American Journal of Plant Sciences, 2011, 2, 120-133
doi:10.4236/ajps.2011.22014 Published Online June 2011 (http://www.SciRP.org/journal/ajps)
Pollen Morphology of Tribes Gnaphalieae,
Helenieae, Plucheeae and Senecioneae (Subfamily
Asteroideae) of Compositae from Egypt
Ahmed Kamal El-Deen Osman
Faculty of Science, Botany Department, South Valley University, Qena, Egypt.
Received October 10th, 2010; revised December 9th, 2010; accepted December 20th, 2010.
ABSTRACT
POLLEN morphology of twenty five species representing 12 genera of tribes Gnaphalieae, Helenieae, Plucheeae and
Senecioneae (Asteroideae: Asteraceae) was investigated using ligh t and scanning electron microscopy. The genera are
Phagnalon, Filago, Gnaphalium, Helichrysum, Homognaphalium, Ifloga, Lasiopogon, Pseudognaphalium, Flaveria,
Tagetes, Sphaeranthus an d Senecio. Two pollen types were recognized viz. Senecio pollen type and Filago pollen type.
Description of each type, a key to the investigated taxa as well as LM and SEM micrographs of pollen grains are pro-
vided.
Keywords: Pollen, Morphology, Asteroideae, Asteraceae, Egypt
1. Introduction
Gnaphalieae, Helenieae, Plucheeae and Senecioneae (As-
teroideae: Asteraceae) are of the well represented tribes
in Egypt, where 12 genera with about thirty five species
are native in the country [1]. Reference [2] separated the
Compositae pollen into two groups based on the gross
morphology of the pollen grain: the liguliflorae-type
characterized by echinolophate pollen, which is generally
found in the Cichorieae; and the tubuliflorae- type, which
is found in many other tribes. Reference [3] recognized
three pollen types, i.e., psilate (this type can not be easily
recognized), echinate, and lophate pollen. Based on pol-
len wall ultrastructure, [4] and [5] reconsidered these
pollen types and recognized five major pollen wall types,
the Helianthoid, the Senecioid, the Arctotoid, the An-
themoid and pollen types without designated patterns.
These pollen types are variously distributed among the
Cichorioideae and the Asteroideae. Pollen of the Barna-
sioideae is similar to the echinolophate pollen of the
liguliflorae-type, but lacks the spines [6]. Reference [7]
observed that the pollen grains of some Compositae
tribes consist of three compound apertures, each one of
them is made up of ecto-, meso- and endo-aperture, the
apertural membrane is formed by a foot layer and
endexine with complete ectexine granules, the ectoaper-
ture involves the tectum and infratectum, the mesoaper-
ture involves the foot layer and the outer layer of the
endexine and the endoaperture involves the inner layer of
the endoxine. The intine is thickened considerably in
Anthemideae near the aperture. Reference [8] described
Gnaphalieae pollen as a two-layered pollen sexine with
an outer baculae and an inner perforated layer. Reference
[9,10] observed that the pollen morphology and anatomy
of the Plucheeae correspond to that of the Inuleae; thus,
the pollen is echinate and caveate with one layer of
baculae between the spines and two baculate layer in the
spines bases. This paper considers pollen grains of the
tribes Gnaphalieae, Helenieae, Plucheeae and Senecioneae
in the flora of Egypt, which may prove of value in sys-
tematic treatments. Pollen types that show all the possi-
ble characteristic features (shape, size, apertures, wall
stratification) are considered, with special reference to
the specific characters of each pollen type. The available
palynological data will be discussed.
2. Materials and Methods
Pollen materials were removed from herbarium speci-
mens identified according to [1,11,12] and [13]. The
specimens were kept in Cairo University Herbarium
(CAI) and Herbarium of South Valley University (QNA,
proposed abbreviation) (Table 1). Light microscopy (LM)
observations were carried out on acetolyzed pollen and

Pollen Morphology of Tribes Gnaphalieae, Helenieae, Plucheeae and Senecioneae (SubfamilyAsteroideae) 121
of Compositae from Egypt
Table 1. Investigated specimens of 25 species (12 genera) of Gnaphalieae, Helenieae, Plucheeae & Senecioneae (Asteraceae)
according to [14].
Species Collector Herb.
Subfamily Asteroideae
I-Tribe Gnaphalieae
1. Unassigned Gnaphalieae
Phagnalon ba rbey a num EL-GARF & A.K.OSMAN, s.n. QNA
P. nitidum FAYED et al. s.n. QNA
P. rupestre EL-GARF& A.K.OSMAN, s.n. QNA
P. schweinfurthii A. K. OSMAN & K.N. ABDEL KHALIK, s.n. CAI
2.Subtribe Gnaphaliinae
Filago contracta EL-GARF & A.K.OSMAN, s.n. CAI
F. desertoru m EL-GARF & A.K.OSMAN, s.n. QNA
F. mareotica EL-GARF & A.K.OSMAN, s.n. QNA
F. prolifera A. K. OSMAN & K.N. ABDEL KHALIK, s.n. CAI
Gnaphalium uligi nosum A. K. OSMAN, s.n. CAI
Helichrysum conglobatum EL-GARF & A.K.OSMAN, s.n. CAI
H. glumaceum A. K. OSMAN & K.N. ABDEL KHALIK, s.n. CAI
H. orientale EL-GARF & A.K.OSMAN, s.n. CAI
Homognaphalium pulvinatum A. K. OSMAN, s.n. CAI
Ifloga spicat a (Forssk.) Sch.Bip. FAYED et al. s.n. QNA
Lasiopogon muscoides (Desf.) DC. EL-GARF & A.K.OSMAN, s.n. CAI
Pseudogn ap hal ium l uteo alb um EL-GARF & A.K.OSMAN, s.n. CAI
II-Tribe Helenieae
1.Subtribe Flaveriinae
Flaveria bidentis EL-GARF & A.K.OSMAN, s.n. CAI
2.Subtribe Pectidinae
Tagetes minut a A. K. OSMAN, s.n. QNA
III-Tribe Plucheeae
Sphaeranthus suaveolens EL-GARF & A.K.OSMAN, s.n. CAI
IV-Tribe Senecioneae
1.Subtribe Senecioninae
Senecio aegyptius A. K. OSMAN, s.n. CAI
S. flavus FAYED et al. s.n. QNA
S. glaucus subsp. coronopifolius EL-GARF & A.K.OSMAN, s.n. QNA
S. glaucus subsp. glaucus EL-GARF & A.K.OSMAN, s.n. CAI
S. hoggariensis A. K. OSMAN & K.N. ABDEL KHALIK, s.n. CAI
S. vulgaris EL-GARF & A.K.OSMAN, s.n. CAI
C
AI = Cairo University Herbarium, QNA = Qena University Herbarium (proposed name), s.n. = collecting number is missing.

Pollen Morphology of Tribes Gnaphalieae, Helenieae, Plucheeae and Senecioneae (SubfamilyAsteroideae) 122
of Compositae from Egypt
prepared according to the method of [15] and SEM ob-
servations were made on acetolyzed grains coated with a
thin layer of gold/palladium for 3 minutes using a
EMITECH K550 sputter according to the Erdtman method
[16]. SEM examined with a JEOL-6300 SEM of Central
Lab., Faculty of Science, South Valley University, Qena,
Egypt. The terminology used is that of [17,18] and [19].
3. Results
The subtribes, genera and species of tribes Gnaphalieae,
Helenieae, Plucheeae and Senecioneae (according to [14])
that are represented in the flora of Egypt are arranged
alphabetically to facilitate consultation. For each species,
the valid scientific name is given followed by the citation
of the authority and the date of publication. Synonymy is
at a minimum to avoid complications. For full synonymy
of the species see [1,12] and [13].
A careful examination of the available pollen material
of the Egyptian species of tribes Gnaphalieae, Helenieae,
Plucheeae and Senecioneae revealed the presence of 2
pollen types, which can be distinguished through the
following key.
1.a. Pollen grains 3-zonocolpororate..Senecio pollen type
b. Pollen grains 3-zonocolporate.........Fil ago poll en ty pe
Main characters of pollen types.
1. Filago pollen type.
LM Figures 1(i)-(p), Figures 2(a), (b), (i), (j), (m-p).
SEM Figures 3(i)-(m), Figures 4(e)-(i). (Tables 2, 3).
Pollen grains 3-colporate, (NPC = 345 ), spheroidal to
prolate-sheroidal (P/E = 1.00-1.09), 20-24 (18-25) × 20-23
(19-25) μm. Apocolpium diameter 5-7 (3-8) μm. Colpi
13-16 (12-18) μm long, 2-4 (2-5) μm slightly wide, wide
or slender at the equator, pointed or tapered towards the
ends. Mesocolpia 12-16 (10-18) μm wide. Ora lolongate
(circular only in Homognaphalium pulvinatum) with
elleptic to ovate-shaped, 4-6 (3-8) μm in diameter. Exine
3-5 (2-6) μm thick (spinules included) at centre of
mesocolpia, decreasing slightly in thickness towards the
aperture margins. Sexine as thick as nexine or thicker
than nexine in Gnaphalium uliginosum and Lasiopogon
muscoides, tectate, has sharp spinules (spines in Gna-
phalium uliginos um) with broad bases, 1-3 (1-4) μm high,
perforated; sculpturing diameter the same on the whole
of the pollen surface or occasionally restrict to the echi-
nae bases that beset all the pollen surface (perforation
increases towards the aperatures margins inGnaphalium
uliginosum); the intratectal elements thin, minute, radial
and supported by fairly thin and short infratectal colu-
mellae. Nexine 1-2 (1-3) μm thick, increasing in thick-
ness towards the aperture margins.
The following taxa belong to this type:
I-Tribe Gnaphalieae
Subtribe Gnaphaliinae Less.
Genus Filago L. Syn. Evax Gaertn (4 species).
Species included:
1. F. contracta (Boiss.) Chrtek & Holub, Preslia 45: 3
(1963).
Syn. Evax contracta Boiss., Diagn. Pl. Orient., ser.
1, 11: 3 (1849).
2. F. desertorum Pomel, Nouv. Mat. Fl. Atlant. 1: 46
(1874).
Syns. Filago spathulata C. Presl forma desertorum
(Pomel) Pamp., Atti R. Ist. Ven. Sci. Lett. Arti 92: 234
(1932).
Evax mauritanica Pomel var. cyrenaica Pamp.,
Nuovo Giorn. Bot. Ital., n. s., 34: 962 (1927).
3. F. mareo tica Delile, Descr. Egypte, Hist. Nat. 274, t.
47, f. 2 (1814).
Syns. Filago mareotica Delile var. floribunda
(Pomel) Maire, Bull. Soc. Hist. Nat. Afr. Nord 26: 209
(1935).
Gifolaria mareotica (Delile) Chrtek & Holub, Pres-
lia 35: 10 (1963).
4. F. prolifera Pomel, Nouv. Mat. Fl. Atlant. 1: 47
(1874).
Syns. Filago germ anica L. subsp. prolifera (Pomel)
Maire, Cat. Pl. Maroc 3: 746 (1934).
Filago prolifera Pomel subsp. taeckholmiana Chrtek
in Tackh., Stud. Fl. Egypt, ed. 2, 554 (1974).
Genus Gnaphalium L. Syns. Amphidoxa DC., Fila-
ginella Opiz (one species).
Species included:
5. G. uliginosum L., Sp. Pl., ed. 1, 856 (1753).
Syn. Filaginella uliginosa (L.) Opiz, Abh. Bohm.
Ges. Wiss., ser. 5, 8 (Sitzungsber. Sect.): 52 (1854).
Genus Homognaphalium Kirp. Syn. Homognaphal-
ium fayed & Zareh non Kirp (one species).
Species included:
6. H. pulvinatum (Delile) Fayed & Zareh, Willdenowia
18: 451 (1989).
Syn. Gnaphalium pu lvinatum Delile, Descr. Egypte,
Hist. Nat. 266, t. 44 (1814).
Genus Lasiopogon Cass (one species).
Species included:
7. L. muscoi des (Desf.) DC., Prodr. 6: 246 (1838).
Syn. Gnaphalium muscoides Desf., Fl. Atlant. 2:
267, t. 231 (1799).
Genus Pseudognaphalium Kirp. Syn. Hypelichry-
sum Kirp (one species).
Species included:
8. P. luteoalbum (L.) Hilliard & B. L. Burtt, Bot. J.
Linn. Soc. 82: 206 (1981).
Syn. Gnaphalium luteoalbum L., Sp. Pl., ed. 1, 851
(175). 3

Pollen Morphology of Tribes Gnaphalieae, Helenieae, Plucheeae and Senecioneae (SubfamilyAsteroideae) 123
of Compositae from Egypt
Table 2. Tabular summary showing the Pollen Grains Dimensions (μm).
Species P. E. P/E.
Ex. th. Incl.
echin.
Ech.
Len.
Nex.
Th. Colp. Len.Colp. wid.
Ora.
diam. Apo. diam.Meso. diam.
I-Tribe Gnaphalieae
1. Unassigned
Gnaphalieae
Phagnalon
barbeyanum 28(25-31) 27(25-30) 1.04 5(4-6) 3(2-4)2(1-3)16(15-18)5(4-6)8(7-9) 8(7-9) 17(15-18)
P. nitidum 25(20-30) 27(25-30) 0.93 5(3-6) 2(2-3)2(1-3)13(12-15)5(3-7)8(7-9) 8(7-9) 12(10-13)
P. rupestre 32(30-35) 28(26-35) 1.10 5(3-7) 3(2-4)2(1-3)15(13-17)5(4-6)8(5-10) 8(7-10) 17(15-20)
P. schweinfurthii 27(23-28) 29(25-33) 0.93 5(3-7) 3(2-4)1(1-2)15(13-17)5(4-6)7(6-8) 10(8-12) 15(12-16)
2.Subtribe
Gnaphaliinae
Filago contracta 21(20-23) 21(20-23) 1.00 3(2-4) 1(1-2)2(1-3)15(13-17)4(2-5)5(4-6) 7(5-8) 12(10-15)
F. desertorum 23(20-25) 23(20-25) 1.00 3(2-4) 1(1-2)1(1-2)16(15-18)3(3-4)5(4-6) 6(5-8) 13(12-15)
F. mareotica 24(22-25) 22(20-25) 1.09 3(2-4) 1(1-2)2(1-3)16(15-18)4(3-5)6(5-8) 7(5-8) 15(12-18)
F. prolifera 23(20-25) 23(20-25) 1.00 4(2-5) 1(1-2)2(1-3)16(15-18)4(2-5)5(3-7) 5(3-7) 16(15-18)
Gnaphalium
uliginosum 21(20-23) 21(20-23) 1.00 5(4-6) 3(2-4)1(1-2)13(12-15)2(2-3)4(2-5) 5(3-7) 12(10-13)
Helichrysum
conglobatum 21(20-23) 21(20-23) 1.00 3(2-4) 2(1-3)1(1-2)13(12-15)3(2-4)5(3-7) 8(7-9) 13(12-14)
H. glumaceum 26(25-28) 26(25-28) 1.00 5(4-6) 3(2-4)2(2-3)18(15-20)4(3-5)5(3-6) 5(4-6) 14(12-15)
H. orientale 26(25-28) 28(25-30) 0.93 5(4-6) 3(2-4)2(1-3)12(10-15)4(3-5)6(5-7) 7(5-8) 15(13-17)
Homognaphalium
pulvinatum 22(20-25) 22(20-25) 1.00 3(2-4) 2(1-3)1(1-2)13(12-15)4(3-5)5(4-6) 7(5-8) 13(12-15)
Ifloga spicata 21(20-23) 21(20-23) 1.00 4(3-5) 2(1-3)1(1-2)13(12-15)3(2-4)4(3-5) 7(5-8) 11(9-15)
Lasiopogon
muscoides 20(19-23) 20(19-23) 1.00 3(2-4) 1(1-2)1(1-2)13(12-14)3(2-5)5(4-6) 5(3-6) 12(10-13)
Pseudognaphalium
luteoalbum 21(18-22) 21(19-22) 1.00 3(2-4) 2(1-3)1(1-2)13(12-15)3(2-4)4(3-5) 5(3-7) 12(10-13)
II-Tribe Helenieae
1.Subtribe
Flaveriinae
Flaveria bidentis 27(25-33) 31(30-33) 0.87 7(6-8) 5(4-6)2(1-3)16(15-18)4(3-5)7(5-8) 6(5-8) 17(15-20)
2.Subtribe
Pectidinae
Tagetes minuta 44(42-45) 44(42-45) 1.00 8(7-9) 5(4-6)2(2-4)21(20-25)5(3-6)8(7-9) 16(15-18)24(22-25)
III-Tribe Plucheeae
Sphaeranthus
suaveolens 23(22-25) 23(22-25) 1.00 4(3-5) 5(3-6)1(1-2)12(10-13)4(3-5)6(5-8) 8(7-9) 10(8-12)
IV-Tribe
Senecioneae
1.Subtribe
Senecioninae
Senecio aegyptius 31(30-33) 32(30-35) 0.97 5(4-6) 3(2-4)2(1-3)18(17-20)5(4-6)6(5-8) 6(5-8) 15(13-17)
S. flavus 29(27-30) 29(25-33) 1.00 6(5-7) 3(2-4)1(1-2)16(15-18)5(4-6)7(5-8) 6(5-10) 14(12-15)
S. glaucus subsp.
coronopifolius 25(23-26) 25(23-26) 1.00 5(4-6) 2(2-3)1 (1-2)15(14-16)4(3-5)5(4-6) 5(3-6) 13(12-14)
S. glaucus subsp.
glaucus 31(30-33) 33(31-35) 0.94 5(4-6) 4(3-5)2(2-3)19(17-20)4(3-5)5(4-6) 8(7-10) 14(12-15)
S. hoggariensi s 37(35-38) 42(40-45) 0.88 6(5-8) 3(2-4)1(1-2)27(25-30)4(3-5)8(7-9) 10(8-12) 18(15-20)
S. vulgaris 35(30-38) 41(35-45) 0.85 7(6-8) 5(3-6)3(1-4)19(15-20)5(4-6)8(7-9) 8(7-10) 16(15-18)
P. = Polar axis, E. = Equatorial diameter, P/E = The ratio of the length of the polar axis (P) to the equatorial diameter (E), Ech. Len. = echinae length, Ex. Th.
incl. echin. = exine thick includes echinae, Nex. Th. = nexine thick, Colp. Len. = colpus length, Colp. wid. = colpus width, Ora. diam. = ora diameter, Apo.
diam. = apocolpium diameter, Meso. diam.= mesocolpium diameter, μm = micrometer.

Pollen Morphology of Tribes Gnaphalieae, Helenieae, Plucheeae and Senecioneae (SubfamilyAsteroideae) 124
of Compositae from Egypt
Table 3. Tabular summary showing the Description of LM & SEM Samples.
Species Pollen
Class
Pollen
Shape
Col. wid.
at eq.
Colpi
Ends Ora ShapeEchinae Type
Sculpture
Type
Sculpture
State
Columellae
State
Nexine
State
I-Tribe
Gnaphalieae
1. Unassigned
Gnaphalieae
Phagnalon
barbeyanum II P.Sh. Wide Tapered La., rect. Spines Perforate 3 + 4 1 +
P. nitidum II O.Sh. Wide Pointed Lo., ovate Spinules Perforate6 1 ++
P. rupestre II P.Sh. Wide Tapered Lo.,ovate. Spines Perforate 6 1 +
P. schweinfurthii II O.Sh. Wide Pointed Lo.,ovate Spines Perforate 6 1 ++
2.Subtribe
Gnaphaliinae
Filago contracta I Sh . ±Wide Pointed Lo. ellep. Spinules Perforate5 1 +
F. desertorum I Sh. ±Wide Pointed Lo., ovate Spinules Perforate6 1 +
F. mareotica I P.Sh. ±Wide Pointed Lo. ellep. Spinules Perforate6 1 +
F. prolifera I Sh. ±Wide Pointed Lo., ovate Spinules Perforate6 1 +
Gnaphalium
uliginosum* I Sh. Wide Tapered Lo., ovate Spines Perforate 6 + 2 1 ++
Helichrysum
conglobatum II Sh. Wide Tapered Lo., ovate Spinules Perforate6 1 +
H. glumaceum II Sh. Wide Pointed La., rect Spines Perforate 6 + 3 1 +
H. orientale II O.Sh. Wide Tapered Circular Spines Perforate 3 1 +
Homognaphalium
pulvinatum I Sh. Wide Pointed Circular Spinules Perforate6 1 +
Ifloga spicata II Sh. ±Wide Tapered Lo. ellep. Spinules Perforate6 1 ++
Lasiopogon
muscoides I Sh. Slender Tapered Lo. ellep. Spinules Perforate6 1 ++
Pseudognaphalium
luteoalbum I Sh. Wide Pointed Lo., ovate Spinules Perforate6 1 +
II-Tribe Helenieae
1.Subtribe
Flaveriinae
Flaveria bidentis II Subob Wide Pointed Lo., ovate Spines Perforate 3 1 +
2.Subtribe
Pectidinae
Tagetes minuta II Sh. Wide Pointed La., rect Spines Perforate 3 1 +
III-Tribe Plucheeae
Sphaeranthus
suaveolens** II Sh. Wide Tapered Circular Spines Perforate 2 1 ++
IV-Tribe
Senecioneae
1.Subtribe
Senecioninae
Senecio aegyptius II O.Sh. Narrow PointedLa., rect Spines Perforate 3 1 ++
S. flavus II Sh. Wide Pointed Lo., ovate Spines Perforate 3 1 ++
S. glaucus subsp.
coronopifolius II Sh. ±Wide Pointed Lo., ovate Spinules Perforate6 1 ++
S. glaucus subsp.
glaucus II O.Sh. ±Wide Pointed Lo. ellep. Spines Perforate 6 1 ++
S. hoggariensis II O.Sh. Slender TaperedLo., ovate Spines Perforate 6 1 ++
S. vulgaris II Subob ±Wide Tapered Lo. ell ep . Spines Perforate 5 1 ++
Col. wid. at eq. = Colpi Width at equator, Ellep. = Elleptic, La. = Lalongate, Lo. = Lolongate, O. = Oblate, P. = Prolate, Rect. = Rectangular, Sh. = Spheroidal,
Subob. = Suboblate, ± Wide = Slightly wide, I = Pollen 3-zonocolporate, II = Pollen 3-zonocolpororate, 1 = Columellae with thin, short and unbranched rods, +
= Sexine as thick as nexine, ++ = Sexine thicker than nexine, 2 = Sculpturing diameter increases towards the aperture margins, 3 = Sculpturing diameter in-
creases towards the echinae bases, 4 = Sculpturing diameter increases towards the pollen pole, 5 = Sculpturing restrict to the echinae bases, 6 = Sculpturing
diameter the same on the whole of the pollen surface, * = Aperture margins perforate, ** = Aperture margins microreticulate.

Pollen Morphology of Tribes Gnaphalieae, Helenieae, Plucheeae and Senecioneae (SubfamilyAsteroideae)
of Compositae from Egypt
125
Key to taxa of Filago pollen type
1.a. Pollen prolate-spheroidal in shape..Filago mareotica
b. Pollen sheroidal in shape….........................………2
2.a. Colpi slender at equator......Lasiopogon muscoides
b. Colpi otherwise………………...………………….3
3.a. Colpi wide at equator……Gnaphalium uliginosum
…..………………….….Homognaphalium pulvinatum
.......................................Pseudognaphaliumluteoalbum
b. Colpi slightly wide at equator.........Filago contracta
.................................................................F. desertorum
.....................................................................F. prolifera
2. Senecio pollen type
LM (Figures 1(a)-(h), Figures 2(c)-(h), (k), (l), Fig-
ures 3(a)-(p), Figures 4(a), (b), SEM Figures 4(c)-(h),
Figures 5(a)-(d), (j)-(o), Figures 6(a)-(j).
Pollen grains 3-colpororate, spheroidal, oblate-spher-
oidal, prolate-spheroidal to suboblate (P/E = 0.85-1.10),
21-44 (20-45) × 21-44 (20-45) μm. Apocolpium diameter
5-16 (3-18) μm. Colpi 12-27 (10-30) μm long, 3-5 (2-7)
μm wide, wide, slightly wide, slender or narrow at the
equator, pointed-tapered towards the ends. Mesocolpium
10-24 (8-25) μm wide. Ora lolongate-lalongate (circular
in both Helichrysum orientale and Sp haeranthus suaveo-
lens), with ovate, rectangular to elleptic-shaped, 4-8
(3-10) μm in diameter. Exine 3-8 (2-9) μm thick (spines
included) at centre of mesocolpia, with the same thick-
ness towards the aperture margins. Sexine thicker than
nexine or as thick as nexine, tectate, with long, pointed
spines or spinules, 2-5 (1-6) μm high, perforated; perfo-
rations number and diameter the same on the whole of
the pollen surface or increase towards the echinae bases
(increase towards the aperature margins in Sphaeranthus
suaveolens or restrict to the echinae bases in Senecio
vulgaris); the intratectal elements thin, minute, radial and
supported by fairly thin and short infratectal columellae.
Nexine 1-3 (1-4) μm thick, increasing in thickness to-
wards the aperture margins.
The following taxa belong to this type:
I-Tribe Gnaphalieae
Gnaphalieae unassigned to Subtribe.
Genus Phagnalon Cass (4 species).
Species included:
1. P. barbeyanum Asch. & Schweinf., Mém. Inst.
Egypt. 2: 87 (1887).
Syn. P. Aegyptiacum Boiss., Fl. Orient. Suppl. 296
(1888).
2. P. nitidum Fresen., Mus. Senckenb. 3: 81 (1839).
3. P. rupestre ( L.) DC., Prodr. 5: 396 (1836).
Syn. Conyza rupestris L., Mant. 113 (1767).
4. P. schweinfurthii Sch. Bip. ex Schweinf., Verh. K.
K. Zool. Bot. Ges. Wien 18: 685 (1868).
Subtribe Gnaphaliinae Less.
Genus Helichrysum Mill. Syn. Leontonyx Cass. (3 spe-
cies).
Species included:
5. H. conglobatum (Viv.) Steud., Nomencl. Bot., ed. 2,
1: 738 (1840).
Syns. Gnaphalium conglobatum Viv., Fl. Libyc.
Spec. 55, t. 3, f. 5 (1824).
Helichrysum siculum Boiss., Fl. Orient. 3: 229
(1875).
6. H. glumaceum DC., Prodr. 6: 197 (1838).
Syn. Achyrocline glumacea (DC.) Oliv. & Hiern in
Oliv., Fl. Trop. Afr. 3: 340 (1877)
7. H. orientale (L.) Gaertn., Fruct. Sem. Pl. 2: 404
(1791).
Syn. Gnaphalium orientale L., Sp. Pl., ed. 1, 853
(1753).
Genus Ifloga Cass. Syn. Comptonanthus B. Nord
(one species).
Species included:
8. I. spicata (Forssk.) Sch.Bip. in Webb & Berthel.,
Phyt. Canar. 2: 310 (1845).
Syns. Chrysocoma spicata Forssk., Fl. Ae-
gypt.-Arab. LXXIII (1775).
Chrysocoma spicatum (Forssk.) Vahl, Symb. Bot. 1:
70 (1790).
II. Tribe Helenieae Benth.
Subtribe Flaveriinae Less.
Genus Flaveria Juss. (one species).
Species included:
9. F. bident i s (L.) Kuntze, Rev. Gen. 3: 148 (1898).
Syns. Ethulia bidentis L., Mant. 110 (1767).
Flaveria contrayerba (Cav.) Pers., Syn. Pl. 2: 489
(1807).
Flaveria bonariensis DC., Prodr. 5: 635 (1836)
Subtribe Pectidinae Less.
Genus Tagetes L. (one species).
Species included:
10. T. minuta L., Sp. Pl., ed. 1, 887 (1753).
III. Tribe Plucheeae Anderb.
Genus Sphaeranthus L. (one species).
Species included:
11. S. suaveolens (Forssk.) Dc., Prodr. 5: 370 (1836).
Syn. Polycephalos suaveolens Forssk., Fl. Ae-
gypt.-Arab. 154 (1775).
IV. Tribe Senecioneae
Subtribe Senecioninae Dumort.
Genus Senecio L. (6 species).
Species included:
12. S. aegypt i u s L., Sp. Pl., ed. 1, 867 (1753).
13. S. flavus (Decne.) Sch. Bip. In Webb & Berthel.,
Phyt. Canar. 3: 317 (1847).
S
yns. Crassocephal umfl avum Decne., Ann. Sci. Nat.

Pollen Morphology of Tribes Gnaphalieae, Helenieae, Plucheeae and Senecioneae (SubfamilyAsteroideae) 126
of Compositae from Egypt
Figure 1. LM observations of acetolysed pollen grains (LM × = 1000). (a)-(b) Phagnalon barbeyanum, (a) Oblique polar view,
(b) Oblique equatorial view. (c)-(d) P. nitidum, (c) Polar view, (d) Oblique equatorial view. (e)-(f) P. rupestre, (e) Polar view, (f)
Equatorial view. (g)-(h) P. schweinfurthii, (g) Polar view, (h) Equatorial view. (i)-(j) Filago contracta, (i) Polar view, (j) Equa-
torial view. (k)-(l) F. desertorum, (k) Polar view, (k) Oblique equatorial view. (m)-(n) F. mareotica, (m) Polar view, (n) Equa-
torial view. (o)-(p) F. prolifera, (o) Oblique polar view, (p) Equatorial view.

Pollen Morphology of Tribes Gnaphalieae, Helenieae, Plucheeae and Senecioneae (SubfamilyAsteroideae) 127
of Compositae from Egypt
Figure 2. LM observations of acetolysed pollen grains (LM × = 1000). (a)-(b) Gnaphalium uliginosum, (a) Polar view, (b)
Equatorial view. (c)-(d) Helichrysum conglobatum, (c) Polar view, (d) Equatorial view. (e)-(f) H. glumaceum, (e) Polar view, (f)
Equatorial view. (g)-(h) H. orientale, (g) Polar view, (h) Equatorial view. (i)-(j) Homognaphalium pulvinatum, (i) Polar view, (j)
Equatorial view. (k)-(l) Ifloga spicata, (k) Polar view, (l) Equatorial view. (m)-(n) Lasiopogon muscoides, (m) Polar view, (n)
Equatorial view. (o)-(p) Pseudognaphaliumluteoalbum, (o) Polar view, (p) Equatorial view.

Pollen Morphology of Tribes Gnaphalieae, Helenieae, Plucheeae and Senecioneae (SubfamilyAsteroideae) 128
of Compositae from Egypt
Figure 3. LM and SEM observations of acetolysed pollen grains (LM × = 1000, SEM × = 5000 - 15000). (a)-(b) Senecio vul-
garis, (a) Oblique polar view, (b) Equatorial view. (c)-(e) Phagnalon barbeyanum, (c) Polar view, (d) Equatorial view, (e) Mag-
nified part of exine. (f)-(h) Phagnalon rupestre, (f) Oblique polar view, (g) Oblique equatorial view, (h) Magnified part of
exine. (j) Filago contracta, (i) Magnified part of exine. (i) (k) Filago prolifera, (i) Oblique polar view, (k) Magnified part of
exine, (l)-(m) Gnaphalium uliginosum, (l) Oblique equatorial view, (m) Magnified part of exine.

Pollen Morphology of Tribes Gnaphalieae, Helenieae, Plucheeae and Senecioneae (SubfamilyAsteroideae) 129
of Compositae from Egypt
Figure 4. SEM observations of acetolyzed pollen grains (SEM × 5000-15000). (a)-(c) Helichrysum orientale, (a) Polar view, (b)
Equatorial view, (c) Magnified part of exine. (d) Ifloga spicata, (d) Magnified part of exine. (e)-(f) Lasiopogon muscoides, (e)
Oblique equatorial view, (f) Magnified part of exine. (g)-(i) Pseudognaphalium luteoalbum, (g) Oblique polar view, (h) Equa-
torial view, (i) Magnified part of exine. (j)-(l) Flaveria bidentis, (j) Oblique polar view, (k) Oblique equatorial view, (l) Magni-
fied part of exine. (m)-(n) Tagetes minuta, (m) Polar view, (n) Equatorial view.

Pollen Morphology of Tribes Gnaphalieae, Helenieae, Plucheeae and Senecioneae (SubfamilyAsteroideae) 130
of Compositae from Egypt
Figure 5. LM observations of acetolysed pollen grains (LM × = 1000). (a)-(b) Flaveria bidentis, (a) Polar view, (d) Equatorial
view. (c)-(d) Tagetes minuta, (c) Polar view, (d) Equatorial view. (e)-(f) Sphaeranthus suaveolens, (e) Polar view, (f) Equatorial
view. (g)-(h) Senecio aegyptius, (g) Polar view, (h) Equatorial view. (i)-(j) S. flavu s , (i) Polar view, (j) Equatorial view. (k)-(l) S.
glaucus subsp. coronopifolius, (k) Polar view, (l) Equatorial view. (m)-(n) S. glaucus subsp. glaucus, (m) Polar view, (n) Equa-
torial view. (o)-(p) S. hoggariensis, (o) Polar view, (p) Equatorial view.

Pollen Morphology of Tribes Gnaphalieae, Helenieae, Plucheeae and Senecioneae (SubfamilyAsteroideae)
of Compositae from Egypt
131
Figure 6. SEM observations of acetolyzed pollen grains (SEM × 5000-15000). (a) Tagetes minuta, (a) Magnified part of exine.
(b)-(d) Sphaeranthus suaveolens, (b) Oblique polar view, (c) Equatorial view, (d) Magnified part of exine. (e)-(g) Senecio fla-
vus, (e) Polar view, (f) Equatorial view, (g) Magnified part of exine. (h)-(i) Senecio vulgaris, (h) Polar view, (i) Magnified part
of exine. (j) Senecio aegyptius, (j) Magnified part of exine.
Bot., sér. 2, 2: 265 (1834).
Senecio dec ai snei DC., Prodr. 6: 342 (1838).
14. S. glaucus L., Sp. Pl., ed. 1, 868 (1753). subsp.
coronopifolius (Maire) C. Alexander, Notes Roy. Bot.
Gard. Edinb. 37: 412 (1979).
Syns. Senecio coronopifolius Desf., Fl. Atlant. 2:

Pollen Morphology of Tribes Gnaphalieae, Helenieae, Plucheeae and Senecioneae (SubfamilyAsteroideae) 132
of Compositae from Egypt
273 (1799), nom. illeg., non Burm.f. (1768).
Senecio desfontainei Druce, Brit. Pl. List, ed. 2, 61
(1928).
15. S. glaucus L., Sp. Pl., ed. 1, 868 (1753). subsp.
glaucus
Syns. Senecio vernalis Waldst. & Kit. var. carnosus
Post. Fl. Syr. Pal. Sinia, ed. 1, 442 (1896).
Senecio joppensis Dinsm. In Post, Fl. Syr. Pal. Sinai.
Ed. 2, 2: 69 (1933).
16. S. hoggariensis Batt. & Trab., Bull. Soc. Bot.
France 58: 671 (1911).
17. S. vulgaris L., Sp. Pl., ed. 1, 867 (1753).
Key to taxa of Senecio pollen type
1.a. Polar area long axis median 21 - 29 μm……….….4
b. Polar area long axis median 31 - 44 μm…..……….. 2
2.a. Polar area long axis 44(42 - 45) μm…Tagetes minuta
b. Polar area long axis median 31 - 37 μm…………….3
3.a. Polar area long axis 31 - 32 μm...Phagnalon rupest re
..……………………………….…... Senecio aegyptius
…..……………………....… S. glaucus subsp. glaucus
b. Polar area long axis 35 - 37 μm…Senecio hoggariensis
...................................................................... S. vulgaris
4.a. Polar area long axis 21 - 25 μm….Phagnalon nitidum
….…………..………..…... Helichrysum conglobatum
…..……………..………………….…… Ifloga spicata
……..…………….……...….Sphaeranthus suaveolens
……………….. Senecio glaucus subsp. coronopifolius
b. Polar area long axis 26 - 29 μm………………………
…………….……………...….. Phagnalon barbeyanum
………...……………………………. P. schweinfurthii
…...................................…… Helichrysum glumaceum
……………..…………………….…....… H. orientale
....…….……….................................. Flave r i a bidentis
..……………………………..…...….… Senecio flavus
4. Discussion
Palynological characters applied to the species of Gna-
phalieae, Helenieae, Plucheeae and Senecioneae in Egypt
proved to be useful in the distinction between two groups.
The pollen grains are usually 3-zonocolporate or 3-zo-
nocolpororate with lolongate or lalongate (rarely circular)
ora.
There is semi-similarity between the pollen grains of
the investigated taxa, as a result it will be difficult to
make accurate distinguishing and to construct true key
between the studied species. Two pollen types are dis-
tinguished according to the pollen class. The Senecio
pollen type, in which the pollen are 3-colpororate, is
characteristic of the species of genera Phagnalon, Heli-
chrysum, Ifloga, Flaveria, Tagetes, Sphaeranthus and
senecio. The Filago pollen type, in which the pollen are
3-colporate, is characteristic of the species of genera
Filago, Gnaphalium, Homognaphalium, Lasiopogon and
Pseudognaphalium.
The smallest pollen grains are those of Lasiopogon
muscoides species 20 (19-23) μm and the largest are
those of Tagetes minuta 44 (42-45) μm. Pollen size of
other taxa ranged between (21-37) μm. Pollen grains are
more or less similar in shape being spheroidal, ob-
late-spher- oidal, prolate-spheroidal to suboblate.
Ora shape contributes to differentiate between the re-
lated species of Helichrysum glumaceum, Ifloga spicata,
Flaveria bidentis, Tagetes minuta, Senecio flavus and
Senecio vulgaris are characterized by lalongate ora shape,
Helichrysum orientale, Homognaphalium pulvinatum
and Sphaeranthus suaveolens exhibit circular ora shape,
while lolongate ora shape is observed in the remaining
studied species. The aperture width also contributes to
differentiate all studied species of genus Filago, Ifloga
spicata, Senecio glaucus subsp. coronopifolius, S. glau-
cus subsp. Glaucus and S. vulgaris which exhibit colpi
are slightly width at equator, Lasiopogon muscoides and
S. hoggariensis which have slender colpi at equator, Se-
necio aegyptius has narrow colpi at equator from the
other investigated taxa which are characterized by wide
colpi at equator. It also points to the close relationship
between the four Filago species which exhibit very close
pollen grains being similar in pollen shape, colpi charac-
ters, sculpture and dimensions.
The pollen sculpture is more uniform in most investi-
gated species with perforate texture. The pollen grains
are spiny to spinulate with perforate echinae bases. In
Gnaphalium uliginosum aperture margins are perforate
while in Sphaeranthus suaveolens species they are mi-
croreticulate, also in both Filago contracta and Senecio
vulgaris perforation restrict to the echinae bases.
REFERENCES
[1] L. Boulos, “Flora of Egypt, Verbenaceae-Compositae,”
Al-Hadara Publishing, Cairo, Vol. 3, 2002.
[2] J. Muller, “Fossil Pollen Records of Extant Angio-
sperms,” Botanical Review, Vol. 47, No.1, 1981, pp.
1-142. doi:10.1007/BF02860537
[3] J. J. Skvarla, D. A. Larson, “An Electron Microscopic
Study of Pollen Morphology in the Compositae with Spe-
cial Reference to the Ambrosinae,” Grana Palynol., Vol.
6, 1965, pp. 210-269. doi:10.1080/00173136509429147
[4] J. J. Skvarla, B. L. Turner, “Systematic Implications from
Electron Microscopic Studies of Compositae Pollen—A
Review,” Annals of the Missouri Botanical Garden, Vol.
53, 1966, pp. 220-256. doi:10.2307/2394944
[5] J. J. Skvarla, B. L. Turner, V. C. Patel and A. S. Tomb,
“Pollen Morphology in the Compositae and in Morpho-
logically Related Families,” In: V. H. Heywood et al.

Pollen Morphology of Tribes Gnaphalieae, Helenieae, Plucheeae and Senecioneae (SubfamilyAsteroideae)
of Compositae from Egypt
133
Eds., The Biology and Chemistry of Compositae, Vol. 2,
London, 1977, pp. 141-265.
[6] M. S. Zavada, S. E. De Villiers, “Pollen of the Asteraceae
from the Paleocene-Eocene of South Africa,” Grana, Vol.
39, 2000, pp. 39-45. doi:10.1080/00173130150503795
[7] R. Tormo, J. L. U. Jimenez, “The Apertural System of
Pollen Grains in Anthemideae and Cardeae (Compositae)
with Special Reference to the Mesoaperture,” Review of
Palaeobotany and Palynology, Vol. 62, 1990, pp. 1-9.
[8] A. A. Anderberg, “Taxonomy and Phylogeny of the Tribe
Gnaphalieae (Asteraceae),” Opera Botanica, Vol. 104, 1991,
pp. 1-195.
[9] P. Leins, “Pollensystematische Studien an Inuleen, I:
Tarchonanthinae, Plucheinae, Inulinae, Buphthalminae,”
Botanishe Jahrbücher für Systematik, Pflanzengeschi-
chte und Pflanzengeographie, Vol. 91, 1971, pp. 91-146.
[10] P. Leins, “Pollensystematische Studien an Inuleen, II:
Filagininae,” Botanishe Jahrbücher für Systematik, Pflan-
zengeschichte und Pflanzengeographie, Vol. 93, 1973, pp.
603-611.
[11] V. Tackholm, “Students’ Flora of Egypt,” 2nd Edition,
Cairo University, Cairo, 1974.
[12] M. N. El Hadidi, A. A. Fayed, “Materials for Excursion
Flora of Egypt (EFE),” Taeckholmia, Vol. 15, 1994/1995,
pp. 96-98.
[13] L. Boulos, “Flora of Egypt Checklist, Revised Annotated
Edition,” Al-Hadara Pblishing, Cairo, 2009.
[14] K. Bremer, “Asteraceae. Cladistics and Classifications,”
Timber Press Inc., Portland, 1994.
[15] G. Erdtman, “The Acetolysis Method, a Revised Descrip-
tion,” Svensk Bot. Tidskr , Vol. 54, No. 1960, pp. 561-464.
[16] G. Erdtman, “Handbook of Palynology, an Introduction
to the Study of Pollen Grains and Spores,” Munksgaard,
Copenhagen, 1969.
[17] G. Erdtman, “Pollen Morphology and Plant Taxonomy,
Angiosperms,” Hafner Publishing Co., New York, 1966.
[18] K. Faegri and J. Iversen, “Textbook of Pollen Analysis,”
3rd Edition, Munksgaard, Copenhagen.
[19] M. T. Dimon, “Problémes Généraux Soulevés par L’étude
Pollinique de Composées Méditerranénnes,” Nat. Monsp.
Sér. Bot., Vol. 22, 1971, pp. 129-144.