Bangladesh J. Plant Taxon. 14(2): 101-115, 2007 (December)
CHARACTERISTICS OF THE PROXIMAL TO DISTAL REGIONS
OF THE PETIOLES TO IDENTIFY 15 TREE SPECIES OF
PAPILIONOIDEAE-FABACEAE
SAMIA HENEIDAK1 AND ABDEL SAMAI M. SHAHEEN2
Botany Department, Faculty of Applied Sciences, Suez Canal University, Suez, Egypt
Key words: Accessory ridge bundles, Crystals, Papilionoideae, Petiole anatomy, Petiole vasculature,
Ridge bundles, Secretory cavities
Abstract
Comparative studies on the structure of the vascular supply of stem-leaf transitional
zone of the petioles were carried out in 15 papilionoid tree species. Anatomical
characteristics and changes in the main vascular trace were recorded. The anatomical
features of significance include outline; epidermal cell; pericyclic fiber patterns; main
petiolar vasculature; presence, number and separation of ridge vascular bundles; presence
of additional accessory ridge bundles; crystal types; secretory elements and multicellular
trichomes. Erythrina variegata and Pterocarpus indicus show no change in the petiole
trace structure throughout their petioles from proximal to distal, while the rest of the
species have minor to major changes. Sophora secundiflora has the highest number of
ridge vascular bundles (5-6), while these are absent in the two Dalbergia species, E.
variegata, Derris robusta, Sophora davidii and S. japonica. Only Derris robusta and
Sophora japonica show unusual petiole trace structure by having two additional
accessory ridge bundles adaxial of the main trace enclosing with it by a complete ring of
pericyclic fibers. The studied species of tribe Millettieae show the presence of secretory
cavities lined by epithelial cells. The usefulness of these character states is shown for
assessing, identifying and delimiting these examined species.
Introduction
Papilionoideae is the biggest subfamily of the three widely recognized subfamilies of
Fabaceae, and consists of 31 tribes (Polhill 1981, Mabberley 1997, Lewis et al. 2005). It
contains approximately 426 genera and about 12,150 species distributed with greatest
diversity in Brazil, Mexico, East Africa and Madagascar; and a few basic stocks in
Mediterranean (Mabberley 1997).
Anatomical data have been used at all levels of taxonomic hierarchy as well as for
identification and assessment of taxonomic relationships among the taxa of flowering
plants (Stuessy 1990). Vesque (1885) studied a number of families and used petiole
anatomy to separate genera and families formerly united. Subsequently, the study of
petiole anatomy has received much attention and has provided many useful characters
1
Corresponding author. E-mail: sheneidak2000@yahoo.com 2Botany Department, Aswan Faculty of
Science, South Valley University, Egypt. E-mail: abdushaheen@yahoo.com
102
HENEIDAK AND SHAHEEN
(Watari 1934, Hare 1943, Metcalfe and Chalk 1950, Howard 1979, Al-Nowaihi et al.
1980, Khalifa and El Gohary 1982). Petiole anatomy has proved to be useful in the
delimitation of the genera Musa (Ennos et al. 2000), Quercus (Maria and Rodriigo 2003),
Cucurbita (Agbaywa and Noukwu 2004) and Ficus (Saquaro 2005).
Regarding the family Fabaceae (Leguminosae), Watari (1934) examined the
vasculature of the petioles and leaves of 133 species of its three subfamilies. Of these, 96
species were from Papilionoideae, 12 from Mimosoideae, and 25 from Caesalpinioideae.
Shaheen (1995, 2006, 2007) studied the anatomy of the stem-leaf transitional zone of the
petiole, and used the results to delimit and identify some species of mimosoid and
caesalpinioid species. Taxonomically useful anatomical characters included pericyclic
fiber patterns, main petiole trace types, number and position of secondary bundles and
additional accessory bundles, as well as crystal and trichome types.
No comprehensive work has been done on petiole anatomy and their proximal to
distal vascular supply in trees of the subfamily Papilionoideae. Thus, the present
comparative anatomical investigation was carried out based on fresh samples from the
proximal, medial and distal regions of the petioles of 15 papilionoid species to obtain a
good guide and additional characters for the identification and delimitation of these
important ornamental trees.
Materials and Methods
Fresh materials of mature petioles of 14 cultivated species of Papilionoideae was
collected from Plant Island, Aswan, Egypt on 15 May 2006, while Sesbania sesban was
collected from Aswan Faculty of Science, South Valley University, Egypt on the same
day (Table 1). All the studied taxa are cultivated plants except Sesbania sesban, which is
wild and cultivated. These species were identified by comparison with herbarium
specimens kept at ASW Herbarium (Aswan Faculty of Science Herbarium, South Valley
University, Egypt), and the herbarium of Plant Island, Aswan, Egypt. Herbarium
specimens were prepared for all species from the collected fresh materials, and are
deposited in the ASW Herbarium and Biological Sciences Department, Suez Faculty of
Education, Suez Canal University, Egypt.
Three samples per species from different individuals and three petioles per sample
were examined. Petiole segments measuring 2-3 mm long were cut from proximal to
distal regions. These segments were fixed and preserved in formalin acetic alcohol
(FAA), then transferred to 70% ethanol (Johansen 1940). Using a hand microtome in Ain
Shams University, Egypt, sections (10-20 µm) were obtained and double stained with
safranin (1% solution in 50% ethanol) and light green (1% solution in absolute ethanol
and clove oil) (Corgan and Widmoyer 1971). These sections were photographed using an
Olympus photomicroscope, Model CH3ORF200, Japan at Public Service Center of
Biological Control, Faculty of Agriculture, Suez Canal University.
CHARACTERISTICS OF THE PROXIMAL TO DISTAL REGIONS OF THE PETIOLES
103
Table 1. The classification, uses and distribution of the studied 15 papilionoid species according to ElHadidi and Boulos (1979), Lock (1989) and Lewis et al. (2005).
Tribe
Dalbergieae
Millettieae
Phaseoleae
Sesbanieae
Sophoreae
Species
Dalbergia lanceolaria L.f. subsp.
paniculata (Roxb.) Thoth.
[Syn: D. paniculata Roxb.]
D. sissoo Roxb. ex DC.
Pterocarpus indicus Willd.
Tipuana tipu (Benth.) Kuntze
[Syn: Machaerium tipu Benth.]
Bolusanthus speciosus (Bolus)
Harms [Syn: Lonchocarpus
speciosus Bolus]
Pongamia pinnata (L.) Pierre
Derris elliptica (Wallich) Benth.
D. robusta (Roxb. ex DC.) Benth.
Cajanus cajan (L.) Millsp.
Erythrina variegata L. [Syn: E.
indica Lam.]
Glycine sinensis Sims.
Sesbania sesban (L.) Fawc. &
Rendle
Sophora davidii (Franchet) Skeels
[Syn: S. viciifolia Salisbury]
S. japonica L. [Syn: Styphnolobium
japonicum (L.) Schott]
S. secundiflora (Gomez-Ortega)
Lag. ex DC.
Use
Ornamental
Distribution
Africa
Fodder & timber
Ornamental
Ornamental
Africa & Asia
Africa & Asia
Africa & South America
Ornamental
Zaire (Africa)
Ornamental
Ornamental
Ornamental
Human food &
fodder
Ornamental
Africa & Asia
Africa
Africa
Africa
Ornamental
Fibre, fodder,
medicinal &
ornamental
Ornamental
Medicinal &
ornamental
Ornamental
Africa, Asia, and Indian &
Pacific Oceans
Africa
Africa & Asia
Kenya, South Africa (Africa)
& Asia
Egypt, Kenya, South Africa,
Zimbabwe (Africa) & Asia
Kenya (Africa), Central &
North America
Results and Discussion
Some recognizable variations as well as similarities were observed among the studied
15 papilionoid species in terms of 18 anatomical characters as summarized in Tables 2 &
3 and illustrated in Figs. 1-5. In the following sections only the salient features, features
showing micro-variations within a category of a character state and the importance of
these features in taxonomy are discussed in detail under two broad headings.
Non-vascular features
Some features, namely epidermal cells shape, cortex structure, pith, crystal type,
mucilage elements showed some specific variations among the studied species. The
results of petiole outline agree with those of Shaheen (2006, 2007) who reported the
usefulness of this character in the assessment of the identification of some species of
Table 2. The characters (non-vascular) of the petiole anatomy of the studied 15 species of Papilionoideae.
Characters →
Species ↓
Figure
Outline (ridges
& adaxial
groove)
1
1
2
2
2
2
2 (circular to
rectangular)
Epidermal
cells
Cortex
1
3
1
3
2
1
1
1
1
1
1
3
1
1
Pericyclic
fibres
Dalbergia lanceolaria
Dalbergia sissoo
Pterocarpus indicus
Tipuana tipu
Bolusanthus speciosus
Pongamia pinnata
Derris elliptica
Figs. 1a-c
Figs. 1d-f
Figs. 1g-i
Figs. 2a-c
Figs. 2d-f
Figs. 2g-i
Figs. 3a-c
Derris robusta
Figs. 3d-f
2 (circular to
oblong)
3
1
Cajanus cajan
Erythrina variegata
Figs. 3g-i
Figs. 4a-c
3
1
2
1
2
1
2
1
3
2
2
2
2 (proximal)
& 4 (medial,
distal)
2 (proximal)
& 4 (medial,
distal)
2
1 (small)
Glycine sinensis
Figs. 4d-f
1
3
2
Sesbania sesban
Figs. 4g-i
2 (circular to
obtriangular)
2 (circular to
oblong)
1
3
Sophora davidii
Sophora japonica
Figs. 5a-c
Figs. 5d-f
1
2
1
3
2
1
Sophora secundiflora
Figs. 5g-i
2 (circular with
small ridge
adaxially)
2
1
3 (proximal)
& 1 (medial,
distal)
2
2 (proximal)
& 4 (medial,
distal)
2
Crystal
type
Secretory
elements
Mucilage
elements
Multicellular
trichomes
1
2
2
1
3
3
3 (hollow
center)
3
3
3
3
1
4
4
0
0
0
0
1
1
1
2
2
2
1
0
2
2
0
1
1
1
1
1
0
3
4
1
1
0
4
1 (solid to
hollow center)
3 (solid to
hollow center)
2
4
2
0
0
1
2
1
0
4
0
1
1
3
0
2
0
4
2
3
4
0
0
2
0
1
0
2
4
0
0
1
Pith
Key to attributes: Outline: 1= wavy circular, 2= wavy circular with two ridges and a shallow to deep adaxial groove, 3= angular to five angled with five angles in between them
five furrows. Epidermal cells: 1= tangentially elongated, 2= radially elongated, 3= tangentially elongated and radially elongated. Cortex: 1= large isodiametric parenchyma cells,
2= large irregular parenchyma cells, 3= outer chlorenchyma and inner parenchyma cells especially in the medial and distal regions. Pericyclic fibers: 1= isolated strands, 2= a
dissected ring, 3= continuous layers, 4= a continuous ring. Pith: 1= large isodiametric parenchyma cells, 2= small isodiametric parenchyma cells, 3= large and small isodiametric
parenchyma cells, 4= large irregular parenchyma cells. Crystal types: 1= numerous druses and rhombohedral solitary, 2= numerous rod-shaped solitary, 3= few to numerous
rhombohedral solitary, 4= numerous rhombohedral and rod-shaped solitary. Secretory elements: 1= present, 0= absent. Mucilage elements: 1= numerous brown mucilage cavities
in the cortical, phloem and pith parenchyma cells, 2= brown mucilage cells present in epidermis, cortical, phloem and pith parenchyma cells, 0= absent. Multicellular trichomes:
1= present, 0= absent.
Table 3. The characters of the petiole vasculature of the studied 15 species of Papilionoideae (v. b. = vascular bundles).
Main petiolar vasculature
Shape
Characters →
Species ↓
Figure
Dalbergia lanceolaria
Dalbergia sissoo
Pterocarpus indicus
Tipuana tipu
Figs. 1a-c
Figs. 1d-f
Figs. 1g-i
Figs. 2a-c
2
2
1
3
In proximal
1
3
6
7
Bolusanthus speciosus
Figs. 2d-f
3
5
Pongamia pinnata
Derris elliptica
Derris robusta
Cajanus cajan
Erythrina variegata
Glycine sinensis
Sesbania sesban
Sophora davidii
Sophora japonica
Figs. 2g-i
Figs. 3a-c
Figs. 3d-f
Figs. 3g-i
Figs. 4a-c
Figs. 4d-f
Figs. 4g-i
Figs. 5a-c
Figs. 5d-f
3
3
3
3
2
3
3
2
2
5
5
5
4
3
4
6
1
1
Sophora secundiflora
Figs. 5g-i
2
2
Type
In medial & distal
1
3 (8-10 v.b.)
5
2 (flat abaxially in
distal)
2 (invaginated
towards pith
adaxially)
4
1 (nearly square)
2
4
3 (12 v.b.)
4
3 (5-8 v.b.)
1
1 (medial) & 2
(distal)
2 (abaxially
compressed)
V. b.
division
Changes
Presence
of ridge
v. b.
No. of
ridge
v. b.
Separation
of ridge
v. b.
Number of
additional
accessory v. b.
3
1 (1-3 v.b.)
4
1 (2-6 v.b.)
2
2
1
3
0
0
3
1
0
0
1
2
0
0
2
2
0
0
0
0
2 (2 v.b.)
3
2
1
2
0
1
2
1
2 (1 v.b.)
4
1 (6-7 v.b.)
1 (3-4 v.b.)
3
3
3
3
3
2
1
3
3
2
2
2
1
0
1
0
1
2
0
0
1
1
0
1
0
1
1
0
0
2
2
0
2
0
2
2
0
0
0
0
2
0
0
0
0
0
2
3
3
1
3
1
0
Key to attributes: Main petiolar vasculature type: 1= open, 2= closed, 3= open in the proximal region and closed in the medial and distal regions. Main petiolar vasculature
shape in the proximal region: 1= excentric cylindrical amphiphloic siphonostele, 2= two closely neighbouring amphiphloic siphonosteles, 3= circular dictyostele, 4= wavy fiveangled dictyostele, 5= continuous bicollateral cup-shape, U-shaped, or O-shaped, 6= small bicollateral arc, 7= Large bicollateral flask-shaped. Main petiolar vasculature shape in
the medial and distal regions: 1= excentric cylindrical amphiphloic siphonostele, 2= wavy angular amphiphloic siphonostele, 3= circular dictyostele, 4= wavy dictyostele, 5= small
bicollateral arc. Main petiolar v.b. division: 1= number of bundles increased laterally and abaxially, 2= number of bundles increased adaxially, 3= xylem division only, 4=
unchanged bundles. Main petiolar vasculature changes: 1= no changes, 2= minor changes, 3= major changes. Presence of ridge bundles: 1= in the three regions of the petioles, 2=
only in the medial and distal regions, 3= only in distal regions, 0= absent. No. of ridge v. b.: 1 = constantly 2 (1 at each ridge side), 2 = 4 bundles (2 at each ridge side), 3 = 5-6
bundles, 0 = absent. Separation of ridge v.b.: 1= from the adaxial sides of the two vascular rings, 2= from both ends of the open main trace, 0= absent. Number of additional
accessory v. b.: 0= absent, 2= presence of two bundles.
106
HENEIDAK AND SHAHEEN
mimosoid and caesalpinioid species. Metcalfe and Chalk (1983) also listed the presence
of mucilage cavities and cells in primary cortex, phloem or pith of the species of the
family Fabaceae. Our findings match their records.
Figs. 1a-i. Cross-sections of the proximal (a, d, g), medial (b, e, h) and distal (c, f, i) regions of the petiole:
a-c. Dalbergia lanceolaria subsp. paniculata, d-f. Dalbergia sissoo, g-i. Pterocarpus indicus (tribe
Dalbergieae). Ab = Abaxial bundles, Ad = Adaxial bundles, C = Cortex, E = Epidermis, H = Multicellar
hairs, I = Inner phloem, L = Lateral bundles, O = Outer phloem, P = Pith, Pf = Pericyclic fibers, Rb =
Ridge bundles, X = Xylem. (Bar = 1 mm)
CHARACTERISTICS OF THE PROXIMAL TO DISTAL REGIONS OF THE PETIOLES
107
Pericyclic fiber forms: The main petiolar vascular supply is strongly supported by the
pericyclic fibers outside the outer phloem in all examined species. These patterns appear
to be a diagnostic character between the studied species (Table 2). Generally, the
characters of pericyclic sclerenchyma in the petioles are diagnostic to Pterocarpus
indicus, Bolusanthus speciosus, Pongamia pinnata, Derris elliptica, D. robusta and
Sophora japonica. In this respect, the presence or absence of pericyclic sclerenchyma in
woody plants appears to have taxonomic value (Petit 1887). Metcalfe (1983) mentioned
that phloem fibers are of considerable taxonomic importance. Ibrahim (1996) reported
that a continuous ring of sclerenchyma cells surrounds the main petiolar vasculature in 27
species of subfamily Caesalpinioideae, and the remaining species have either a dissecting
ring or isolated strands.
Crystal type: Crystal is a characteristic of Bolusanthus speciosus (numerous druses
and rhombohedral solitary crystals) and Erythrina variegata (numerous rod-shaped
solitary crystals). In this respect, Metcalfe and Chalk (1950) reported the presence of rodshaped solitary crystals (styloids) in the palisade tissue of Derris, Pongamia and Sophora
species. Also, Ibrahim (1996) recorded the presence of druses in 11 species, solitary
crystals in 10 species, solitary and druses crystals in 6 species, and crystals absent in 6
species of subfamily Caesalpinioideae.
Secretory elements: These elements are restricted to the cortical region of the petiole
of Bolusanthus speciosus, Derris elliptica (small in these two species), Pongamia pinnata
and Derris robusta (large in these two species, Figs. 2g & 3d) of tribe Millettieae. Each
secretory element is formed of a large intercellular space surrounded by a layer of
tangentially flattened papillose epithelial cells. Earlier, Metcalfe (1983) recorded the
presence of these secretory elements in Bolusanthus species (Papilionoideae), and their
presence is of diagnostic rather than of taxonomic value.
Multicellular trichomes: Sparsely distributed multicellular-uniseriate trichomes were
found in nine studied species (Table 2). In Bolusanthus speciosus and Cajanus cajan
these are, however, dense and long (Figs. 2d-f & 3g-i). Shaheen (2006, 2007) also
showed trichomes seemed to be of taxonomic interest for the distinction of some of
mimosoid and caesalpinioid species.
Vascular bundle features
Some features, namely tendency of bundles to divide in the main petiole vasculature,
changes within the petiole of the same species, presence of ridge (secondary) vascular
bundles, number of ridge vascular bundles and separation of ridge bundles are
summarized in Table 3. However, a few features need detailed discussion as illustrated
below.
Main petiole vasculature type: Within the taxa studied, petiole vasculature type
(Table 3) seems to be diagnostic to the two Dalbergia species, Pterocarpus indicus,
108
HENEIDAK AND SHAHEEN
Erythrina variegata and Sophora davidii. In addition, the unique type of petiole
vasculature in Bolusanthus speciosus and Pongamia pinnata may support a close
relationship between them belonging to tribe Millettieae. In this respect, De Candolle
(1879) proposed two types of petiole vasculature: open - where the bundles are arranged
in an arc, and closed - where the bundles form circular comparable to that of the stem.
His proposals were incorporated in the work of Metcalfe and Chalk (1950).
Figs. 2a-i. Cross-sections of the proximal (a, d, g), medial (b, e, h) and distal (c, f, i) regions of the petiole:
a-c. Tipuana tipu (tribe Dalbergieae), d-f. Bolusanthus speciosus, g-i. Pongamia pinnata (tribe
Millettieae). G = Adaxial groove, I = Inner phloem, M = Brown mucilage elements, R = Ridges,
Rb = Ridge bundles, S = Secretory elements. (Bar = 1 mm)
CHARACTERISTICS OF THE PROXIMAL TO DISTAL REGIONS OF THE PETIOLES
109
Tendency for the bundles to divide: The increased number of vascular bundles in the
medial and distal regions than the proximal one of the petioles (Table 3) seems to be a
diagnostic feature of Dalbergia sissoo, Tipuana tipu and Cajanus cajan. In addition, the
unique pattern of bundle division in Dalbergia sissoo and Tipuana tipu account for the
close relationship between them belonging to tribe Dalbergieae, similar relationship was
seen in Derris elliptica and Bolusanthus speciosus belonging to tribe Millettieae. These
results agree with those of Shaheen (2006, 2007) who reported the importance of this
character in the assessment of the identification and delimitation of some mimosoid and
caesalpinioid species.
Changes in vascular bundles from proximal to distal region: Three cases of change
are recorded from proximal to distal region (Table 3). The main petiolar vascular supply
has many variations in types, shapes and tendency of its bundles to divide from the
proximal to distal regions. It is diagnostic to Dalbergia lanceolaria (large amphiphloic
siphonostele), Dalbergia sissoo (small dictyostele of 7 bicollateral bundles), Sophora
japonica, S. davidii (amphiphloic siphonostele) and S. secundiflora (an abnormal
structure of two closely neighbouring amphiphloic siphonosteles; one much bigger than
the other). The complexity in the structure of the petiole vasculature is also recorded in
Faidherbia albida (Shaheen 1995) and in Quercus (Maria and Rodriigo 2003). In
contrast, Shaheen (2006, 2007) reported the importance of that character in the
assessment of the identification of some mimosoid and caesalpinioid species. In this
respect, homogeneously in the structure of the petiole vasculature is also recorded in
Acacia phyllodes and Acacia podalyriifolia (Duarte and Wolf 2005) and in some
caesalpinioid species (Shaheen 2007).
Main petiole vasculature shape in the proximal region: The shape, number and
topography vary greatly within the petiole of the same species and between the studied
species (Table 3). In the proximal region, seven types were seen:
1) excentric cylindrical amphiphloic siphonostele, in Dalbergia lanceolaria, Sophora
davidii and S. japonica (Figs. 1a, 5a, d).
2) two closely neighbouring amphiphloic siphonosteles (one much bigger than the other,
in Sophora secundiflora) (Fig. 5g).
3) circular, dictyostele, in Dalbergia sissoo (small, excentric of 7 bundles; three adaxial,
two lateral and two abaxial, Fig. 1d) and Erythrina variegata (large of 12 small
bundles; three adaxial, six lateral and three abaxial, Fig. 4a).
4) wavy five angled dictyostele, in Cajanus cajan (9 bundles of one abaxial and 8
laterals; four big and four small in between, Fig. 3g) and Glycine sinensis (5 bundles
of one big abaxial and four laterals two big and two small, Fig. 4d).
110
HENEIDAK AND SHAHEEN
5) continuous bicollateral cup-shape with two incurved ends adaxially, in Bolusanthus
speciosus (Fig. 2d), Pongamia pinnata (Fig. 2g), Derris elliptica (U-shaped, Fig. 3a)
and Derris robusta (open O-shaped of small 5 bundles; four laterals and one abaxial,
Fig. 3d).
Figs. 3a-i. Cross-sections of the proximal (a, d, g), medial (b, e, h) and distal (c, f, i) regions of the petiole:
a-c. Derris elliptica, d-f. Derris robusta (tribe Millettieae), g-i. Cajanus cajan (tribe Phaseoleae).
Ar = additional accessory ridge bundles, I = Inner phloem, M = Brown mucilage elements, Pf = Pericyclic
fibers, S = Secretory elements. (Bar = 1 mm)
CHARACTERISTICS OF THE PROXIMAL TO DISTAL REGIONS OF THE PETIOLES
111
6) small bicollateral arc in Pterocarpus indicus (continuous, Fig. 1g) and Sesbania
sesban (5 bundles, Fig. 4g).
7) large bicollateral flask-shaped; round basal part with five large bundles, and two long
incurved ends adaxially separating each end to two ridge bundles in Tipuana tipu
(Fig. 2a).
Figs. 4a-i. Cross-sections of the proximal (a, d, g), medial (b, e, h) and distal (c, f, i) regions of the petiole:
a-c. Erythrina variegata, d-f. Glycine sinensis (tribe Phaseoleae), g-i. Sesbania sesban (tribe Sesbanieae).
Ch = Chlorenchyma cells, I = Inner phloem. (Bar = 1 mm)
112
HENEIDAK AND SHAHEEN
Figs. 5a-i. Cross-sections of the proximal (a, d, g), medial (b, e, h) and distal (c, f, i) regions of the petiole:
a-c. Sophora davidii, d-f. S. japonica, g-i. S. secundiflora (tribe Sophoreae). Ar = additional accessory
ridge bundles, I = Inner phloem, Pf = Pericyclic fibers, R = Ridge bundles. (Bar = 1 mm)
Generally, this feature is characteristic to the two species of the two Dalbergia and
Derris species, Petrocarpus indicus, Sophora secundiflora, S. javanica and Tipuana tipu.
In addition, this result seems to be accounted for the close relationship between Cajanus
cajan and Glycine sinensis. In this connection, Hare (1943) and Metcalfe and Chalk
(1950) recorded main petiole vasculature strand shape composed of separate bundles or
CHARACTERISTICS OF THE PROXIMAL TO DISTAL REGIONS OF THE PETIOLES
113
continuous in the form of U-shaped, cylindrical or open arc. Ibrahim (1996) also
summarized the types of petiole vasculature in the medial regions of the petioles as
siphonostele, dictyostele and complete crescent shape.
Main petiole vasculature shape in the medial and distal regions: The major petiole
vasculature shapes in the medial and distal regions are tabulated in Table 3. However, the
wavy dictyostele show some variations among the species: Pongamia pinnata (fiveangled and invaginated towards pith adaxially, Figs. 2h-i), Cajanus cajan (five-angled
with five big bundles lying in the five angles in between them five small bundles lying in
the five furrows, Figs. 3h-i) and Glycine sinensis (triangular and invaginated towards pith
adaxially, Figs. 4e-f).
Ridge (secondary) vascular bundles: These are situated at the ridges of the adaxial
side of the petiole in its cortex with certain taxonomic significance in family
Leguminosae (Watari 1934, Ibrahim 1996). There are four cases identified in the present
study (Table 3). In case of Pterocarpus indicus, the ridge is present only in the distal
region.
Separation of ridge bundles: In terms of the characters of ridge bundles (Table 3),
our findings agree with those of Watari (1934) who recorded the presence of the ridge
bundles in Pterocarpus indicus, Derris elliptica and Sophora japonica, and their absence
in Dalbergia sissoo. In contrast, Watari (1934) reported the presence of the ridge bundles
in 53 species of subfamily Papilionoideae, 11 species of Mimosoideae, and 18 species of
Caesalpinioideae. Ibrahim (1996) recorded also the presence of the ridge bundles in the
medial regions of the petioles of 14 species of Caesalpinioideae.
Additional accessory ridge bundles: There are two such ridge bundles which lie
adjacent to the main petiole strand adaxially enclosing with it by a complete ring of
pericyclic fibers in Derris robusta and Sophora japonica (Figs. 3e-f & 5d-f). They
separate in the medial region in Derris robusta (Fig. 3e), or in the proximal region in
Sophora japonica (Fig. 5d). Shaheen (1995, 2007) reported the presence of the accessary
bundles in the core of the main trace in some mimosoid and caesalpinioid species.
Ibrahim (1996) recorded also the presence of the additional accessory ridge bundles in
the medial regions of the petioles of seven species of subfamily Caesalpinioideae.
In this study, we found major changes in the petiolar supply from proximal to distal
regions in the most of the studied species, contrary to Metcalfe and Chalk (1979) who
reported that the middle of the petiole is the most reliable position from which a single
section can be taken for comparative purposes. Our findings also suggested that petiolar
outline shape, the number of ridge bundles (two), crystal type, secretory elements and the
unique patterns in the main petiolar vasculature (especially open main vascular type in
the proximal regions and closed main vascular type in the medial and distal regions)
account for close relationship among the species of tribe Millettieae.
114
HENEIDAK AND SHAHEEN
Acknowledgements
The authors appreciate the corrections and suggestions of the referees who much
contributed to the improvement of this manuscript.
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(Manuscript received on 19 September 2007; revised on 6 November 2007)