Thaiszia - J. Bot., Košice, 21: 73-83, 2011
http://www.bz.upjs.sk/thaiszia
Systematic value of foliar epidermal morphology in
some taxa of the tribes: Urticeae and Parietariae of
the West African Urticaceae
1*
2
2
AKEEM BABALOLA KADIRI , BOLA OBOH & CHIMA OHA
1
Department of Botany, University of Lagos, Akoka Yaba, Lagos, Nigeria;
abkadiri2001@yahoo.com
2
Department of Cell Biology and Genetics, University of Lagos, Akoka Yaba, Lagos,
Nigeria; bolaoboh@yahoo.co.uk
*Corresponding author
Kadiri A. B., Oboh B. & Oha C. (2011): Systematic value of foliar
epidermal morphology in some taxa of the tribes: Urticeae and
Parietariae of the West African Urticaceae. – Thaiszia – J. Bot. 21:
73-83. – ISSN 1210-0420.
Abstract: The foliar epidermal characteristics of all West African
species of the tribe Parietarieae and some taxa in the tribe Urticeae
were investigated by the means of light microscopy for the
purposes of easy identification and justification of recent taxonomic
merging of Fleurya with Laportea. In the tribes, cell number is
higher on the adaxial surface (52 to 110) than on the abaxial
surface (19 to 96), stomata number varies from 31 to 42 per mm²
while cell size ranged from 60.8 - 70.4 µm x 22.4 - 32.0 µm on the
adaxial surface and 32.0 - 40.0 µm x 16.0 - 22.4 µm on the abaxial
surface. Generally, all the species have hypostomatic leaves and
anomocytic stomatal type but paracytic and anisocytic types are
diagnostic for Laportea aestuans, Laportea ovalifolia, Parietaria
laxiflora, and Laportea alatipes. Glandular and simple trichomes of
different sizes were observed in all the taxa except in Girardinia
heterophylla and L. ovalifolia. The cell shape, anticlinal wall pattern,
and stomatal type which correlate to a reasonable extent have
supported the amalgamation of Laportea and Fleurya. Crystal
sands of calcium oxalate were found in L. ovalifolia, Girardinia
condensata, and Parietaria laxiflora while peltate scales
distinguished P. laxiflora from other species. The species have
medicinal and economic values. An artificial indented dichotomous
taxonomic key for separating the taxa is presented.
Keywords: Urticaceae, Urticeae, Parietariaeae, leaf epidermis,
morphology, West Africa.
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Introduction
Urticaceae, the nettle plant, is a family of flowering plants consisting of herbs,
shrubs and trees (HUTCHINSON & DALZIEL 1963; HUTCHINSON 1967; HEYWOOD
1978). The family has 45 genera with about 700 species (HEYWOOD 1978).
There are eleven genera in West Africa and four of these which are grouped in
two tribes (Urticeae and Parietarieae) were selected for the present study. The
taxa were specifically selected for study because of the recent taxonomic
change, lack of sufficient information on leaf endo-morphological characteristics
and synonymy problem. Fleurya Gaud. has been merged with Laportea Gaud.
(LOWE & SOLADOYE 1990). There is a very thin line of distinction between
Parietarieae and Boehmerieae (HEYWOOD 1978) and perhaps because of the
ecological preference (montane areas) of Parietarieae, taxonomic information
about it is scanty in West Africa; furthermore, the two species of Girardinia are
often regarded as synonyms and Droguetia is poorly represented in and
taxonomic information about it is almost lacking (HUTCHINSON & DALZIEL 1963).
The following taxa were investigated, Urticeae: Droguetia iners (Forsk.)
Schweinf., Laportea aestuans (Linn.) Chew., syn. Fleurya aestuans (Linn.) Miq.,
Laportea ovalifolia (Schumach.) Chew., syn. Fleurya ovalifolia (Schum. &
Thonn.) Dandy, Laportea alatipesHook f., Laportea mooreana (Hiern.) Chew.,
Girardinia heterophylla (Vahl.) Decne, Girardinia condensata (Hochst. ex Steud.)
Wedd., syn. Girardinia diversifolia (Link.) Friis, and Parietarieae: Parietaria
debilis Forst. f. and Parietaria laxiflora Engl. were studied. These species have
culinary, clothing and aesthetic uses (BURKILL 1997; HUTCHINSON & DALZIEL
1963). Based on exo-morphology, BENTHAM & HOOKER (1965) grouped all taxa
in the family in one tribe Urereae while HEYWOOD (1978) subdivided the family
into six tribes and our genera were grouped into two tribes Urticeae and
Parietarieae. In the later account documented by LOWE & SOLADOYE (1990), the
genus Fleurya Gaud. was changed to Laportea Gaud. and all the species were
transferred based on exo-morphological traits. The present investigation
therefore aims at justifying the merging of Fleurya with Laportea using endomorphological features of the leaf and also to document the leaf epidermal
features of other species in order to facilitate their delimitation and ease their
recognition even if leaf samples are available in fragments. The only existing
report on leaf endo-morphology in the group is the general family account by
METCALFE & CHALK (1950, 1979) with scanty information on few taxa of the
tribes: Urticeae and Parietarieae, which are the major focus of the present work;
however, other taxa in family are not also sufficiently accounted for. The present
study will therefore contribute more taxonomic data that will enhance taxonomic
understanding of the group. In order to facilitate this, an artificial indented
dichotomous taxonomic key is therefore presented to assist in taxa delimitation.
Material and methods
Preserved specimens obtained from the herbarium of the Forestry Research
Institute of Nigeria, Ibadan (FHI) and fresh specimens collected from field trips
74
were used for the study. Specimens deposited in the herbarium of the University
of Lagos (LUH) were visited for authentication and study. Herbarium
abbreviations follow HOLMGREN et al. (1990). The herbarium indices of the
investigated species are given as follow: D. iners (FHI 8889), F. eastuans (FHI
33365), F. ovalifolia (FHI 51098), L. alatipes (FHI 29958), L. mooreana (LUH
3163), L. ovalifolia (FHI 101554), G. heterophylla (FHI 97877), G. condensata
(FHI 42715), P. debilis (FHI 97893) and P. laxiflora (FHI 39217). Specimens of
synonymous taxa were compared.
Methods follow ADEDEJI (2004), AKHIL & SUBHAN (1997), KADIRI et al. (2007),
KOTRESHA & SEETHARAM (2000), OGUNDIPE & W UJEK (2004), REJDALI (1991).
Portions of 2–5 cm² were cut from the standard median part of the leaf lamina
near the mid-rib, or the whole leaf was used. Dried leaves were boiled in water
for thirty minutes and subsequently soaked in either concentrated trioxonitrate (v)
acid (HNO3) in capped specimen bottles for about 8–24 hrs to macerate the
mesophyll, or irrigated in sodium hypochlorite solution (commercial bleach) for
30–120 minutes to bleach the leaf portions. Tissue disintegration was indicated
by bubbles and the epidermal layers were separated and transferred into Petri
dishes containing water for cleansing. In case of fresh materials, the leaves were
scraped with razor blade to separate epidermis. Tissue debris was cleared off
the epidermis with fine-hair brush and washed in several changes of water.
Drops of different grades of ethanol, 50 % – 100 %, were added in turn to
dehydrate the cells. The preparations were later stained with Safranin O in 50 %
alcohol for about five minutes before being mounted in glycerine on glass slides.
The epidermal layers were mounted on glass slides with the uppermost surfaces
facing up, covered with cover-slips and ringed with nail varnish to prevent
dehydration. Photographs of epidermal features of the leaf were obtained with
the aid of digital Motic camera attached to a microscope and Pentium IV
computer while line diagrams of structures were made with camera lucida.
Results
D. iners has the highest number of trichomes on the adaxial surface.
Trichomes occur on both surfaces of the leaves of D. iners, L. aestuans, L.
alatipes, G. condensata and Parietaria spp. while L. ovalifolia and G.
heterophylla are glabrous (Plates 1 and 2, Tab. 1). The epidermal cell shape is
polygonal on both surfaces of D. iners, G. condensata and Parietaria spp., but it
is irregular on both surfaces of L. ovalifolia and L. aestuans. It is
polygonal/irregular on the surfaces of L. alatipes, L. mooreana and specimens of
L. ovalifolia (Fig. 1, Plate 1, Tab. 1). The two species of Parietaria showed limited
variations in their foliar epidermal features. The leaf is pubescent on both
surfaces; the anticlinal wall is undulate while the cell shape is also uniform on
both surfaces (Fig. 1, Plates 1 and 2, Tab. 1). Mean cell size is 45.2 - 107.8µm x
20.5 -40.6µm and stomatal size is 18.9 µm x 7.7 - 10.6 µm. Cell wall thickness
varies from 4.3 – 8.8 µm on both surfaces (Tab. 2). The species grouped under
the tribe: Urticeae are more variable in their foliar epidermis than the species of
Parietarieae. The leaf is either glabrous on both surfaces in G. heterophylla and
75
L. ovalifolia or pubescent on both surfaces in D. iners, G. condensata, L.
aestuans, L. alatipes, L. mooreana and Parietaria spp. (Tab. 1). Intra-generic
differences in these features were also recorded. Mean cell size is 27.0 - 57.8
µm x 10.5 - 27.2µm and stomatal size is 13.4 - 20.8 µm x 6.7 – 11.9 µm. Cell
wall thickness varies from 3.1 – 4.8µm on both surfaces (Tab. 2). Stomatal types
recorded included the anomocytic type which is present in all species (Plate 1),
anisocytic which is restricted to Laportea species: L. alatipes, L. aestuans and L.
ovalifolia and paracytic type which occurred only in P. laxiflora (Tab. 1).
Cystoliths filled the cell lumen in all Laportea species including specimens
regarded as Fleurya (Fig. 1 D, E, F, H, K).
B
A
G
F
E
I
D
C
K
J
H
L
Fig. 1: Line drawings of some taxonomically useful foliar epidermal features of
some members of West African Urticaceae. A: D. iners (short conical trichomes),
B: P. debilis (bulbous base glandular trichome), C: P. laxiflora (large cell size in
Parietaria spp.), D, E (L. aestuans ), F (L. ovalifolia), G, H (specimen designated as
Fleurya ovalifolia at FHI) and K (L. alatipes) have cystoliths within cell lumen; C, D
(L. aestuans); F, H, J (G. heterophylla) and L (L. alatipes): show anomocytic
stomata, the common stomatal type in the two tribes. The leaf is hypostomatic. I:
P. laxiflora (scar of bulbous base glandular trichome). Adaxial surface: A, B, E, G, I
and K; abaxial surface: C, D, F, H, J, and L. Scale is 50µm.
76
Tab. 1: Relative quantitative and qualitative characteristics of foliar epidermis of the species of the tribes: Urticeae and
Parietarieae of the West African Urticaceae.
Species
Tribe: Urticeae
Droguetia iners
Laportea aestuans
syn. Fleurya aestuans
Laportea alatipes
77
Laportea mooreana
syn. Fleurya mooreana
Laportea ovalifolia
syn. Fleurya ovalifolia
Girardinia heterophylla
Girardinia condensata
Tribe: Parietarieae
Parietaria debilis
Parietaria laxiflora
Surface Cell shape
Wall pattern
Stomatal type
Trichome type
Cell no.
2
(x) per mm
Stomatal no.
2
(x) per mm
Trichome no.
2
(x) per mm
adaxial
abaxial
adaxial
abaxial
adaxial
abaxial
adaxial
abaxial
adaxial
abaxial
adaxial
abaxial
adaxial
abaxial
polygonal
polygonal
irregular
irregular
polygonal/irregular
polygonal/irregular
polygonal/irregular
irregular
irregular
irregular
irregular
polygonal/irregular
polygonal
polygonal
curved
curved
curved
curved
straight/curved
straight/curved
straight/curved
straight/curved
curved
curved
straight/curved
straight/curved
straight
straight
absent
anomocytic
absent
anomocytic, anisocytic
absent
anomocytic, anisocytic
absent
anomocytic
absent
anomocytic, anisocytic
absent
anomocytic
absent
anomocytic
present
present
present
present
present
present
present
present
absent
absent
absent
absent
present
present
110
62
41
33
39
27
30
26
46
19
34
49
45
97
absent
3
absent
15
absent
14
absent
10
absent
32
absent
42
absent
10
16
3
1
8
7
7
7
6
absent
absent
absent
absent
14
55
adaxial
abaxial
adaxial
abaxial
polygonal
polygonal
polygonal
polygonal
undulate
undulate
undulate
undulate
absent
anomocytic
absent
anomocytic, paracytic
present
present
present
present
64
24
16
20
absent
12
absent
5
1
75
7
6
Tab. 2: Relative quantitative characteristics of foliar epidermis of the species of the tribes: Urticeae and Parietarieae of the
West African Urticaceae.
Species
Tribe: Urticeae
Droguetia iners
Surface
78
adaxial
abaxial
Laportea aestuans
adaxial
syn. Fleurya aestuans abaxial
Laportea alatipes
adaxial
abaxial
Laportea mooreana
adaxial
syn. Fleurya mooreana adaxial
Laportea ovalifolia
adaxial
syn. Fleurya ovalifolia abaxial
Girardinia heterophylla adaxial
abaxial
Girardinia condensata adaxial
abaxial
Tribe: Parietarieae
Parietaria debilis
adaxial
abaxial
Parietaria laxiflora
adaxial
abaxial
S.E= Standard error
Epidermal cell length
Epidermal cell width
Epidermal cell thickness Stomatal length
Stomatal width
Min(Mean±S.E)Max (µm) Min(Mean±S.E)Max (µm) Min(Mean±S.E)Max (µm) Min(Mean±S.E)Max (µm) Min(Mean±S.E)Max (µm)
32.0(36.6±4.12)48.0
32.0(35.5±1.72)40.0
29.0(36.1±4.0)48.0
32.0(39.6±4.4)52.0
29.0(37.7±4.2)48.0
19.2(27.0±5.3)42.0
27.0(35.7±4.1)45.0
16.2(25.0±5.0)40
48.0(50.9±2.2)56.0
35.0(41.9±3.4)48.0
38.0(43.5±7.1)64.0
38.0(45.1±3.1)51.0
22.0(29.1±4.1)42.0
22.0(31.3±1.3)32.0
16.0(17.6±0.8)19.0
21.0(10.5±0.5)13.0
8.0(11.2±1.1)13.0
10(11.8±1.1)14.4
13.0(17.9±2.3)24.0
10.0(13.4±2.3)19.0
11.0(16.9±2.2)22.0
8.0(10.2±2.1)16.0
22.4(25.6±2.1)32.0
16.0(17.9±2.4)26.0
16.0(21.7±2.3)25.6
14.0(18.5±1.6)22.4
8.0(10.6±1.2)12.8
13.0(17.6±2.4)24
4.0(4.7±0.2)5.1
4.0( 4.6±0.2)4.8
1.6(3.4±0.8)4.8
3.2(3.6±0.2)4.8
3.2(3.5±0.7)4.4
2.9(3.3±0.2)4.0
3.0(2.9±0.6)4.0
2.4(3.0±0.3)4.0
3.2(4.3±0.7)6.0
3.2(3.7±0.8)4.0
3.7(4.3±0.8)4.8
4.0(4.6±0.2)4.8
3.2(3.5±0.1)3.8
2.4(3.4±0.5)4.8
absent
12.8(15.3±1.0)16
absent
12.8(15.4±1.0)16.0
absent
11.2(13.4±0.7)14.0
absent
10.2(11.4±0.8)13.0
absent
16(16.6±1.2)18.0
absent
16.0(17.9±1.2)21.0
absent
11.0(16±3.0)24.0
absent
6.4(6.7±0.1)8
absent
8.0(9.3±0.7)9.6
absent
6.2(8.3±0.8)9.6
absent
5.2(6.3±0.7)7.8
absent
8.0(9.3±0.8)11.0
absent
8.0(11.5±1.7)16.0
absent
6.4(8.6±0.9)11.2
37.0(45.2±3.2)51.0
35.0(40.3±3.2)48.0
69.0(91.9±8.7)112.0
96.0(107.8±9.0)136.0
19.2(20.5±2.8)29
16(21.1±4.5)25
32(40.6±6.1)56.0
32(40.0±4.6)51.0
4.8(5.2±0.3)5.6
3.2(4.3±0.8)4.8
6.4(8.8±1.4)12.8
4.8(7.2±1.3)9.6
absent
11.2(18.9±1.1)16.0
absent
14.4(18.9±2.1)22.4
absent
6.4(7.7±0.7)8.0
absent
8.0(10.6±1.1)13.0
Plate1: Foliar epidermal surfaces of some species of Urticaceae. A, B: Girardinia
heterophylla; C, D: Droguetia iners; E, F: Laportea mooreana. A, C, and E: adaxial
surface, B, D, F: abaxial surface. Scale bar is 25µm.
79
Plate 2: Trichome types recorded within the tribes: Urticeae and Parietarieae. A:
Simple multiseriate acicular trichome found in Girardinia condesata, B:
Multicellular glandular trichome reported in Laportea spp., C: Unicellular nonglandular acicular type recorded in Drougetia iners, D: Unicellular tip-bent
glandular trichome found in Parietaria spp.. Scale is 75µm.
Discussion
In the general anatomical account of Urticaceae, METCALFE & CHALK (1950,
1979) reported the occurrence of laticiferous elements, glandular, non glandular
and stinging hair types, hypostomata, presence of hydathodes, fluid-loaded
epidermis, silicified and calcified cell walls which occur as cystoliths of different
forms. Lumen-filled cystoliths were encountered in all the Laportea species.
There were variations in the length and type of trichomes observed among these
species; simple multiseriate acicular trichomes were found in Girardinia,
80
multicellular glandular type was recorded in Laportea spp., simple non-glandular
long acicular type was found in D. iners and multicellular tip-bent glandular type
was recorded in Parietaria spp. and L. alatipes has the longest trichomes of
60.1µm. These features were found to have reliable systematic value. The
amalgamation of Laportea and Fleurya is supported by the foliar epidermal
characteristics especially all the qualitative features as these features correlate to
a large extent. DAVIS & HEYWOOD (1963) have described qualitative features as
those characters which are limitedly influenced by environmental factors as they
are under strict genetic control. However, DAVIS & HEYWOOD (1963); HEYWOOD
(1973) and DILCHER (1947) cautioned the application of trichomes in making
taxonomic decisions as its expression can be influenced by environmental
factors. In the quantitative characters, there was a considerable overlap in all
measurements; the mean values of epidermal cell and stomatal size are
reasonably close; therefore, the merging of these species as documented by
LOWE & SOLADOYE (1990) is strongly supported. Generally, all the species have
hypostomatic leaves and anomocytic stomatal type but paracytic and anisocytic
types were also recorded in L. aestuans, L. alatipes, L. ovalifolia and P. laxiflora.
The taxonomic relevance of trichomes in Urticaceae and the systematic
relevance of epidermal features has been articulated in angiosperms (AKHIL &
SUBHAN 1997, KADIRI et al. 2007, KOTRESHA & SEETHARAM 2000, METCALFE &
CHALK 1950 1979, OGUNDIPE & W UJEK 2004 and REJDALI 1991). Crystal sands of
Calcium oxalate was found in L. ovalifolia, G. condensata, L. ovalifolia, and P.
laxiflora while peltate scales distinguish P. laxiflora from other species. However,
the variations recorded in these features have pointed to the fact that these
species cannot be lumped together in a single tribe as done by BENTHAM &
HOOKER (1965). The species of Parietaria share many features in common;
therefore their grouping in the same tribe Parietariaeae is supported. An artificial
indented dichotomous taxonomic key for separating the taxa is presented as
follows:
Artificial intended dichotomous key for separating the species of the two
West African tribes: Urticeae and Parietariae of the family Urticaceae
1. Cell shape polygonal/irregular, anticlinal wall pattern straight/curved. Stomatal
types anomocytic, anisocytic and paracytic. Leaf pubescent on either surface,
mean cell size 27.0 - 57.8 µm x 10.5 - 27.2 µm, stomatal size 13.4 - 20.8 µm x
6.7 – 11.9µm. Cell wall thickness varies from 3.1 – 4.8 µm, cystoliths
present…………………………………………………………………Tribe Urticeae
1. Cell shape polygonal, anticlinal wall pattern undulate, stomatal types
anomocytic and paracytic. Leaf pubescent on both surfaces, Mean cell size
45.2 - 107.8µm x 20.5 - 40.6µm and stomatal size is 18.9 µm x 7.7 - 10.6 µm.
Cell wall thickness varies from 4.3 – 8.8 µm on both surfaces, cystoliths
absent………….………………………………………………..….Tribe Parietariae
Tribe Urticeae:
1. Epidermal cell shape undulate, leaf surface glabrous….Girardinia heterophylla
1. Epidermal cell shape polygonal/ irregular , leaf surface pubescent
81
2. Stoma more than twice as long as wide, mean trichome number on the
abaxial surface 55 per mm²………………………………Girardinia condesata
2. Stoma twice as long as wide, mean trichome number on the abaxial surface
3 per mm²…………………………………………………………Droguetia iners
3. Cell shape irregular on both surfaces,
4. Trichome present………………………………..…..…Laportea aestuans
4. Trichome absent…………………………………….….Laportea ovalifolia
3. Cell shape irregular/polygonal on both surfaces
5. Anisocytic stomata present………………….……….Laportea alatipes
5. Anisocytic stomata absent……………….…….….Laportea mooreana
Tribe Parietariaeae:
1. Paracytic stomata absent, trichome number greater than 50, mean cell length
almost uniform on both surfaces……………..…………...……..Parietaria debilis
1. Paracytic stomata present, trichome number less than 10, mean cell length
longer on the adaxial than abaxial surface……………………Parietaria laxiflora
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