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Acta Bot. Croat. 69 (1), 83–92, 2010 CODEN: ABCRA 25
ISSN 0365–0588

Pollen morphology of the genus Lathyrus (Fabaceae)

section Cicercula in Thrace (European Turkey)

FATMA GUNES1*, ALI CIRPICI2

1 Kafkas University, Faculty of Arts and Sciences, Department of Biology, Kars, Turkey

2 Marmara University, Faculty of Arts and Sciences, Department of Biology, Göztepe,
Istanbul, Turkey

We examined the pollen morphology of four taxa from the Cicercula section of Lathyrus,
grown in the Thrace region (European Turkey), including L. annuus L., L. gorgoni
Parl. var. pilosus C. C. Townsend, L. cicera L. and L. hirsutus L. The pollen grains are
3-zonocolporate of subprolate and prolate types (P/E=1.2378-1.4491), medium to large
sized, elliptical or rectangular-obtuse-convex (equatorial view) and circular to slightly tri-
angular-obtuse-convex (polar view). The ornamentation is reticulate.

Keywords: pollen, morphology, Lathyrus, Cicercula, Turkey

Introduction

Pollen morphology is not affected by environmental conditions; therefore it has been
accepted as a reliable taxonomic marker in the identification of higher plants (AYTUG
1959). The genus Lathyrus L. (Fabaceae), comprising nearly 200 species with annual and
perennial plants, is centered in Mediterranean countries and mainly distributed in the
northern hemisphere and the highlands of tropical Africa (SEEN 1938; DAVIS 1970, 1988;
HEYWOOD 1978; KUPICHA 1983). The flora of Turkey contains 64 species belonging to the
sections Orobus, Platystylis, Pratensis, Orobastrum, Orobon, Lathyrus, Cicercula, Aphaca,
Nissolia and Clymenum; a total of 24 of these are endemic (DAVIS 1970, 1988; GUNER et al.
2000).

The pollen morphology have been already investigated in Lathyrus digitatus (AYTUG
1967, AYTUG et al. 1971), L. undulatus, L. sylvestris and L. ochrus (GUNES and CIRPICI
1998). L. sylvestris, L. pratensis, L. maritimus, L. nissolia and L. montanus (MOORE et al.
1991), L. grandiflorus, L. latifolius, L. sylvestris, L. tuberosus, L. alpestris, L. aureus, L.
linifolius, L. niger, L. palustris, L. transsilvanicus, L. venetus and L. vernus (TOSHEVA et al.
2004, TOSHEVA and TONKOV 2005), L. emodii, L. cicera, L. humulis and L. pratensis
(PERVEEN and QAISER 1998). The authors indicated that the pollen grains are mainly
tricolporatae. In addition to pollen type, a tectate type of exine structure and a reticulate

ACTA BOT. CROAT. 69 (1), 2010 83

* Corresponding author, e-mail: drgunes@gmail.com

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type of sculpture were also reported in all Lathyrus species. Furthermore, MOORE et al.
(1991) also noted that the endexine thickens 2–3 times around the colpus and the porus and
form a large costae formation.

Material and methods

Four taxa of Lathyrus were collected from their natural habitats in the Thrace region of
Turkey between 1996 and 1998. All collected specimen were identified and deposited at
the herbarium of the Biology Department, Faculty of Arts and Sciences of the Marmara
University (MUFE).

Pollen grains for LM examination were prepared following the standard procedure of
WODEHOUSE (1959) and the acetolysis method (ERDTMAN 1960). Fifteen pollen characters
were investigated by light microscope (LM). P (polar diameter), E (equatorial diameter)
were measured at a magnification of 400 times and clg (colpus length), clt (colpus width),
plg (porus lenght regarding the poles), plt (porus width regarding the equatorial diameter), t
(one edge of polar triangle), exine thickness, intine thickness and costa were also measured
under a magnification of 1000´. Structure (exine), sculpture (exine ornamentation) were
also shown.

For SEM examination, pollen grains were coated with gold. The microphotographs
were obtained using JEOL-JSM-5200 scanning electron microscope (SEM) at a magnifi-
cation of 2000–5000´. The arithmetic means, standard deviation and variations were cal-
culated for each characteristic. Standard deviations are not given for characteristics having
no requirement for multiple measurements. In table 1, fresh pollen measurements are given
in light and the fossilized pollen measurements are given in bold. To determine the struc-
ture and sculpture characteristics, the specimens specifically prepared with the acetolysis
method were analyzed under LM microscopy. Reticulate type of sculpture was shown by
photographs taken in SEM (Fig. 2). The pollen morphological descriptions follow the ter-
minology of IVERSEN and TROELS-SMITH (1950), REITSMA (1970), MOORE et al. (1991) and
PUNT et al. (1994).

Results

The following shapes of pollen were found in the four Lathyrus species: 3-zonocol-
porate type, a tectate type of exine structure and a reticulate type of sculpture, subprolate
and prolate. Colpus width in fossilized grains appears to have decreased based on the val-
ues determined as described above. As the pollens fossilize they increase in the direction of
the poles, that is they increase in length. The colpus measurements were done at the regions
close to the pores. Although the colpus above the pores in L. gorgoni enlarged as much as
the pore itself, the colpus above the pore narrowed to a line in L. cicera.

Lathyrus annuus L. (Tab. 1, Figs. 1, 2) A1(E). Edirne: Saçlimüsellim-SigH ircik village
road, 1 km from rice-field edges, 30 m, 25.05.1997, F. Gunes, MUFE 5392.

Pollen class: 3-zonocolporate.
Pollen group: Subprolate [P/E=1.112 (Wodehouse) and P/E=1.114 (acetolysis)]
Dimensions: Medium size [P ´ E = 36.679 ´ 33.085 (Wodehouse), 39.676 ´ 35.620

(acetolysis)].

84 ACTA BOT. CROAT. 69 (1), 2010

GUNES F., CIRPICI A.

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Apertures: Ectoapertures – colpi: short, narrow, deep, with acute ends, thick costae
along the margins of the colpi, colpus membrane covered by clearly visible granules; clg:
20.300 (Wodehouse), 32.5 (acetolysis) mm, clt:1.8 (Wodehouse), 2.3 (acetolysis) mm.
Endoapertures – pori : lalongate (Wodehouse), lolongate (acetolysis) methods., plg: 7.0
(Wodehouse), 12.6 (acetolysis) mm, plt: 9.0 (Wodehouse), 9.9 (acetolysis) mm and plg/plt=
0.8 (Wodehouse), 1.3 (acetolysis).

Outlines: Equatorial view – elliptic to rectangular-obtuse-convex; polar view circular

Ornamentation: reticulate; reticules distinct, big, irregular. Heads of single columellae
are visible inside the lumina. Colpus area, porus and apocolpium are slightly reticulate.
Ex/int (Wodehouse): 1/1

Exine (acetolysis): 2 mm thick, tectate – infrastructure, nexine/sexine = 1/1

Lathyrus gorgoni Parl. var. pilosus (Tab. 1; Fig. 1–4). A1(E) I
·
stanbul: Büyükçek-

mece-highway way out, through road, meadow, 30 m, 11.05.1996, F. Gunes, MUFE 5007.

Pollen class: 3-zonocolporate.

Pollen group: subprolate [P/E=1.1426 (Wodehouse)] – prolate [P/E=1.3944 (acetolysis)].

Dimensions: Medium size [P ´ E = 41.228 ´ 36.081 (Wodehouse), 56.992 ´ 40.872
(acetolysis)].

Apertures: Ectoapertures – colpi: long, narrow, deep, nearly reaching the poles, with
acute ends, thick costae along the margins of the colpi, colpus membrane covered by
clearly visible granules; clg: 34.266 (Wodehouse), 55.240 (acetolysis) mm, clt: 2.737
(Wodehouse), 3.383 (acetolysis) mm. Endoapertures – pori: lalongate to slightly lalongate,
plg: 9.790 (Wodehouse), 13.440 (acetolysis) mm, plt: 11.437 (Wodehouse), 14.252 (aceto-
lysis) mm and plg/plt= 0.856 (Wodehouse), 0.943 (acetolysis).

Outlines: Equatorial view – elliptic to rectangular-obtuse-convex; polar view circular

Ornamentation: reticulate; reticules distinct, big and irregular. Heads of single columellae
are visible inside the lumina. Colpus area, porus and apocolpium are slightly reticulate.

Ex/int (Wodehouse): 1/2

Exine (acetolysis): 3 mm thick, tectate – infrastructure, nexine/sexine = 1/1

Lathyrus cicera L. (Tab. 1; Figs. 1–6). A1(E) Çanakkale: Gelibolu-Eceabat Road,
20 km. 0–10 m, 24.04.1998, F. Gunes and A. Cirpici, MUFE 5646.

Pollen class: 3-zonocolporate.

Pollen group: subprolate [P/E=1.321 (Wodehouse)) – prolate (P/E=1.574 (acetolysis)].

Dimensions: medium size [P ´ E = 41.707 ´ 31.581 (Wodehouse), 60.320 ´ 38.324
(acetolysis)].

Apertures: Ectoapertures – colpi: long, slightly wide, nearly reaching the poles, with
acute ends, thick costae along the margins of the colpi, colpus membrane covered by small
granules; clg: 31.390 (Wodehouse), 58.804 (acetolysis) mm, clt: 4.031 (Wodehouse), 3.864
(acetolysis) mm. Endoapertures – pori: lalongate, plg: 4.379 (Wodehouse), 10.366 (aceto-
lysis) mm, plt: 8.120 (Wodehouse), 14.868 (acetolysis) mm and plg/plt= 0.539 (Wode-
house), 0.6972 (acetolysis).

Outlines: Equatorial view – elliptic to rectangular-obtuse-convex; polar view circular
to slightly triangular

ACTA BOT. CROAT. 69 (1), 2010 85

POLLEN MORPHOLOGY IN LATHYRUS

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86
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Tab. 1. Mean pollen characteritics (M), structure and sculpture (ornamentation). s – standard deviation, var. – variations of line numbers of polen lamela, W
– Wodehouse-metod, A – asetolysis-metod, P – polar diameter, E – equatorial diameter, P/E – pollen shape, Ex – exine thickness, Ex/Int – the ratio of
exine to the intine, clg – colpus length, clt – colpus width, plg – porus length regarding the poles, plt – porus width regarding the equatorial diameter,
plg/plt – porus shape, t – one edge of polar triangle.

Lathyrus annuus Lathyrus gorgoni var. pilosus Lathyrus cicera Lathyrus hirsutus

M s var. M s var. M s var. M s var.

P W 36.679 ±1.617 11–13 41.228 ±1.597 13–16 41.707 ±1.354 13–15 38.976 ±2.504 13–14.5

A 39.676 ±2.590 12–17 56.992 ±2.257 20–24 60.320 ±3.247 20–25 52.832 ±2.463 18–23

E 33.085 ±1.462 10–13 36.081 ±1.476 11–14 31.581 ±0.974 10–11.5 32.799 ±1.227 10–11.5

35.620 ±2.031 12–16 40.872 ±2.146 14–17 38.324 ±2.108 13–16 30.784 ±1.198 10–14

P/E 1.112 Subprolate 1.143 Subprolate 1.321 Subprolate 1.188 Subprolate

1.114 Subprolate 1.394 Prolate 1.574 Prolate 1.716 Prolate

Ex Not
measured

Not
measured

I
·
nce Not

measured
Not

measured
Not

measured
Not

measured
Not

measured

2 3 3 2

Ect/end

1/1 1/1 1/1 1/1

Ex/int 1/1 Not
measured

½ Not
measured

2/1 Not
measured

1/1 Not
measured

clg 20.300 ±1.477 16–21 34.266 ±1.317 27–32 31.390 ±1.631 24–30 27.882 ±1.211 22–2 6

32.523 ±2.343 20–26 55.240 ±3.553 35–44 58.804 ±3.493 39–47 47.796 ±2.761 29–38

clt 1.856 ±0.568 1–2 2.7377 ±0.950 1–5 4.031 ±0.733 2–5 3.712 ±0.843 2–4

2.275 ±0.500 1–2 3.383 ±1.063 1–4 3.864 ±1.548 1–5 1.960 ±0.686 1–2

plg 7.018 ±0.498 5–7 9.790 ±0.903 7–10 4.379 ±0.710 3–5 8.294 ±0.611 6–8

12.593 ±1.565 7–11 13.440 ±1.313 7–12 10.366 ±1.249 5–9 10.556 ±0.978 6–9

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Tab. 1. – continued

Lathyrus annuus Lathyrus gorgoni var. pilosus Lathyrus cicera Lathyrus hirsutus

M s var. M s var. M s var. M s var.

plt 9.048 ±0.696 7–9 11.438 ±0.957 8–12 8.120 ±0.935 5–9 10.701 ±0.661 8–10

9.870 ±1.744 5–10 14.252 ±1.810 8–13 14.868 ±1.768 8–13 10.136 ±0.856 6–8

plg/plt 0.776 Not
measured

Not
measured

0.856 Not
measured

0.539 Not
measured

Not
measured

0.775 Not
measured

Not
measured

1.276 Not
measured

Not
measured

0.943 Not
measured

0.697 Not
measured

Not
measured

1.041 Not
measured

Not
measured

t Not
measured

Not
measured

Not
measured

14.500 ±1.249 10–15 17.110 ±1.264 13–16 Not
measured

Not
measured

Not
measured

Structure Tectate-
infra-

structure

Tectate-
infra-

structure

Tectate-
infrastruc-

ture

Tectate-
infra-

structure

Sculpture Reticulate,
distinct,

big, irreg-
ular

Reticulate,
distinct,

big and ir-
regular

Reticulate,
distinct,

big and ir-
regular

Reticulate,
sligylt,
distinct,
regular

and small

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Ornamentation: Reticulate; reticules slightly distinct, big and irregular. Colpus area,
porus and apocolpium are slightly reticulate.

Ex/int (Wodehouse): 2/1
Exine (acetolysis): 3 mm thick, tectate – infrastructure, nexine/sexine = 1/1

Lathyrus hirsutus L. (Tab. 1, Figs. 1 – 8). A1(E) Edirne: Edirne-Doyran village en-
trance, through water channels, 60 m, 31.05.1998, F. Gunes and A. Cirpici, MUFE 5739.

Pollen class: 3-zonocolporate.

88 ACTA BOT. CROAT. 69 (1), 2010

GUNES F., CIRPICI A.

Fig. 1. Pollen grains of Lathyrus annuus (1, 2), L. gorgoni (3, 4), L. cicera (5, 6) and L. hirsutus (7, 8).
1, 3, 5, 7 – equatorial view; 2, 4, 6, 8 – polar view (LM). Bar denotes 15 mm.

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Pollen group: Subprolate [P/E = 1.188 (Wodehouse)) – prolate (P/E=1.716 (aceto-
lysis)].

Dimensions: Medium size [P ´ E = 38.976 ´ 32.799 (Wodehouse), 52.832 ´ 30.784
(acetolysis)].

Apertures: Ectoapertures – colpi: long, slightly wide (Wodehouse), narrow (acetolysis),
nearly reaching the poles (Wodehouse), with acute ends, thick costae along the margins of
the colpi, colpus membrane covered by small granules; clg: 27.882 (Wodehouse), 47.796

ACTA BOT. CROAT. 69 (1), 2010 89

POLLEN MORPHOLOGY IN LATHYRUS

Fig. 2. Ornamentation in pollen grains of Lathyrus annuus (1, 2); L. gorgoni (3, 4); L. cicera (5, 6);
L. hirsutus (7, 8) (SEM). 1, 3, 5, 7 – equatorial view; 2, 4, 6, 8 – polar view.

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(acetolysis) mm, clt: 3.712 (Wodehouse), 1.960 (acetolysis) mm. Endoapertures – pori:
lalongate to circular, plg: 8.294 (Wodehouse), 10.556 (acetolysis) mm, plt: 10.701 (Wode-
house), 10.136 (acetolysis) mm and plg/plt= 0.775 (Wodehouse), 1.041 (acetolysis).

Outlines: Equatorial view – eliptic to rectangular-obtuse-convex; polar view circular
Ornamentation: reticulate; reticules sligylt, distinct, regular and small. Colpus area is
psilate. Porus and apocolpium are slightly reticulate.

Ex/int (Wodehouse): 1/1
Exine (acetolysis): 2 mm thick, tectate – infrastructure, nexine/sexine = 1/1

Discussion

Pollen dimensions varied in three of the taxa; the exception is L. annuus. The subprolate
type of pollen shape (P/E) was found in fresh pollens of L. gorgoni, L. cicera and L.
hirsutus, but the prolate shape was found in fossilized pollens of those three species. In
contrast to L. gorgoni, L. cicera, L. hirsutus, subprolate shape (P/E) was observed in both
fresh and fossilized pollens of L. annuus (Tab. 1).

Pollen dimensions and shape differed in fresh and fossilized pollens (Tab. 1). The ratio
of plg to plt in fresh pollens was 0.775 in L. annuus, 0.856 in L. gorgoni, 0.539 in L. cicera,
and 0.775 in L. hirsutus. The values found in fossilized pollens were 1.276 in L. annuus,
0.943 in L. gorgoni, 0.697 in L. cicera, and 1.041 in L. hirsutus. Looking at the pore length,
the ratio of plg to plt in fossilized pollens revealed an increase in the four species.

Although a tectate-infrastructure (exine) and a reticulate ornamentation were seen in all
four taxa (IVERSEN and TROELS-SMITH 1950, AYTUG et al. 1971, MOORE et al. 1991,
TOSHEVA and TONKOV 2005), our study showed that the reticulate size and its arrangement
were distinct from each other (Tab. 1, Fig. 2) and that L. annuus possessed a distinct, big,
and evenly shaped reticulation. A very distinct and unevenly shaped reticulation was also
seen in L. gorgoni. Similar to these results in L. gorgoni, a very big distinct and uneven
shaped reticulation was observed in L. cicera. A slightly distinct, evenly shaped and small
sized reticulation was found in L. hirsutus. Although a reticulate pattern in all Lathyrus
species is clearly distinct in the center of its pollens, a distinguishable reticulate character-
istic appeared to be diminished in the split and holes of pollens. Reticulation was clearly
distinguishable in the polar regions of L. annuus, L. gorgoni and L. cicera, whereas L.
hirsutus (Fig. 2). Operculum above pores and colpus had a reticulated appearance.

The contribution of this study to previously reported data on the study of pollen mor-
phology can be summarized as follows:

1. The pollen grains are 3-zonocolporate of subprolate – prolate type (P/E = 1.112 – 1.188
(Wodehouse), 1.114 – 1.716 mm (acetolysis)), medium to large in size. The smallest
pollen grains belong to L. annuus (P´E = 36.679 ´ 33.085 mm in wodehouse and
Asetolysis) and the biggest to L. cicera (P/E = 41.707 ´ 31.581 [Wodehouse) – (P/E =
60.320 ´ 38.324 mm (acetolysis)]. Equatorial view is elliptic to rectangular-obtuse-con-
vex. The polar view in all pollen grains is circular.

2. The aperture system consists of ectoapertures – colpi and endoapertures – pori. The
colpi are generally straight, narrow, with acute ends and thick costae along the colpus
regions. There are small granules in colpus membrane. The pori are large, circular to
lalongate.

90 ACTA BOT. CROAT. 69 (1), 2010

GUNES F., CIRPICI A.

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3. The thickness of the exine is 1 mm in Wodehouse and 2–3 mm in acetolyse, the nexine as
thick as the sexine.

4. The ornamentation is reticulate in all four taxa, but the size and the arrangement of the
lumina are different from each other. The reticulate ornementation is clear in meso-
colpium, while the apocolpium, the colpus area and the porus are psilate or small.

Although pollens in observed taxa seemed to be of a type 3-zonocolporate characteris-
tic, the shapes and largeness of apertures, the largeness of the polar triangle, the clearness
of the aperture borders, the shape of the pollens (P/E), the thickness of the exine and the
intine, played an important key for the diagnostic of the differentiation of the taxa AYTUG
(1959).

In conclusion, it can be stated that the four taxa discussed, although belonging to the
same section, can clearly be differentiated based on the palynological findings as de-
scribed.

Acknowledgements

This work is a part of the project supported by the Institution of Science at Marmara
University (1996 FEN-16).

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GUNES F., CIRPICI A.

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Composite  150 lpi at 45 degrees