size
and
with
sharp
in
± 3
regular rows;
apices;
m arginal face
slightly grooved, cells raised into blunt tubercles; radicle
87
equaling the cotyledons; hilum sunken in the hilar notch; pads
large and globular, cells elongate or elliptic; surface granular.
M e a s u re m e n ts :
lateral face cells
76-133 pm long and c. 24
pm wide; spurs number ± 12 - 20 , c. 6
pm long , pad cells 33 -
66 pm long.
Specimens examined: Appendix 2 No 566, 567.
Distribution: Mediterranean, extending to S.W. Europe .
S .n o c tu rn a L.
F I.L ib .
Seeds
rounded-reniform
c.
0.72-1.2
mm
(Plate
29)
long,
with
concave faces and wide, obtusely grooved, tuberculate back, un
winged .
Seed shape, size and colour: Reniform, <1 mm, dark brown to
black.
Testa
surface:
elliptic or
Lateral faces deeply biconcave, cells narrowly
elongate in regular rows, each cell raised into many
blunt,
distinct or indistinct warts, with
simple or bifid
ends;
spurs,
regularly spaced, with sharp apices; spurs with sharp
apices, cell ends simple or bifid; marginal face plane or grooved,
cells, arranged in ± 5 rows, narrow or elliptic, raised up into
long or small, blunt warts; radicle equalling cotyledons; hilum
sunken in the hilar notch; pads large and globular, cells mostly
elongate, spurs indistinct; surface finely granular.
Measurements:
Lateral face cells 41-119 pm long and 15-25
88
pm wide; spurs ± 16-22, c. 6 pm long; pad cells
36-45 pm long.
Specimens examined: Appendix 2 No 575, 576.
Distribution:
Mainly Mediterranean.
S .ru b e !la L.
F I.L ib .
(Plate 29)
Seeds
rounded
grooved
on
moderately
wide
back,
tubercled .
Seed shape, size and colour: Reniform, <1 mm, red brown.
Testa
surface:
Lateral
faces
biconcave,
cells
narrow ly
elongate with tapering ends, each cell raised into many blunt or
rarely into distinct, round warts; spurs short, regularly spaced,
with sharp apices; marginal face slightly concave, cells bluntly
tuberculate; radicle equalling the cotyledons;
hilar notch;
pads slightly flattened,
hilum sunken
cells elliptic
in
or elongate,
dark brown; surface minutely granular.
Measurements:
Lateral face cells 40-185 pm long and 15-25
pm wide; spurs 8-22 , 5-10 pm long;
pads 10-50 pm long, 10-
30 pm wide; marginal face tubercles c. 20 pm long.
Specimens examined: Appendix 2 No 590.
D istribution:
S .s e d o id e s
F I.L ib .
Mediterranean.
Poiret
(Plate
29)
Seeds reniform, 0.3 - 0.5 mm long, faces plane or ±
concave, striate, back obtusely grooved, black .
89
Seed shape, size and colour: Reniform, <1 mm, dark-brown .
Testa
surface: Lateral faces plane, slanted toward the hilum,
cells narrowly or broadly elliptic, arranged in ± 4 regular rows,
all the cells raised into one distinct papilla at one end; spurs
regularly spaced, with sharp apices; marginal face grooved, cells
arranged
in ± 4 rows, elliptic or elongate, with one distinct
papillae on each cell; radicle equalling the cotyledons;
sunken in
hilum
the hilar notch; pad cells mostly ovate, small and few
in number ± 3, one papilla raised on each cell; surface granular.
M easu rem ents: Lateral face cells 41-75 pm long and 20-25 pm
wide; spurs 18-20, 5-10 pm long; papillae 3-8 pm long, pads
cells 3-8 pm long, 15 pm wide; marginal
face cells 15-48 pm
long and 15-21 pm wide .
Specimens examined: Appendix 2 No 594, 595.
Distribution: Widely distributed in the Mediterranean area.
S .s u c c u le n ta
Forsskal
(Plate
30)
FI.Lib. Seeds rounded-reniform, c. 1 mm long, ± smooth or finely
striate,
Seed
brown.
shape,
size
and
colour: C ircular-reniform ,
<1
mm,
brown.
Testa
surface:
elliptic without
Lateral faces
plane,
papillae and lacking
cells faint,
spurs; marginal
elongate
or
face flat or
slightly concave; radicle equaling cotyledons; hilum sunken in
90
the
hilar notch;
pad cells
mostly elongate,
slightly flattened;
surface smooth.
Measurements:
Lateral surface cells 53 - 66 pm long and 10-
25 pm wide, pad cells 33 -54 urn long and c. 18 pm wide
D is t r ib u t io n :
S.
Europe,
N. Africa,
Egypt, Lebanon, Crete,
Sardinia, Corsica.
Specimens examined: Appendix 2 No 600, 601.
Distrbution:
Mediterranean.
S .tr id e n ta ta
Desf.
(Plate
30)
F I.L ib .Seeds rounded-reniform, c. 0.8 mm long, dark brown,
faces concave, striate, back wide with 1 or sometimes 2 shallow
grooves .
Seed shape, size and colour: Reniform, <1 mm, dark brown.
Testa
surface: Lateral faces deeply biconcave, cells elongate
or elliptic,
raised
into many warts;
marginal face concave,
spurs
blunt or indistinct;
cells broadly elliptic
and tuberculate;
radicle equalling cotyledons; hilum sunken in the hilar notch,
surrounded by finger-shaped papille; pad cells mostly ovate, flat
or
raised
into
distinct
papillae;
surface
finely
granular
or
smooth.
M e a s u re m e n ts :
Latteral face cells 20-100 pm long and10-20
pm wide; pad cells
papillae c. 26 pm long.
28-36 pm long,
12-20 pm wide;
hilum
91
Specimen examined: Appendix 2 No 602.
D is trib u tio n :
Spain,
N. Africa, W.
Irano-Turanian
region,
extending towards W. Mediterranean territories.
S .v illo s a
F I.L ib .
Forsskal
Seeds
rounded-reniform ,
c.
0.75
mm
(Plate
30,31)
long,
brown,
reticulate ± convex, grooved on the back, winged.
Seed
shape,
size
and colour:
C ircular-reniform , <1
mm,
orange.
Testa
the
surface:
m arginal
Lateral faces
face,
cells
plane, slightly slanted towards
ellip tic,
ovate
or
round,
papillae
indistinct; spurs lacking, cells surrounded by very thin, thread
like
structures
concave;
notch;
(probably
artefacts);
marginal
face
shallow ly
radicle equalling cotyledons; hilum sunk in the hilar
pad cells resembling
lateral face cells and difficult to
distinguish; surface smooth.
Measurements :
Lateralface elliptic cells 55-88 pm
long and
33 pm wide, ovate cells 44-77 pm long and c. 44 pm wide ,
round cells 41-100 pm diam.; spurs ± 16-22 , c. 3 pm long.
Specimens examined: Appendix 2 No 606, 607.
Distribution: N. Africa, Saharo-Arabian region.
The SEM in Melzheimer (1988) does not show the testa in great
detail but the cells appear elongate in contrast to the Libyan
92
material, with short, broad cells.
S . v iv ia n ii Steudel
(Plate
31)
FI. Lib. There in no seed description.
Seed shape, size and colour: Circular-reniform, < 1 mm, with
undulate wings, brown.
Testa surface: Lateral faces plane, slanted towards the hilum,
cells elongate, each cell raised into many blunt, round warts
concentrated in the midzone area; spurs regularly spaced, blunt
or indistinct; cells of the undulate wings arranged in 2 regular
rows, each cell raised
concave;
into many blunt warts; marginal face
radicle equalling the cotyledons; hilum sunken in the
hilar notch; pad cells distinct and globular in shape; surface
wrinkled, smooth.
Measurements:
Lateral face cells 75-150 pm long and 8-20 pm
wide; marginal tubercles c. 30 pm long.
Specimens examined: Appendix 2 No 610, 611.
Distribution: N. Africa, Jordan and Palestine.
S .vulga ris ( Moench ) Garcke
(Plate
31)
FI.Lib.Seeds rounded - reniform , c. 1.5 mm long, tuberculate.
Seed
shape,
size
and
colour:
C ircula r-ren iform ,
1-2
mm,
black.
Testa
surface: Lateral faces biconvex,cells elliptic, arranged
93
in ± 6 rows, each cell raised into one distinct, conical papilla in
the middle, uniform in shape, size and position; spurs distinct
with regularly spaced and sharp apices; marginal face convex,
cells
mostly round-stelliform ,
tubercle;
hilar
each
cell raised
into a conical
radicle equalling the cotyledons; hilum sunken in the
notch;
pad
cells
elongate
and
globular;
surface
finely
granular.
M e a s u r e m e n ts :
Lateral face cells 69-207 pm long and 35-69
pm wide; lateral face papillae and marginal face tubercles c. 68
pm long; spurs ± 11-20, 7-30 pm long; pad cells 69-75 pm long,
c. 23 pm wide.
Specim ens exam ined: Appendix 2 No 615, 616.
D is tr ib u tio n :
Europe, Mediterranean area, Middle East, Central
Asia eastwards to Kamtschatka .
The SEM of testa cell papillae and spurs in Barthlot (1981)
shows strong sim ilarities with the Libyan
material
particularly
the cells near the marginal face.
A g ro s te m m a
g ith a g o . L.
(P la te
31)
Agrostemma L., Sp. PI.: 453 (1753).
FI.
L ib .
Seeds ± reniform, c. 3 mm or more long, acutely
tubercled, black.
Seed shapes, size and co lo u r: Reniform, 1-2 mm, dark brown.
Testa surface: Lateral face plane, slightly slanted near the
94
hilar notch, cells mostly elongate, arranged ± 7 regular rows,
each cell raised into highly distinct, conical papillae; marginal
face broad, plane or slightly concave, cells arranged in many
regular rows, each cell raised into long tongue-shaped tubercles;
radicle ± equalling cotyledons, hilum sunken in the hilar notch;
surface granular.
M easu rem ents: Lateral face cells c. 100-280 pm long and c. 70
pm wide, papillae c. 30-60 pm long; marginal face tubercles c.
175 pm long.
Specimen examined: Appendix 2 No 1,2.
Distribution: Europe, Canary Island, N. Africa and Asia.
3.3 D ISC U SSION
There are examples within the SEM survey of Libyan seed morphology
which are very relevant to some taxonomic problems at the subspecies,
species, generic and subfamily/ family levels. At the infraspecific level
Silene apetala and Silene colorata subspecies colorata are good cases,
as are Arenaria serpyllifolia and Sagina apetala. For species, the endemic
taxa of Silene are especially good, as is the Stellaria media group. At the
subfamily level the precise rank of Paronychioideae and the family postion
of the genus Telephium have long been controversial. At the generic level
the status of Gymnocarpos is considered and Polycarpon, Polycarpaea and
Robbairea
have been much discussed as has the subgeneric status of
Minuartia geniculata .
95
Though not indigenous in Libya Sderanthus annuus and Corrigiola
littoralis have been examined for seed morphology. The seeds are very
similar and this has consequencies for the tribal divisions.
AREN ARIA SERPYLLIFOLIA
Group
This group of taxa has been often considered
regarding specific or
subspecific status. McNeil (1963) regarded seed size as the most
satisfactory character separating the tetraploid A. serpyllifolia
from the
diploid A. leptoclados, both automatically self-pollinating species. Perring
and Sell (1967) used seed characters but not those of the testa to separate
the subspecies leptoclados , serpyllifolia and macrocarpa (Lloyd) Perring &
Sell. In the first edition of Flora Europaea A. serpyllifolia and A. leptoclados
are treated as species but in the second edition they are regarded as
subspecies. Greuter et al. (1984) recognised as species the following in
the Mediterranean area. A. argaea Rech. fil., A. leptoclados (Rechenb.)
Guss., A. marschlinsii Koch, A. ? minutiflora Loscos [sic],
A. peloponnesiaca Rech. fil. and A. serpyllifolia L. For Britain Stace (1991)
accepts subspecific status: serpyllifolia, leptoclados (Reichenb.) Nyman and
lloydii (Jordan) Bonnier {A. macrophylla). For Libya Abdul Ghafoor lists
only A. serpyllifolia without any discussion of infraspecific taxa.
Soon after the SEM was available Godeau (1973) published
photographs of seeds of Breton origin. On the basis of seemingly only three
specimens he considered the value of testa features, including some visible
only at x 10,000 or x 30,000; these few observations were scarcely enough
to reveal the full range of variation. In his paper on Arenaria from the USA
Wofford (1981) published an SEM of A. serpyllifolia but did not indicate
which precise taxon. British, North African and Tenerife material showed
that the mid zone cells can vary in shape from more or less isodiametric to
96
very elongate, even in the seeds from one capsule of A. serpyllifolia s.s.
However, in most of populations of A. leptoclados examined most mid zone
cells are narrowly elongate.
The specimen collected by Davis (50344) from Wadi Derna in
Cyrenaica is very noteworthy. The specimen is well into the fruiting
condition and no petals can be seen. However, it has conspicuously
glandular hairiness on the stems, the small leaves and the sepals which
are c. 3.5 mm long and narrowly acuminate. These are some of the
diagnostic features of A. minutiflora Loscos, described from northeastern
Spain; see map in Jalas and Suominen (1983). Loscos (1877) in his
description of the taxon made no mention of seed characters. Lindberg
(1932) made A. minutiflora a subspecies of A. serpyllifolia and Monserrat
(1981) considered it a subspecies of leptoclados. Jahandiez and Maire
(1934) listed it from Morocco, Ozenda (1977) from Algeria and Maire (1963)
gave all of these countries and added Libya. Maire treated A. serpyllifolia
as having 3 subspecies. These are typica (with two varieties), leptoclados
(also with two varieties) and minutiflora. Like Maire Greuter eta!. (1983)
listed Algeria, Libya, Morocco and Tunisia but the species minutiflora is
qualified by an interrogation mark. In Flora Iberica Catroviejo et al. (1990)
lumped A. minutiflora with subspecies leptoclados
which is also the
treatment in the second edition of Flora Europaea .
As made clear in the seed description in 3.2 the seeds of Davis
(50344) are highly distinctive in their marked papillae. In the absence of
examination of Spanish material and in particular Loscos’ type specimen it
is not known if the seeds are papillate. Should such seeds prove to be
papillate the case for recognition of minutiflora as a distinct taxon would be
strengthened.
97
About 40 sheets labelled A. serpyllifolia s.s. and 18 labelled A. leptoclados
from North Africa, Tenerife and Britain were studied thoroughly. With the
use of a scale lupe 10 X, 36 specimens were measured for sepal length,
capsule width and seed diameter. These are three of the characters
considered important by Stace (1991) for the separation of these two taxa.
See tables 3 and 4. These measurements fit Stace's criteria and were
assumed to be adequate grounds for the distinction.
Distinct papillae had been noticed on the capsules but the papillae
were not uniformly distributed over the capsules in the two taxa There
appears to be a total separation: A. serpyllifolia s.s. has the distinct papillae
on the capsule body but not on the teeth whereas A. leptoclados has the
papillae on both the body and the teeth. These characters can be
recognised using a dissecting microscope and be seen very clearly in
capsule impressions (Plate 47). Furthermore the teeth of serpyllifolia are
glossier and blunter than those of leptoclados. All these characters of the
capsule teeth, unremarked by previous authors, were found in all the
material examined, whatever the geographical origin. It should be
emphasized that fully ripe capsules are necessary for clear observation.
A
necessary next step would be to consult the type specimens. If that of A.
serpyllifolia s.s. lacks distinctly papillose teeth and that of A. leptoclados
has papillose teeth then this would add weight to the recognition of the
specific status of the two taxa. Study of these features in other taxa listed in
the first paragraph has yet to be carried out.
The capsular teeth of Davis (50344) are papillate right to the apices
and this is further confirmation that the specimen is subspecies leptoclados,
if it does not prove to be a distinct taxon.
98
Sepal
Seed
Capsule
Capsule
size mm
size mm
size mm
±3.0
0.3 x 0.4
2.5 x 1.2
feature
*
±2.5
0.3 x 0.4
3.0 x 2.0
*
±3.0
~
3.0 x 1.9
*
±3.0
0.3 x 0.4
3.0 x 2.0
*
±3.0
0.4 x 0.4
3.5 x 1.5
*
±2.5
0.3 x 0.4
2.5 x 1.0
★
±3.5
0.3 x 0.4
3.0 x 1.7
★
±3.0
0.4 x 0.4
3.0 x 1.8
*
±3.5
0.3 x 0.4
2.5 x 1.1
*
±3.0
0.3 x 0.4
3.0 x 1.4
*
±2.5
0.3 x 0.4
3.0 x 2.0
*
±3.0
0.3 x 0.4
3.0 x 2.0
*
±3.0
0.3 x 0.4
3.0 x 2.0
★
±4
0.4 x 0.5
3.0 x 2.0
*
Table 3. Measurements of British in (GL) and Mediterranean specimens
of Arenaria leptoclados.
* = Capsule covered with distinct papillae including the teeth.
~ = No seeds present.
90
Sepal
Seed
Capsule
Capsule
size mm
size mm
size mm
feature
±3.0
0.3 x 0.4
2.5 x 1.2
+
±2.5
0.3 x 0.4
3.0 x 2.0
+
3.0 x 1.9
+
±3.0
~
±3.0
0.4 x 0.4
3.0 x 2.0
+
±3.0
0.4 x 0.4
3.5 x 2.0
+
±3.5
0.3 x 0.4
3.0 x 1.7
+
±3.5
0.3 x 0.4
3.0 x 1.7
+
±3.0
0.4 x 0.4
3.0 x 1.8
+
±3.5
0.3 x 0.4
2.5 x 1.1
+
±3.0
0.3 x 0.4
3.0 x 1.4
+
±2.5
0.3 x 0.4
3.0 x 2.0
+
±3.0
0.3 x 0.4
3.0 x 2.0
+
±3.0
0.3 x 0.4
3.0 x 2.0
+
±4.0
0.4 x 0.5
3.0 x 2.0
+
±3.5
0.5 x 0.6
3.5 x 2.5
+
±3.5
0.5 x 0.6
3.5 x 2.0
+
±4.0
0.5 x 0.6
~
3.5 x 2.0
+
3.5 x 2.5
+
3.5 x 2.5
+
±3.5
±3.0
~
±3.0
0.5 x 0.6
3.0 x 2.0
+
±4.0
0.5 x 0.6
4.0 x 2.5
+
±4.0
0.5 x 0.6
3.5 x 2.5
+
Table 4. Measurements of British in (GL) and North African specimens of
Arenaria serpyllifolia.
+ = Capsule covered with distinct papillae except the teeth .
- = No seeds present.
100
SAGINA APETALA Group
There has been much discussion of infraspecific taxa. The first edition of
Flora European gives subsp. apetala (S. ciliata Fries) and subsp. erecta
(Horneman.) F. Hermann (S. apetala auct.). The second edition adds (p.
178) “ Intermediates
are not uncommon, especially in the
Mediterranean region where the subspecific distinction is difficult to
maintain." In Flora of Turkey there had already been the statement (p.92) as
follows: “The differences between S. apetala and S.ciliata are inconstant,
and the two cannot be maintained as separate taxa. Stace (1991)
recognised the two subspecies but then adds (p. 208) “Possibly these 2
ssp. should be recognised as 2 or 3 vars”. None of these works makes any
detailed reference to seeds and certainly not to testa micromorphology.
Crow (1979) produced SEM of seeds of 15 species of the genus
from North America, Europe and eastern Asia. He included S. apetala Ard.
(two separate collections from California) but he made no mention of
infraspecific taxa. However, he showed SEMs of papillate and nonpapillate seeds. The description given in 3.2 deals with non-papillate seeds
from Libya, Morocco and Afganistan which closely resembles that in Crow,
Fig. 3 d.
Material from high altitude on Tenerife gathered by J. H. Dickson and
C. Rodriguez Pinero has markedly papillate seeds (Plate 20). The papillae
are sparse but prominent on the lateral face and, in that, the seeds are
similar to the seed of the second Californian material illustrated by Crow. It
is clear that the seeds of S. apetala sensu lato are variable but whether
this variation relates discretely to particular infraspecific taxa remains to be
seen.
101
SILENE APETALA
Silene apetala is a polymorphic species; Maire (1962) listed four varieties
and four forms. This polymorphism is clearly shown by the marked
differences in testa ornamentation in the specimens from different localities
in Libya. Most of the specimens examined have the ornamentation of the
kind here designated morphotype A (Morocco, Algeria and Libya). The two
other morphotypes, B and C, have been encountered in a few specimens,
all from Libya, (Plates 23, 24). It may be that the full range of variation in
testa micomorpholgy has not yet been found. Much more work would be
needed to formalise this variation taxonomically, if indeed it proved
possible.
SILENE ARTICULATA
AND S. GALLIC A
Silene articulata is an endemic species in Libya. It was treated in many
sources as a variety of Silene gallica, e.g. Durand and Barratte (1910) and
Keith (1965) and as a separate species in other sources e.g. Corti (1942),
Maire (1963) and Abdul Ghafoor (1978). Durand and Barratte attached
importance in separating var. articulata from S. gallica on calyx length.
Abdul Ghafoor gave differences in carpophore length. In specimens 494 to
496 (Appendix II) and 533 to 539 (Appendix II) the calyx measures 13-15
mm. In var articulata the seeds are reniform 1.4 X 1.6 mm, with undulate
wings, and in S. gallica the seeds are reniform 0.5 x 0.9 mm, without wings.
The difference in seed shape between these two species can be
recognised very quickly even without a hand lens. The SEM pictures show
great differences, particularly in shape, size and ornamentation of the
lateral and marginal face cells as listed in 3.2. Thus, it seems reasonable
to claim that the seed results obtained in the present study support the
separation of S. articulata from S. gallica . On the other hand it is worth
t
I
102
pointing out that the seed shape and testa features of S. gallica resemble
those of S. nocturna and S. cerastoides
and those of
S. articulata
resemble S. apetala, S. colorata and S. cyrenaica.
SILENE CYRENAICA
AND S. COLORATA
Silene cyrenaica is another endemic species in Libya and like S. articulata,
it has been given different ranks. It is very closely related to S. colorata . It
was treated as a variety of S.colorata by Durand and Barratte (1910) but it
was classified as a separate species in Corti (1942), Maire (1962), Keith
(1967) and Abdul Ghafoor (1978). These two have been recognized as
separate species on the basis of floral morphological characters as given
by Abdul Ghafoor (1978).
+Flowers in helicoid cymes with zigzag axis.
Calyx antrorsely appressed hairy throughout
S. cyrenaica.
- Flowers in scorpioid cymes, axis not zigzag.
Calyx antrorsely appressed hairy on nerves alone.....................S. colorata.
The descriptions of the seeds of these two species were given in
Flora of Libya as follows. S. colorata, “ seeds rounded-reniform, 1-1.5 mm
long, dark brown, faces plane and smooth to somewhat tuberculate, deeply
grooved with 2 undulate or wavy wings”. S. cyrenaica, “seeds roundedreniform, c. 2 mm long, faces plane, back grooved with 2 undulate wings”.
However, Maire (1963 p.112-115) described these species as follows.
S. colorata “seeds many, dark chestnut brown or dark brown-black, roundreniform, very compressed, 1.3-1.8 mm long, slanted, with undulate wings,
faces plane or slightly concave, with very fine radiate striation, sometimes
with a few papillae on the surfaces” and S.cryenaica “ seeds many, brownblack, round-reniform, c. 2 mm long, very compressed, with subplane faces,
smooth, at back slanted, with undulate wings, elegantly
103
striated radially, With papillae in rows”.
Abdul Ghafoor (1978 p. 75) said “ [Silene colorata is] a very common
and widely distributed species, very variable in habit, hairiness, calyx
length and petal colour. These characters have insufficient correlation, at
least in our area, and hence no infraspecific taxa are worth recognizing
here”. According to the material studied now which was collected from
Libya in 1939 and 1952, and some more recently, there appear to be two
different types of seeds belonging to the infraspecific taxa S. colorata,
subsp. colorata, and subsp. trichocalycina Fenzl., var. lasiocalyx S.-W. &
Godr. The separation between subspecies and varieties of S. colorata are
based on the presence or absence of hairs on the calyx and seed
characters (Maire, 1963). Seed features separate
these infraspecific taxa
and S. cyrenaica as follows.
IA. All lateral face cells with clear sinuate margins
subsp.
trichocalycina...var. lasiocalyx.
IB. Many lateral face cells lacking sinuate margins, mostly with straight or
slightly sinuate margins.
2A. Undulate wing cells, in indistinct rows and with blunt warts....subsp.
colorata.
2B. Undulate wing cells in distinct rows and with conspicuous
warts............................................................................S. cyrenaica.
The shape of the seeds and the SEM observations show very
considerable similarities between S. cyrenaica and S. colorata subsp.
colorata. Therefore the micromorphological characters, being only two
(wing cell arrangement and ornamentation) give little help in separating
these taxa and might be taken as supporting only subspecific rank for
S.cyrenaica.
104
SILENE MARMARICA AND S. ITALICA
Silene marmarica is the third endemic species of Silene in Libya. It is
treated as a separate species by authors such as Pampanini (1931), Corti
(1942), Maire (1962), Abdul Ghafoor (1978) and Greuter etal. (1984). This
plant is distributed in only one region, Cyrenaica. The seeds show very
great resembalance to those of S. italica which has been reported by Maire
(1963) from Cyrenaica. The seeds of these two species are alike in the
broad reniform to fan-like shape and the lateral faces being plane with cells
arranged in regular rows with distinct spurs and globular pads. The
difference between these two taxa lies in the lateral face cells which are
mostly elliptic and papillate in S. italica and elongate and smooth in S.
marmarica (Plates 27,28). It appears therefore that use of (SEM) to reveal
testa features can aid identification of these closely related species and
supports specific rank for S. marmarica.
STELLARIA MEDIA
GROUP
The taxonomy and nomenclature of the Stellaria media group has caused
much discussion for many years and in Libya, as elsewhere, different
authors have advocated its division into different taxa. Just before SEM was
available, Whitehead et al. (1967) in a detailed morphological study,
considered that seed length and weight, length of testa tubercles and
pollen diameter were the most important characters in separating the three
species Stellaria media, Stellaria pallida and S. neglecta. Using SEM,
Morton (1972) discussed the differences in seed morphology particularly in
the separation of North American S. media and S. pallida . He showed
marked differences between the seeds of these species, particularly the
tubercules and cells of the midzone area.
105
Berggren (1981) provided a key for the three species based on seed
size and ornamentation. Stamen number and seed size are the best
characters to separate S. media and S. neglecta according to Stace
(1991). In the most recent assessment, the second edition of Flora
Europaea recognised S. media (with two subsp.), S. neglecta and S.
pallida. For the separation of the three species and of the subspecies of S.
media seed size and colour as well as tubercle shape are used.
With regard to Libya, Durand and Barrate (1910) mentioned some
characters including smallness of seeds and followed by Corti (1942)
recorded only one species S. pallida (as S. apetala Ucria) from different
localities. In the Flora of Libya, however, Abdul Ghafoor (1978) took a very
wide view of S. media, incorporating both S. pallida
and S. neglecta .
Greuter et al. (1984) listed the presence of just one species of Stellaria in
Libya namely S. pallida (Dumort.) Pire . By using light microscopy and
SEM, two species are now confirmed in Libya, one is S. media subsp.
media and the other is S. pallida . This separation was based initially on
morphological characters particularly, the sepal length, stamen number,
and presence or absence of petals. The new (SEM) pictures show marked
characters of the seeds to distinguish the two species (Plates 16,17).
M in u a rtia g e n ic u la ta
and Status of R h o d a ls in e
McNeill (1962 p.135) discussed the geographical distribution and variation
of the species Minuartia geniculata as follows ” plants of subgenus
Rhodalsine are common throughout the more southerly Mediterranean
coasts extending to Portugal and the Canary Islands and with a distinctive
species in Somaliland. Throughout the greater part of its range, however,
the subgenus is only represented by one rather variable species for which
M. geniculata is the correct name”. This species has a variable leaf shape,
106
size and pubescence (Flora Europaea ; Abdul Ghafoor, 1978).
McNeill and Bassett (1974) made a thorough study of the pollen
morphology of Minuartia with the aim of elucidating the position of M.
geniculata, one of four species in subgenus Rhodalsine. The pollen of that
species differed from that of all other species of the subfamily Alsinoideae
but resembled that of Spergularia of subfamily Paronychioideae. M.
geniculata and Spergularia share similarities of habit and petal colour and
the subgenus had formerly been regarded as a genus by Gay (1845) and
Williams (1898). Despite their findings McNeill and Bassett rejected the
transfer of M. geniculata to the Spergularia but thought that the case for
raising the subgenus to generic status was strong. Greuter et al. (1984)
accept the generic status of Rhodalsine. MacNeill and Bassett were aware
that the seeds of M. geniculata resemble those of other species of the
genus Minuartia. The SEM survey of specimens from Libya, Morocco and
Spain makes this point very clearly but it does show a certain
polymorphism if only in the amount and arrangement of warts and perhaps
also in the size of testa cells showing differinences between Libyan
material and that from other countries.
Polycarpon, Polycarpaea and Robbairea
Polycarpaea and Robbairea are very closely related genera. The
characters used for their separation differ from one Flora to another, e.g.
bracts and stipules with or without a thick green midrib, and glabrous or
hairy plants. Zohary (1966, vol.I p. 128) mentioned this resemblance
“[Robbairea] resembling Polycarpaea but differing from it mainly by the
clawed petals and absence of staminodes”. Robbairea delileana was listed
under the genus Polycarpaea as P. prostrata (Oliver, 1868); Med-checklist
gives Robbairea delileana as Polycarpaea robbairea (Greuter et al., 1984)
107
but other
Floras
accept the
separate
genus
Robbairea (Post,
1883;Tackholm,1974; Abdul Ghafoor,1978; Hazim and Daoud,1985). The
genus R o bb aire a may be considered as linking Polycarpaea and
Polycarpon.
Post (1883, p. 158-159) mentioned the resemblance of
Robbairea to Polycarpon by describing the former as “Herbs resembling
Polycarpon” and he described Polycarpaea as having “seeds pearshaped, somewhat incurved - Herbs or shrubs, resembling Polycarpon
Pax (1889) in Pflanzenfamilien divided the genus Polycarpon into two
sections (Eupolycarpon, Robbairea) and genus Polycarpaea into three
sections (Aylmeria, Polycarpia, Planchonia), whereas Pax and Hoffman
(1934) classified P o lyca rp o n as a single genus but they divided
P olycarpaea into four sections (A y lm e ria , Polycarpia, Robbairea,
Planchonia) . Recently Bittrich (1993) has listed the two genera Polycarpon
and Polycarpaea in his classification but he did not accept the genus
Robbairea .
Seeds of six species of Polycarpaea were studied morphologically
by using light microscopy and SEM. Five of them are endemic species from
Tenerife (P. carnosa, P. divaricata, P. latifolia, P. smithii, P. tenuis). Also
studied were the Libyan species Polycarpaea repens, Polycarpon
prostrata, P. tetraphyllum and Robbairea delileana. Seeds of these species
show strong similarity in shape, colour and size (Table 5 ), (Plates 3 to 6).
The SEM allow the division of these spp. into two groups on the basis of
testa surfaces, the first group Polycarpaea repens, P. carnosa,Robbairea
delileana having more or less smooth or slightly wrinkled surfaces.The
second group Polycarpaea divaricata
Polycarpon prostrata
P. latifolia, P. smithii, P. tenuis,
and P. tetraphyllum. has papillate surfaces.This
second group can be further divided in to two, one with smooth papillae e.g.
Polycarpon prostratum and P. tetraphyllum. and the other with rough
108
granular papillae e.g. Polycarpaea divaricata P. latifolia, P. smithii and P.
tenuis . A key was made from the SEM pictures to differentiate between
Polycarpon, Polycarpaea and Robbairea species.
IA. Lateral face smooth or wrinkled but not papillate.
2A. Lateral face smooth...............................................Polycarpaea robbairea.
2B. Lateral face wrinkled............................... Polycarpaea carnosa, P. repens.
IB. Lateral face papillate.
3A. Lateral face papillae smooth.
4A. Papillae on lateral faces dense, mostly of uniform size..............................
............................................................................Polycarpon prostratum.
4B. Papillae on lateral face more widely spaced, less uniform in size
.......................................................................... Polycarpon tetraphyllum.
3B. Lateral faces papillae rough, granular.
5A. Papillae very closely spaced............ Polycarpaea divaricata.
5B. Papillae widely spaced
Polycarpaea latifolia, P. smithii, P. tenuis
Robbairea, Polycarpaea andPolycarpon
share a common seed shape.
With Polycarpaea the testa varies fromstrongly papillate to somewhat
wrinkled . Though much of the testa is smooth, Robbairea delileana has
indistinct wrinkles at the hilar end. Therefore there are insufficient seed
characters to support the generic status of Robbairea. The strikingly
papillate testa of Polycarpon
is very like those of many species of
Polycarpaea and so that generic separation is not supported by the seed
features. However, without a thorough investigation of all the 50 or so
species of Polycarpaea and the 16 of Polycarpon to suggest lumping these
genera would be unjustified.
109
Species
Shape
Colour
name
Polycarpaea
caronsa
Elongate
Brown
Size
Position
Lateral
Dorsal
L. xW.
R&H.
face
face
0.4x0.2
divaricata
99
0.6x0.3
latifolia
99
0.6x0.4
T. V.
Convex
99
99
99
99
99
99
99
99
99
99
repens
O.c.
Creamy
0.8x0.3
smithii
Cuneate
Brown
0.5x0.3
99
99
tenuis
Elongate
99
0.6x0.3
99
99
99
Grooved
99
99
99
99
99
Polycarpon
prostratum
tetraphyllum
19
99
Creamy
0.5x0.3
99
99
0.5x0.3
99
99
0.4x0.2
99
99
99
99
99
99
99
99
99
99
Robbairea
delileana
B. e.
Table 5. Comparison between, Polycarpaea, Polycarpon and Robbairea,
seeds.
O. c. = Oblong curved, B. e. = Broad elongate, H. = Hilum, L. =
Length, W. = Width, R. = Radicle, T. = Terminal, V. = Ventral
G ym nocarpos: Generic Status
Bittrich (1993) has incorporated the monotypic genus Gymnocarpos into the
much larger Paronychia with c. 50 spp according to Mabberly (1987). Five
spp of the latter have been examined and they proved very alike each other
in seed shape, circular to broad elliptic, with an annular embryo according
to Abdul Ghafoor, but unlike Gymnocarpos which has obovate seeds or in
the words of Zohary (1966, p. 129) " oblong-reniform, with a horseshoe
shaped embryo." On this basis bearing in mind that many spp of
Paronychia remain unexamined the genus Gymnocarpos seems worthy of
recognition.
110
TELEPHIUM : Family position
The genus Telephium with five species has been treated differently
regarding family position by many authors. Bentham & Hooker (1867)
placed it in tribe Mollugineae of the Ficoideae. Post (1883) in Flora of Syria,
Palestine and
Sinai placed this genus and the genus Glinus inOrder
Mollugineae (= Molluginaceae). In Flora of Cyprus Meikle (1977) listed it
under the tribe Telephieae of the family Aizoaceae. But Pax and Hoffmann
(1934 ), Davis (1967) and the second edition of Flora Europaea included
Telephium in subfamily Paronychioideae of Caryophyllaceae.
Meikle (1977) described the seeds of some species of
the
Aizoaceae including Telephium imperati. The testas of these species with
their verruculose or papillose cells show marked similarities as described
below:
Glinus lotoides : Seeds numerous , reniform, testa rich brown, regularly
papillose-verruculose.
Mesembryanthemum nodiflorum:
Seeds numerous, reniform, testa rich
brown, with distinct, bluntly verruculose dorsal ridges. [ Veruculosus =
covered with small wart-like outgrowth (Stearn 1993). Meikle is here
referring to the overall shape of the testa cells and not the minute granules
on the surface.]
Telephium imperati : Seeds compressed-subglobose, testa black or dark
brown, closely and regularly verruculose.
In a note Meikle (1977, p. 686) discussed the “ obscure” affinities of
Telephium and stressed the importance of the capsular, many-seeded fruit
in putting the genus in the Aizoaceae. Gilbert (1987) mentioned that most
authors who consider Telephium as belonging to the Caryophyllaceae
ignore the 3-4 locular capsules, which are anomalous in that family.
Recently Bittrich (1993) explained that “ The close relationship of the two
111
families is also demonstrated by the fact that a transfer of certain genera
from Caryophyllaceae (Sub. Paronychioideae) to Molluginaceae has
occasionally been proposed, recently by Gilbert (1987) for Telephium and
Corrigiola . Both genera show a reduction of the septa although only in the
upper part of the ovary in Telephium, and perigynous flowers, if weakly in
Telephium, and lack the bar-like thickenings in the endotegmic walls.
Therefore, a transfer would markedly increase the heterogeneity of the
Molluginaceae. Unfortunately, the sieve element plastids, which could
provide important evidence for or against an inclusion of Corrigiola and
Telephium in Caryophyllaceae, have not
been studied yet”. However,
Behnke (1994, p. 107) examined these plastids and considered that their
size (average dismeter 0.77 pm) and their polygonal crystals fit these two
genera into the Caryophyllaceae. Although Hoffman (1994, p .132)
indicated that " Telephium lacks the thickened and lignified apical ovary
walls of most Caryophyllaceae", she stated (p. 164) "The ontogeny of the
androecium clearly represents a variant of the caryophyllaceae pattern,
similar to that of Drypis. Therefore, Telephium and Corrigiola belong to the
Caryophyllaceae."
Telephium sphaerospermum with its globular and carunculate seeds
and
regular distinctive testa cells differs from all other species of the
Caryophyllaceae of whatever subfamily. The genus Vaccaria has globular
seeds as do some other small genera such as P leineuria
and
O chotonophila and species of Silene such as S. pendula and S.
pseudoatiocion have subglobular seeds. However in these cases and
certainly \r\Vaccaria and Silene the testa cells are obviously of the types
characteristic of the Caryophyllaceae. The genus Moehringia is well known
to have carunculate (strophiolate) seeds but again the testa cells are
spurred as in so many Caryophyllaceae. Furthermore, the large caruncules
112
in all the 12 species well illustrated by Pignatti (1982) consist of elongate
cells very different from the cells of Telephium. The eight Pyrenean species
of Petrocoptis have large caruncles in the form of tufts of hairs, again very
unlike Telephium.
This distinctive combination of seed characters supports the removal
of Telephium from the Caryophyllaceae. However, not all the species of
Telephium have globular seeds. Bittrich (1993, p. 225) describes the
seeds of T e le p h iu m as " globular to reniform." There is a fine
drawing of a seed of T. imperati L. in Castroveijo et al. (1990, p.102) and
the seeds are described as “ovaod-reniformes
finamente granulosas.” In
his monograph of the genus, Williams (1904) gave brief descriptions of the
seeds of four of the species such as that for T. oligospermum Boiss."
reniform-compressa, punctata vel subtiliter granulata, umbrinia." (p. 301).
In order to have the best evidence from the seeds, all five members of the
genus will need to be examined by SEM. Only then would be made a case
for or against removal of the genus from the family be strongest.
Subfamilies
As outlined in Chapter 1 the subdivision of the Caryophyllaceae has long
been controversial, particularly the status of subfamily Paronychioideae.
The subfamily Paronychioideae is separated from the other
subfamilies mostly by the presence or absence of stipules. A typical key to
the subfamilies is that of Komarov (1936, p.297).
1- Leaves stipulate.................................. Subfamily Paronychioideae
+ Leaves exstipu late.................................2
2- Sepals free or, if sometimes to the middle, then perianth
single..................................................Subfamily Alsinoideae
113
+ Calyx always with connate sepals, often tubular; perianth
always double........................................Subfamily Silenoideae
All the previous authors rely heavily on macromorphological characters of
vegetative and reproductive organs in their subdivisions and keys and they
have often ignored the importance of seeds.
In his subfamily diagnosis
Bittrich (1993) makes no mention of seed characters.
Seed shape. The reniform shape of the seeds of many members
of the Caryophyllaceae was commented on as early as Linnaeus.
This
shape
and
variations
of
it,
with
a
vertical
plane
of
symmetry through the centre of the concave (or notched) and
convex margins, is typical of the large genera Silene, Arenaria,
Stellaria, Minuartia and Cerastium as well as many small genera
of the
Caryophylloideae
reniform
shape
does
and Alsinoideae.
not
occur
in
the
However, a strictly
Paronychioideae
as
represented by the genera studied in this thesis. In considering
the shapes of the seeds it is most important to understand the
significance of the position of the hilum. In the vast majority of
the species within the Caryophyllaceae sensu lato the hilum is
sym m etrically
placed
at the deepest part of a more or less
distinct hilar notch so that it often appears sunken. This is a
statem e nt
Alsinoideae
which
and
pertains
p a rticu la rly
C aryophylloideae
and
to
not
the
to
sub fa m ilie s
the
subfam ily
Paronychioideae in which the hilum is more or less superficial
(and never appears sunken) and is asymmetrically placed as seen
11 4
in the outline drawings of Fig. 2.
There are a few exceptions to this generalisation. D ia n th u s
and
clo se ly
d o rsive ntrally
superficial
related
genera
flattened
but
and
are
in
sym m etrically
very
these
placed.
d istin ct
genera
in
the
Vaccaria
being
hilum
also
is
has a
superficial hilum and is unusual in the globose shape of its
seeds. In Alsinoideae Sagina cannot be said to have a notch but
the hilum is almost centrally placed on the concave margin. In
Paronychioideae the hilum may appear in some cases to be in a
very shallow groove but this groove runs along the ventral face
and not across it as does the hilar notch of the Caryophylloideae
and Alsinoideae.
Seed
Size. The seeds were placed into three categories: less
than 1mm, 1-2 mm and more than 2mm. All the members of
Alsinoideae studied here fall into the smallest category. Only a
few species of the other two subfamilies have seeds in the
largest and in the Libyan flora D ianthus
crin itu s , P te ra n th u s
dichotom ous and Sclerocephaius arabicus are examples.
Seed Colour and Lustre. So many members of Paronychioideae
have
glossy
seeds
that
this
fea tu re
can
be
considered
characteristic of the subfamily. Glossiness, however, is not
115
exclusive
to that subfamily.
M inuartia
and the monotypic
In Alsinoideae
some
species
of
Honkenya have shiny seeds. No
particular colour can be thought of as so typical of any subfamily
as to be important. Brown, orange and black occur in all the
subfamilies. Black or dark brown is charcteristic of Dianthus and
related genera, as has often been stated. The seeds of L o e flin g ia
h ispa nica
have
P te ra n th u s
species
a very
d ich o to m u s
in the
distinct
is
Libyan flora
greyish
utterly
white
different
in its dark brown
colour,
from
all
and
other
spot centrally
placed on the dorsal face. Meikle (1977, p. 285) describes this
feature as " a very prominent red blotch on one side of the seed".
The non-Libyan Scleranthus annuus has yellowish seeds with a
brown spot at the base of the radicle.
Lateral
with
Faces. Features of the lateral faces have been dealt
in detail
in section
Alsinoideae these faces
strongly
subfam ily
concave,
there
unlike
are
all
3.2.2.
In the
Paronychioideae
may be plane or convex
the
types
Caryophylloideae.
of face
and
but
never
In the
latter
including
the
deeply
concave as in S iie n e \ see Plates 31 (1), 27 (3) and 30 (6). In
Petrorhagia
the dorsal face can be strongly convex and the
ventral face strongly concave as in P. velutina; see Plate 23
(3,5). In the Paronychioideae as shown in Plates 1 to 11, a more
116
or less clearcut groove separates the lateral and marginal faces
or runs along the dorsal face. This is particularly obvious in
S pergularia
(Plate
P a r o n y c h ia
1,4),
in
S clerocephalus
Plate 8 (1), in
H erniaria
Plate
Plate
8
7
(3),
in
(7)
and
in
P o ly c a rp a e a Plate 3 (5). The groove is discernable even if only
faintly in all the other examples displayed, except P te ra n th u s
which is dorsiventrally compressed.
Testa
Cell
Shape.
In
Paronychioideae
these
are
smooth,
particularly in tribe Paronychieae. The cells may appear wrinkled
or very faint. In a few genera the cells may have jigsaw-like
shapes e.g Spergularia, Plate 2 (2).
Telephium with its globular
cells is very distinct, Plate 12 (40). In the other two subfamilies
the cells are almost always very distinct and often in more or
less
c o n c e n tric
rows.
Good
exam ples
of
c o n c e n tric ,
predominantly uniform, cells whether long or short are round in
Arenaria,
Plate
13 (3,4),
Cerastium,
Plate
14
(5,6),
and
M inuartia, Plate 17 (7,8). The cells can be of mixed shapes as in
A re n a ria , Plate 13 (2), C e rastiu m , Plate 15 (4), M in ua rtia, Plate
17 (5,6), Silene, Plate 28 (1,2), and Petrorhagia, Plate 23 (1.2).
The cells can be few in number as in Sagina, Plate 20 (3), S ilene,
Plate 29 (7) or a much greater number as in C erastium , Plate 15
(3), Vaccaria, Plate 22 (1) and Silene, Plate 26 (5). Exceptionally
the cells the testa are smooth as in Silene succulenta.
117
Spurs. These are universal in Alsinoideae and Caryophylloideae.
They can be long or short and even very short. Particularly well
developed spurs are found in A r e n a ria , Plate 13 (2), S te lla ria ,
Plate 15 (8), M in u a rtia , plate 17 (6), S a gin a, Plate 20 (4), and
S ilen e, Plate 28 (2). By contrast in Paronychioideae spurs are
absent
from
tribe
Paronychieae
and
only
present
in
some
members of tribe P olycarpeae especially Spergula, Plate 1 (2).
P a pillae ,
lacking
T u b e rcle s
and
altogether from
W arts.
the
These
seeds
of
ornam entations
are
Paronychioideae,
tribe
Paronychieae. In tribe Polycarpeae conspicuous if sparse papillae
are found in Spergula and Spergularia and very well ornamented,
numerous papillae occur in Polycarpea. However, no warts occur
in these genera nor is there ever the development of large
conspicuous tubercles as, for instance, in Stellaria.
P ads. This remarkable feature is to be seen only in Silene
not in
any other genus in any subfamily. This permanent character can
be
useful
as
a
good
taxonom ic
tool
at
the
specific
and
subspecific levels; see Plate 25 (2), 27 (5,7), 28 (5,8).
M a rg in a l
m arginal
fa c e .
face
In
Paronychioideae,
differs
strongly
in
tribe
Polycarpeae
m orphology:
broad
the
e.g
S p e rg u la ria , Plate 1 (7) and Loeflingia, Plate 6 (7), or expanded
into a membranous wing e.g. Spergula, Plate 1 (1), or with a deep
narrow furrow in the dorsal face e.g.
Polycarpaea, Plates 3 (5)
and 4 (8). However, in tribe Paronychieae the marginal face has
118
only
two
types,
broad
e.g.
P te ra n th u s , S cle roce ph alus and
G y m n o c a rp o s , Plate 7 (1,3 and 6) or strongly compressed e.g.
Herniaria,
Plate
10 (3), and
P a r o n y c h i a , Plate 9 (6). In
subfam ilies Alsinoideae and Caryophylloideae the marginal face
has a variety of shapes: convex e.g.
A renaria, Plate
13 (7),
C e ra stiu m , Plate 14 (1), S agina, Plate 20 (1), and S ilen e, Plates
30 (4) and 31 (5), concave in Cerastium, Plate 14 (5), M inuartia',
Plate 18 (7), and Silene, Plate 29 (2). Strongly concave marginal
faces extended
into undulate but not membranous wings are
found only in Silene e.g. Plate 23 (7) and 27 (5). The marginal
cells in these two subfamilies are arranged mostly in regular
rows, with cells sim ilar to or different from the cells of the
lateral face, see Plates 19 (9), 29 (4), 30 (2), and 21 (4).
R a d ic le . The radicle is the embryonic first root of a seed. Its
tip lies immediately below a more of less distinct pit on the
testa called the micropyle. In this study it is the prominence and
shape of the radicle, wihout the removal of the testa, that has
been
considered.
prom inent,
prom inent
In the
straight
radicle
and
usually
Paronychioideae the
pointed
curved,
in
radicle
is very
S c le ro c e p h a lu s but very
also
occur
in
Alsinoideae
(some species of Minuartia) and in Caryophylloideae (G ypsophila
and D ianthus). Because of the collar cells e.g., Plates 6 (7), 8 (6),
and 11 (5,6), the micropyle has a clearcutness in H erniaria,
119
P a ro n y c h ia and Lo e flin g ia
which is never the case in the other
two subfamilies which lack collar cells.
120
T
H
Fig 1. A-B = Length of a seed, C-D = Width of a seed,
E = Lateral face, F = Marginal face, G = Radicle ,
H = Hilum, I = Particular place of lateral face investigation
(midzone), J = Concentric arrangement of cells on lateral face.
*
Fig.2.
Range o f outline shapes in the Subfamilies o f the
Caryophyllaceae. * = Position o f the hilum.
Fig.2A. Subfamily Paronychioideae.
A = Broadly e lliptic e.g.
P a r o n y c h ia c h lo r o t h y r s a .
B = Broadly elongate e.g.
C = C ircular e.g.
P a r o n y c h ia a r a b ic a
D = Circular-obovate e.g.
E = Crescent-shape e.g.
F = Cuneate e.g.
P o ly c a r p a e a r o b b a ir e a .
H e r n ia r i a g la b r a .
P o ly c a r p a e a r e p e n s .
P o ly c a r p o n p r o s t r a t u m .
G = Globose e.g.
T e le p h iu m s p h a e r o s p e r m u m .
H = Obovate e.g.
S p e r g u l a r i a s a li n a .
1 = Narrow-obovate e.g.
P t e r a n t h u s d ic h o t o m u s .
Fig.2B. Subfamily Alsinoideae.
A = Broadly-cuneate e.g.
C e r a s tiu m g lo m e r a tu m .
B = Circular-reniform e.g.
C
= Circular-obovate e.g.
S t e lla r ia m e d ia
D = C ircular-retortiform e.g.
E = Elliptic-retortiform e.g.
F = Elongate-reniform e.g.
G = Reniform e.g.
subsp.
S t e lla r ia m e d ia
subsp.
m e d ia .
c u p a n in a .
M i n u a r t ia m e d ite r r a n e a .
M in u a r t ia h y b r id a .
S a g in a a p e t a la .
M in u a r t ia g e n ic u la t a .
H = Retortiform e.g.
M i n u a r t i a c a m p e s tr is .
I = Triangular-reniform e.g.
S a g in a m a r it im a .
Fig.2C.Subfamily Caryophylloideae.
A = Broadly-elliptic e.g.
D ia n t h u s c r in it u s .
B = Cricular-reniform e.g.
C —
Globose e.g.
G y p s o p h i l a e le g a n s .
V a c c a r i a h is p a n i c a .
D = Reniform e.g.
S i le n e c e r a s t o id e s .
E = Triangular-reniform e.g.
A g r o s t e m m a g it h a g o .
E
Fig.2B
A
E
F
Fig.2C.
A
E
D
B
Fig.2A.
B
B
H
I
C
D
H
I
C
D
122
Fig.3. Range of radicle outlines in the Subfamilies of the Caryophyllaceae.
Radicle shown by shading.
Fig.3A. Subfamily Paronychioideae.
A = e.g. Herniaria glabra.
B = e.g. Loeflingia hispanica.
C = e.g. Paronychia argentea.
D = e.g. Pteranthus dichotomus.
E = e.g. Sclerocephalus arabicus.
F = e.g. Telephium sphaerospermum.
Fig.3B. Subfamily Alsinoideae.
A = e.g. Cerastiumpumilum.
B = e.g. Minuartia campestris.
C = e.g. Sagina apetala.
D = e.g. Stellaria media.
Fig.3C. Subfamily Caryophylloideae.
A = e.g. Dianthus crinitus.
B = e.g. Gypsophila elegans.
C = e.g. Silene conoidea
D = e.g. Vaccaria hispanica.
Fig.3A.
Fig.3B.
B
B
D
Fig.3C.
B
D
123
LIST OF PALTES
The plates from no. 1 to 31 are SEMs of the seeds showing
details of the testa ornamentation located near the midzone of
the lateral face, the marginal face, the radicle and the hilar
notch as following.
PLATE 1.
1. Spergula fallax whole seed.
2.
„
„
lateral face (midzone).
3.
„
„
near the wing.
4. Spergularia bocconii whole seed.
5.
„
„
lateral face (midzone).
6.
„
„
near the hilum.
7. Spergularia diandra whole seed.
8.
„
„
lateral face (midzone).
9.
„
„
near the hilum.
PLATE 1.
PLATE 2.
1. Spergularia maritima whole seed.
2.
„
„
lateral face (midzone).
3.
„
„
near the marginal face and the wing.
4. Spergularia rubra whole seed.
5.
„
„
lateral face (midzone).
6. Spergularia salina (winged seed) whole seed.
7.
„
,, lateral face (midzone).
8.
„
„
near the marginal face and the wing.
I’ L A TE 2.
125
PLATE 3.
1. Spergularia salina (without wing) whole seed .
2.
„
„ lateral face (midzone).
3. Polycarpaea carnosa whole seed, ventral face.
4.
„
„ near the hilum and the radicle.
5.
„
„ whole seed, dorsal face.
6.
„
„ lateral face (midzone).
7. Polycarpaea divqricata whole seed.
8.
„
„ lateral face (midzone).
P L A T E 3.
PLATE 4.
1. Polycarpaea divaricata papillae.
2. Polycarpaea repens whole seed.
3.
„
„ lateral face (midzone).
4.
„
fJ whole seed.
5.
„
„ lateral face (midzone).
6.
„
„ ventral face.
7. Polycarpaea robbairea whole seed, ventral face.
8.
„
„ lateral face (midzone).
PLATE 4.
PLATE 5.
1. Polycarpaea simithii whole seed.
2.
„
„ near the hilum and the radicle.
3.
„
„ lateral face (midzone).
4.
„
„ papillae.
5. Polycarpaea tenius whole seed.
6.
„
,, near the radicle.
7.
„
,, lateral face (midzone).
8.
„
,, papillae
PLATE 5.
128
PLATE 6.
1. Polycarpon prostratum whole seed.
2.
„
„ lateral face (midzone).
3. Polycarpon tetraphyllum whole seed.
4.
„
„ lateral face (midzone).
5. Loeflingia hispanica whole seed.
6.
„
7.
8.
„ lateral face (midzone).
,, ventral face.
„
,, near the radicle.
P L A T E 6.
PLATE 7.
1. Pteranthus dichotomus whole seed.
2.
,, lateral face (midzone).
3. Sclerocephalus arabicus whole seed.
4.
„
„ lateral face (midzone).
5.
„
„ near the hilum and the radicle.
6. Gymnocarpos decander whole seed.
7.
„
,, lateral face (midzone).
8.
„
,, near the hilum and the radicle. .
PLATE 7.
PLATE 8.
1. Paronychia arabica whole seed.
2.
„
„ lateral face (midzone).
3.
„
„ near the hilum and the radicle.
4. Paronychia argentea whole seed.
5.
„
„ lateral face (midzone).
6.
„
„ near the hilum and the radicle.
7. Paronychia capitata whole seed.
8.
,,
,, lateral face (midzone).
PLA TE 8.
----
PLATE 9.
1.Paronychia capitata near the hilum and the radicle.
2.Paronychia chlorothyrsa whole seed.
3.
„
„ lateral face (midzone).
4.
„
„ near the hilum and the radicle.
5.
„
„ near the margin.
6. Paronychia kapela whole seed.
7.
„
„ lateral face (midzone).
8.
„
„ near the hilum and the radicle.
P L A T E 9.
PLATE 10.
1. Herinaria cinerea whole seed.
2.
„
„ near the hilum and the radicle.
3. Herniaria cyrenaica whole seed.
4.
„
,, near the hilum and the radicle.
5. Herniaria ericifolia whole seed.
6.
„
„ near the hilum and the radicle.
7.
„
,, lateral face (midzone).
8.
„
„ near the margin.
PLATE 11.
1. Herniaria fontanesii whole seed.
2.
„
„ lateral face (midzone).
3.
„
„ near the hilum and the radicle.
4. Herinaria glabra whole seed.
5.
„
„ near the hilum and the radicle
6.
„
„ near the hilum.
7.
„
,, near the radicle
8. Herinaria hemistemon whole seed.
PLATE 11.
PLATE 12.
1. Herniaria hemistemon lateral face (midzone).
2.
„
„ near the hilum and the radicle.
3. Telephium sphaerospermum whole seed x 300
4.
„
„
lateral face (midzone) x 1200.
5.
„
„
near the hilum x 600.
6. Arenaria serpyllifolia s. s. whole seed.
7.
„
„
lateral face (midzone).
8.
„
„
near the marginal face.
PLATE 12.
PLATE 13.
1. Arenaria serpyllifolia s.s.whole seed.
2.
„
„
lateral face (midzone).
3.
„
„
whole seed, x 300.
4.
„
„
lateral face (midzone), x 1200.
5. Arenaria leptoclados whole seed.
6.
„
„
lateral face (midzone).
7. Arenaria serpyllifolia (Davis 50344) whole seed.
8.
„
„
lateral face (midzone).
9.
„
„
near the hilum and theradicle.
PLATE 13.
PLATE 14.
1. Cerastium dichotomum whole seed.
2.
„
„
lateral face (midzone).
3. Cerastium glomeratum whole seed.
4.
„
„
lateral face (midzone).
5. Cerastium iliyricum whole seed.
6.
„
„
lateral face (midzone).
7. Cerastium ligusticum whole seed.
8.
,, lateral face (midzone).
PLA TE 14.
PLATE 15.
1. Cerastium pumilum whole seed.
2.
„
„
lateral face (midzone).
3. Cerastium semidecandrum whole seed.
4.
„
„
lateral face (midzone).
5. Cerastium siculum whole seed.
6.
„
„
lateral face (midzone).
7. Steilaria media whole seed.
8. Steilaria media lateral face.
9.
„
„
marginal face.
PLA TE 15.
PLATE 16.
1. Steilaria media subsp. cupanina. whole seed, x 150.
2.
„
„
lateral face (midzone), x 1200.
3.
„
„
near the hilum and the radicle, x 1200.
4.
„
„
marginal face, x 1200
5. Steilaria media whole seed, x 150.
6.
„
„
lateral face (midzone), x 1200.
7.
„
„
near the hilum and the radicle, x 600.
8.
„
„
marginal face, x 1200.
PLATE 16.
PLATE 17.
1. Steilaria pallida whole seed, x 150.
2.
„
„
lateral face (midzone), x 1200.
3.
„
„
near the hilum and the radicle, x 600.
4.
„
„
marginal face, x 1200.
5. Minuartia campestris whole seed.
6.
„
„
lateral face (midzone).
7. Minuartia geniculata whole seed, x 300.
8.
„
„
lateral face (midzone), x 2400.
PLATE 17.
PLATE 18.
1. Minuartia geniculata whole seed.
2.
„
„
lateral face (midzone).
3.
„
„
whole seed.
4.
„
„
lateral face (midzone).
5.
„
„
near the hilum and the radicle.
6. Minuartia geniculata marginal face.
7.
„
„
whole seed.
8.
„
„
near the hilum and the radicle.
PLATE IS.
PLATE 19.
1. Minuartia geniculata lateral face (midzone).
2.
„
„
marginal face.
3. Minuartia hybrida whole seed.
4.
„
„
lateral face (midzone).
5. Minuartia mediterranea whole seed.
6.
„
,,
lateral face (midzone).
7.
„
„
near the hilum and the radicle.
8. Minuartia montana whole seed.
9.
„
„
marginal face.
PLATE 19.
142
PLATE 20.
1. Sagina apetala (with papillae) whole seed, x 150
2.
„
„
marginal face showing the tubercles, x 4800
3. Sagina apetala (without papillae) whole seed.
4.
„
„
lateral face (midzone).
5. Sagina martima whole seed.
6.
„
„
lateral face (midzone).
7. Gypsophila elegans whole seed.
8.
„
„
lateral face (midzone).
PLATE 20.
PLATE 21.
1. Gypsophila pilosa whole seed.
2.
„
„
lateral face (midzone).
3. Gypsophila pilosa the radicle and marginal face cells.
4.
„
„
marginal face.
5.
„
„
near the hilum.
6. Vaccaria pyramidata near the hilum.
7.
„
„
whole seed.
8.
„
„
lateral face (midzone).
PLA TE 21.
PLATE 22.
1. Vaccaria pyramidata whole seed, x 72.
2.
„
„
lateral face (midzone), x 1200.
3. Dianthus crinitus (dorsal face) whole seed.
4.
„
„
5.
„
6.
„
„
„
lateral face (midzone).
7.
„
„
„
near the hilum.
8.
„
„
„
dorsal face near the middle line.
„
„
lateral face (midzone)
(ventral face) whole seed.
PLATE 22.
PLATE 23.
1. Petrorhagia illyrica whole seed.
2.
„
„ lateral face (midzone).
3. Petrorhagia velutina (dorsal face) whole seed.
4.
„
„
,,
5.
„
„ (ventral face) whole seed.
6.
„
„
„
lateral face (midzone)
near the hilum.
7. Silene apetala morphotype A. whole seed.
8.
„
„
„ lateral face (midzone).
PLATE 23.
146
PLATE 24.
1. Silene apetala morphotype B. whole seed.
2.Silene apetala morphotype B. lateral face.
3.
„
„
„
„
near the hilum.
4. Silene apetala morphotype C. whole seed.
5.
„
„
„
„
lateral face (midzone).
6.
„
„
„
„
near the hilum.
7. Silene articulata whole seed.
8.
„
„
lateral face (midzone).
147
PLATE 25.
1. Silene articulate lateral face (midzone).
2. Silene behen whole seed.
3.
„
„
lateral face (midzone).
4. Silene cerastoides whole seed.
5.
„
„
lateral face (midzone).
6. Silene colorata subsp. colorata whole seed.
7.
„
„
„
„
lateral face (midzone).
8.
„
„
„
„
near the hilum.
PLATE 25.
PLATE 26.
1. Silene colorata var.lasiocalyx whole seed.
2.
„
„
„
„
lateral face (midzone).
3. Silene conoidea whole seed.
4.
„
„
lateral face (midzone).
5. Silene cyrenaica whole seed.
6.
„
„
lateral face (midzone).
7.
„
„
near the hlium showing the callus.
8. Silene fruticosa marginal face.
PLATE 27.
1. Silene fruticosa whole seed.
2.
„
„
lateral face (midzone).
3. Silene fuscata whole seed.
4.
„
„
lateral face.
5. Silene gallica whole seed.
6.
,,
„
lateral face (midzone).
7. Silene italica whole seed.
8.
„
„
lateral face (midzone).
PLATE 27.
150
PLATE 28.
1. Silene longipetala whole seed.
2.
„
„
lateral face (midzone).
3.
„
„
near the hilum.
4.
„
„
marginal face.
5. Silene marmarica whole seed.
6.
„
„
lateral face.
7.
„
„
near the hilar notch.
8. Silene muscipula whole seed.
PLATE 28.
PLATE 29.
1. Silene muscipula lateral face.
2. Silene nocturna whole seed.
3.
„
„
lateral face (midzone)
4.
„
„
marginal face.
5. Silene rubella whole seed.
6.
„
„
lateral face (midzone)
7. Silene sedoides whole seed.
8. Silene sedoides near the hilar notch.
PLA TE 29.
PLATE 30.
1. Silene sedoides lateral face (midzone).
2.
„
„
marginal face.
3.
„
„
near the hilum.
4. Silene succulenta whole seed.
5.
„
„
lateral face.
6. Silene tridentata whole seed.
7.
„
„
lateral face.
8. Silene villosa lateral face.
PLA TE 30.
PLATE 31.
1. Silene villosa whole seed.
2.
„
„
lateral face (midzone).
3. Silene vivianii whole seed.
4.
„
„
lateral face (midzone).
5. Silene vulgaris whole seed.
6.
„
„
lateral face.
7. Agrostemma githago whole seed.
8.
„
„ lateral face (midzone).
P LA TE 31.
154
Chapter 4.
Crystals
4.1 Introduction:
According to Arnott (1981, p.225) “The structure of the crystalline deposits
of calcium oxalate in plants has been of interest to botanists since early in
the 19th century. Although first discovered by Leeuwenhoek in 1675, an
intensive study of plant crystals began as soon as light microscopy (LM)
developed to an adequate stage”. Al-Rais et al. (1971, P. 1217) stated “ The
crystals of different forms occuring in a wide variety of flowering plants
growing under normal conditions can be confidently identified as consisting
almost entirely of calcium oxalate”. The formation of calcium oxalate
crystals is very common in a great range of plant families across the world
as made clear by McNair (1932). He listed 77% of tropical families and
78% of temperate ones as crystal producers.
These crystals occur mainly in five major forms: druses, crystal sand,
prisms, raphides and styloids (Metcalfe and Chalk, 1950; Eames and
Macdaniels,1947; Cutter, 1969; Esau, 1977; Franceschi and Horner, 1980;
Fahn,1982). The crystals can occur in tissues of the leaf laminas, petioles,
flowars, stems, fruits and seeds; they can be associated with specific
tissies, e.g. epidermis, cortex, xylem, phloem and pith (Bohn, 1925; Scott,
1941; Price,1970; Laurance, 1976; Buttrose and Lott, 1978; Horner and
Frarceschi, 1978; Horner and Wagner, 1980; Arnott,1981; Kausch and
Horner, 1983).
In tieir important summarising paper Franceschi and Horner (1980)
considered several reviews published in the previous two decades. They
discjssed the significance of calcium oxalate and oxalic acid in plants in a
variety of ways, but did not consider taxonomy in detail. There are detailed
155
accounts of the importance of the shape, size and number of crystals within
the idioblasts as well as the development and ultrastructure of idioblasts
and of the function of oxalate crystals.
The occurrence and abundance of crystals in specific tissues of
various plants is often so constant as to be useful as a taxonomic tool
(Metcalfe and Chalk, 1950). The observation that crystals of calcium oxalate
are found associated with some bast fibres but not with others has been
used as a guide to identification by Jarman and Kirby (1955) who were
concerned with separating jute (Corchorus capsularis) from jute substitutes.
Agreeing with Metcalfe and Chalk, Al-Rais et al. (1971, p.1213) stated
“many flowering plant species produce relatively unreactive intracelluar
crystals, usually referred by anatomists to calcium oxalate.These can vary
considerably in form from species to species and sometimes also from one
region of the plant
to another; but in general the crystal forms or
combinations of forms are characteristic of species or higher taxonomic
groupings, so that they constitute useful classificatory criteria...” . SEM was
used to study the shape and location of the crystals in the perennial woody
stems of many families by Scurfield et al. (1973).
The detailed and direct application of crystals to taxonomic
problems was made by Dormers (1961,1962). He (1961) recorded crystals
of differing forms in the ovary wall of many Compositae, some of which
were of very restricted taxonomic distribution. In a more extensive study of
the genus Centaurea , Dormer (1962) studied 112 species, and on the
basis of crystal forms, made suggestions for taxonomic changes at the
subgeneric and sectional levels. Franceschi and Horner (1980)
briefly
consider a variety of other papers dealing with the taxonomic impotance of
crystals. Patterns of calcium oxalate were evaluated as a taxonomic tool for
studying Camellia sasanqua X sinensis and Agrimonia by Umemoto
156
(1981) and Murata and Umemoto (1983). Koteshwar and Ramayya (1984)
showed the importance of crystal size, shape and distribution in 26 species
of F icus; their key for identification was based on the crystals and
crystalliferous elements. Investigating the crystals of the corm tunics of
Crocus bulbs, Wolter (1990) found that the prismatic shape was restricted
to a few closely related taxa. He made taxonomic assessments at the
infrageneric level.
In the Caryophyllaceae, Amar (1904) recognised three types of
distribution of crystals within roots, stems and leaves and she discussed
particularly Tunica saxifraga, Dianthus carthusianorum and Saponaria
officinalis. Bohn (1925) pointed out the presence of calcium oxalate in the
epidermal cells of Lychnis fios-jovis, Silene dioica and Spergula arvensis;
in the leaves of these three species different forms, shapes and sizes of
crystals were observed. Haberlandt (1914, p.531) stated that “genuine
sphaerocrystals” had been reported from “ Silene cucubalus and certain
other Caryophyllaceae (according to Hegelmaier)". In the few previous
studies of crystals in the Caryophyllaceae no taxonomic conclusions had
been made (Amar, 1904; Bohn, 1925; Metcalfe and Chalk, 1950) for any
genus, not even Minuartia which has now been investigated in detail. For
the Caryophyllaceae Metcalfe and Chalk (1950, p. 150) stated." Calcium
oxalate commonly present in the form of large, conspicuous cluster crystals
in many genera and species including Arenaria, Corrigiola, Gymnocarpos,
Minuartia sp., Pteranthus, Scleranthus, Silene. The abundance of the
crystals sometimes varies within a single species in specimens from
different localities. Crystal-sand also recorded in Dysphania, Gymnocarpos,
Habrosia, and other genera.”
In the present investigation, the results of studies on type,
157
shape,
size and
distribution
of crystals
in mature
leaves of
Arenaria (86 spp), Moehringia (15 spp) and M inuartia (60 spp) are
presented
and their taxonom ic significance evaluated.
follow ing
assessm ent
the
results
are
com pared
In the
with
the
taxonomic treatment of the generic and infra-generic groups of
McNeill
(1962-1963).
Some
nom enclatural
changes
made
by
Rabeler (1993) have been followed.
4.2 Morphology and Distribution of Foliar Crystals in the Genera
Arenaria, Moheringia and Minuartia
Within the three genera the crystal were found to be druses, sand and
elongate. No raphides or styloids were encountered, either singly or
aggregated.
Druses: These are spheroidal aggregates of prismatic crystals found in all
three genera, ( c. 8-97 pm) diam. See Plates 32, 33 (1 to 6), 36 (1 and 2).
Crystal sand: Very small spheroidal or prismatic crystals in ellipsoidal or
spheroidal masses, found in Arenaria and Minuartia but not in Moehringia,
(c. 10-29 pm) long See Plates 33 (7 and 8), 34 (1,2 and 4), 35, 36 (3 to 8).
Elongate crystals: These resemble very rough-skinned cigars. They
appear to be aggregates, large to very large, up to 230 pm in length. In
overall shape and size these crystals are unlike anything reported in the
literature cited above, (c. 39-233 pm) long. Though their formation is not
understood they have taxonmic significance as discussed below. They are
found in Arenaria and very strikingly in Minuartia but not in Moehringia.
Plates 34 (5 to 8), 35 (1 to 3,5 to7), 37.
158
The patterns of crystal distribution discussed in the next section are
illustrated in Figure 4.
4.2.1 A re n a ria L.
I.Subgenus Leiosperm a McNeill
Distribution: Is confined to the New World, and has its great centre of
diversity in the Andes. Probably all the South and Central American,
species of Arenaria belong to one of two subgenera, Leiosperma or
Dicranilla .
A. aisinoides: Druses, dense, scattered
irregularly throughout the leaf,
except the veins, mostly spheroidal with irregular margins and very sharp
points; size mostly large c. 29-68 pm diam., Plate 32 (2,3), plate 36 (1).
A. decussata: Druses, dense, scattered irreguarly throughout the leaf,
except the veins, spheroidal with irregular margins, with sharp or blunt
points; size mostly large 29-68 pm diam.
A. guatemalensis: Druses, dense, scattered irregularly throughout leaf,
except the veins, spheroidal with irregular margins, with very sharp points;
size mostly large 29-68 pm diam.
A. lanuginosa: Druses, dense, scattered irregularly throughout the leaf,
except the veins, with mixed shapes and sizes, mostly spheroidal,
ellipsoidal or polygonal, margins irregular with sharp or blunt blunt points;
size 10-78 pm diam.
A. lycopodioides : Similar to A. guatemalensis; size c. 0.2-0.5 mm diam. .
A. paludicola : Druses, mostly few, scattered irregularly throughout the leaf,
except in the veins, mostly spheroidal, with regular margins and blunt
points, size mostly medium or large, 19-68 pm diam., Plate 33 (3,4,5).
A. reptans : Similar to A. guatemalensis .
II. Subgenus Dicranilla (Fenzl) William
159
D istribu tion: The distribution is similar to the previous subgenus
Leiosperma.
A. boliviana : Druses, few, scattered irregularly throughout the leaf, along
veins, mostly with irregular margins and blunt points; size mostly medium or
large, 29-68 pm diam.
A. bryoides : Similar to A. boliviana .
A. pycnophylla : Druses, dense, scattered irregularly throughout the leaf
except in the veins, margins irregular, with sharp points, mixed sizes, 10-68
pm diam.
III. Subgenus Porphyrantha (Fenzl) McNeill
Pyrenees and the Cantabrian Mountains
A. purpurascens : No material studied.
IV. Subgenus Arenaria
A. Sectio Rariflorae Williams
Distribution: Is widely distributed Arctic-Alpine group occuring throughout
the northernmost parts of Euroupe and America and extending south into
the mountains of C. Euroupe, Spain, the Balkans, Anatolia and Iran.
A. ciliata : Druses, dense, scattered irregularly throughout the leaf, except in
the veins, spheroidal, margins irregular, with sharp points; size 19-78 pm
diam.
A. humifusa : Druses, few, scattered irregularly throughout the leaf, except
the veins, spheroidal, margins irregular with sharp points, size mostly large
19-58 pm diam.
A. huteri. Similar to A. humifusa.
A. pseudofrigida „
„
160
B. Section Grandiflorae McNeill
Distribution: Section Grandiflorae, which is closely related to section
Rariflorae, comprises two species complexes, one in S. W. and C. Europe
and one in Turkey .
A. grandiflora: Druses, dense, scattered irregularly, mostly concentrated
along
veins, spheroidal, margins mostly irregular with sharp points; size
10-49 pm diam..Plate 32 (5,6).
A. incrassata: Druses, dense,Scattered irregularly throughout the leaf
except the veins, margins irregular with sharp points; size 19-58 pm diam.
C. Sectio Plinthine (Reichb.) McNeill
Distribution: It is a very distinctive group of plants endemic to the Iberian
Penisula and North Africa .
A. armerina : Druses, very dense, scattered throughout the leaf except on
veins, spheroidal, margin regular with blunt points; size 10-29 pm diam.
A. lithops : Druses, dense, scattered irregularly throughout the leaf except
the veins, spheridal, margins regular with sharp points, mostly small size
10-49 pm diam.
A. tetraquetra: Similar to A. lithops size 10-29pm diam. .
D. Sectio Rotundifoliae McNeill
Distribution: Greece, Turkey, Aremenia and Georgia.
A. biflora : Druses, few in number,scattered throughout the leaf area except
the veins, spheroidal, margins regular with blunt points; size 19-58 pm
diam.
A. halacsyi: Similar to A. biflora', but margins irregular, with sharp points;
size 29-116 pm diam.
161
E. Sectio Planosepalae McNeill
Distribution: A single species of the sections, is confined to the Iberian
Penisula and South-west France and appears to be taxonomically rather
isolated .
A. montana: Druses, very dense, scatered throughout the leaf area except
the veins, spheridal, margins irregular, with sharp points; size 19-78 pm
diam., Plate 32 (7,8).
F. Sectio Orientales McNeill
Distribution: The centre of distribution of the section is the Eastern
Mediterranean .
F. (I). Series Anomalae McNeill
A. bertolonii: Druses, dense, scattered throughout the leaf except in the
veins , spheroidal, margins irregular with sharp or blunt points; size 19-97
pm diam.
F. (II). Series Graecae McNeill
A. filicaulis : Druses, dense, distributed throughout the leaf area except in
the veins, sheroidal, margins irregular, with sharp points; size 10-39 pm
diam.
A. teddii: Similar to A. filicaulis.
F. (III). Series Deflexae McNeill
A. deflexa: Druses, very dense, scattered throughout the leaf except in the
veins, spheroidal, margins irregular, with very sharp points; size mixed, 1949 pm diam.
F. (IV). Series Hispidae McNeill : No material studied
F. (V). Series Orientales
A. retusa : Druses, slightly dense, scattered throughout the leaf except in
the veins, spheroidal, margins irregular, with sharp points; size 10-49 pm
162
diam.
A. rhodia : Similar to A. retusa .
G. Sectio Pseudosabulina McNeill
Distribution: This single species of Section Pseudosabulina is confined to
the semidesert region on the borders of Turkey, Syria and Iraq.
A. sabulinea : Druses, very dense, scattered throughout the leaf except in
the veins , spheroidal , margins irregular, with sharp points; size 19-58 pm
diam.
H. Sectio Occidentales McNeill
Distribution: Occours in west Mediterranean.
A. loscosii: Druses, mostly few, scattered throught the leaf except in the
veins, concentrated near the upper third of lamina, spheroidal, margins
regular, with blunt points; size variable though mostly large, 19-145.5 pm
diam.
A. ciliaris : Druses Similar to those of A. loscosii, except size 19-78 pm
diam.
J. Sectio Africanae McNeill
Distribution: Comprises a small group of Spanish and North African plants
which are very uniform in habit and general appearance .
J. (i). Series Africanae
A. cerastioides : Druses, dense, scattered throughout the leaf except in the
veins, spheroidal, margins irregular with sharp points; size 10-78 pm diam.
J. (ii). Series Papillospermae McNeill
A. hispanica : Druses, dense, distributed irregularly throughout the leaf
area except in the veins, margins irregular with sharp points; size 10-78 pm
163
diam.
K. Sectio Arenaria
Distribution: Section Arenaria is a fairly homogeneous group of annual
plants, two of which (A serpyllifolia & A. leptoclados) are very widespread
throughout Eurasia. The remaining species are more localised, five
occuring in the Eastern Mediterranean area, one in North America and one
in Spain .
K. (i). Series Arenaria
A. conferta : Druses, dense, scattered throughout the leaf, except in the
veins, spheroidal, margins irregular with a very sharp points , size 19-97
pm diam ..
A. leptoclados : Similar to A. conferta .
A. serpyllifolia:
,,
K. (ii). Series Saponarioides McNeill
A. saponarioides : Druses, dense, scattered throughout the leaf except in
the veins, spheroidal, margins irregular with sharp points; size variable 1097pm diam.
K. (iii) Series Cylindricae McNeill
A. guicciardii. Druses, dense, scattered throughout the leaf except in the
veins, spheroidal, margins irregular, with sharp points, size 10-78 pm diam.
L. Sectio Compressae McNeill
Distribution: An extremely distinctive monotypic section from the western
Himalayas and Afghanistan.
A. compressa : No material studied.
V. Subgenus Arenariastrum Williams
164
Distribution: The subgenus is confined to small area in the South of France.
A. gouffeia : Druses, dense, scattering in whole leaf except onveins,
margins irregular, with sharp points, size 19-78 pm diam. .
VI. Subgenus Eremogoneastrum Williams
Distribution: North America and Sino-Himalaya
A. franklinii: Druses, dense, scattered irregularly throughout the leaf except
in veins, margins irregular with blunt points, size 10-39 pm diam.
A. hookeri: Druses, dense, scattered irregularly throughout the leaf except
in veins, margins mostly regular with blunt points, size 10-29 pm diam. .
A. festucoides: Similar to A. hookeri.
A. kansuensis : Similar to A. hookeri .
VII. Subgenus Eremogone (Fenzl) Fenzl in Ledebour
Distribution: Its main centres of diversity are the mountains ofCentral and
South-West Asia and Western North America .
A. Sectio Capillares McNeill
A. capillaris : Druses, scattered irregularly, mostly dense in the lower third
near leaf base, or scattered in or along leaf veins, spheroidal, margins
mostly regular with blunt points; size variable 10-29pm diam.
A. fendleri : Druses or spheroidal masses of crysatal sands, very dense,
scattered irregularly or in regular short rows in or along veins, mostly
spheroidal, margins regular with blunt points; size 10-39 pm diam.
A. formosa : Similar to A. capillaris , the upper half of the leaf with absent or
very rare crystals.
A.lychnidea : Druses, very dense particularly near the leaf base scattered
irregularly, or spheroidal masses of crystal sands, mostly few in the upper
165
part of the leaf, concentrated in or along veins, spheroidal, margins regular
with blunt points; size variable mostly small 19-29 pm diam.
B. Sectio Monogone Maxim.
Distribution: A monotypic section endemic to the Tien Shan and Altai
regions. The single species, A. potaninii has not been studies.
C. Sectio Eremogone
Distribution: From Central and Eastern Europe across to Central Asia and
extends south into the Caucasus, Armenia and probably S. Turkey .
A. steveniana : Druses or spheroidal mases of crystal sand, dense,
arranged in regular rows or scattered irregularly mostly in veins, with
regular margins and blunt points; size variable, 10-49 pm diam..Plate 36
(3,4).
A. graminea : Druses or spheroidal masses of crystal sand, very dense
arranged regularly in rows or irregularly in or along veins, round, regular
with blunt points; size variable 10-49 pm diam.
A. macradenia : Druses or spheroidal masses of crystal sand, very dense
over whole leaf though concentrated mainly in the leaf veins, scattered
irregularly or in short rows, mostly spheroidal with blunt points; size variable
10-29 pm diam.
D. Sectio Glomeriflorae Fenzl ex Williams
Distribution: The Section is confined to the Caucasus, E. Turkey and Iran .
A. dianthoides : Spheroidal masses of crystal sand, somewhat dense,
arranged in regular rows only in veins, no crystals in the intercostal area,
mostly spheroidal or ellipsoidal or short elongate,with regular margins,
spheroidal size mostly small 10-19 pm diam.
166
A. cucubaloides : Similar to A. dianthoides, but mostly spheroidal masses of
crystal sand; size 19-49 pm diam., Plate 32 (4).
A. gypsophiloides : Spheroidal masses of crystal sand or ellipsoidal,
regular in rows or slightly irregular in the veins, shape mixed
mostly
spheroidal or elongate with blunt points; size variable and mixture of
spheroidal 19-78 pm diam., or elongate length c. 58 pm , width c. 19 pm ,
intercostal area filled with spheroidal druses of small size 10-19 pm diam.
E. Sectio Rigidae McNeill
Distribution: The main centre of distribution in E. Europe, S. Russia and the
Caucasus.
E. (i). Series Rigidae
A. holostea : Spheroidal masses of crystal sand, dense, arranged
in
regular rows from leaf base to the top in the veins, with regluar margins
and blunt points; size mostly 19-49 pm diam.
A. szowitsii : Similar to A. holostea
E. (ii). Series Setaceae McNeill
A. angustisepala : Druses, mostly very dense near leaf base with sharp or
blunt points; druses or spheroidal masses of crystal sand in the upper side
arranged along or in the veins with round and blunt points, size 10-29 pm
diam.
F.Sectio Scariosae McNeill
Distribution: In North Iran and Turkish Armenia, but absent from the
Caucasus .
F. (i). Series Polycnemifoliae McNeill
A. polycnemifolia: Druses,dense near leaf base, mostly scattered, irregular;
size small c. 10 pm diam; spheroidal masses of crystal sand with regular
167
margins and blunt points in the upper side arranged in short regular rows,
they joined 3-5 crystals in each row in the veins with variable shapes but
mostly spheroidal 19-58 pm diam. or ellipsoidal length c. 58 pm and width
c. 39 pm, or elongate c.19 pm length .
A. pseudacantholimon : Druses very dense at the base scattered
irregularly, spheroidal with regular margins and blunt points, size c. 19 pm
diam.; few spheroidal masses of crystal sand and druses in general
scattered in or along the veins, rare or no crystals in the upper third of the
leaf; size c. 19 pm diam.
A. zargariana : Druses, very dense near leaf base scattered irregularly or in
short rows, size c. 19 pm diam.; spheroidal masses of crystal sand and
druses, scattered irregualrly or in short rows in the upper side of the leaf in
or along the veins, mostly spheroidal with blunt points,size c. 49 pm diam.,
or ellipsoidal, size c. 58 pm length, 29 pm width.
F. (ii). Series Scariosae
A. armeniaca : Druses, near leaf base mostly scattered irregularly,
spheroidal with blunt or sharp points; spheroidal masses of crystal sand,
ellipsoidal or elongate,arranged irregularly or arranged in regular short with
small mostly c. 10 pm diam., crystals on the upper side of the leaf
concentrated in main veins scattered irregularly, or arranged in short rows,
spheroidal, 10-58 pm diam, or elongate to ellipsoidal, length c. 34 pm,
length, width c. 19 pm with regular margins and blunt points.
A. scariosa ; Druses concentrated at leaf base, mostly spheroidal with blunt
margins; in the upper side of the leaf crystals scattered irregularly or
arranged in short
regular rows in or along veins, mostly spheroidal
masses of crystal sand with regular margins and blunt points, size c. 49 pm
diam., or elongate length c. 78 pm, width 49 pm
168
G. Sectio Sclerophyllae (Boiss.) McNeill
Distribution: Occurs in two disjunct areas, the steppes of Central and SouthWest Asia and the dry regions of the Westren United States .
A. acerosa: Druses, very dense at the leaf base, with sharp points, small
size c. 10 pm diam.; spheroidal masses of crystal sand or druses, mostly
with blunt points scattered or arranged in short rows in or along the veins;
size 10-39 pm diam. .
A. acutisepala
: Druses, dense, scattered irregularly at the leaf base,
mostly small with sharp points; spheroidal masses of crystal sand or druses
on the upper side, in regular short rows mostly c. 4-8 crystals in each row
arranged in or along the veins, with blunt points; size mostly large 10-39 pm
diam. Plate 33 (1, 2).
A. davisii : Druses or spheroidal masses of crystal sand, very dense,
scattered irreguarly throughout the leaf in or along veins with the same
density from base to the top, with regular margins and blunt points; size
10-49 pm diam.
A. drypidea : Druses, dense near the leaf base with sharp points, size IQ19 pm diam.; spheroidal crystal sand masses arranged in more longer
regular and dense rows in veins; size 10-78 pm diam.
A. griffithii: Druses, very dense near leaf base with sharp points, size mostly
small, c. 10 pm diam.; spheroidal masses of crystal sand
and druses
arranged in or along the vein with blunt points; size 10-78 pm diam.
A.insignis : Druses and spheroidal masses of crystal sand, dense scattered
irregularly throughout the leaf mostly with the same density along veins
rarely in the veins, spheroidal with regular margins and blunt points; size
10-29 pm diam. .
A. kingii : Similar to A. insganica\ size 10-49 pm diam., (Plate 33,1,2).
A. ledebouriana : Similar to A. acutisepala ; size 19-58 pm diam.
169
A. aculeata: Similar to A. kingii.
A. macradenia : Spheroidal masses of crystal sand and druses, very dense
scattered irregularly throughout the leaf mostly with the same density
concentrated
mostly in veins, with regular margins and blunt points; size
10-49 pm diam.
A. persica : Spheroidal masses of crystal sand and druses, dense scattered
irregularly throughout the leaf with the same density from base to top, with
regular margins and blunt points; sizes variablie 19-58 pm diam. .
A. tetrasticha : Similar to A. persica ; size 10-49 pm diam.
H. Sectio Pungentes McNeill
Distribution: Spain and North Africa, is a very isolated group whose nearst
affinities are not appearent.
A. pungens : Druses and spheroidal masses of crystal sand, very dense,
scattered irregularly throughout the leaf, size very variable, spheroidal,
margins regular with sharp or blunt points, mostly medium size 19-87 pm
diam., Plate 33 (6).
VIII. Subgenus Dolophragma (Fenzl) McNeill
Distribution: The subgenus is probably made up of seven species, all in the
sino-Himalya region .
A. denissima : No crystals observed in the leaves.
A. oreophila : Druses, very rare, occasionally found in small aggregated
groups near leaf base with small size c. 10 pm diam. with irregular margins
and sharp points, or in the upper side, with variable sizes 10-49 pm diam.
A. polytrichioides : Similar to A. oreophila .
170
IX. Subgenus Solitaria McNeill
Distribution: In the eastern Himalya and the mountains of South-Western
C hina.
A.ciliolata : Druses, very few in number c. 40-100, scattered irregularly
throughout the leaf except the veins, spheroidal with regular margins
slightly sharp or blunt point; size mostly large 10-78 pm diam ..
A. forrestii: Similar to A. ciliolata.
A.napuligera: Druses, dense, scattered irregularly throughout the leaf
except in veins, spheroidal, margins irregular with very sharp points; size
19-87 pm diam.
A. trichophora: Similar to A. napuligera ; size variable; 10-97 pm diam.
A yunnanensis: „
„
„
„
,,
19-68 pm diam.
4.2.2 M oehringia L.
A. Sectio Pseudomoehringia McNeil
Distribution: Spain and Norht Africa.
M. intricata : Druses, very dense, scattered irregularly throughout the leaf
except in veins, spheroidal, with regular margins and blunt points; size 1997 pm diam.
M. tejedensis : Druses, dense, scattered irregularly throughout the leaf
except in the veins, spheroidal, mostly with blunt margin, size variable but
mostly medium or big 19-49 pm diam.
B. Sectio Latifoliae Nyman ex Graebner
M. trinervia : Druses, dense, scattered irregularly throughout the leaf
except in veins,
spheroidal, with sharp points and regular margin; size
variable, mostly medium or large 19-97 pm diam., Plate 36 (2).
M. lateriflora : Druses, dense, scattered irregularly throughout the leaf
171
except in the veins, spheroidal with regular margins and sharp points; size
variable mostly medium or big 10-49 pm diam.
M. radiolata : Druses,dense, scattered throughout the leaf except in the
veins, spheroidal, with irregular margins and sharp points; size mostly
medium or large 10-49 pm diam.
M. stellarioides: Similar to M. trinervia, size mostly medium c. 58 pm diam.
Plate 32 (1).
C. Sectio Diversifoliae Nyman ex Graebner
M. diversifolia : Druses, dense, scattered irregularly throughout the leaf
except in the veins, spheroidal, with irregular margin and sharp points, size
mostly small, 10-19 pm diam. .
M. jankae : Druses, dense, scattered irregularly throughout the leaf except
in the veins, spheroidal with irregular margins and sharp points; size mostly
large or medium, 19-97 pm diam.
M. pendula : Similar to M. diversifolia .
D. Sectio Moehringia :
M. sedifolia: Druses, very dense, scattered irregularly concentrated
particularly along veins, spheroidal, with irregular margins and blunt points,
size meduim or large 19-68 pm diam. .
M. ciliata : Druses, dense, scattered irregularly throughout the leaf
particularly along main veins, spheroidal with irregular margins and sharp
points; size mostly medium or large, 19-68 pm diam.
M. glaucovirens : Druses, very rare or without crystals, scattered irregularly
throughout the leaf mostly concentrated along main vein, spheroidal, with
irregualr margins and sharp points; size mostly big 19-39 pm diam.
M. muscosa : Druses, dense, scattered irregularly throughout the leaf,
172
concentrated particularly along main vein, spheroidal, mostly with regular
margins and blunt points;size mixed, 19-68 pm diam.
M. tommasinii: Druses, very dense, scattered irregularly throughout the
main vein, spheroidal, with regular margins and blunt points; size 10-78 pm
diam.
4 .2 .3 Minuartia L.
I. Subgenus Rhodalsine: (J. Gay) Graebner
Distribution: It is common throughout the more southerly Mediterranean
coasts extending to Portugal and the Canary Islands and with a distinctive
species in Somaliland .
M. geniculata : Druses, dense, scattered irregularly throughout the leaf,
except in the veins, with different shapes but mostly round, margins
irregular, with sharp or blunt points, size variable, 10-97 pm diam., Plate 34
(3).
II. Subgenus Spergella
Distribution: Very distinctive groups of two sympatric species in the IranoTuranian Sharo-Sindian regions of the Levant, Southern Turkey and Iraq .
M. formosa : Druses not dense scattered irregularly or arranged, in regular
rows along main veins, shape mostly round, with regular margins and blunt
points; size 19-78 pm diam.
M. picta : Druses c. 97 pm diam. or spheroidal masses of crystal sand but
mostly elongate length 49-194 pm , width 29-78 pm.
III.Subgenus Hymemella (Moc. & Sesse ex Ser.) McNeill
Distribution: is only known from Central Mexico .
M.moehringioides : Druses dense, scattered irregularly throughout the leaf
173
except in veins, round with regular margins and blunt points, size 10-49 pm
diam.
IV. Subgenus Minuartia
A.Sectio Spectabiles (Fenzl) Hayek
Distribution: On the mountains of Eurasia and throughout the Arctic .
A. a. Subsectio Spectabiles
A. a. (i) Series Laricinae
Distribution: Widely distributed in Western North Ameriaca and Arctic Asia,
extending south to japan .
M. colchica : Druses, very dense, scattered irregularly throughout the leaf
except in the veins, mostly spheroidal, with irregularly margins and sharp
points, size 10-49 pm diam.
M. imbricata : Similar to M. colchica\ size 10-29 pm diam.
M. inamoena:
,,
A. a. (ii). Series Spectabiles
[new series Biflorae of Rabeler, 1993]
Distribution: A series of three species throughout Arctic Eurasia and
extending southwards into the western United States and Central Asia (to
the Himalayas).
M. arctica : Druses, dense, scattered irregularly throughout the leaf except,
veins, spheroidal, with regular margin and blunt points; size 10-49 pm
diam.
M. biflora : Druses, slightly dense, scattered irregularly throughout the leaf
except in the veins, spheroidal, margins mostly regular with blunt points;
size mostly small 10-29 pm diam.
M. obtusiloba : Druses, few, scattered
irregular particularly along both
sides of the veins, spheroidal, with regular margins and blunt points; size
variable 10-39 pm diam.
174
A. b. Subsectio Cherleria (L.) McNeill
Distribution: A monotypic group restricted to the mountains of Central
Europe and the Scottish Highlands .
M. sedoides : Druses, not dense, scattered irregularly, mostly arranged
along both sides of veins, mostly spheroidal, with regular margins and
sharp points ; size 19-49 pm diam.
A. c. Subsectio Laricifoliae (Mattf.) McNeill
A.c. (i). Series Caucasicae Mattf. in Fedde Rep. Beih.
Distribution: A series of two species in western Anatolia and the Caucasus .
M. aizoides : Druses, dense, scattered irregularly throughout the leaf
except in the veins, mostly spheroidal, with irregular margins and blunt or
sharp points; size mostly small, 10-29 pm diam ..
A. c. (ii). Series Laricifoliae
Distribution: On the mountains of Southern Europe, extending into N. W.
Anatolia and in the Lebanon mountains .
M. baldaccii : Druses, and spheroidal masses of crystal sand, dense,
scattered irregularly throughout the leaf including veins, round with regular
margins, sharp or blunt points; size 10-49 pm diam.
M. capillacea : Similar to M. baklaccii
M. laricifolia :
,,
„
,, ; size 10-29 pm diam.
B. Sectio Plurinerviae McNeill
Distribution: On the mountain of Central and Southern Europe and S. W.
Asia.
M. bulgarica : Spheroidal masses of crystal sand, arranged in 4-5 regular
rows in each vein, no crystals in intercostal area, mostly spheroidal, with
entire margin; size mostly uniform on veins, c. 19 pm diam., Plate 33 (8),
175
Plate 36 ( 6, 7, 8).
M. hirsuta : Similar to M. bulgarica .
M. recurva : Similar to M. bulgarica, but the intercostal area filled with
various crystals, shape mostly elongate length c. 97 pm , or spheroidal, c.
39 pm diam. .
C. Sectio Lanceolatae (Fenzl) Graebner in Ascherson & Graebner
C. (i). Series Graminifoliae Mattf.
Distribution: A series of three species in South-Eastern Europe (centred in
the Balans) extending into N.W. Anatolia .
M. graminifolia : Spheroidal masses of crystal sand and elongate, arranged
in the veins each vein has c. 4-5 rows, spheroidal c. 49 pm diam., elongate
length c.78 pm , width c. 49 pm , the intercostal area covered with mixed
crystals have different shapes mostly elongate length c. 19 pm .
M. saxifraga : Spheroidal masses of crystal sand and druses, arranged in
seven veins, several rows of regular crystals arranged along each vein,
spheroidal or ellipsoidal, mostly small c. 19 pm diam. ; intercostal area
filled with mixed crystals mostly small, spheroidal Plate 35 (4).
M. stellate : Smilar to M. saxifraga .
C. (ii). Series Dianthifolia
M. dianthifolia: Spheroidal masses of crystal sand, arranged in c. 7 veins in
regular rows with regular margins, mostly spheroidal and small size c. 10
pm diam., intercostal area filled with small druses variable shapes and
sizes, mostly regular wih blunt or sligthly sharp points.
M. acuminata: Spheroidal masses of crystal sand, arranged regularly in
c.10 veins in regular rows, spheroidal, with regular margins and blunt
points
mostly the same size c. 10 pm ; intercostal area covered with of
variable sizes, mostly druses with blunt points.
176
M. pestalozzae: Spheroidal masses of crystal sand, arranged in c. 10 veins
in regular rows and regular shape mostly ellipsoidal, size 10-29 pm diam.;
intercostal area filled with a very dense small crystals of variable shape and
size, mostly spheroidal and ellipsoidal (Plate 35, 8).
C. (iii). Series Lanceolatae
[new name series Cerastifoliae]
Distribution: Restricted distribution in Asia Minor, Nakhichevan & Northern
Iran.
M. cerastiifolia : Spheroidal masses of crystal sand and elongate crystals,
arranged in the three veins, c.7 regular rows in each vein, mostly elongate
length c. 49 pm , width c. 29 pm , or ellipsoidal c. 29 pm length, intercostal
area filled with small mostly spheroidal crystals 10-29 pm diam., Plate 36
(5).
M. rupestris: Similar to M. cerastifolia .
C. (iv). Series Grigneenses
Distribution: A monotypic series endemic to the Bergamo Alps in Northern
Lombardy.
M. grigneensis: Material not studied.
D. Sectio Aretioideae (Fenzl) Mattf.
Distribution: A section of one species, endmic in the Alpas.
M. aretioides : Spheroidal masses of crystal sand, arranged in c. 4-5 rows
in each vein, mostly spheroidal, small 10-110 pm diam. or ellipsoidal; no
crystals in the intercostal area.
E. Sectio Sclerophylla Mattf.
Distribution: In eastern and western North America (Chiefly U.S.A.) .
M. crotiniana : Druses,very rare, mostly concentrated near the lower half of
the leaf, scattered irregularly in the intercostal area but not in veins,
177
spheroidal, with regular margins and blunt points; size c. 19 pm diam.
M. dawsonensis : Spheroidal masses of crystal sand, arranged regularly in
the middle vein only,
c. 4 rows, mostly spheroidal, c. 49 pm diam., or
elongate with variable length, length 29-49 pm in , width c. 29 pm , rarely
with crystals spheroidal found in the intercostal area, c. 49 pm diam.
F. Sectio Acutiflorae (Fenzl) Hayek
Distribution: Throughout Central Europe and Central and South-West Asia .
F. (i). Series Acutiflorae
[new name series Flaccidae]
Distribution: Occuring on the mountains of Central Europe and Central
Asia from the Pyrenees to Himalayas-extending southwards into the steppe
lands of South-West Asia .
M. austriaca : Elongate and spheroidal masses of crystal sand, arranged in
regular rows in three veins mostly elongate length c. 78 pm , width c. 29 pm
; no crystals in the intercostal area.
M. flaccida : Similar to M. austriaca , size length c. 49 pm , width c. 29 pm .
F. (ii). Series Pichleriae Mattf.
Distribution: Throughout Southern and Eastern Anatolia .
M. rimarum : Spheroidal masses of crystal sand, arranged in regular rows
in veins , with regular margin, mostly ellipsoidal length c. 29 pm , c. 19 pm,
width or spheroidal; no crystals in the intercostal area., Plate 33 (7).
F. (iii) Series Umbelluliferae McNeill
M. umbeliulifera : Elongate and spheroidal masses of sand crystals,
arranged in regular rows in three veins, of mixed shapes but mostly
elongate, length c. 19-68 pm , width c. 29 pm ; crystals scattered in the
intercostal area but not dense, mostly elongate .
178
G. Sectio Tryphane (Fenzl) Hayek
Distribution: Throughout Eurasia and North America and extending South
wards onto the mountains of the Mediterranean region .
M. rubella : Spheroidal masses of sand crystal, arranged in regular rows in
three veins, mostly ellipsoidal, length c. 39 pm , width c. 29 pm.
M. verna : Similar to M. rubella, but there are elongate crystals near leaf
base .
H. Sectio Alsinanthe (Fenzl) Graebener
Distribution: Occuring in the Alps and on the mountains of C. Asia.
M. ro ssii: Spheroidal masses of sand crystal, arranged in regular rows in
the middle vein only, mostly spheroidal with regular margins, c. 29 pm
diam.; crystals in the intercostal area variable in shape, mostly spheroidal,
10-49 pm diam.
M. stricta : Similar to M. rossii; but crystals in the intercostal area variable in
shape and size particularly near leaf base, mostly spheroidal, 10-97 pm
diam., or elongate length c. 19 pm, width c. 10 pm Plate 35 (5).
J. Sectio Uninerviae (Fenzl) Mattf.
Distribution: An eastern North American section, extending to Greenland .
M. brevifolia : Druses, scattered irregularly throughout the leaf except in
veins, mostly spheroidal with regular margins and blunt points, c.19 pm
diam.
M. glabra : Druses, very rare, small spheroidal, with regular margin and
blunt points, scattered irregularly in the intercostal area except in the veins,
size c. 19 pm diam.
M. groenlandica : Similar to M. glabra .
M. patula : Elongate and druses, scattered irregularly throughout the leaf,
179
but not in veins, with different shapes and sizes, but mostly elongate length
39-155 pm , width c. 19 pm.
K. Sectio Greniera (Gay) Mattf.
Distribution: A section comprising two annual species confined to western
North America.
M. douglasii: Spheroidal masses of sand crystal and elongate, mostly few,
concentrated in veins in short rows, mostly spheroidal near leaf base and
more elongate in the upper half, length 78-232.8 pm , width c. 29 pm,
crystals very rare in the intercostal area., Plate 35 (7).
M. howellii: Elongate and spheroidal masses of sand crystal, mostly few,
arranged in veins in short rows, mostly elongate,
length c. 0.5-2 mm, width
c. 29 pm.
L. Sectio Minuartia
L. a. Subsectio Minuartia
L. a. (i) Series Montanae Mattf.
Distribution: Centred in the Eastern Mediterranean region but extending to
northern India and occuring in Spain and western N. Africa .
M. montana : Elongate and spheroidal masses of crystal sand, arranged
regularly in three veins c. 5 rows of crystals in each vein, mostly spheroidal
or ellipsoidal, intercostal area mixed mostly elongate, length 39-339.5 pm,
width 29-68 pm ( cigar shape), or ellipsoidal, length c. 39 pm , width c. 29
pm Plate 35 (3), Plate 37 (5,6).
M. globulosa : Spheroidal masses of crystal sand, arranged in many
regular veins, spheroidal c. 19 pm diam., or elongate, 97-175 pm length, c.
29 pm in width, mostly concentrated in the upper half of the leaf;
intercostal
area filled with crystals different shapes and sizes but mostly spheroidal,
180
10-49 (um diam. Plate 34 (4).
L. a. (ii). Series Minuartia
Distribution: All species of this series native in the Mediterranean region .
M. dichotoma : Spheroidal masses of crystal sand and elongate crystals,
arranged regularly in the veins, mostly spheroidal, c. 19 pm diam.,
intercostal area covered with small granules sandy structure Plate 35 (6).
M. hamata : Elongate and spheroidal masses of crystal sand, arranged
regularly in three veins, c. 5 rows of crystals in each vein, mostly spheroidal;
intercostal area covered with crystals mixed shapes, mostly spheroidal or
elongate (cigar shape), length 39-194 pm , width 19-29 pm, Plate 35 (2).
L. b. Subsectio Xeralsine (Fourr.) McNeill
L. b. (i). Series Leucocephalae Mattf.
M. leucocephala : No material studied.
L. b. (ii). Series Setaceae Mattf. in Fedde Rep.
Distribution: Occuring in C.Europe and the Mediterranean region (incl.
Anatolia and the Caucasus).
M. anatolica : Spheroidal and elongate crystals, arranged regularly in three
veins,
c. 5 rows in each vein, mostly spheroidal, c. 39 pm diam., or
ellipsoidal, intercostal area filled
with mixed sizes and shapes mostly
elongate, length c. 78 pm, widthc. 19 pm.
M. confusa : Similar to M. anatolica .
M. mutabilis: Similar to M. anatolica .
M. restrata\
M. setacea:
L. b. (iii). Series Xeralsine
[ new name Series Fasciculata]
Distribution: Usually on the mountains of Southern Europe, extending into
N. A frica.
181
M. fasciculata : Spheroidal masses of crystal sand, arranged in three veins,
with c. 5 rows of crystals on each vein, mostly spheroidal, 10-29 pm diam.,
intercostal area filled with mixed crystals that are mostly spheroidal, or
polygonal.
M. funkii: Similar to M. fasciculata
L. b. (iv). Series Campestres Mattf.
Distribution: A series of one annual species in Spain and North Africa .
M. campestris : Spheroidal masses of crystal sand and druses, arranged in
three veins, with c. 3 rows of crystals on each vein, mostly round; intercostal
area filied with mixed crystals that are mostly spheroidal or druses Plate 34
( 1,2).
M. Sectio Sabulina (Reichb.) Graebn.
M. (i). Series Sabulina
Distribution: Centred in S. Europe and the Mediterranean area.
M. hybrida: Elongate and spheroidal masses of crystal sand arranged in
three regular veins in the leaf base, crystals mostly elongate, length c. 39
pm , width c. 29 pm , these crystals in veins started joining gradualy from
the base to the top making a long (cigar shape) crystals in the upper half of
the leaf, no crystals in the intercostal area, Plate 34 (5,6).
M. mediterranea : Similar to M. hybrida; crystals in the upper half of the leaf,
length c. 268.6 pm, width c. 68 pm , Plate 34 (7,8), Plate 37 (1,3,4).
M. mesogitana : Elongate and spheroidal masses of crystal sand,
arranged in three veins with variable sizes, mostly elongate, length 49-78
pm , width c. 29 pm , very rare crystals in the intercostal area mostly
spheroidal with blunt points, c. 19 pm diam., Plate 35 (1).
M. tenella : Elongate, spheroidal masses of crystal sand and druses,
arranged in regular rows, crystals mostly elongate from short near the lower
182
half to long in the upper half, length 116-223 pm , width c. 29 pm ; very few
crystals in the intercostal area mostly round with sharp or blunt points, c. 49
pm diam.
M. viscosa: Similar to A. mesogitana
M. urumiensis : Spheroidal masses of crystal sand and elongate crystals,
mainly arranged in three regular rows, with regular shape mostly
spheroidal, c. 19 pm diam., intercostal area covered with few different
shapes but mostly spheroidal, 19-78 pm diam., or elongate, length 49-116
pm , rarely elongate, 194 pm long
M. (ii). Series Californicae Mattf.
Distribution: In California and in Chile .
M. californica : Spheroidal masses of crystal sand, elongate and druses,
crystals mostly arranged in short rows of c.4-6, mostly spheroidal, 19-39
pm diam, attached together, or scattering irregularly, mostly in or along the
main veins .
183
4.3 DISCUSSION
4.3.1 ARENARIA L.
McNeill (1962, p.89) stated "In Arenaria itself very similar species such as
A. lychidea and A. capillaris (treated as conspecific by Regal, 1862) have
minute and prominent glands respectively, and Williams consequently
places the one in subgenus Euarenaria and the other in subgenus
Pentadenaria." In his classification McNeill listed these two species under
one section Capillares. As found during this investigation, both species
show great similarities in crystal shape, margin, size, density and
distribution. This agrees
with the morphological characters to keep them both under one section.
McNeill (1962,p.89) added "In the same way A.cucubaloides and A.
gypsophiloides, which are satisfactorily distinguished only by flower size
and the length of the leaf sheath, are widely separated from one another in
his [Regel] classification." McNeill kept these two species under one section
Glomerifiorae. Both these two species show differences in crystal size and
distribution. In A. cucubaloides the crystals are arranged in regular rows in
the veins with no crystals only in the intercostal areas, and the size is mostly
small ( c. 10-19
pm diam.) and the shape round. However in A.
gypsophiloides the crystals are round and larger (c.19-78 pm diam.) or
elongate length (c. 58-116 pm in, (width c. 19 pm) and the intercostal area
contains dense small crystals (c. 10-19 pm). Therefore this crystal evidence
supports Regalel’s (1862)
placement of these two species of different
subgenera.
Arenaria sabulinea is the sole species in section Pseudosabulina. McNeill
(1962, p.115) said "The section shows a strong superficial resemblance to
Minuartia Section Sabulina (particularly in the sepal structure) but its true
affinities lie with Arenaria Section Orientates Series Orientates (particualrly
184
A. kurdica)." Davis (1967,p.28) in Flora of Turkey also mentioned the
superficial resemblance of A. sabulinea to M inuartia Sect. Sabulina,
especially M. mesogitana. According to the crystal studies there is no
resemblance between these two taxa. In all the species of Section
Sabulina of Minuartia the crystals are arranged in regular rows in the veins
(c. 49 pm in length, c. 78 pm in width); whereas in M. mesogitana the
crystals are mainly arranged in the three veins and are variable in size,
(mostly 49-78 pm in length, c. 29 pm in width). There are also crystals of
round shape ( c. 19 pm diam.) in the intercostal area. In A. sabulina
however the crystals are scattered throughout the leaf except in the veins.
The shape and distribution of the crystals point to a very close relationship
between A. sabulina and the species in Series Orientales Section
Orientates as claimed on morphological ground in by McNeill with
particular A. kurdica.
McNeill (1962, p.125) said" The Section [Rigidae] is closely related to
section Capillares with which it was linked by Fenzl in Ledebour (1842)
(both in his "Chromolemmae") and from which it has probably evolved by
the sepals becoming more hardened, coriaceous, scarious, and acuminate.
In the Series Setaceae it shows some affinity with Sections Sclerophyllae
and Scariosae." According to the crystal study, Series Rigidae shows
crystals in very regular rows arranged in the veins all the way from the base
to the top {A. szowitsii, A. holostea ). This is different from section Capillares
with its dense irregularly scattered crystals in or along the veins, with
greatest density near the lamina base.
Section Capillares resembles
Series Setaceae in Section Rigidae particularly the section Scariosa of
section Sclerophylla with regard to crystal shape and distribution. McNeill
(1962) explained that, in Section Africanae with its small group of Spanish
and North African plants, Series Africanae has seeds of the type that is
185
normally found in Subgenus Arenaria. But A. hispanica, in Series
Papillospermae, has a seed testa structure showing strong similarities with
Subgenus Leiosperma of Arenaria and genus Moehringia. Because of its
distinctive seeds A. hispanica stands in a very isolated position. The crystal
shape, margins and distribution show marked similarities between the two
series of Section Africanae and many species in Subgenus Leiosperma
and in Moehringia. Using only the crystal evidence Series Africanae and
Papillospermae would be amalgamated.
However, in A. hispanica, with its resemblance in vegetative and floral
characters to Section Africanae suggests that it would be better to include
A. hispanica as a separate Series within Section Africanae.
McNeill (1962, p.119) said "The subgenus [Arenriastrum] is confined to a
small area in the South of France; it appears to be a bicarpillary derivative
of Subgenus Arenaria and in habit and sepal structure closely resembles
such species as A. sabulinea and A. capiliipes. The crystals results show
no difference in type, shape, margins and size
between A. gouffeia
(subgenus Arenriastrum), and A. sabulinea (Subgenus Arenaria).
According to McNeill (1962, p.123) " The geographically isolated
Caucasian A. lychnidea [ Section Capillares] is distinctive among Eurasian
plants in its very weakly developed staminal glands and in its sepal
structure shows an approach to Section Glomeriflorae". The Crystals show
a big difference between A. lychnidea (with its very dense crystals
concentrated irregularly in the lamina base and mostly without crystals in
the upper half) and the species in the in Section Glomeriflorae in which the
crystals are arranged regularly in rows in the veins only. Here the crystal
evidence argues against the inclusion of A. lychnidea
Glomeriflorae.
in Section
186
McNeill (1962, p.123) added " In North America, in the Colorado-New
Mexico region, there are two species (A. eastwoodii and A. fendleri) with
very narrow pointed sepals; this characteristic is also found in a Tibetan
species (A. acicularis), and makes these species readily distinguishable
from the rest of the section. A. acicularis was not studied.
McNeill (1962, p. 126) claimed th a t" the species [in Section Sclerophyllae]
have been very confused taxonomically, and like Section Capillares but
unlike most groups of subgenus Eremogone, its members seem to be in a
state of active evolution and speciation". The crystals of 12 species of
Section Sclerophyllae were studied. According to their shape, size and
distribution two groups can be recognised; the first group A.acerosa, A.
acutisepala, A. drypidea, A. griffithii, A. kingii, A. ledebouriana, and
A.tetrasticha. has dense crystals concentrated and scattered irregularly
near the leaf base, mostly small in size (c. 10 pm diam.) with sharp or blunt
margins. It also has short regular crystals, round with blunt margins and
variable sizes (10-97 pm diam.) arranged in veins throughout the leaf.
The second group A. darisii, A. insignis, A. persica, and A.
macradenia . has dense or very dense crystals scattered irregularly
throughout the leaf mostly concentrated along the veins, with round blunt
margins and variable sizes (10-58 pm diam.) eg.
McNeill(1962, p.127)
added" the small and not very spiny-leaved A. tetrasticha and A. davisii (S.
Iran and E. Turkey) perhaps form a third group to be separated from the
main aggregation of rather spiny caespitose or suffruticose species. The
results obtained from the crystals separate those two species. A. tetrasticha
falls into group one and A. davisii into group two .
McNeill (1962, p. 127) wrote " Section Pungentes, confined to Spain
and North Africa, is a very isolated group whose nearest affinites are not
apparent. The type species has usually been placed in Eremogone and
187
because of its semi-terete pungent leaves and superficial resemblance to
A. stenomeres and A. persica, it is retained for the present in that subgenus;
it differs however from the members of the other sections in that its
cotyledons are incumbent and not accumbent. It is even more distant from
Subgenus Arenaria (in which cotyledons are always incumbent) and with
more evidence may come to be placed in a subgenus of its own". Many
specimens from different localities in Morocco were checked. This revealed
the crystals to be very dense and scattered irregularly througout the leaf
area, mostly round with blunt or sharp margins and sizes variable (19-87
pm diam.). The crystals in density, shape, size and distribution in A.
pungens
give a good support for McNeill's suggestion that A. pungens
should placed in a separate subgenus.
4.3.2MOEHRINGIA L.
The species in Moehringia can be classified into different groups according
to density of the crystals: very dense e.g. M. bavarica , dense e.g. M. ciliata,
few or absent crystals e.g. M. glaucovirens.
They can also be grouped by size, small ( 10-19 pm diam.) eg. M.
diversifoliae, medium (10-49 pm diam.) eg. M. radiolata and big (19-97 pm
diam.) e.g. M. trinervia. They can also be grouped according to margins:
with points sharp eg. M. trinervia or blunt e.g. M. muscosa, and according
to crystal distribution: throughout the leaf area except in veins M. pentandra
or including veins e.g. M. tommasimii.
All members of Moehringia show great similarities with the genus
Arenaria except the Subgenus Eremogone of Arenaria, and also display
great similarities with Minuartia except Subgenus Minuartia but within that
Subgenus the Sections Spectabiles, Sclerophylla and Uninerviae do
resemble Moehringia.
188
4.3.3MINUARTIA L.
McNeill (1962, p. 136) wrote" The distinctive facies of the single species of
this subgenus [Hymenella] makes its retention within Minuartia open to
question but in the absence of any more definite characters than the
quadrangular stem and the spreading calyx and capsule, it has been
thought wisest to follow Mattfeild's treatment, but raising it to subgeneric
rank. M. moehringioides is only known from Central Mexico. The crystals in
this monotypic subgenus with their round shape, blunt margins, scattered or
arrangment in short rows along veins resemble those of other subgenera.
Therefore the crystals lend no support to the separation of the subgenus
from the genus Minuartia, McNeill (1962, p. 143) stated "Fenzl named two
"species prototyp" for his infrageneric group Tryphane_Alsine recurva and
A. verna. When he divided the group into two Sections Mattfeld chose
recurva as lectotype, ignoring the fact that Boissier, more than 50 years
before, had by excluding that species (and placing it in the Lanceolatae)
effectively typified Tryphane by Alsine verna. A new name is, therefore,
required for this very homogeneous section which inculdes M. recurva and
M. hirsuta; the name Plurinerviae, which has been given (above), refers to
the 5-7 nerved sepals, the major distinguishing feature between this section
and Tryphane s.s. (=Polymechana Mattf.)."
The crystals of three species from Section Plurinerviae were studied and it
was found that the two species M. bulgarica and M. hirsuta have the same
system of crystal shape and distribution; the crystals are arranged in 4-5
rows in each vein but there are no crystals in the intercostal area. However,
in the third one M. recurva, the crystals are basically arranged in regular
rows in the veins and the intercostal area is filled with variable shapes and
sizes, mostly elongate (c. 97 pm in length) or round (c. 39 pm diam.). In
Section Tryphane, two species were investigated (M. verna, M. rubella ) it
189
was seen that their crystal shape and distribution were similar to those of M.
bulgarica and M. hirsuta.. In Section Lanceolatae (M. graminifolia, M.
saxifraga, M. stellata, M. cerastifolia, and M. ruperstris) the crystals were
studied. This revealed strong similarities with M. recurva. According to
these results M. recurva should be retained in Section Lanceolata this
supports the system of Boissier.
According to McNeill (p. 145) " A Section [Sclerophylla] of three to six
species in eastern and western North America (chiefly U.S.A.). The three
species known to Mattfeld were each placed by him in monotypic series.
They are very distinct species and the section appears rather
heterogenous." The crystal shape and distribution between the two species
in this section are quite variable, in M. caroliniana the crystals are scattered
irregularly along veins, are round with blunt margins and in size are mostly
small (c. 19 pm diam.). However, in M. dawsonensis, they are arranged
regularly in only the main vein, have variable shapes and sizes, and are
mostly round (c. 49 pm diam.) or elongate (29-49 pm in length, 29 pm in
width). Therefore, the evidence of the crystals emphasies the heterogeneity
and suggests that splitting may be a reasonable course.
McNeill (1962, p. 146) stated " Mattfeld based the Series Pichleriae on M.
pichleri, a species endemic to the Peloponnese, but it is clear that M.
rimarum which he placed in his Series Flaccidae should be included with it.
M. rimarum which is widely distributed throughout Southern and Eastern
Anatolia shows some resmblance to
M.
um bellulifera
(Series
Umbelluliferae) but differs in the more spreading equally three-nerved
leaves". The arrangement type and size of crystals in series Pichieriae,
Umbellulifera and Acutiflorae are closely similar but M. rimarum of series
Pichieriae shows greater resemblance to series Acutiflorae than to Series
Umbellulifera. In the last named there are intercostal crystals but in M.
190
rimarum and Acutiflorae there are none. The four examined species from
Section Uninerviae, distributed in eastern North America, show strong
similarities
in their crystals to Series Spectabiles in Subsection
Spectabiles,Section Spectabiles. They both have round crystals with blunt
margins scattered irregularly through the leaf except in the veins and they
have seeds mostly
obscurely tuberculate. While in Series Laricinae in
Subsection Spectabiles, Section Spectabiles, the crystals have the same
distribution as in the previous group but with both round and sharp points,
and the seeds have a fimbriate crest on the dorsal ridge. From these
observations, it can be suggested that Section Uninerviae might well be
united with Section Spectabiles as Subsection Uninerviae.
According to McNeill (1962,p. 148) "The type series [Series Minuartia]
comprises the three highly specilised annual members of the genus, all
native in the Mediterranean region. M. hamata with its recurved fruiting
bracts has a very distinctive facies and is often maintained in a separate
monotypic genus (Queria hispanica). In fact, as Mattfeld (1922 p. 69) has
pointed out, it is merely a reduced derivative of Minuartia and is very close
to M. dichotoma the type species of the genus." The crystals in M.
dichotoma and M. hamata show big differences in distribution. They both
have mostly round crystals arranged in regular rows in the veins. The
difference concerns the intercostal areas which are filled with small sandy
particles in M. dichotoma but filled with elongate crystals (39-194 pm in
length, c.
19 pm in width) in M. hamata . These crystal shapes and
distribution in M. hamata resemble those in the species in Series Montanae
in the same Subsecton Minuartia. Therefore, it might be best to place M.
hamata in Series Montanae than retain it in Series Minuartia.
191
Fig 4. Different types of crystal forms and distributions in the genera
Arenaria, Moehringia and Minuartia.
1) Arenaria deflexa showing a more or less uniform high density of crystals
in theintercostal areas but no crystals in the veins.
2) Arenaria ledebouriana showing a high density of druses in the leaf base
and an irregularly spread crystal sand in the midrib and sparse
crystal sand in the intercostal areas.
3) Arenaria lanuginosa showing irregularly scattered druses in the
intercostal areas throughout the leaf.
4) Minuartia hamata showing dense, regularly spaced crystal sand
throughout the veins and crystal sand, elongate crystals and druses
in the intercostal areas.
5) Minuartia hybrida showing elongate crystals spread throughout the veins
without any crystals in the intercosal areas.
6) Moehringia pendula showing druses widely scattered through the leaf
but only in intercostal areas.
I
T
*
*
*
r *♦ *,> r ’'<•*\
^ *L **
' l ***
"
/
f
!
4
8
192
The plates from no 32 to 37 are photomicrograhs and SEMs of crystals in
tissues of mature leaves.
PLATE 32.
1. Moehringia stellarioides x 100
2. Arenaria alsinoides x 100
3. „
„
x400
4. A. cucabaloides x 100
5.A grandifiora x 100
6. „
„
x400
7. A. montana x 100
8. A. montana x 400.
PLATE 32
PALTE 33.
1. Arenaria kingii x 100. (near the base)
2.
„
„
x 400. (in the middle)
3. A. paiudicola x 100.
4. „
„
x 160.
5. „
„
x400.
6. A. pungens x 100.
7. Minuartia rimarum x 100.
8. M. bulgarica x 160.
P LA TE 33
194
PLATE 34.
1.Minuartia campestris x 100.
2.
„
„
x 100.
3. M. geniculata x 100.
4. M.
„
x 100.
5. M. hybrida x 100 (near the base).
6. „
„
x 100 (in the upper half).
7. M. mediterranea x 100.
8. M.
„
x 400.
P LATE 34
•VI*
*
PLATE 35.
1. Minuartia mesogitana x 100.
2. M. hamatax 100.
3. M. montana x 100.
4. M. saxifraga x 100.
5. M. stricta x 100 (near the base).
6. M. dichotoma x 100.
7. M. douglasiix 100.
8. M. pestalozzae x 100.
PLA TE 35
0, m• mw+ * + 4 ^ i»
"•**•*** •
jnNrm p-
Otf
> *
« „,
dpMfc . . * » * » * •* *^ - ^
• a » »- * * *
&K&*4*£$3
*
*
« ♦
*
2
4
\'V
PLATE 36.
SEMs of the different types of calcium oxalate crystals in Arenaria,
Moehringia and Minuartia as following.
1. Arenaria alsinoides.
2. Moehringia trinervia.
3. Arenaria steveniana.
4.
„
„
5. Minuartia cerastifolia.
6. M. bulgarica.
PLATE 36
197
PLATE 37.
SEMs of the different types of crystals of calcium oxalate crystals in
Minuartia .
1. Minuartia mediterranea.
2. M. tenetla
3. M. mediterranea.
4. „
,,
5. Minuartia montana.
PLATE 37
198
Chapter 5
Capsule and Nutlets
5.1 Introduction
The work presented here which concerns the epidermal morphology of
capsules and of nutlets has taxonomic significance in each of the three
subfamilies. The capsules of the Arenaria serpyllifolia group have already
been discussed in section 3.3. Arenaria leptocaldos can be distinguished
from Arenaria serpyllifolia sensu stricto on the basis of papillae on the
capsular teeth, Plate 47 (1,2). However, there are no detailed treatments
given of the other studies of Silene, Spergula, Spergularia, Polycarpaea,
Polycarpon, Herniaria, Paronychia and Minuartia.
5.2 Si lene
The results are to be considered provisional especially with regard to the
very large genus Silene for which the superglue technique (Chapter two)
was used. In many of the Silene species SEMs were also made and can be
compared with the impressions, Plates 38 to 44. In all, 66 Mediterranean
species of the genus were studied. According to Greuter etal. (1984) there
are 60 species of the genus in North Africa.
These 66 species have been arranged into 13 groups according to the
shape, size and ornamentation of the cells from the middle of the capsule,
as shown in the Plates. The Plates reveal a very considerable diversity of
these features. Some species have smooth cells while others have very
elaborate ornamentation; see particularly S. colirosa, Plate 44 (1,4).
Group A. Epidermal cells, thin walled, mostly elongate with tapering ends,
50-250 pm long, about 20 pm wide, smooth. S. acaulis (Plate 38).
199
Group B. Epidermal cells, thin walled, assymmetricaly polygonal-oval, 90180 pm long and 40-100 pm wide, lacking papillae, arranged irregularly.
S. aegyptica, S.fruticosa, S. heterodonta, S. ibosii, S. otites, S. littorea, S.
niceensis, S.marmarica,S. pseudontiocion, S. sedoides, S. succulenta,S.
villosa , S. vulgaris (Plate 38). S. vulgaris is very variable in features of the
epidermal cells and same specimens can be placed in groups D, G and H.
Group C. Epidermal cells, thin walled, sharply distinct, mostly irregularly
pentagonal and hexagonal, more or less isodiametric to about twice as
long as wide, each cell raised into distinct papillae. S. armeria, S. atlantica
(Plate 38).
Group D. Epidermal cells, appearing imbricated, mostly regularly ovate,
with or without necks, faintly papillate at the swollen base. S. alba, S.
dioica, S. latifolia, S. vulgaris (Plate 39)
Group E. Epidermal cells narrow, elongate with long necks, faintly
papillate. S. noctiflora, S. nutans (Plate 39).
Group F. Epidermal cells, regular mostly narrowly elongate, of different
sizes, one small faint papilla at one end of each cell. There are also
irregularly scattered papillae of varied size and with rough surfaces (Plate
40). Encountered in specimens from Spain, Algeria, Libya and Saudi
Arabia, these papillae are unique to S. conoidea.
Group G. Epidermal cells oval or rectanglar, margins faint,
each cell
raised into a distinct, dome shaped papilla, papillae scattered irregularly. S.
apetala, S. articulata, S. colorata, S. corregata, , S. cyrenaica, S. fuscata,
200
S. nocturna, S. ramosissima, S. rubella, S. secundiflora,
S.sericea, S.
vulgaris (Plate 40).
Group H, Epidermal cells elongate, with distinct or very faint walls,
papillae dome shaped, arranged in regular rows of a few to many
(occasional
single
papillae),
number of rows
10-24 per
1mm.
S.arenaroides, S.behen, S. divaricata, S.glabrescense, S.krem eri,
S.im bricata,
S.
m icropetala,
S.
inaperta,
neglecta,
S .longicaulis,
S. scabriflora,
S.
m ekinensis,
S. stricta,
S.
S.vivianii,
S. vulgaris (Plate 41).
Group I. Similar to group G, but the epidermal cells in sligthly
curved
and raised up into structurs like a barriers which may
be thin or thick, papillae arranged on the top of the barriers,
number of rows 10-16 per 1 mm.
S. boryi,
S. claryi, S. italica, S.
longipetala, S. mollissima, S. pratensis (Plate 41, 42).
Group J. Similar to group H, but the barriers are very thick with
very faint cell walls, this group distinguished from group I by
the
density of irregularly arranged
papillae on each barrier,
number of rows 5-13 per 1 mm.S. bellidifolia, S. cerastoides, S.
disticha, S. gallica, S. ghiavensis, S.muscipula, S. obtusifolia, S.
oropedrium, S. pomelii, S. tridentata, , S. tuberculata (Plate 42,
43).
201
Group
K. Epidermal cells lumped in regular groups of mostly
about 10 or more cells, each group curved and making a regular
barrier, walls between the cells are very distinct, each barrier
cell joined with a small cell and with a very distinct papilla on
the top, number of rows
Group
about 10 per 1 mm. S. conica (Plate 43).
L. Epidermal cells thin walled, pentagonal, most cells
raised up into bunches with different sizes of smooth
S. coiirosa
(Plate 44).
Group M. Epidermal cells characterized
shape,
papillae.
by a specific rod-like
of different sizes, with a papilla at on one end of each
cell.S. laeta, Lychnis
flos-cuculi (Plate 44).
The only species which falls into more than one group of the
above groups is Silene vulgaris (in groups B, D and G). Perhaps
there is no surprise in that because of the notorious variation of
this species, as described in
M arsden-Jones
and
Turrill
numerous publications such as
(1957)
and
the
many
papers
by
Aeschimann.
Appendix V is a dichotomous key to fruiting plants of Libyan
species of Silene', only one species, S. biappendiculata, is lacking
from the key.
202
5.3 Other Genera
Plates 45
shows the
P o ly c a rp o n and two of
r o b b a ir e a .
capsular walls
of six species
of
Polycarpaea, namely P. repens and P.
The great sim ilarity
of the
latter two
is further
evidence for the lack of distinctiveess of the genus R o b b a ria
Plate 46 shows all five species of the genus Spergula and three
of 11 studied species of the genus Spergularia which has 40 in
all. Four of the S p e rg u la species stand out in their cells with
very marked arms but S. viscosa has only slightly wavy cells,
many of which are short. The three species of Spergularia by
contrast have long, narrrow cells with no arms or with only very
slightly wavy walls.
Spergula
arvensis
is
a
very
widespread
species.
Its
variability in habit, pubescence, size of seeds and testa pattern
is mentioned in many Floras such as Tutin et al. (1993) and Davis
(1965). The capsular epidermis is totally distinct from those of
the other species in the transverse fibral-like thickenings. This
feature
was found
in specimens
from
14 different
localities
from Australia to Atlantic islands (Appendix II) and so appear to
be a good specific character.
Spergula
m orisonii and S. pentandra are alike in habit and
are separated on floral and seed characters. They can also be
differentiated
by the epidermal cells which have rounder, less
tapered arms of more equal lengths in S. morisonii than in
203
S. pentandra, Plate 46 (3,4).
In M inuartia subgenus M in u a rtia
it was noticed that the
capsules have delicate but clear cuticular striations in the upper
third of the valves. This feature appears to be confined to that
subgenus and furthermore the striations are not found in those
sections
in which the crystals are only intercostal. Therefore
this correlation is found in sections
A re tio id e a e ,
Minuartia.
P lu rin e rvia e , Lanceolatae,
Sclerophylla, T ryphane, Alisanthe,
The
sole
section
in
which
it
is
Greniera and
not
found
is
S pectabiles.
Plate 47 (3,8) shows five species of P a r o n y c h i a . In all
cases and also in those on the last Plate, 48 the SEMs are of the
upper
ends
of
nutlets.
These
are
very
striking
patterns
of
ornamentation to be seen in both Paronychia and Herniaria. H.
cyrenica, an endemic of Libya (and perhaps
also of Egypt), has
very elaborate features which are very like those of the very
widespread H. glabra.
204
The plates from 38 to 44 are SEMs of capsule wall (midzone) and LMs of
impressions of capsule wall of Silene.
PLATE 38.
1. SEM of Silene acaulis.
2. LM of
„
„ x 400.
3. LM of Silene fruticosa x160.
4. LM of S. niceensis x160.
5. LM of S. succulenta x160.
6. LM of S. villosa x160.
7. LM of S. armeria x160.
8. LM of S. atlantica x160.
PLATE 38
PLATE 39.
1. SEM of Silene dioica.
2
99
99
99
99
3. LM of Silene latifolia x160.
4. LM of S. dioica x160.
5. SEM of Silene nutans.
6*
»>
>1
>>
»»
7. LM of Silene noctiflora x160.
8. LM of S. nutans x160.
PLATE 39
•I*
»
PLATE 40.
1. SEM of Silene conoidea.
2
t- '
9t
ff
ft
3. SEM of Silene colorata.
4
~
m
ft
ft
ff
5. LM of Silene apetala x160.
6. LM of S. colorata x160.
7. LM of S. cyrenaica x160.
8. LM of S. rubella x160.
PLATE 40
t
*
■
I
^
^ i l 'r ^
ii '
\«-v
k
‘
c v v
.
»_w
X
^W w
VC
w ,\ * '
*
4
*
sV* .V wW
v
- '7l t^S >v W
>
^ v 7
V
*
b4r ' • C4w^ k
o * l f ’« >
i
^
4>‘ ^
*
\-*--»> *
,, ?*£Vi*>VlX>Tv'•*
^1
i,V V ^ v
'/
* V * * * V ■ * ' ’ * * v - w' \ ' i , .
* * *
V > ^
* * V*" w *•
«*Vk
CT"
C w^
^
c. ^. . V v
f c ^ ^ ^^ ^ ^ ~
J* V * r , “ v
PLATE 41.
1. SEM of Silene longicaulis.
2
3. LM of
„
, , /x160.
4. LM of S. neglecta x160.
5. SEM of Silene longipetala.
6.
m
7. LM of
ii
n
„
,, x160.
8. LM of S. claryi x160.
PLATE 41
208
PLATE 42.
1. SEM of Silene atlaica .
2
If
If
If
3. LM of S. boryi x160.
4. LM of S. italica x160.
5. LM of S. mollissima x160.
6. LM of S. protensis x160.
7. SEM of Silene pomelii.
Q
n
i>
n
P L A T E 42
209
PLATE 43.
1. SEM of Silene tridentata.
2
£ -■
ft
ft
ft
ft
3. LM of
„
„x160.
4. LM of Silene gallicax 160.
5. SEM of Silene cerastoides,
6. LM of Silene conica x160.
7. SEM
PLATE 43
PLATE 44.
1. SEM of Silene colirosa.
2
*-■
99
99
99
99
3. L M of
„
„ x400.
4.,,
„
„x160.
„
5. SEM of Silene laeta.
6. ft
if
7. LM of
ft
ft
„
„ x400.
8. LM of Lychnis flos-cuculi x400.
PL A T E 44
211
The plates no 45 and 46 LMs of capsule walls showing cell shape and
ornamentation in the midzone areas of Polycarpon andPolycarpaea
PLATE 45.
1. P. arabicum x160.
2. P. bivonae x160.
3. P. depressum x 160.
4. P. peploides x160.
5. P. polycarpoides x160.
6. P. tetraphyllum x160.
7. Polycarpaea repens x160.
8. Polycarpaea robbairea x160.
P L A T E 45
PLATE 46.
1. Spergula arvensis x160.
2. S. fallaxx 160.
3. S. morisonii x160.
4. S. pentandra x160.
5. S. viscosa x160.
6. S. salina x160.
7. Spergularia maritima x160.
8. S. rubra x160.
PLATE 46
213
The plates no 47 and 48 are SEMs of capsule wall showing surface
ornamentation of Arenaria, Paronychia and Herniaria
PLATE 47.
1. Arenaria serpyllifoiia.
2. A. leptoclados.
3. Paronychia arabica.
4. P. argentea.
5. P. capitata.
6. P. chiorothyrsa.
7. Paronychia kapela.
PLATE 47
r.r '
214
PLATE 48.
1. Herniaria cinerea.
2. H. cyrenaica.
3. H. glabra.
4. „
„
5. Herniaria fontanesii.
6.
7. Herniaria hemistemon.
PLATE 48
215
Chapter 6
General Discussion and Conclusions
6.1 Chapter 2 Methods
Light microscopy and standard SEM techniques were used to
produce the results and data concerning seeds discussed in this
thesis and such a statement largely suffices for the work on the
fo lia r
crystals.
However,
the
sligh tly
m odified
version
of
Bokhari's technique allowed the much speedier production of
good preparations. In examining the capsular surfaces of Silene
the superglue method proved to be swift, easy and inexpensive
and gave very satisfactory impressions and photographs which
can be compared with the SEM results.
6.2 Chapter 3
SEM techniques give excellent images of details of the testa
surfaces, as has been known for the few decades since their
first applications
Libyan
species
in late
and
also
1960s. The survey of seeds of the
of
those
few
Canarian
endemics
(P o lyca rp a e a ) bears this statement out in striking fashion. Some
particularly beautiful examples among the Paronychioideae are
the Tenerife specimens of P o lycarp ae a, notably the details of the
papillae (Plates 3,4 and 5), among the Alsinoideae the spurred
cells of Cerastium (Plate 14 and 15) and among the
216
Caryophylloideae
figures
of
Petrorhagia
S ile n e
velutina
longipetala
(Plate 23) and the four
(Plate 28, 1 to 4). Chapter 3
includes very detailed descriptions of the seeds of almost all the
Libyan species of Caryophyllaceae and there are discussions of
particular examples of the use of seed characters to distinguish
species, genera and tribes. The complex details of the seed
characters are summarised in Tables 6 to 9 (pages 228-231) to
facilitate the understanding of the sections 6.2 to 6.2.6 which
are the concluding statments on taxonomy from the species to
the subfam ilies.
6.2.1
Testa
M icrom orphology:
Species
and
Infraspecific
T axa
The Davis specimen 50344 of Arenaria s e rp y llifo lia
whole
question
of
the
taxonom ic
value
of
raises the
papillosity
(and
verrucosity). It has papillate seeds and the many other British
and North African specimens of the Arenaria serpyllifolia g ro u p
that were examined do not. It would be useful to know if the type
specimen of A. minutiflora has papillate seeds or not but this is
not
cru cia l
to
the
argum ent
here
which
is
"Can
the
presence/absence and abundance of papillae/warts be to some
degree under environmental control?" The work of New (1958,
1959,
1961)
dealt with
papillate
and
non-papillate
seeds
British Spergula arvensis and she advanced strong arguments
of
217
that these features of the testa were correlated with latitude
and altitude and also with germinability. These papillae are well
illustrated by Kowal (drawings, 1966) and by Stace (SEMs, 1991).
Mentioning Clapham et at. (1952), New attached no taxonomic
significance
to
the
differences
in
testa
m icrom orphology.
However, despite the evidence for some environmental control,
Stace (1991) gives two varieties of this species based on the
testa features. In the presence or absence of papillae and warts,
Sagina apetala, Silene apetala and Minuartia geniculata
a re
variable but in all cases further study is needed.
The
Plates
3 to 5 reveal the
great diversity of testa
ornamentation within a small number of species of P o lyca rp a e a ,
a genus with some 50 species according to Mabberley (1987).
This is a genus well worth revision and any revision cannot
ignore these striking seed characters.
6.2.2 Seed Shape and Testa Micromorphology Genera/
Subgenera
On
gross
m orphological
grounds the
genera
Robbariea
and
Polycarpaea are very similar. In both the overall shape seeds and
the details of the micromorphology the seeds of Robbariea are so
similar to those of P olycarpaea that there are no grounds for the
generic separation made by some authors. Similarly the seeds of
Minuartia geniculata are not distinct in
218
any marked way from those of other species of the genus and
hence
there
is
no support for
a genus
or even
subgenus
R h o d a ls in e though characters of pollen and other features may
give good reason for such a separation. See also the final
discussion of foliar crystals.
The very large genus S ile n e
has species with reniform
seeds with no wings or with double, undulate wings, reniform
seeds with single thin wings [ S. uralensis (Rupr.) Bocquet and S.
furcata Rafn.] and even globular seeds (as already mentioned in
3.3 T e le p h iu m ).
The shape of the lateral faces varies from
species to species in convexity and concavity. There are many
shapes of testa cells as shown by Rechinger et al. (1988) and in
this thesis. Any attempt to revise the sections or perhaps even
to split the genus should take account of these seed attributes.
6.2.3 Seed Shape and Testa M icro m o rp h o lo g y:
The case was made
in 3.3 that
Telephium
T e le p h iu m has seeds very
different from all other seeds in the fam ily and the tentative
argument made for removal of the genus from that family. In the
B ittrich
trib e
classification
C o rrigiolae
T e le p h iu m
Dumort.
(1927).
is joined with C o rrig io la in
However,
the
C o r r ig io la greatly resemble those of Scleranthus
seeds
of
annuus
in
subfamily Alsinoideae. A description of British material of seeds
of the last named is as follows:
219
Seeds
more
indistinct,
or
with
ornamentation,
less
thin
subglobular,
som ewhat
marginal
pale
yellow,
sinuous
face with
w alls,
a broad,
testa
cells
lacking
any
sharply delimited
darker yellow brownish band all round the seed, radicle terminal,
prominent and
pointed, hilum
a subterminal dark patch, seed
surface smooth and the marginal band somewhat glossy.
Such
Corrigiola
a
description
littoralis
almost
except that the
pointed. Clearly more work is
Corrigiola and
suffices
all 10 of
for
radicle
the
seeds
is obtuse
of
not
needed on all the 11 species of
Scleranthus. Though
as stated in 3.3 it
would be premature to suggest the removal of Telephium from
the family , it is clear that the evidence from seed morphology is
already strong enough for the placement of the genus in its own
trib e .
6 .2 .4 Seed Shape: the Dorsiventral Genera
That the monotypic genus Pteranthus of the Paronychioideae is
som ewhat
dorsiventrally
flattened
has
perhaps
received
no
previous comment and it is at least of interest in showing that
such flattening is not confined to Dianthus and related genera of
the Caryophylloideae. Nor is flattening lacking totally from the
Alsinoideae. The small genus Holosteum has "seeds longitudinally
keeled,
peltate",
according
to
Bittrich
(1993,
p.
227)
but
"asym m etrically reniform and laterally compressed", according
220
to Flora Europaea
(p. 164). Berggren (1981,p. 79) described the
seeds of H. umbellatum L. as "shield-formed, with a slight apical
notch". Her illustration shows the seed well.
In
Dianthus,
P e tro rh a g ia ,
Kohlrauschia
(sometimes
subsumed in P e tro rha gia, as in Flora Europaea) and Velezia the
flattening at the greatest can produce very thin, seeds of that
very
diagnostic
shape
often
called
scutate.
Last century
taxonomists attached importance to this shape as for instance in
Genera Plantarum of Bentham and Hooker. These authors linked
the genera
142)
Velezia, Dianthus and Tunica under the heading (p.
" Semina
peltata,
hilo
faciali.
Embryo erectus. Styli
2.
(Diantheae)".
20th century taxonomists have ignored this distinction at
the
level
above
the
genus
with
Bittrich
(1993)
the
latest
example. There is little if anything in the testa cell shape and
ornamentation of Dianthus and related genera to separate them
from
other
members
of
Caryophylloideae.
dorsiventral
flattening
into the
scutate
Nevertheless,
and
boat-like
the
shapes
deserves formal taxonomic recognition which would have to be
at tribal rank.
6.2.5
Seed Characters:
the Tribes
In the two previous sections
dealing with
Telephium and the
dorsiventral genera the erection of tribes based on seed
221
characters has been considered and this matter can be taken
further. In his tribal diagnoses Bittrich (1993) makes no mention
of seeds other than the number of seeds per fruit.
The genera studied for this thesis are shown in italicised bold.
Many of the unstudied genera are very small or even monotypic
and Bittrich gives no seed characters.
Tribes
1.
and
Subfam .
1. T rib e
Genera a c c o rd in g
to B ittric h .
P a ro n ych io id e a e
Polycarpeae
Drymaria,
Spergula, Spergularia, Sanctam brosia,
Pol ycarpaea, Polycarpon,
Ortegia,
Loeflingia, Haya, Xerotia,
Krauseola, Polytepalum, Stipulicida, Cerdia, Microphyes, Pirinia
2.
T ribe
P aronychieae
Co met es ,
Di cher ant hus, P t e r a n t h u s , S p h a e r o c o m a ,
Sclerocephalus,
Lochia,
Paronychia
G y m n o c a r p o s ) , H e r n i a r i a , P h ilip p ie lla ,
(including
C h a e to n y c h ia ,
Achyronychia, lllecebrum, Cardionema, Scopulophila, Pollichia
3.
T ribe
C o rrig io le a e
Tel ep hi u m, Corrigi ol a,
II. S ubfam .
1.
T ribe
A ls in o id e a e
A ls in e a e
A r e n a r i a , T h yla co sp e rm u m ,
M o e h r i n g i a , Brachystemma,
Thurya, Bufonia, Lepyrodiclis, Cerastium, Moenchia, Stellaria,
2 2 2
Pseudostellaria,
H onkenya,
Hol osteum, Myosoton, Minuartia, Withelmsia,
S a g i n a , C olobanthus,
A lsinidendron,
Schiedea,
Reicheella, Plettkea, Pycnophyllopsis
2.
T ribe
P ycn o p h ylle a e
Pycnophyllum
3. T ribe
Geocarpeae
Geocarpon
4. T ribe
Habrosieae
Habrosia
5.
T rib e
S cle ran th ea e
Scleranthus,
Pentastemonodiscus
III.
C a ry o p h y llo id e a e
1.
S ubfam .
T ribe
C a ryop hyllea e
A canthophyllum ,
A llochrusa,
D ia p h a n o p te ra ,
Cyathophylla,
S cle ran th op sis,
Gypsophila,
Bolanthus,
Ochotonophila,
V a c c a r i a , Pleioneura,
Ankyropetalum ,
Phrynella,
Dianthus,
Kohlrauschia, Petrorhagia, Velezia
2. T ribe
Dryideae
Drypis
3.
T ribe
Silene,
U ebelinia
Sileneae
Lychnis,
P e tro c o p tis ,
Cucubalus,
Agrostemma,
223
If seed characters as discussed in this thesis are given great
importance
then
these
tribes
would
be
rearranged
into
the
following grouping, consisting only of these genera studied here.
Polycarpeae would be split in two:
Spergula with
Spergularia
: Seed shape including wings; well-
marked spurs, ornamentation.
Polycarpaea
wi t h
characteristic
Polycarport
and
L o e flin g ia
: Seed shape;
papillae.
Paronychieae into three:
Paronychia with Herniaria : Seed shape; indistinct,smooth
Pteranthus : dorsiventrality;
Sclerocephalus
: Seed
cells
dark spot.
shape; well-marked furrow;
prominent
radicle.
Corrigioleae into two:
Telephium : Seed shape; cell shape; caruncle
C orrigiola with S cle ra n th u s (from the Alsinoideae): seed shape;
cells indistinct; marginal band; dark spot.
Alsineae stays undivided.
A r e n a r ia
Sagina
with
Minuartia, M o e h r in g ia , C e r a s tiu m , S te lla ria ,
224
Caryophylleae into
D ia n th u s with P e tr o r h a g ia , K o hlra usch ia ,
dorsiventrality;
Velezia :
P articular
prominent straight radicle.
Gypsophila with Vaccaria : reniform to globular shape
Silenoideae stays undivided
Silene with Agrostem m a : reniform shape (a few spp globular)
These
informal
groupings
are
put
forward
with
some
confidence even though many genera were not examined during
the course of this work. They do little violence to the Bittrich
scheme
except
Alsinoideae
and
for
its
the
removal
placement
of
in
S cle ra n th u s
the
from
the
Paronychioideae
with
Corrigiola. The main key character in splitting Paronychioideae
from
the
other two
subfam ilies
is the
presence/absence
of
stipules. Why should stipules be of such importance? Should not
seeds be considered a more deep-seated attribute? In any case
Davis (1967, p. 245) has "The leaves of Scleranthus
do, however,
have a scarious flange towards the base which may be stipular in
nature, and here emphasis is placed rather on the nature of
flower and fruit in recognising the lllecebraceae as a separate
fa m ily ."
225
6.2.6 Seed Shapes and Testa M ic ro m o rp h o lo g y:
S u b fa m ilie s
and
llle c e b ra c e a e
The Alsinoideae and Caryophylloideae appear very similar in seed
characters in their reniform shapes with sym m etrically placed
and often sunken hilums. Though it encompasses a diversity of
shapes, by contrast the Paronychioideae never has such precisely
reniform shapes as defined in this thesis. Similarly, testa cell
shape and the types of ornamentation which are such strong
features of the first two subfamilies are lacking from most if
not all of the Paronychoideae. Therefore it may be claimed that
on the totality of seed features there is a distinction between
the Alsinoideae and Caryophylloideae on the one hand and the
Paronychoideae on the other.
S p e rg u la and
Spergularia, here recognised as a distinct
grouping on seed features, are genera interesting in that they
breach
the
above
generalisation
in
large
m easure.
The
delimitation of the testa cells can be very conspicuous and is
never unclear. Some species have
well marked ornamentations
very strongly developed spurs as well as papillae and granules.
Furthermore, in overall seed shapes the species of Spergula may
be thought of as little different from reniform.
In addition, in
gross morphological attributes such as opposite and decussate
leaves and conspicuous corollas, these two genera resemble
A lsinoideae.
226
The concept of lllecebraceae of Bentham and Hooker is
followed by Davis (1967) in Flora of Turkey where the following
genera are included
1. Leaves all alternate (spirally arranged)
3.
C o rrig io la
1. Leaves essentially opposite (sometimes appearing alternate
when one of a pair fails to develop, but never spirally
arranged)
2. Leaves without distinct stipules, though with a scarious
margin towards the base
4. S c le r a n th u s
2. Leaves with distinct scarious stipules
3. Bracts very conspicuous, exceeding and often concealing the
flowers
2. P a ro n y c h ia
3. Bracts not very conspicuous, not exceeding the flowers
4. Annual, erect; leaves reddish, aristate, with a membranous
m argin
2. P a ro n ych ia
4. Annual or perennial, procumbent to ascending; leaves green,
not aristate, without a membranous margin
3. H e r n ia r ia
These four genera have all been discussed in this thesis as
have been P te ra n th u s , Sclerocephalus
and G y m n o c a rp o s , all
Libyan genera included in the family by Abdul Ghafoor. As well as
the presence of stipules, the members of the lllecebraceae lack a
227
corolla and have perigynous flowers which develop into oneseeded
te sta
indehiscent fruits. These seven genera share smooth
c e lls
and
never
have
protuberances. However, they
papillae
or
other
c e llu la r
show a considerable diversity of
seed shapes. Furthermore, lllecebrum verticillatum has elongate
seeds distinctly pointed at one end and with a very high gloss
see Berggren (1981, Plate 31). These seeds in some features
resem ble
those
of
some
P o lyca rp a e a spp. H erniaria
and
P a ro n y c h ia seeds have collar cells but the other five genera do
not. Therefore the recognition of lllecebraceae
by the testa features.
is supported only
convex
brown
H
R
None
y
«
n
elongate
R
Long
R
f
or
Dense
R
granular
Finely
u
eniform
Stelliform
Sunker
Curved
Convex
Slightly
Dark
II
None
granular
a
Circular
circular
surface
curved
grooved
concave
Elongate or
brown
Near
Slightly
Convex
Convex,
spaced
elongate
H
concave
Light
Regularly Varied
irregular
Ovate or
ii
convex
or short
elongate
or
Minutely
Granular
With or
without
Surface
W arts
d
St ell a ria
or stelliform
Blunt
Long
Mostly
Elongate ovate Regular
spaced
regularly
Blunt
None
Mostly
Tu bercles
Papillae
Spurs
L ib y a ).
n
reniform
Cuneate
Curved
curved
u
Sagina
Convex or
grooved
mostly
Mostly
D
form
Flat or
convex
Convex,
grooved
or elliptic
Sunker Oval,elongate
Cell shape
in
fea tu re s
found
Testa
(genera
i
Orange
H
brown
orange
Orange o Flat or
brown
convex or
convex
dark
Curved
Hilum
Alsinoideae
«
R etorti-
cuneate
Broadly
reniform
Flat
Flat or
face
face
Radicle
Subfamily
Light or
Marginal
the
L ateral
of
«
M in u a rtia
C erastium
A renaria
circular
mm
C o lo u r
characteristics
A ls in o id e a e
Seed
S h a p e Size
6.
V
S u b f a m ily
Table
228
R
7.
Seed
Herniaria
P a ro n yc h io id ea e
S ubfam ily
Table
V
C\i
1
<D
+-J
03
-Q
>
o
n
<0
o
e-
03
O
o
c
E
&
v
M ostly
ii
Creamy
Convex
Faint
Indistinct
ii
ii
ii
ii
curved
ii
Slightly
None
Indistinct
None
irregular
Smooth,
None
None
ii
if
ii
ii
ii
ii
ii
ii
ii
without
granular
Smooth or
ii
With or
Smooth
Smooth
glossy
Surfaces
W arts
Libya).
Tubercles
fea tu re s
in
Smooth
None
Papillae
Testa
found
ii
elongate
v
Polycarpaea
gray
white
.
Elongate
keeled
Curved
Mostly
Convex
surface
Near
ii
Loeflingia
ii
elliptic
ii
1 -2
Curved
Convex
Flat
curved
Keeled
Convex
Slightly
face
face
(genera
Cell shape Spurs
Paronychioideae
Radicle Hilum
subfamily
L ateral Marginal
the
ii
Circular
Brown
Colour
of
ii
Paronychia
circular
Mostly
mm
Shape size
characteristics
229
ii
ii
ii
8.
Mostly
Telephium
only.
ii
or ovate
curved
Slightly
ii
Convex
Membranous
Broad or
wing
Membranous Curved
surface
Near
ii
Caruncle found in
ii
ii
Brown
ii
ii
Globular
ii
Spherical
Irregular
Elongate or
None
None
Irregulai Conical
spaced
Blunt
None
Conical
None
ii
Telephium
ii
Rough
Granular
Smooth
ii
None
ii
Conical
ii
Regular
n
Mostly elliptic
ii
shaped
ii
surface
ii
curved
ii
obovate
Mostly
Spergularia
ii
Flat
None
Surface
ii
Straight
Dome
Papillae Tubercles W arts
Smooth
Spurs
Libya )
1fe a tu re s
in
None
shape
Testa
found
ii
None
Near
Slightly
Cells
(genera
ii
Indistinct
Hilum
Radicle
Paronychioideae
ii
brown
Convex
Dark
1 -2
Circular
Spergula
Convex
Creamy Convex
Flat
face
Marginal
Subfamily
face
Light
Sclerocephal us
2
the
Colour Lateral
of
ii
Faint
Obovate
>
v
i
pbovate
mm
Size
v
P e te ra nthu s
Polycarpon
Shape
characterstics
cuneate
Seed
Paronychioideae
Subfam ily
Table
230
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ii
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6.3 C hapter
4.
According to the results obtained
fundam ental
Arenaria,
points
concerning
Moehringia,
different characteristic
160 taxa
within
these
in the present study some
the
infrageneric
and Minuartia
morphologies
three
can
be
of the
genera
have
groupings
clarified.
crystals
been
in
The
of about
thoroughly
described in Chapter 4, section 3. It was concluded that shape,
size
and
distribu tion
of
the
crystals
were
taxo no m ically
siginificant at the subgeneric level. These can be summarised
briefly, always with the proviso that not all species in these
large genera were examined.
1. According to these studies of the crystals, the great majority
of the groupings of McNeill can be supported.
2. Subgenera of Arenaria (Dolophragma and Solitaria) are distinct
from all the other subgenera of that genus and also from all
members of M oehringia and of Minuartia of whatever subgenera
by the absence or great scarcity of crystals.
3. All members of M o e h rin g ia show great similarities with the
genus Arenaria, except the subgenus Eremogone of Arenaria, and
also display great similarities with Minuartia, except
subgenus
Minuartia but within that subgenus the section Spectabiles does
re se m b le
Moehringia. In M oehringia all the crystals are druses;
no sand has been encountered.
4. The subgenus Eremogone of Arenaria
has characteristic
233
distributions
of
crystals
which
makes
every other subgenus of A r e n a ria . Its
it
very
crystals
distinct
are
from
arranged
mostly irregularly but sometimes regularly in short rows in both
the intercostal areas and in the veins (in the 27 spp. examined)
or in the veins only (in two spp). By contrast all other members
of Arenaria of whatever subgenus (44 spp) have the crystals only
in the intercostal areas and never in the veins. Eremogone is the
sole subgenus to have crystal sand; the other subgenera have
druses only.
5. The subgenus E re m o g o n e of
su b ge nu s
Arenaria
gre atly
resem bles
Minuartia of M inuartia in the distribution of crystals in
that the crystals are present in both the intercostal areas and in
the veins. However in subgenus Minuartia the costal crystals are
mostly continuous in the cells whereas in subgenus E re m o g o n e
the costal crystals are discontinuous.
6. Druses and sand both occur in Minuartia except
subgenera
R h o d a lsin e and H y m e n e lia which only have druses, subgenus
S p e r g e lia with only two species has druses only in one ( M .
form osa) but druses, sand and elongate crystals in the other (M
picta). All sections in subgenus Minuartia have sand and druses
except section Spectabiles with only druses. Series
Laricifoliae
of section Spectabiles, has both druses and sand.
7. Giant crystals up to 233 pm long are found in most sections of
subgenus Minuartia of Minuartia and in no other members of
234
M in u a rtia , A r e n a r ia
or M oe hring ia. In subgenus E re m o g o n e
of Arenaria there are large crystals but they never reach 100 pm
long. These giant crystals appear not to have been reported
elsew here.
8.
By the application of this easy, swift technique for crystal
exam ination,
distribution
these
can
features
be used
of the
crystal
in assigning
morphology
and
sterile plants of these
genera to a genus. A specimen thought to belong to subgenus
M in u a rtia of Minuartia could have its identity confirmed by the
crystals except if it were of section S p e c ta b ile s .
Sim ilarly a
specimen thought to be of subgenus Eremogone of Arenaria could
have its identity confirmed although there is the possibility that
the specimen might be of subgenus M inuartia happening to have
some discontinuity of the crystals.
6.4 C hapter 5.
The results from
light microscopy and
m icromophology seem very
SEM of the capsular
promising for the
large,
complex
genus Silene and should be taken much further by comparing with
the sectional divisions of past authors. The results from the
other genera seem also to be valuable but again there is a need
for more detailed investigations.
235
6.5 Some General Concluding Points:
Within the Libyan Caryophyllaceae there are the endemic species
of Silene (3 spp.), Petrorhagia (1 sp.), and Herniaria (1 sp.). The
seeds of most of these taxa have been examined from admittedly
a small sample of specimens in all cases but nevertheless the
seed micromorphology bears on the validity of these special
taxa. The number of examined specimens needs to be increased
for greater thoroughness and the gathering of new material from
the field would be desirable. Such fresh collecting would be best
carried out in northernmost Cyrenaica. This would then allow a
complete taxonomic reassessment not just from seed characters
but from many mophological and other appropriate aspects.
The survey of Caryophyllaceous seeds from Libya and of some
non-Libyan species deals only with a small proportion of the
seeds of all the many species in the family. All the large genera
of the fam ily are represented in the survey and many of the
smaller genera as well. Therefore the survey gives a very good
idea of the scope of seed macro- and micromorphology.
The range of seed characters as discussed in this thesis
and by many previous workers shows that such features can be
helpful at the specific and infraspecific levels. At the tribal and
subfamilial levels seed characters are also helpful and may be
more helpful than has hitherto been fully realised. It, however
appears that at the generic level seed characters are less useful,
236
though not totally without importance. Seeds are therefore, of
little use in resolving the problems of generic delimitation, a
well-known
difficulty
in the fam ily
as
pointed
(1967) and by Bittrich (1993, p. 207) who stated
out
by Davis
"Many genera in
the fam ily are ill-defined and difficult to distinguish".
It would be a finite task to complete the survey of all the
genera, many of which are of only one or of few species. After
that the subfamilial and tribal divisions could be revised with
com plete
confidence
to
take
full
advantage
of
the
very
considerable value of seed characters. Then formal proposals
could be made. Similarly the extension of the studies of crystals
and
of the
fruit
epiderm is
to
cover
all the
fam ily
may
be
considered as well worthwhile.
It is highly desirable that a phylogenetic analysis of the family
with the application of the most up-to-date methods be carried
out. Such an analysis should use the very detailed seed, fruit and
crystal characters as described in this thesis.
237
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Appendix I. List of Taxa with Authorities.
Agrostem m a L.
githago L.
Arenaria L.
acerosa Bioss.
aculeata S. Wats.
acutisepala Williams
alsinoides Schlecht.
angustisepala McNeill
armeniaca Boiss.
armerina Bory
bertolonii Fiori & Paol.
biflora Griseb.
boliviana Williams
bryoides Schlecht.
capillaris Poir.
cerastoides Poir.
ciliaris Loscos
ciliata L.
ciliolata Edgew.
conferta Boiss.
cucubaloides J. E. Smith
davisii McNeill
decussata Eilld. ex Schlecht.
deflexa Decaisne
densissima Edgew.
dianthoides J. E. Smith
drypidea Boiss.
fendleri A. Gray
festucoides Benth.
filicaulis Fenzl
formosa Ser.
forrestii Diels
frankinii Dougl. ex Hooker
gouffeia Chaub.
graminea C. A. Mey.
grandiflora L.
griffithii Boiss.
guatemalensis Stand ley & Steyerm.
guicciardii Heldr.
gypsophiloides L.
halacsyi Bald.
hispanica Sreng.
holostea M. Bieb.
hookeri Torr. & Gray
humifusa (Swartz) Wahlb.
huteri Kern.
incrassata Lange
insignis Litiv.
kansuensis Maxim.
kingii (S. Wats.) M. E. Jones
lanuginosa (Michx.) Rohrb.
ledebouriana Fenzl
leptoclados Guss.
lithops Heywood
loscosii Texid.
lychnidea Schlecht.
lycopodioides Schlecht
macradenia S. Wats.
montana L.
napuligera Franch.
oreophila Hook.
oresbia W. W. Sm.
paludicola Robinson
Arenaria L.
palustris Naud.
persica Boiss.
polycnemifolia Boiss.
polytrichoides Edgew.
pseudacantholimon Bornm.
pulvinata Huter
punges Clem.
purpurascens DC.
pseudacantholimon Bornm.
pseudofrigida Shischk.
pycnophylla Rohrb.
reptans Hemsl.
retusa Boiss.
rhodia Boiss.
sabulinea Fenzl
saponarioides Boiss. & Bal.
scariosa Boiss.
serpyllifolia L.
steveinana Boiss.
szowitsii Boiss.
teddii Turrill
tetraquetra L.
tetrasticha Boiss.
trichophora Franch.
yunnanensis Franch.
zargariana Pasa
Cerastium L.
comatum Desv.
dichotomum L.
glomeratum Thu ill
ligusticum Viv.
pumilum Curtis
semidecandrum L.
siculum Guss.
Corrigiola L.
littoralis L.
Dianthus L..
crinitus Sm.
Gymnocarpus Forsk.
decander Forsk.
Gypsophila L.
elegans Bieb.
pilosa Hudson
Herniaria L.
cinera DC.
cyrenaica F. Hermann
ericifolia Town sen d
fontanesii Gay
glabra L.
hemistemon Gay
Loeflingia L.
hispanica L.
Lychnis L.
flos-cuculi L.
Mmuartia L.
acuminata Turrill
aizoides (Boiss.) Bornm.
anatolica (Boiss.) Woronow
arctica (Steven) Graebn.
aretioides (Somerauer) Schinz & Thell. austriaca (Jacq.) Hayek
baldaccii (Hal.) Mattf.
biflora (L.) Schinz & Thell.
brevifolia (Nutt.) Mattf.
bulgarica (Velen.) Graebn.
californica (Gray) Mattf.
campestris L.
capillacea (All.) Graebn.
caroliniana (Walt.) Mattf.
cerastifolia (DC.) Graebn.
colchica Kharadze
confusa (Boiss.) Maire & Petitm.
dawsonensis (Britton) House
dianthifolia (Boiss.) Hand-Mzt.
dichotoma L.
douglasis (Fenzl) Mattf.
fasiculata (L.) Reichb.
flaccida (L.) Reichb.
formosa (Fenzl) Mattf.
funkii (Jord.) Graebn.
geniculata (Poir) Thellung
glabra (Michx.) Mattf.
globulosa (Labill.) Schinz & Thell.
graminifolia (Arduino) Javorka
groenlandica (Retz.) Ostenf.
hamata (Hausskn.) Mattf.
hirsuta (M. Bieb.) Hand.-Mzt.
howellii (Wats.) Mattf.
hybrida (Vill.) Schischk.
imbricata (M. Bieb.) Woronow
inamoena (C.A. Mey.) Woronow
laricifolia (L.) Mattf.
mediterranea (Ledeb.) Maly
mesogitana Mattf.
moehringioides (Ser.) Mattf.
montana L.
mutabilis (Lapeyr.) Becherer
obtusiloba (Rydb.) House
patula (Michx) Mattf.
pestalozzae (Boiss.) Bornm.
picta (Sibth. & Sm.)
recurva (All.) Schinz. & Thell
rimarum (Boiss. & Bal.) Mattf.
rossii (R.Br.) Graebn.
rostrata (Pres.) Reichb.
rubella (Wahlb.) Hiern.
rupestris (Scop.) Schinze & Thell
saxifraga (Friv.) Graebn.
sedoides (L.) Hiern.
setacea (Thuill.) Hayek
stellata (Clarke) Maire & Petitm.
stricta (Siv.) Hiern
tenetla Mattf.
umbellulifera (Boiss.) McNeill
verna (L.) Hiern.
viscosa (Schreber) Schinz & Thell.
M oehringia L.
ciliata (Scop.) Dalla Torre.
diversifolia Dolliner
glaucovirens Bertol.
intricata Willk
255
Moehringia L.
jankae Janka
laterifora (L.) Fenzl
muscosa L.
pendula (W. & K.) Fenzl
radiolata Pancic
sedifolia
stellarioides Cosson
tejedensis Willk.
tommasinii Marchesetti
trinervia (L.) Clairv.
Paronychia Miller
arabica (L.) DC.
argentea Lam.
capitata (L.) Lam.
chlorothyrsa Murb.
kapela (Hacq.) A. Kenner
Petrorhagia (Ser.) Link
illyrica (Ard.) P. W. Ball & Heywood
velutina (Guss.) P. W. Ball & Heywood
Polycarpaea Lam.
carnosa Chr. Sm. ex Buch
divaricata (Ait.) Poir
repens (Forsskal) Ascherson & Schweinf.
robbiarea (O. Kunze) Greuter & Burdet
smithii Link
tenuis Wibb ex Christ
Polycarpon Loefl. ex L.
arabicum Boiss
bivonae Gay
depressum Rohrb.
indica Lam.
ioeflingiae Benth. & Hook.
peploides DC.
polycarpoides (Biv.) Jahandiez & Maire prostratum (Forsk.) Ascherson & Schweinf.
succulentum (Delile) Gay
tetraphyllum (L.) L.
Pteranthus Forsk.
dichotomus Forsk.
Sagina L.
apetala Ard.
maritima G. Don.
Scleranthus L.
annuus L.
Sclerocephalus
arabicus Boiss.
Silene L.
acaulis L.
aegyptiaca (L.) L. fit.
apetala Willd.
arenaroides Desf.
armeria L.
articulata Viv.
atlantica Cosson & Durieu
behen L.
S ilene L.
bellidifolia Jacq.
boryi Boiss.
cerastoides L.
claryi Batt.
coelirosa (L.) Godron
colorata Poiret.
conica L.
cono idea L.
corrugata Ball
cyrenaica Maire & Wei Her
dioica (L.) Clairv.
disticha Willd.
divaricata Lag.
fruticosa Otth
fuscata Brot.
gallica L.
ghiavensis Batt.
glabrescens Cosson
heterodonta F. N. Williams
ibosii Emberger & Maire
imbricata Desf.
inaperta L.
italica (LA Pers.
kremeri Soyer-Wiliemet & Godron
laeta (Aiton) Godron
latifolia Poiret
littorea Brott.
longicaulis Lag.
longipetala Vent.
marmarica Beguinot & Vacc.
mekinensis Cosson
micropetala Lag.
mollissima (L.) Pers.
muscipula L.
neglecta Ten.
nicaeensis All.
noctiflora L.
nocturna L.
nutans L.
obtusifolia Willd.
oropediorum Fenzl
otites (L.) Wibel
pomeli Batt.
pratensis L.
pseudoatiocion Desf.
ramosissima Desf.
reticulataDest
rubella L.
scabriflora Brot.
secundiflora Otth
sedoides Poiret
sericea All.
stricta L.
succulenta Forsk.
tridentata Desf.
tuberculata (Ball) Maire & Weiller
uniflora Roth
villosa Forsk.
vivianii Steudel
vulgaris (Moench) Garcke
Spergula L.
arvensis L.
fallax (Lowe) E. H. L. Krause
morisonii Boreau
pentandra L.
viscosa Lag.
Spergularia (Pers.) J. & C. Persl.
bocconei (Scheele) Graebner
cerastoides Fouc.
diandra (Guss.) Boiss.
fimbriata Boiss. & Reuter
grandis Cambess.
levis Cambess.
maritima (All.) Chiov.
nicaeensis Burnat
ramosa Camess.
rubra (L.) J. Presl & C. Presl
salina J. Presl & C. Presl
Stellaria L.
cupaniana (Jordan & Fourr.) Beguinot
pallida (Dumort.) Pire'
Telephium L.
sphaerospermum Boiss.
Vaccaria Medik.
pyrmidata Medik.
media (L.) Vill.
258
Appendix II. Details of the Seed, Crystal and Fruit Capsule samples Studied;
Herbaria abbreviations were taken from Patricia (1981). Material collected from
Libya written in bold.
No
Taxon
01
02
03
04
05
06
07
08
09
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
Agrostemma githago
A. githago
Arenaria acerosa
A . aculeata
A..acutisepala
A .alsinoides
A. alsinoides
A.alsinoides
A. angustisepala
A .angustisepala
A .armeriacea
A . armerina
A. armerina
A . armerina
A. armerina
A. armerina
A. armerina
A. armerina
A. bertolonii
A. biflora
A. biflora
A. boliviana
A. bryoides
A. capillaris
A. cerastoides
A. cerastoides
A. cerastoides
A. cerastoides
A. cerastoides
A. cerastoides
A. cerastoides
A. cerastoides
A. cerastoides
A. cerastoides
A. cerastoides
A. cerastoides
A. cerastoides
A. cerastoides
A. ciliaris
A. ciiiata
A. ciiiata
A. ciiiata
A. ciliolata
A. conferta
A. conferta
A. cucubaloides
A. davisii
Locality
Syria
Turkey
s. I.
Canada
Turkey
Mexico
Mexico
Mexico
Turkey
Turkey
Turkey
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Algeria
Italy
Britain
Britain
Bolivia
Mexico
Canada
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Morocco
Morocco
Algeria
Morocco
Asia
Switzerland
France
Spain
Tibet
Yugoslavia
Greece
Turkey
Turkey
Date
Collector
1906
1952
1845
1959
1957
1902
1907
1908
1954
1934
1957
1973
1924
1936
1951
1973
1973
1975
1818
1901
1903
1860
1905
1956
1830
1832
1850
1855
1891
1851
1852
1884
1891
19 07
1912
1927
1939
1972
1947
18 58
1874
1889
1947
s. d.
1966
1934
1961
Manoog
s. n.
Davis
19320
J. Uelreide
s. n.
W. Bird
5083
Davis & Hedge 31477
C. A. Purpus
2627
C. A. Purpus s. n.
C. A. Purpus
s. n.
22224
Davis et al.
E. K. Balls
s. n.
Davis & Hedge 51885
E. Jahandiez 402
E. Jahandiez 626
E. K. Balls
3045
D. H. N. Spence594
Davis
55011
Davis
55037
Davis
58951
Adri Fiori
1032
P. Ewing
15
P. Ewing
95
G. Mandon
960
C. A. Purpus
1655
J. A. Calder
17508
W. Schimper 8
W. Schimper
128
P. Jamin
22
G. L. Durando s. n.
John Ball
54
B. Balansa
96
B. Balansa
453
O. Debeaux
s. n.
Warion
200
A. Faure
s.
C. J. Pitard
2702
E. Jahandiez 22
A. Faure
s.n.
F. K. Kupicha 278
Sesse & Mociro 11186
Balfour
s. n.
Jee' Jaas
26
E. Reverchon s. n.
F. Idlow
15647
S. H.Lecnhouts .157
J. C. Archibald . 322
E. K. Balls
1471
Davis
694
No.
Herb?
(E)
(E)
(BM)
(BM)
(BM)
(E)
(E)
(E)
(E)
(BM)
(BM)
(E)
(E)
(E)
(E)
(E)
(E)
(E)
(E)
(E)
(E)
(BM)
(BM)
(BM)
(E)
(E)
(E)
(E)
(E)
(E)
(E)
(E)
(E)
(E)
(E)
(E)
(E)
(E)
(ULT)
(E)
(E)
(E)
(BM)
(L)
(L)
(BM)
(E)
259
No
Taxon
Locality
Date
Collector
48
49
50.
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
A. decussata
A. decussata
A. deflexa
A. densissima
A,, dianthoides
A. dianthoides
A. drypidea
A. fendieri
A. festucoides
A. filicaulis
A. formosa
A. formosa
A. forrestii
A. frankinii
A. gouffeia
A. gouffeia
A. graminea
A. grandiflora
A. griffithii
A. guatemalensis
A. guicciardii
A. gypsophiloides
A. halacsyi
A. holostea
A. holostea
A. holostea
A. hispanica
A. hookeri
A. humifusa
A. huteri
A. incrassata
A. insignis
A. kansuensis
A. kingii
A. lanuginosa
A. lanuginosa
A lanuginosa
A. lanuginosa
A. lanuginosa
A. lanuginosa
A. lanuginosa
A. lanuginosa
A. lanuginosa
A. lanuginosa
A. lanuginosa
A. lanuginosa
A. lanuginosa
A. ledebourinana
A. leptoclados
A. leptoclados
A. leptoclados
A. leptoclados
A. leptoclados
Mexico
Mexico
Syria
Tibet
Armenia
Turkey
Turkey
U. S. A.
North India
Finland
U. S. A.
Asia
China
U. S. A.
s. I.
France
U. S. S. R.
Spain
Afghanistan
Guatemala
Greece
Turkey
Jugoslavia
Turkey
Iran
Iran
Spain
Nebraska
Newfound land
Italy
Spain
Iran
E. Tibet
U. S. A.
Mexico
Mexico
Guatemala
s. I.
Guatemala
Mexico
Mexico
Nicaragua
Nicaragua
Costarica
Nicaragua
Nicaragua
Mexico
s. I.
Greece
Serbia
Crete
Morocco
Morocco
1894
1948
1863
1935
1957
1966
1952
1961
1881
1887
1935
1936
s. d.
1873
1825
1839
1836
1970
1956
1967
1975
1954
1900
1957
1960
1962
1900
1947
1958
1900
1885
1966
1926
1956
1928
1935
1947
1959
1974
1976
1977
1979
1979
1979
1981
1981
1991
1855
1896
1897
1940
1969
1969
6479 (E)
C. G. Pringle
2967 (E)
F. G. Meyer
(BM)
B. T. Lowne
s. n.
Ludlow et al.
3928 (BM)
(BM)
s. n.
s. c.
44024 (E)
Davis
Dodds & Cetibe20190 (BM)
C. M. lit
19119 (BM)
1391 (BM)
J. F. Dutlis
24
(E)
S. Graeca
W. J. Eyerdam s. n.
(BM)
E. W. Tisdale 40367 (BM)
Y. Lichiang
15375 (C)
(BM)
C. C. Parry
35
s. c.
388
(E)
H.de Larambergue442 (L)
R. F. Hohenackers. n. (BM)
3501 (E)
P. Aichibold
W. Thesiger
1394 (BM)
A. Molina
21016 (BM)
E. A. Menneg s. n.
(E)
David & Polunin23944 (BM)
L. Vaccri
(E)
s. n.
Davis & Hedge 29365 (BM)
95
(E)
P.Furse
P.Furse
2290 (E)
E. Reverchon s. n.
(E)
W. Kiener
22100 (BM)
T. T. Elkieton 130
(BM)
Andri Fiori
s. n.
(E)
Parta et Figo
s. n.
(E)
J. C. Archibald 2480 (E)
J. F. Rock
14222 (BM)
E. K. Balls
10899 (BM)
522
(M)
E. Lyonnel
H. Lesueure
(M)
465
A. Molina
(M)
419
J. Duke
1672 (M)
30024 (M)
A. Molina
D. E. Breedlore 42776 (M)
(M)
W .Bennet etal. 10
W . Douglas et al. 14954 (M)
W . Douglas
15977 (M)
W. D. Stevens 13406 (M)
P. P. Moreno 10347 (M)
P. P. Moreno 9692 (M)
J. A. Soule
2918 (M)
B. Balansa
(BM)
s. n.
J. Dorfler
196
(E)
s. c.
15025 (E)
Davis
1338 (E)
Davis
48969 (E)
Davis
48969 (E)
No
Herb;
260
No
Taxon
101
A. leptoclados Libya
A. leptoclados Algeria
A. leptoclados Algeria
A. leptoclados Tunisia
A. leptoclados Britain
A. leptoclados Britain
A, leptoclados Britain
A. leptoclados Britain
A. leptoclados Britain
A. leptoclados Britain
A. leptoclados Britain
A. leptoclados Britain
Spain
A. lithops
Spain
A. loscosii
A. lychnidea
s. I.
A. lychnidea
Caucasus
A. lycopodioides Mexico
A. macrodenia U. S. A.
A. montana
Spain
A. montana
Spain
A. montana
Spain
A. napuligera China
A. oreophila
China
A. oresbia
Jalisco
A. paludicola Mexico
A. palustris
South America
A. persica
s.l.
A. persica
Iran
A. polycnemifolia Iran
A. polytrichoidesT\beX
.A. pseudacantholimon Turkey
A. pseudacantholimon Turkey
A.pseudofrigida Greenland
Spain
A. pulvinata
A. pungens
Morocco
A. pungens
Morocco
A. pungens
Morocco
A. pungens
Morocco
A. pungens
Morocco
A. pungens
Morocco
A. pycnophylla Armenia
A. reptans
Mexico
A. reptans
Mexico
A. reptans
Mexico
A. reptans
Mexico
A. reptans
Mexico
A. retusa
Spain
A. rhodia
Turkey
A. sabulinea
s. I.
A. sabulinea
Turkey
A. saponarioides Cyprus
A. saponarioides Cyprus
A.scariosa
Turkey
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
Locality
Date
Collector
No
Herbaii
1970
1971
1975
1975
1889
1890
1897
1908
1932
1944
1949
1984
1970
1894
1933
1979
1932
1943
1852
1861
1889
1910
1932
1952
1899
s. d.
1842
1934
1902
1938
1957
1957
1982
1948
1936
1962
1973
1973
1973
1985
1937
1905
1907
1965
1966
1971
1973
1958
1888
1957
1880
1940
1957
Davis
Davis
Davis
Davis
J. Andrew
R. Withie
R. Kidston
D. Patton
T. Wise
T. Megrouther
E. R. Wise
C. Rodiguaz
E. Dominguez
E. Reverchon
E. R. Balls
Z. Grinianidze
G. B. H.
A. Davidson
E. Bourgeau
J. Ball
E. Reverchon.
G. Forrest
J. Rock
R. Mcvaugh
L. Robinson
H. F. Comber
R. F. Hohenacker
Davis
A. Bornmuller
Ludlow et al.
Davis & Hedge
Davis & Hedge
C. Bay e ta l.
V. H. Heywood
E. K. Balls
J. C. Archibald
Davis
Davis
Davis
C. Blanche et al.
s. c.
C. A. Purpus
C. A. Purpus
D. E. Breedlove
D. E. Beedlove
D. E. Beedlove
E. Dominguez
Davis & Hedge
O. Stapf
Davis
s. c.
Davis
Davis & Hedge
50344
52652
58386
57561
245
240
234
155
s. n.
5137
1940
1984
s. n.
4027
557
s. n.
2458
s. n.
1706
s. n.
21
6509
24751
13817
233
1039
597
796
6406
5030
31631
31659
1349
1178
2738
165
55231
55359
55484
31042
207
1665
2769
8106
15304
22737
1611
1958
825
28524
1816
1858
s. n.
(E)
(E)
(E)
(E)
(GL)
(GL)
(GL)
(GL)
(GL)
(GL)
(GL)
(GL)
(E)
(E)
(E)
(E)
(BM)
(BM)
(E)
(E)
(E)
(BM)
(BM)
(BM)
(E)
(E)
(BM)
(E)
(BM)
(BM)
(BM)
(E)
(BM)
(E)
(E)
(E)
(E)
(E)
(E)
(E)
(BM)
(E)
(E)
(M)
(M)
(M)
(E)
(BM)
(E)
(E)
(E)
(E)
(E)
261
Locality
No
Taxon
154
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
A. serpyllifolia Morocco
A. serpyllifolia Morocco
Aserpyllifolia Morocco
A. serpyllifolia Libya
A. serpyllifolia Tripoli
A.serpyllifolia Algeria
A. serpyllifolia Morocco
A. serpyllifolia Algeria
A. serpyllifoia Algeria
A.serpyllifolia
Algeria
A.serpyllifolia Tunisia
A. serpyllifolia Algeria
A. serpyllifolia Algeria
A. serpyllifolia Algeria
A. serpyllifolia Morocco
A. serpyllifolia Britain
A. serpyllifolia Britain
A. serpyllifolia Britain
A. serpyllifolia Britain
A. serpyllifolia Britain
A. serpyllifolia Britain
A. serpyllifolia Britain
A. serpyllifolia Britain
A. serpyllifolia Britain
A. serpyllifolia Britain
A. serpyllifolia Britain
A. serpyllifolia Britain
A. serpyllifolia Britain
A. serpyllifolia Britain
A. steveiana
Himilaya
A. szowitsii
Iran
A. teddii
Greece
A. tetraquetra Europe
A. tetrasticha s. I
A. trichophoraq China
A. yunnanensis China
A. zargariana Iran
Cerastium. comatum Beida
C. comatum
Morocco
C. comatum
Greece
C. dichotomum Greece
C. dichotomum Spain
C. dichotomum Spain
C. dichotomum Britain
C. glomeratum Tripoli
C. glomeratum Tripoli
C. glomeratum Tripoli
C. ligusticum Italy
C. ligusticum Italy
C. ligusticum Italy
C. pumilum
Italy
C. pumilum
Spain
C. pumilum
Cechoslovacae
Date
Collector
No
Herba
1924
1932
1951
1970
1976
1971
1973
1975
1971
1971
1975
1975
1975
1975
1981
1889
1882
1883
1892
1902
1905
s. d.
1912
1912
1948
1980
1985
1985
1987
1914
1962
1936
1825
s. d.
1987
1890
1966
1970
1912
1883
1884
1900
1891
1892
1970
1976
1977
1844
1851
1907
1897
1962
1962
E. Jahandiez
A. Faure
D.H. Spence
Davis
S. M. Jafri
Davis
Davis
Davis
Davis
Davis
Davis & Lamond
Davis
Davis
Davis
Davis
J. Andrew
W. Gourlie
J.Wglie
Ahorollywod
s. c.
Horwards
P. Ewing
P. Ewing
P. Ewing
K. W. Praid
J. Dickson
J. Dickson
H. J. Noltie
s. n.
W. Foiber
P. Furse
K. H. Rechinger
J. Ganillo
Dalm Kow
D. Chamberlain etal.
A. E. Pratt
J. C. Archibald
Davis
G. Orphanides
P. Ascherson
s. c.
E. Reverchon
E. Reverchon
P. Ewing
s. c.
S. M. Jafri
A. Gafoor
T. Caruel
J. Ball.
Andri Fiori
A. Kneuker
R. K. Brummitt et. al
F. Slavonovsky
615
1932
5239
50344
6424
52537
55434
59389
53105
53105
57537
59262
59074
59466
67628
s. n.
s. n.
s. n.
234
s. n.
245
243
391
392
1948
42
s. c.
s. c.
s. c.
28722
2237
10317
885
4262
1305
155
2540
50519
188
143
s. n.
s. n.
556
375
s. n.
6452
298
8
s. n.
s. n.
4710
654
1422
(E)
(E)
(E)
(E)
(ULT)
(E)
(E)
(E)
(E)
(E)
(E)
(E)
(E)
(E)
(E)
(GL)
(GLI)
(GL)
(GL)
(GL)
(GL)
(GL)
(GL)
(GL)
(GL)
(GL)
(GL)
(GL)
(GL)
(BM)
(E)
(E)
(E)
(BM)
(BM)
(BM)
(E)
(ULT)
(E)
(E)
(E)
(E)
(E)
(E)
(ULT)
(ULT)
(ULT)
(E)
(E)
(E)
(E)
(E)
(E)
262
No
Taxon
207
208
209
210
211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
252
253
254
255
256
257
258
259
260
C. pumilum
Tripoli
C. semidecandrum Hungaria
C. semidecandrum France
C. semidecandrums. Italy
C. semidecandrum s. n.
C. siculum
Italy
C. siculum
Spain
Corrigiola littoralis Britain
C. littoralis
Britain
D. crinites
Libya
D. crinites
Afghanistan
Gymnocarpus decander Libya
G. decander
Libya
G. decander Tagma
G. decander
Libya
Gypsophila elegans Tripoli
Gypsophila pilosa Gharian
Hemiaria cinera Tripoli
H. cinera
GarIan
H.cinerea
Tripoli
H. cyrenaica
Benghazi
H. ericifolia
Libya
H. ericifolia
Libya
H. ericifolia
Tilel
H. fontanesii Morocco
H. fontanesii Kabao
H. fontanesii Zentan
H. fontanesii Nalut
H. fontanesii Mezda
H. fontanesii Morocco
H. glabra
Spain
H. glabra
Libya
H. glabra
Libya
H. hemistemon Al kararim
H. hemistemon Egypt
H. hemistemon Libya
H. hemistemon Shershara
H. hemistemon Qatar
Loeflingia hispanica Tripoli
L hispanica
Tripoli
L. hispanica
Tripoli
L. hispanica
Tripoli
Lychnis flos-cuculi Britain
L. flos-cuculi Britain
Minuartia acuminata s. I.
M. aizoides
Turkey
M.anatolica
Turkey
M. arctica
U. S. A.
M. aretioides Italy
M. austriaca
Austria
M. baldaccii
Albania
M. biflora
Greenland
M. brevifolia
U. S. A.
M. bulgarica
Bulgaria
Locality
Date
Collector
No
1983
1876
1904
1961
1963
1908
1978
1868
s. d.
1974
1969
1972
1974
1974
1975
1967
1974
1973
1974
1977
1972
1970
1970
1976
1933
1972
1974
1974
1974
1985
1967
1970
1970
1968
1967
1975
1976
1977
1967
1967
1967
1970
1913
1963
1963
1932
1957
1891
1905
1879
1918
1962
1893
1975
M. A. Siddiqi
s. n.
Braun.
3249
F. Sennen
32
K. H. Rechinger
21902
W. Greuter
1919
S. Sommier
s. n.
P. Harrold & R. J. McBeath 478
P. Ewing
2107
C. A. Johns
4379
B. Faris
597
s. c.
9080
S. I. Ali
236
S. I. Ali
1821
S. El. Jaley
199
S. I. Ali
2843
s. c.
s. n.
M. Godeh
269
S. I. Ali & Faruqui
1178
S. I. Ali
2095
A. Gafoor
175
S. I. Ali
484
Davis
49480
Davis
49769
A. Ghafoor & S. lavi
46
A.Faure
s. n.
S. I. Ali
487
S. Elzualy
351
Bashir Faris
577
S. I. Ali
1927
Blanche et al.
20
s. c.
1133
Davis
50243
Davis
50422
L. Boulos
1898
Tackholm etal.
s. n.
Z. Abou Raya
78
M. A. Haleem
s. n.
L. Boulos
10934
L. Boulos
1675
L. Bolous
1722
M. A. Siddiqi
99
Davis & Bolous
50584
J. Ramsay
4710
H. McAllister
20926
N. Jardine
667
s. c.
s. n.
Davis & Hedge
32718
Ben J. Hertage etal
s. n.
L. Vaccri
528
T. Tichler
s. n.
Dorfler
401
G. Argent
s. n.
John K Small
s. n.
A. Petrove
903
Herbarii
(ULT)
(E)
(E)
(E)
(ULT)
(E)
(E)
(E)
(E)
(ULT)
(E)
(E)
(E)
(ULT)
(E)
(ULT)
(ULT)
(ULT)
(ULT)
(ULT)
(ULT)
(E)
(E)
(ULT)
(E)
(ULT)
(ULT)
(ULT)
(ULT)
(E)
(ULT)
(E)
(E)
(E)
(ULT)
(ULT)
(ULT)
(E)
(ULT)
(ULT)
(ULT)
(ULT)
(GL)
(GL)
(E)
(E)
(E)
(E)
(E)
(k)
(E)
(E)
(K)
(K)
263
No
Taxon
Locality
Date
Collector
No
261
262
263
264
265
266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
289
290
291
292
293
294
295
296
297
298
298
299
300
301
302
303
304
305
306
307
308
309
310
311
312
313
314
M. califomica
M. campestris
M. capillacea
M. caroliniana
M. caroliniana
M. cerastifolia
M. colchica
M. confusa
M. dawsonensis
M.dawisonensis
M. dianthifolia
M. dichotoma
M. douglasis
M. douglasis
M. fasiculata
M. fasciculata
M. fasciculata
M. fasciculata
M. fasciculata
M. fasciculata
M. fasciculata
M. flaccida
M. flaccida
M. flaccida
M. formosa
M. formosa
M. formosa
M. funkii
M. funkii
M. funkii
M. geniculata
M. geniculata
M. geniculata
M. geniculata
M. geniculata
M. geniculata
M. geniculata
M. geniculata
M. geniculata
M. geniculata
M. geniculata
M. glabra
M. globulosa
M. graminifolia
M. graminifolia
M. groenlandica
M. hamata
M. hamata
M. hamata
M. hirsuta
M. hirsuta
M. hirsuta
M. howelii
M. hybrida
U. S. A.
Algeria
s. I.
U. S. A.
U. S. A.
Pyrenes
Caucasius
Italy
Canada
s. I.
Turkey
Spain
U. S. A.
U. S. A.
s. I.
s. I.
s. I.
Slovakia
s. I.
Italy
s. I.
Italy
Italy
France
Palaestine
Palaestine
Turkey
Spain
Morroco
Spain
Switzerland
Spain
Spain
Libya
Libya
Morocco
Libya
Somalia
Tunisia
Canary Island
Tunisia
U. S. A.
Greece
Italy
Italy
U. S. A.
Greece
Spain
Green land
Macedonia
Greece
Algeria
U. S. A.
s. I.
1866
1852
1877
1849
1891
1880
1977
1945
1957
1905
1947
1854
1905
1967
1886
1901
1925
1936
1949
1961
1962
1926
1961
1976
1911
1911
1969
1892
1923
1909 ‘
1837
1879
1888
1970
1970
1971
1972
1973
1975
s. d.
1984
1843
1857
1842
1898
1918
1855
1903
1992
1956
1977
1975
1928
1959
H. N. Bolander
Balansa
E. Reverchon
N. Ferscoh
Ben J. Heritage et al.
Timba Lagrave et al.
Meorugze
s. c.
Mrs Eva Beckett
J. Murr.
Davis
E. Bougeau
Louis Krautters
T. Voronova
Louis Keller
J. Murr.
Br. Bl.
J. Weher
P. Vermeulen
S. J. Van Ooststroom
P. Lizler
F. Sennen
S. J. Van Ooststroom
G. M. Lokhorstetal.
Fred S. Meyers
S. Meyers et al.
Davis
E. Reverchon
E. Jahandiez
F. Sennen
s. c.
s. c.
E. Reverchon
Davis
Davis
Davis
S. I. Ali
J. J. Lavranos
Davis & Lamond
O. Burchard
Davis
Torr& Gray
G. Orphanides et al.
G. Rigo
G. Rigo
M. L. Feranald et al.
s. c.
Province De Jaen
W. Burri & F. Rendl
K. H. Rechinger
E. A. Menneg
Davis
J. W. Thompson
Davis
4684 (K)
(E)
s. n.
(E)
5
(K)
s. n.
19105 (E)
(E)
s. n.
(K)
s. n.
(E)
s. n.
152
(E)
4849 (E)
694
(E)
2273 (E)
s. n.
(K)
6018 (L)
(E)
s. n.
4221 (E)
1145 (L)
934
(L)
5677 (L)
22908 (L)
s. n.
(L)
4372 (L)
22984 (L)
68
(L)
411
(E)
820
(E)
1034 (E)
s. n.
(E)
448
(BM)
855
(E)
s. n.
(E)
86
(E)
38
(E)
49772 (E)
50208.(E)
51315 (E)
(ULT)
780
10315 (E)
56920 (E)
288
(E)
60990 (E)
s. n.
(K)
930
(E)
4028 (E)
s. n.
(E)
(K)
358
s. n.
(E)
703
(E)
s. n.
(E)
17350 (E)
140
(E)
58974 (E)
4591 (K)
33345 (E)
Herba
No
Taxon
Locality
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
355
356
357
358
359
360
361
362
363
364
365
366
367
368
M. hybrida
Corsica
M. hybrida
Sicily
M. hybrida
Britain
M. hybrida
Britain
M. hybrida
Britain
M. hybrida
Britain
M. hybrida
Britain
M. hybrida
Algeria
M. hybrida
Britain
M. hybrida
Britain
M. hybrida
Britain
M. imbricata
Turkey
M. inamoena Caucasus
M. laricifolia
s. I.
M. laricifolia
s. I.
M. laricifolia
s. I.
M. mediterranea s. I.
M. mesogitana Turkey
M. mesogitana Turkey
M. moehringioides Mexico
M. moehringioides Mexico
M. montana
Algeria
M. montana
Spain
M. montana
Spain
M. montana
Spain
M. montana
Algeria
M. montana
Morocco
M. montana
Morocco
M. mutabilis
Italy
M. mutabilis
Algeria
M. mutabilis
Italy
M.obtusiloba Alaska
M. patula
U. S. A.
M. patula
U.S. A.
M. pestalozzae s. I.
M. picta
s. I.
M. picta
Palaestine
M. picta
Palaestine
M. picta
Cyprus
M. picta
Syria
M. picta
Jordan
M. recurva
Switzerland
M. recurva
s. I.
M. rimarum
Turkey
M. rossii
USSR
M. rostrata
Algeria
M. rubella
Britain
M. rubella
Britian
M. rupestris
Alps
M. saxifraga
Greece
M. sedoides
s. I.
M. sedoides
s. I.
M. sedoides
Swizerland
M. sedoides
s. I.
Date
Collector
1971
1979
1806
1869
1880
1881
1883
1937
1938
1949
1953
1963
1975
1835
1876
1898
1882
1967
1967
1890
1908
1852
1835
1888
1890
1891
1927
1929
1904
s. d.
s. d.
1979
1844
1967
1949
1835
1911
1911
1941
1943
1945
1873
1983
1963
1979
1938
1888
1930
1860
1978
1833
1881
1889
1951
M. Me Callum
Davis
s. c.
s. c.
H. Searle
R. M. Hay
L. Watt
A. Faure
J. Ramsay
T. Wise
E. C. Wallace
Davis & Hedge
Tarmngze
Dole
Karl Richter
Otto Krebs
L. Thalos
Davis
Davis
Pringle
C. A. Purpus
B. Balansa
Schimper
E. Reverchon
E. Reverchon
Warion
E. Jahandiez
E. Jahandiez
A. Beguinot
A. Faure
Adr. Fiori etal.
Marjorie Rees
Torr& Gray
T. Voronova
Davis
W. Schimper
Fred. S. Meyer et al.
s. c.
Davis
Davis
Davis
Johen Ball
M. A. Siddiqi
Davis & Hedge
V. Petrovsky
A. Faure
L. Watt.
G. Ghreag
J. Ball
W. Greuter
G. Stewart
A. Mermods
Archibald Dickson
Davis
No
Taxon
369
370
371
372
373
374
375
376
377
378
379
380
381
382
383
384
385
386
387
388
389
390
391
392
393
394
395
396
397
398
399
400
401
402
403
404
405
406
407
408
409
410
411
412
413
414
415
416
417
418
419
420
421
422
M. setacea
1962
Turkey
Greece
1937
M. stellata
Britain
M. stricta
1844
M. stricta
Britain
1884
Canada
M. tenella
1893
U.S. A.
1862
M. tenella
1894
M. thomesiana s. I.
M. umbellulifera Turkey
1949
M. verna
1863
s. I.
Britain
1869
M. verna
M. verna
1896
s. I.
M. verna
Morocco
1975
Spain
M. verna
1978
1979
M. verna
Sicily
M. viscosa
Italy
1930
M. viscosa
France
1974
M. viscosa
s. I.
1984
Moehiringia ciiiata Italy
1854
M. diversifolia China
1966
M. glaucovirens Italy
1844
Spain
M. intricata
1903
M. jankae
Bulgaria
1872
U.S. A.
M. lateriflora
1956
M. muscosa
s. I.
1876
Greece
M. pendula
1896
M. radiolata
1887
Thailand
M. sedifolia
France
1886
M. stellarioides Algeria
1898
1971
M. stellarioides Algeria
M. tejedensis Spain
1903
M. tommasinii Italy
1844
M. tommasinii Italy
1908
M. trinervia
Algeria
1971
M. trinervia
Algeria
1975
Paronychia arabica Wadi Al-Kouf 1967
P. arabica
1971
Algeria
P. arabica
1974
Tarhuna
P. arabica
T ripoli
1976
P. arabica
Wadi Sultan 1978
P. arabica
Wadi Ash Shati1978
B ugreen
P. arabica
1988
P. argentea
Algeria
1909
P. argentea
Algeria
1912
P. argentea
Messa
1972
Tokra
P. argentea
1973
T ripoli
1967
P. capitata
P. capitata
Tripoli
1970
1977
P. capitata
Tripoli
P. chlorothyrsa Morocco
1936
P. chlorothyrsa Morocco
1969
P. chlorothyrsa Garian
1973.
P. chlorothyrsa Garian
1974
P. kapela
France
1887
P. kapela
Algeria
1968
Locality
Date
Collector
Davis et al.
E. K. Ball etal.
J. Backhouse
H. S. Mennell
John Macoun
s. c.
A. Locatelli
Davis
Fraser
D. Stuart
Hausskno
Davis
Davis
Davis & D. S. Sutton
Erik Asplund
J. Vicherek
J. P. Theurillat
TH. Caruel
A. K. Schind.
P. Porta
E. Reverchon
s. c.
E. K. Balls
P. Chenevard
J. Dorfler
J. F. Maxwell
E. Reverchon
s. c.
Davis
Province De Jaen
P. Porta
C. Marchesetti
Davis
Davis
Bolous
Davis
K. Milad
S. M. Jafri
C. Rween
s. c.
A. El-Gadi
A. Faure
A. Faure
S. I. Ali
S. I. Ali et al.
Boulos et al.
Davis
Abdul Gafoor
E. K. Balls
Davis
S. I. Ali
El Jaly
E. R.everchon et. al
M. N. Chaudhri
No
Taxon
423
424
425
426
427
428
429
430
431
432
433
434
435
436
437
438
439
440
441
442
443
444
445
446
447
448
449
450
451
452
453
454
455
456
457
458
459
460
461
462
463
464
465
466
467
468
469
470
471
472
473
474
475
476
Petrorhagia illyrica Italy
P. illyrica
Abughilan
Pillyrica
Abughilan
P. illyrica
Bulgaria
P. velutina
Spain
P. velutina
Algeria
Polycarpaea carnosa Tenerife
P. carnosa
Tenerife
P. carnosa
Tenerife
P. divaricata
La Palma
P. divaricata
La Palma
P. repens
Algeria
P. repens
Morocco
P. robbairea
Iraq
P. robbairea Gado
P. robbairea
Sebha
P. robbairea Ghat
P. smithii
La Palma
P. tenuis
Tenerife
Polycarpon arabicum Palestine
P. arabicum
Palestine
P. bironae
Algeria
P. bivonae
Algeria
P. bivonae
Morocco
P. depressum U.S. A.
P. indica
Thailand
P. loeflingiae India
P. loeflingae Indo-China
P. peploides s. I.
P. peploides s. I.
P. polycarpoides Algeria
P. prostratum India
P. prostratum Nigeria
P. prostratum Leptus
P. prostratum Libya
P. succulentum Palaestine
P. succulentum Palaestine
P. succulentum Kuwait
P. tetraphyllum Australia
P. tetraphyllum U. S. A.
p. tetraphyllum Algeria
P. tetraphyllum West Indies
P. tetraphyllum Tenerife
P. tetraphyllum Leptus
P. tetraphyllum Shahat
Pteranthus dichotomus Mizda
P. dichotomus Morocco
Sagina apetala Australlia
S. apetala
Tripoli
S. apetala
Morocco
S. apetala
Afganistan
S. apetala
Tenerife
S. apetala
Tenerife
S. maritima
Libya
Locality
Date
Collector
1969
1975
s. d.
s. d.
1903
1938
1948
1948
s. d.
1944
1966
1982
1936
1957
1970
1973
1977
1966
1975
1897
1910
1854
1856
1929
1906
1887
1874
1891
1900
1903
1933
s. d.
1962
1968
1970
1846
1908
1981
1893
1906
1936
1942
1969
1970
1972
1977
1969
1890
1977
1929
1962
1981
1982
1970
s. c.
S. M. Jafri
S. I. Ali
C. Baenitz
E. Reverchon
A. Faure
E. R. Sventenius
E. R. Sventenius
E. R. Sventenius
E. R. Sventenius
s. c.
s.c.
E. K. Balls
K. H. Rechinger
Davis
S. I. Ali
M. A. Siddiqi
E. R. Sventenius
E. K. Sventenius
J. Bornmuller
s. c.
s. c.
s. c.
E. Jahandiez
G. B. Grant
J. F. Maxwell
J. Ball
B. Balansa et al.
H. Ross
C. Bucknall
A. Falure
S. Stainton et al.
J. T. Swarbrick
L. Bolous
Davis
Duchair
s. c.
s. c.
Alex Morrison
Harriet Walker
Herbier
L. R. Holdrige
D. Bramwell
Davis
S. I. Ali
M. A. Siddiqi
Davis
s. c.
A. Ghafoor
s. c.
s. c.
C. Rodriguez
J. Dickson
Davis
257
Locality
No
Taxon
477
478
479
480
481
482
483
484
485
486
487
488
489
490
491
492
493
494
495
496
497
498
499
500
501
502
503
504
505
506
507
508
509
510
511
512
513
514
515
516
517
518
519
520
521
522
523
524
525
526
527
528
529
530
Greece
S. maritima
Scleranthus annuus Britain
Scleranthus annuus Britain
Sclerocephalus arabicus Hun
Silene acaulis Britain
Britain
S. acaulis
S. aegyptiaca Lebanon
S.apetala
Algeria
Libya
S. apetala
Tripoli
S. apetala
Libya
S. apetala
Libya
S. apetala
Morocco
S. apetala
Tripoli
S. apetala
S. arenaroides Tunisia
Britain
S. armeria
Britain
S. armeria
S. articulata
Cyrenaica
S. articulata
Libya
S. articulata
Rasalhilal
S. atlantica
Algeria
S. behen
Libya
S. bellidifolia
s. I.
S. boryi
Spain
S. boryi
Morocco
S. boryi
Morocco
S. cerastoides Algeria
S. cerastoides Algeria
S. cerastoides Sabrata
S. claryi
Morocco
Italy
S. coelirosa
S. coelirosa
Morocco
Libya
S. colorata
S. colorata
Gargrese
S. colorata
Rass Hilal
S. colorata
Arabia
S. conica
Britain
Britain
S. conica
S. conica
Britain
S. conoidea
Spain
Tripoli
S. conoidea
Gossen
S. conoidea
Algeria
S. conoidea
S. conoidea
Saudi Arabia
S. corrugata
Morocco
Libya
S. cyrenaica
S. cyrenaica
W
adi M
ahbool
S. cyrenaica
Derna
Britain
S. dioica
Britain
S. dioica
Britain
S. dioica
S. disticha
Morocco
S. divaricata
Morocco
S. fruticosa
Libya
Date
Collector
No
Herbarit
1973
1890
1908
1973
1869
1949
1988
1936
1952
1933
1970
1970
1972
1977
19 77
1935
1931
1939
1981
1954
1916
1970
1819
1851
1973
1974
1910
1937
1976
1975
1898
1912
1939
1952
1979
1991
1869
1869
1907
1909
1969
1974
1975
1982
1972
1970
1973
1939
1939
1949
1985
1972
1901
1970
s. c.
L. Watt
G. B. Nielson
S. I. A.
D. Steuart
D. Patton
J. C. Archibald
Lieux
K. M. Guichard
s. c.
Davis
Davis & Boulos
I. B. R. Richardson
Abdul Gafoor
Davis
T. wise
W. Gourlie
s. c.
M. Bhadri & A. Ghafoor
K. M. Guichard
A. Faure
s. c.
s. c.
Bourgeau
Davis
s. c.
A. Faure
A. Faure
S. A. Jafri
Davis
B. Coll.
C. J. Pitard
N. Y. Sandwith
K. M. Guichard
Fauzy Ouheda
s. c.
D. Steuart
G. A. Arnott
G. B.
F. Sennen
A. Ghafoor
G. Faris
Davis
s. c.
Davis
Davis
S. I. Ali e ta l
N. D. Simpson
J. Walton
R. Wise
J. H. Dickson
Davis
C. J. Pitard
Davis
s. n.
1386
4359
1518
4589
230
s. n.
s. n.
1265
s. n.
49943
50587
454
280
61236
203
225
392662
7113
s. n.
s. n.
s. n.
s. n.
185
55370
849
121/92
s. n.
6711
58725
111
2695
2518
s. n.
s. n.
s. n.
4642
4643
4645
850
s. n.
373
58726
s. n.
54203
50228
1076
39299
4691
s. n.
34865
54311
s. n.
50473
(E)
(GL)
(GL)
(ULT)
(GO
(GL)
(BM)
(E)
(K)
(K)
(E)
(E)
(E)
(ULT)
(BM)
(GL)
(GL)
(BM)
(E)
(BM)
(BM)
(E)
(BM)
(BM)
(BM)
(BM)
(E)
(E)
(ULT)
(BM)
(BM)
(E)
(K)
(BM)
(ULT)
(E)
(GL)
(GL)
(GL)
(E)
(ULT)
(ULT)
(E)
(E)
(BM)
(BM)
(ULT)
(BM)
(G
L)
(GL)
(GL)
(E)
(BM)
(E)
No
Taxon
Locality
Date
Collector
531
532
533
534
535
536
537
538
539
540
541
542
543
544
545
546
547
548
549
550
551
452
453
454
555
556
557
558
559
560
561
562
563
564
565
566
567
568
569
570
571
572
573
574
575
576
577
578
579
580
581
582
583
584
S. fuscata
S. fuscata
S. gallica
S. gallica
S. gallica
S. gallica
S. gallica
S. gallica
S. gallica
S. ghiavensis
S. glabrescens
S. heterodonta
S. ibosii
S. imbricata
S. inaperta
S. italica
S. italica
S. italica
S. italica
S. kremeri
S. laeta
S. latifolia
S. latifolia
S. latifolia
S . littorea
S. longicaulis
S . longicilia
S. longipetala
S. longipetala
S . marmarica
S. marmarica
S. mekinensis
S. micropetala
S. micropetala
S. mollissima
S. muscipula
S. muscipula
S. muscipula
S. neglecta
S. neglecta
S. niceensis
S. noctiflora
S. noctiflora
S. noctiflora
S. nocturna
S. nocturna
S. nutans
S. nutans
S. nutans
S. obtusifolia
S. oropediorum
S. otites
S. otites
S. pomeli
Morocco
Algeria
Britain
Britain
Britain
Libya
Algeria
Tripoli
Saudi Arabia
Algeria
Morocco
Morocco
Morocco
Algeria
s. I.
Algeria
s. I.
Britain
Britain
Algeria
Spain
Britain
Britain
Britain
Spain
Portugal
Morocco
Turkey
Greece
Libya
Libya
Morocco
Spain
Spain
Morocco
Spain
Morocco
Morocco
Algeria
Algeria
Morocco
Britain
Britain
Britain
Algeria
Algeria
Britain
Britain
Britain
Morocco
Algeria
Britain
Britain
Algeria
1936
1937
1869
1931
1949
1970
1971
1978
1981
1939
1972
1973
1970
1975
1888
1975
1983
1987
s. d.
1880
1970
1860
1876
1958
1978
1853
1985
1957
1896
1913
1913
1929
1853
1889
1973
1890
1929
1988
s. d.
1910
1970
1869
1891
1903
1897
1936
1839
1880
1894
1974
1938
1878
1952
1975
E. K. Balls
A. Faure
G. A. Arnott
W. Gourlie
R. Wise
Davis
s. c.
A. Gafoor
s. c.
A. Faure
Davis
Davis
Davis
Davis
Abbe H. Coste
Davis
M. Diltice
G. Steven
D. Kent
E. Cosson
B. M. Allen
s. c.
P. Ewing
J. Walton
Davis
Boureau
J. Molero
s. c.
J. Dorfler
K. Musei Florentini
A.Vaccari
E. Jahandiez
E. Reverchon
E. Reverchon
Davis
Wilkoman
E. Jahandiez
s. c.
Davis
A. Faure
Davis
G. A. Arnott
R. Kidston
T. Wise
Elisee Re.
A. Faure
T. G. Rylands
L. watt.
S. Devon
B. M. Exped.
A. Faure
R. Kidston
B. W. Ribbons
Davis
No
Taxon
585
586
587
588
589
590
591
592
593
594
595
596
597
598
599.
600
601.
602
603
604
605
606
607
608.
609
610
611
612
613
614
615
616
617
618
619
620
621
622
623
624
625
626
627
628
629
630
631
632
633
634
635
636
637
638.
S. pomeli
S. protensis
Locality
Date
Morocco
s. d.
Morocco
1974
S. pseudoatiocion Algeria
1975
S. ramosissima Algeria
1852
S. reticulata
Algeria
1975
S. rubella
Tunisia
1975
S. rubella
Morocco
1962
S. scabriflora
Morocco
1929
S. secundiflora Algeria
1975
S. sedoides
Italy
1840
Greece
S. sedoides
s. d.
Italy
S. sericea
s. d.
S. sericea
Europe
1904
S. stricta
Algeria
1910
S. succulenta Dariana
1972
S. succulenta Lulida
1975
S. succulenta Aulad Mohamed 1977
Tunisia
S. tridentata
1975
S. tridentata
Morocco
1952
S. tuberculata Morocco
1927
Britain
S. uniflora
1875
S. villosa
Algeria
1938
S. villosa
Morocco
1969
S. villosa
Sebha
1973
S. villosa
Saudi Arabia
1991
S. vivianii
Morocco
1936
S. vivianii
Egypt
1945
S . vulgaris
Britain
1869
S. vulgaris
Britain
1896
S. vulgaris
Britain
1954
S. vulgaris
Italy
1970
Britain
S. vulgaris
1983
Britain
S.vulgairs
1985
Spergula arvensis Atlantic Islands 1891
S. arvensis
China
1965
S. arvensis
Australia
1939
Japan
S. arvensis
1957
S. arvensis
North Asia
1965
S. arvensis
s. I.
1967
S. arvensis
Portugal
1971
s. I.
S. arvensis
s. d.
S. arvensis
South Africa
1973
S. arvensis
Tanzania
1989
S. fallax
Palestine
1935
S. fallax
Sharshara
1966
S. fallax
Libya
1970
S. morisonii
s. I.
1891
S. morisonii
Berlin
1900
S. pentandra
Morocco
1923
Kashmir
S. pentandra
1956
Asia
S . pentandra
1877
Spain
S. viscosa
1978
Spergularia bocconei Chile
1924
Libya
S. bocconei
1970
Collector
Jamais
B. M. Exped.
Davis
B. Balansa
Davis
s. c.
J. C. Archibald
E. Jahandiez
Davis
s. c.
Th. Kaiis
s. c.
J. W. White
A. Fame
S. I. Ali
A. Fuzi
M. A. Siddiqi
Davis
D. H. Spence
s. c.
R. Kidston
A. Faure
Davis
S. I. Ali et al.
s. c.
E. K. Balls
s. c.
D. Steuart
J. B. Nielson
J. Walton
J. Damblon
J. Dickson
J. Smith et al
J. F. Hamilton
A. K. Schind.
Milos Deyl
Markino
M. Mizushima
Sumike Kobayashi
Davis
Chris Parker
O. M. Hilliard
C. M. Taylor et al.
Amdursky et al.
s. c.
Davis
John Ball
s. c.
s. c.
O. Polunin
J. Ball
s. c.
E. Werdermann
Davis
No
Taxon
639
640
641
642
643
644
645
646
647
648
649
650
651
652
653
654
655
656
657
658
659
660
661
662
663
664
665
666
667
668
669
670
S. bocconei
S. bocconei
S. cerastoides
S. diandra
S. diandra
S. diandra
S. fimbriata
S. grandis
S. levis
S. maritima
S. maritima
S. maritima
S. nicaeensis
S. ramosa
S. rubra
S. rubra
S.saiina
S. salina
S. salina
S. salina
Locality
Date
1992
Chile
Algeria
1989
1905
Chile
1928
Asia
1967
Egypt
Azizia
1970
s. I.
1845
Brazil
s. d.
Australia
1964
Algeria
1934
Tarhona
1976
W
adi Alshati 1978
Iran
1974
s. I.
1831
Ras al Hilal 1972
Talil
1976
1924
Asia
Askadda
1973
Tripoli
1977
Tripoli
1991
Stellaria cupaniana Corfu
1993
S. media
Tenerife
1981
S. media
Tripoli
1993
Britain
S. media
1950
S. media
Tripoli
1992
S. pallida
Tripoli
1992
S. pallida
Tripoli
1992
S. pallida
Ganzoor
1992
Telephium sphaerospermum Derna1967
Vaccaria pyrmidata Jefren
1974
V. pyramidata Gabel Akhder 1978
V. pyramidata Gusbat
1979
Collector
M. E. Gardner etal.
s. c.
Otto Buchtien
Popor et Vredensky
Tackholm etal.
s. c.
s. c.
s. c.
E. M. Barron
Sebkhas
A. Gafoor et al.
s. c.
Davis & Bokhari
Charles Darwin
S. I. Ali
S. M. Jafri
Kultiassov
S. I. Ali
Abdul Gafooor
s. c.
J. Dickson
C. Rodriguez
M. Magrabi
J. Riddell
W. Fashloom
M. Magrabi
S. Fashloom
W. Fashloom
L. Boulos
S. El. Jaly
A. Gafoor
El-Gadi
271
Appendix III.
An artificial key to the wild Libyan species of Caryophyllaceae
based only on seed characters. Only three species of Caryophyllaceae
(including lllecebraceae) are omitted. No ripe seeds of Dianthus serrulatus,
Petrorhagia illyrica and Silene biappendiculata were readily available.
1A. Lateral faces of seeds smooth, cells faint or indistinct, seeds without
wings.
2A. Seeds round-reniform................................................... Silene succulenta.
2B. Not so.
3A. Seeds curved-oblong or cuneate.
4A. Oblong-curved (crescent shaped), surface wrinkled..................................
...............................................................................Polycarpaea repens.
4B. Cuneate, surface smooth.
5A.Dense papillae surrounding the radicle base and dorsal side...................
.................................................................................Loeflingia hispanica.
5B. Few papillae surrounding the radicle only
Polycarpaea robbairea.
3B. Seeds obovate, circular-obovate or broadly elliptic.
6A. Micropyle pointed or compressed laterally.
7A. Keel strongly compressed laterally, hilum surrounded by faint cells........
..................................................................................Paronychia kapela.
7B. Keel slightly compressed, hilum surrounded by distinct cells....................
................................................................................ Paronychia capitata.
8A. Space between radicle and hilum shallow
8B. Space between radicle and hilum deep
Paronychia chlorothyrsa.
Herniaria cyrenaica.
6B. Micropyle blunt circular.
9A. Hilum very close to radicle.........................................Paronychia argentea.
272
9B. Hilum distinctly separated from radicle.
10A. Deep cavity between hilum and radicle.
11 A. Micropyle filled with indistinct thread like structure, collar cells faint......
Paronychia arabica.
11B. Micropyle filled with distinct cells mostly curved elongate ( star
shaped), collar cells faint.........................................Herniaria fontanesii.
10B. Shallow depression between hilum and radicle.
12A. Radicle discoid but the hilum strongly compressed laterally.....................
..............................................................................Herniaria hemistemon.
12B. Radicle and hilum with discoid shape.
13A. Hilum surrounded by double collar of special rectangular cells..............
.......................................................................................Herniaria cinera.
13B. Hilum ended by smooth dome shape.
14A Cells surrounding the micropyle small rectangular with distinct walls
.................................................................................... Herniaria ericifolia.
14B. Cells surrounding the micropyle large with faint or indistinct
walls...............................................................................Herniaria glabra.
1B. Lateral faces seeds rough (or if smooth cells distinct), cells faint or
distinctly protruding, seeds with or without wings.
15A. Seeds globular or subglobular.
16A. Lateral face cells mostly spherical, each cell covered with dense
granules......................................... Telephium sphaerospermum.
16B. Lateral face cells polygonal with blunt sinuate margins,
papillate.....................................................................Vaccaria pyrmidata.
15B. Seed shapes various often reniform but not globular.
17A. Radicle ± straight.
18A. Seed light brown or faint yellow.
19A.Seeds light brown, obovate-reniform, lateral faces flat, slightly slantes
273
radicle gradually tapering to acute apex... Sclerocepahalus arabicus.
19B. Seeds faint yellow, elongate-obovate, dorsally convex, radicle
flatten, broaded round at apex.......................... Pteranthus dichotomus.
18B. Seed black.
20A. Seed obovate, boat shaped
Petrorhagia velutina.
20B. Seed obovate, plane.
21 A. Dorsal face cells narrow, elongate, arranged in regular rows, spurs
regular undulate with blunt apices
Dianthus crinitus.
21B. Dorsal face cells narrow elongate, arranged irregularly, spurs with
irregular shapes and sizes........................................Petrorhagia illyrica.
MB. Radicle slightly or strongly curved.
22 A. Seeds with membranous wings.
23A. Wings membranous, mostly entire, spurs regularly spaced, ending in
pore-1ike epression..........................................................Spergula fallax.
23B. Not so.
24A. Seeds winged, wings margins slightlysinuate, lateral face cells with
irregular jagsaw shape......................................... Spergularia maritima.
24B. Seeds winged or not, wings erose to lanciniate, lateral face cells
elongate or ovate.................................................................. Spergularia salina.
22B. Seed without membranous wings.
25A. Seeds obovate, with shallow or deep groove separating the lateral
and marginal faces.
26A. Radicle long, strongly curved and compressed at apex, lateral faces,
slightly concave................................................ Gymnocarpos decander.
26B. Radicle short, slightly curved,longer than cotyledons, lateral faces
slightly convex.
27A. Shallow groove between lateral and marginal faces, cells distinct, with
jagsaw shape, raised into blunt papillae, spurs few.
274
.................................................................................Spergularia diandra.
27B. Deep groove between lateral and maginal faces, cells mostly
indistinct, elongate, raised into discoid papillae, spurs many.
28A.Radicle thick, broad, round, c. 70 pm wide near apex..............................
........................................................................... Spergularia rubra.
28B. Radicle compressed laterally, slightly tapering, c. 30 pm wide near
apex........................................................................ Spergularia bocconii.
25B. Seeds not obovate, lacking grooves between lateral and marginal
faces.
29A. Seeds mostly retortiform, radicle ± strongly curved, like a hook.
30A. Mid-zone cells mostly long, narrowly elongate.
31A.Seed retortiform-elliptic.................................................. Minuartia hybrida.
31B. Seed reniform to retortiform...............................Minuartia mediterranea.
30B. Mid-zone cells mixed short or elongate, elliptic or ovate.
32A. Hilar notch area covered mostly with dense small warts..........................
.................................................................................Minuartia geniculata.
32B. Hilar notch area covered mostly with few distinct papillae.
33A. Seed retortiform, radicle cells 13-54 pm long............................................
Minuartia campestris.
33B. Seed round -reniform,radicle cells 33-65 pm long.....................................
...................................................................................Minuartia montana.
29B. Seed mostly cuneate or reniform, radicle± slightly incurved or
equalling cotyledons.
34A. Seed cuneate.
35A. Lateral face cells without spurs.
36A. Papillae mostly of regular size and dense,not on clearly defined raised
bases....................................................................Polycarpon prostratum.
275
36B. Papillae mostly of irregular size and less dense, many with raised
bases.................................................................Polycarpon tetraphyllum.
35B. Lateral face cells with distinct spurs.
37A. Lateral face cells mostly round or mixed stelliform, elliptic or elongate.
38A. Lateral face flat slightly concave................................ Cerastium siculum.
38B. Lateral face slightly or strongly convex
39A. Lateral face cells mostly round,raised into sharp or blunt, smooth
papillae.............................................................. Cerastium dichotomum.
39B. Lateral face cells mostly elongate or elliptic,raised into blunt,
granular papillae ........................................ Cerastium semidecandrum.
37B. Lateral face cells mostly narrow-elongate.
40A. Lateral faces cells straight, in regular rows, raised into sharp ridges,
cell length 8-10 times that of the spurs.................. Cerastium illyricum.
40B. Lateral face cells straight or curved, regular or irregular, raised into
blunt or conical papillae, cell length 4-5 times that of the spurs.
41 A. Lateral face cells arranged in c. 4 regular rows, cells straight or
curved, spurs arranged ± pinnately.................. Cerastium glomeratum.
41B. Lateral face cells irregular, slightly curved, spurs not appearing
pinnate.
42A. Lateral face raised into narrow or broad cells that broader than spur,
marginal face with short conical tubercles ± 16 pm
length..................................................................... Cerastium ligusticum.
42B. Lateral face raised into narrow cells ± equal spurs in thickness,
marginal face with long conical tubercles ± 35 pm length
.................................................................................. Cerastium pumilum.
34B. Seeds reniform.
43A. Radical ± equaling cotyledons.
276
44A. Lateral faces deeply concave.
45A. Mid-zone spurs undulate or indistinct.
46A. Lateral face cells broad, elliptic,raised into 2-3 warts, pads large and
globular ........................................................................... Silene gallica.
46B. Lateral face cells narrow; elongate or elliptic, raised up to 8 warts;
pads flat
Silene tridentata.
45B. Mid-zone spurs distinct, markedly tapering.
47A. Mid-zone cells covered with many round warts, pads scarcely
distinct........................................................................Silene cerastoides.
47B. Mid-zone cells covered with many inconspicuous and a few
conspicuous warts; pads distinct with elongate or elliptic cells.
48A. Seed broad-reniform + 0.9
mm long, ±
0.5 mm wide,lateral faces
cavity ±0.57 mm long, ± 0.43 mm wide, mid-zone cells 40-185 pm
long, pads cells rising up
into conical
48B. Seed round-reniform ± 0.8 mm long, ±
papillae.......... Silene rubella.
0.4 mm wide,lateral face cavity
± 0.44 mm long, ± 0.25 mm wide, mid-zone cells 41-119 pm long,
pads cells smooth without papillae............................ Silene nocturna.
44B. Lateral faces plane or slightly concave.
49A. Seeds with large undulate wings.
50A. Mid-zone cells with 1 to several warts on each.
51 A. Mid-zone area cells nearest the hilar notch with only one wart
each......................................................... Silene apetala. Morphotype A.
51B. Mid-zone cells with more than one papilla or wart on each.
52A. Mid-zone cells with an elongate papillae and small warts, which can
be acute,conical or blunt....................... Silene apetala Morphotype C.
52B. Mid-zone cells with irregular round or blunt warts
.................................................................. Silene apetala Morphotype B.
52C. Mid-zone cells with warts mostly arranged in row s
Silene vivianii.
277
50B. Mid-zone cells smooth or rarely with blunt warts.
53A. All mid-zone cells with distinct margins and blunt or sharp
spurs........................................................ Silene colorata var. lasicalyx.
53B. Mid-zone cell margins straight or slightly undulate.
54A. Cells of wings distinct and clearly in 3 regular rows.................................
.....................................................................................Silene cyrenaica.
54B. Cells of wings less distinct and not clearly in rows.
55A. Mid-zone cells narrow, c. 14 pm wide.......................... Silene articulata.
55B. Mid-zone cells narrow, c. 8 pm wide..........................................................
.......................................................
Siienecoloratasubsp.colorata.
49B. Seeds without wings.
56A. Seeds triangular-reniform.
57A. Seeds > 2 mm long, lateral and marginal face cells raised into long
papillae and tubercles, spurs indistinct....Agrostemma githago.
57B. Seeds < 1 mm long, lateral and marginal face cells raised into blunt
papillae and tubercles, spurs regular spaced or cogwheel like sharp
apex
Sagina maritima.
56B. Seeds elongate-reniform or round.
58A. Seeds elongate-reniform, lateral face cells few ±30................................
.......................................................................................Sagina apetala.
58B. Seeds round-reniform, lateral face cells many.
59A. Lateral faces ± plane, marginal face ± concave.
60A. Lateral face cells rising up into distinct globular or elongate papillae.
61 A. Lateral face cells raised into uniform, globular papillae, arranged in
concentric raws................................................................ Silene behen.
61B. Lateral face cells raised into irregularly shaped, even elongate
papillae, one or two papillae per cell not clearly concentric.............
............................................................................................Silene Mica.
278
60B. Lateral faces plane, with cells lacking papillae (or only a few
inconspicuous papillae) .
62A. Lateral face cells smooth or slightly wrinkled (lacking distinct
granules).........................................
Silene marmarica.
62B. Lateral face cells distinctly granular.
63A. Seeds < 1 mm, spurs granular..........
.Silene muscipula.
63B. Seeds > 1mm, spurs smooth............
Silene longipetala.
59B. Lateral faces mostly convex, marginal faces not or slightly concave.
64A. Lateral face cells raised into regularly arranged papillae or warts.
65A. One wart, always situated at one apex, only three pad cells, flat.........
Silene sedoides.
65B. One papilla, large and globular, arranged in the middle of each cell,
pads large and with many cells...................................... Silene vulgaris.
64B. Lateral face cells lacking clearly defined papillae..
66A. Lateral face cells mixed ovate, elliptic or round, spurs lacking, pad
cells similar to mid-zone cells........................................... Silene villosa.
66B. Lateral face cells elliptic, elongate or rarely ovate, spurs normal with
sharp apices, pad cells plane or globular different from the other
mid-zone cells.
67A. Pads globular, mid-zone cells broad-elliptic or ovate................................
Silene conoidea.
67B. Pads plane, mid-zone cells narrow elliptic or elongate...Silene fuscata.
43B. Radicle longer than cotyledons slightly or strongly curved.
68A. Marginal face raised into long conical tubercles.
69A. Lateral face cells stelliform, with long spurs each bearing 1-4
warts,
Stellaria media.
69B. Not so.
70A. Lateral face concave
Gypsophila pilosa.
279
70B. Lateral face plane or slightly convex .
71 A. Seed size > 1mm.......................................................Gypsophila elegans.
71B. Seed size < 1mm............................ Arenaria leptoclados (Davis 50344).
68B. Marginal faces raised into short blunt tubercles.
71 A. Several lateral face, particularly midzone, cells broad, some almost
isodiametric............................................................ Arenaria serpyllifolia.
71B. Lateral face, including mid-zone, cells all ± elongate, 3 or more as
long as wide........................................................... Arenaria leptoclados.
280
A p p e n d ix
IV.
An artificial key to Libyan species of S i l e n e
using fruiting plants. Only one species, Silene biappendiculata,
was omitted; there was no ripe capsule.
1A. Calyx 10 veined.
2A. Capsule epidermal cells not papillate.
3A. Calyx (6-7) mm long; carpophore ± 1mm long.......... S. sedoides
3B. Calyx (15-27) mm long; carpohore (5-10) mm long.
4A. Calyx (15-20) mm long.
5A. Testa cells faint
S. succulenta
5B. Testa cells with distinctive papillae.
6A. Seed size ± .8 mm long............................................................ S. villosa
6B. Seed size ± 1.4 mm long........................................................S. fruticosa
4B. Calyx (23-27) mm long....................................................... S. marmarica
2B. Capsule epidermal cells papillate.
7A. Capsule epidermal cells papillae mostly scattered
irre g u la rly .
8A. Seeds not winged.
9A. Seeds slightly slanted near midzone; carpophore (5-7)
mm long.................................................................................. S. fuscata
9B. Seeds deeply grooved near midzone; carpophore (2-3.5)
mm long
8B. Seeds winged.
10A. Testa cells covered with papillae near midzone;
S. rubella
281
carpophore ± 1.5 mm long................................................... S. apetala
10B. Testa cells lacking papillae near midzone;
carpophore ± (3-9) mm long
11 A. Carpophore 3mm or less long......................................... S. articulata
11B. Carpophore more than 4 mm long.
12A. Calyx appressed hairy throughout; seeds ± 1.2 mm long............
........................................................................................... S. cyrenaica
12B. Calyx appressed hairy on nerves alone; seeds ± 1.5 mm long..
................................................................................................S. colorata
7B. Capsule epidermal cells mostly arranged in rows.
13A. Capsule epidermal cells walls distinctive or faint,
papillae
closely spaced or fused in lines.
14A. Seeds slightly slanted, 3-4 regular rows of acute conical
papillae....................................................................................S. behen
14B. Seeds plane or slightly slanted, papillae mostly irregular.
15A. Seed epidermal cells narrow elongate, raising into many
small warts, spurs blunt....................................................... S. viviani
15B.
Seed epidermal cells elongate orelliptic, raising
into few
papillae or warts, spurs acute.
16A. Seed size 1-1.5 mm long........................................................S. italica
16B. Seed size 1.8-2.2 mm lo n g
S. longipetala
13B. Capsule epidermal cells walls distinctive, papillae arranged
on thick ridge.
17A. Calyx contracted with capsule in fruit.
2 82
18A. Calyx apically contracted teeth acuminate or linear 4-5
mm long; carpophore 1-2 mm long............................ S. tridentata
18B. Calyx contracted both above and below capsule, teeth
linear-lanceolate acute teeth, 3-3.5 mm long, carpophore
2-4 mm long
S.cerastioides
17B. Calyx not contracted with capsule in fruit.
19A. Seed lateral faces plane or slightly slanted
S. muscipula
19B. Seed lateral faces deeply grooved.
20A. Seeds brown, lateral face cells withmany blunt
warts, marginal face grooved
or distinct
S. nocturna
20B. Seeds dark brown to black, lateral face cells raised into 23 distinct warts, marginal face plane or slightly shallow
..................................................................................................S. gallica
1B. Calyx 15-30 veined.
21A. Calyx ± 15-20 veined, glabrous; capsule with wide neck,
capsule epidermal cells very variable in shape.............................
................................................................................................ S. vulgaris
21B. Calyx ± 30 veined, glandular hairy; capsule with long narrow
neck, capsule epidermal cells constant in shape..........................
.............................................................................................. S. conoidea
283
Appendix IV. An artificial key for the identification of the subgenera,
sections and series of the genera Arenaria, Moehringia and Minuartia has
been constructed using the crystal shape and distribution.
1A. Leaves without crystals, or very rare in small groups, scattered
irregularly throughout the leaves.
Arenaria
VIII. Subgenus Dolophragma (a
denissima, A. oreophila,
A. polytrichioides)
IX. Subgenus Solitaria {A. ciiiolata, A. forrestii)
1B. Leaves with crystals few, dense or very dense, scattered irregularly or
regularly in rows throughout intercostal and veins or just on veins.
2A. Crystals scattered irregularly throughout the leaf except veins.
3A. Druses with sharp points, [see 3B and 3C]
Genus A re n a ria .
I. Subgenus Leiosperma (A. aisinoides,A. guatemalensis,
A. lanuginosa, A. reptans).
II. Subgenus Dicranilla {A. pycnophylla).
IV. Subgenus Arenaria
A. Sectio Rariflorae {A. ciliata,A. humifusa,A. pseudofrigida).
B. Sectio Grandiflorae(A grandiflora, A. incrassata).
C. Sectioi Plinthine (A. lithops, A. tetraquetra).
D. Sectio Rotudifoliae {A. halacsyi).
E. Sectio Planosepalae (A. montana).
F. Sectio Orientales
F. (ii) Series Graecae (A. filicaulis, A. teddii)
F. (iii). Series Deflexae (A. deflexa).
284
F. (iv). Series Hispidae {A. retusa, A. rhodia)
G. Sectio Pseudosabulina (A. sabulinea).
J. Sectio Africanae
J. (i). Series Africanae
J. (ii). Series Papillospermae {A. hispanica).
K. Sectio Arenaria
K. (i). Series Arenaria (A. conferta, A. leptoclados, A. serpyllifolia).
K. (ii). Series Saponarioides (A. saponarioides).
K. (iii). Series Cylindricae {A. guicciardii).
V. Subgenus Arenariastrum {A. gouffeia).
X. Subgenus Odontostemma (A. trichophora, A. napuligera,
A. yunnanensis).
M oehringia
B. Sectio Latifolae (M. trinervia, M. lateriflora, M. radiolata).
C. Sectio Diversifolia (M. diversifolia, M. jankae, M. pendula).
D. Sectio Moehringia (M. glaucovirens).
M inuartia
IV. Subgenus Minuartia
A. Sectio Spectabiles
A. a. Subsectio Spectabilies
A. a. (i). Series Laricinae (M. colochia, M. imbricata,
M. inamoena).
A. b. Subsectio Cherleria (M. sedoides).
3B. Druses with blunt points.
A renaria
I. Subgenus Leiosperma (A. paludicola).
II. Subgenus Dicranilla (A. boliviana).
285
IV. Subgenus Arenaria
C. Sectio Plinthine (A. armerina)
D. Sectio Rotundifoliae (A. biflora).
H. Sectio Occidentales (A. conbricensis, A. ciliaris).
VI. Subgenus Eremogoneastrum (A. franklinii, A. hookeri,
A. festucoides)
M oehringia
A. Sectio. Pseudomoehringia (M. intricata, M. tejedensis).
D. Sectio. Moehringia (M. muscosa, M. sedifolia, M. tommasinii).
Minuartia
II. Subgenus Spergella (M. formosa, M. picta) in M. picta are
druses or elongate
III. Subgenus Hymemella (M. moehringioides).
IV. Subgenus Minuartia
A. Sectio Spectabiles
A.a. (ii) Series Spectabiles [New name Series Biflorae]
(M. arctica, M. biflora, M. obtusiloba).
E. Sectio Sclerophylla (M. caroliniana).
J. Sectio Uninerviae (M. brevifolia, M. glabra, M. groenlandica,
M. patula).
3C. Druses with sharp and blunt points.
Arenaria
I. Subgenus Leiosperma (A. decussata, A. lanuginosa).
IX. Subgenus Solitaria {A. ciliolata).
M inuartia
I. Subgenus Rhodalsine (M. geniculata).
IV. Subgenus Minuartia
286
A. Sectio Spectabiles
A. C. Subsectio Laricifoliae
A. C. (i). Series Caucasicae (M. aizoides).
2B. Crystals scattered throughout leaves including veins or just on veins.
4A. Crystals throughout leaves including veins.
5A. Druses, crystal sands but not elongate.
6A. Druses with sharp and blunt points.
A renaria
VII. Subgenus Eremogone
H. Secto Pungentes (A. pungens).
M inuartia
IV. Subgenus Minuartia
A. Sectio Spectabiles
A. c. Subsectio Laricifoliae
A. c. (i). Series Caucasicae (M. baldaccii, M. capillacea,
M.laricifolia).
6B. Druses with blunt points.
7A. Dense crystals near leaf base.
A renaria
VII. Subgenus Eremogone
A. Sectio Capillares (A. capillares, A. fendleri, A. lychnidea).
F. Sectio Scariosae
F. (i). Series Polycnemifoliae (A. polycnemifoiia, A.
pseudacantholimon, A. zargariana).
F. (ii). Series Scariosa (A. armeniaca, A. scariosa).
G. Sectio Sclerophyllae (A. acerosa, A. aculeata, A. acutisepala,
A. davisii, A. drypidea, A. griffithii, A. insignis, A. kingii,
A. ledebouriana)
287
7B. No crystals near leaf base.
A renaria
VII. Subgenus Eremogone
C. Sectio Eremogone {A. steveniana, A. graminea, A. koriniana, A.
macradenia).
G. Sectio Sclerophyllae (A. macradenia, A. persica, A. tetrasticha)
5B. Druses, crystal sands and elongate.
Arenaria
VII. Subgenus Eremogone
D. Secto Glomeriflorae (A. gypsophiloides).
M inuartia
IV. Subgenus Minuartia
B. Sectio Plurinerviae (M. recurva).
C. Sectio Lanceolatae
C. (i). Series Graminifoliae (M. gramminifolia, M. saxifraga,
M. stellate).
C. (ii). Series Dianthifolia ( M. dianthifolia, M. acuminata, M.
pestalozzae).
C. (iii). Series Lanceolatae ( M. cerasitifolia, M. rupestris).
E. Sectio Sclerophylla (M. dawsonensis).
F. Sectio Acutiflorae
F. (iii) Series Umbelluiferae (M. umbellulifera).
H. Sectio Alsinathe (M. rossii, M. stricta)
K. Sectio Greniera (M. douglasii, M. howellii).
L. Sectio Minuartia
L. a. (i). Series Montanae (M. montana, M. globulosa).
L. a. (ii). Series Minuaria (M. dichotoma, M. hamata).
L. b. Subsectio Xeralsine
288
L. b. (ii). Series Setaceae (M. anatolica, M. confusa, M. mutabilis)
L. b. (iii). Series Xeralsine (M. fasciculata, M. funkii).
L. b. (IV). Series Cam pestres (M. campestris).
M. Sectio S abulina
M. (i). Series Sabulina (M. mesogitana, M. tenella, M. urumiensis).
M. (ii) Series Californicae (M. californica).
4B. Crystals in veins only.
8A.Crystals mostly round ovate with blunt points but not elongate.
VII. Subgenus Eremogone
E. Sectio R igidae
E. (i). Series Rigidae {A. holostea, A. szowitsii).
M inuartia
IV. Subgenus Minuartia
D. Sectio Aretioideae (M. aretioides).
B. Sectio Plurinerviae (M. bulgarica, M. hirsuta).
F. Sectio Acutiflorae
F. (ii). Series Pichleriae (M. rimarium).
G. Sectio Tryphane (M. rubella).
8B. Crystals mixed, mostly elongate.
Arenaria
VII. Subgenus Erem ogone.
D. Sectio Glomeriflorae (A.dianthoides, A.cucubaloides,
A. gypsophiloides).
Minuartia
F. Sectio A cu tiflorae
F. (i). Series Acutiflorae [New name Series Flaccidea].
(M.austriaca, M. flaccida).
G. Sectio Tryphane (M. verna).
IV. Subgenus Minuartia.
M. Secto Sabulina
M. (i). Series Sabulina (M. hybrids, M. mediterranea).
Mohamed Abuhadra