Proc. Indian Acad. Sci., Vol. 87 B, No. 2, February 1978, pp. 55-64, 9 Printed in India.

Morphological studies in Meliaceae. II. A reinvestigation of

floral anatomy of members of Swietenieae and Trichilieae

Y S M U R T Y and S U S H M A G U P T A
Botany Department, Institute of Advanced Studies, Meerut University, Meerut 250 001

MS received 10 September 1977

Abstract. The floral anatomy of Swietenia mahogoni Jacq., Soymldafebrtfuga Juss.,

Chukrassia valutina A. Juss., Dysoxylum binectariferum Hook., Aphanamixis rohituka
W. & A. and Heynea trijuga Roxb. are discussed. Calyxis supplied by two whorls
of traces in Swietenia, Aphanamixis, Chukrassia and Heynea and only one whorl of
traces in Soymida and Dysoxylum. In all the species, except Heynea, petals receive
their supply independently. In Heynea petals receive their supply from compound
strands. Staminal tube receives its supply from compound strands in Swietenia,
Chukrassia, Heynea, Aphanamixis and from independent strands in Soymida and
Dysoxylum. Anthersare syngenesious in Dysoxylum.
Ovary is supplied only by carpellary ventrals in Soymida, carpellary ventrals and
ovary wall traces in Swietenia, carpellary ventrals, ovary wall traces and carpellary
dorsals in Chukrassia and carpellary ventrals, ovary wall traces, carpellary dorsals and
secondary marginals in Aphanamixis, Dysoxylum and Heynea. Disc is considered
receptacular. Ovary is 2-5 locular at the base but becomes unilocular above. In
Chukrassia it remains tetralocular throughout. Ovules show attachment to axile
placentae in Chukrassia, Heynea and Aphanamixis and to parietal placentae in Heynea,
Soymida, Dysoxylum and Swietenia. Non-vascular, 8-20 teeth-like structures are
formed from staminal tube. These are interpreted as fused or interpetiolar stipules.

Keywords. Floral anatomy; Meliaceae; Swietenieae; Trichilieae.

1. Introduction

A preliminary reinvestigation (Murty and Gupta 1976, 1978a,b,c) of the floral ana-
tomy of some species of Meliaceae revealed important differences from earlier wor-
kers (Saunders 1937; Narayana 1958a,b, 1959; Nair 1962, 1963). Hence a detailed
reinvestigation of Swietenia mahogoni Jacq., Soymida febrifuga Juss., Chukrassia
valutina A. Juss., belonging to Swietenieae and Dysoxylum binectariferum Hook.,
Aphanamixis rohituka W & A. and Heynea trijuga Roxb, belonging to Trichilieae
has been undertaken which has revealed interesting differences from the earlier
studies.

2. Materials and methods

The materials for the present study were collected from Saharanpur and Forest Re-
search Institute of Dehra Dun. Customary methods of dehydration, clearing, em-
bedding and sectioning were followed and stained with crystal violet and erythrosine
combination.

55
56 Y SMurty and Sushma Gupta

3. Observations

3.1. Swietenia mahogoni

As described earlier by Narayana (1958b) and Nair (1963) thepedicel shows 8-10
bundles and the short calyx tube is supplied by a whorl of five sepal medians and
another whorl of five conjoint sepal laterals, all of which show some branching
(figures 1, 2).
Petal traces are independent and the petals are single traced (figure 3). Due to
branching each petal shows 8-10 bundles. Petals show twisted aestivation (figures 6-10).
I0 compound strands move from the central vascular ring. Each of these strands
divide tangentially into three (figures 4, 5). The peripheral part goes into the staminal
tube, middle one supplies the disc and the inner one the ovary wall (figures 5, 6).
The conspicuous, thick many lobed disc is present between the staminal tube and
ovary (figures 6, 7). The ten carpellary ventrals situated on septal radii are derived
from the central vascular tissue (figure 6). They fuse in pairs to form 5 inverted
carpellary ventrals (figure 7). Ovary is pentalocular at the base and apex and is
otherwise unilocular (figures 6-8). Ovules are on parietal placentae (figure 7). Each
capellary ventral bundle divides into two above the level of ovular supply (figure 8).
The ovary wall traces fuse with the ventrals and continue up to the stigma (figure 8).
Staminal tube becomes dilated at a higher level. The anther lobes extend down-
wards below the level of attachment with the filament. The remaining part of the
staminal tube above the level of filament splits into ten thick non-vascular lobes
arranged in one whorl on alternate radii with anthers (figures 9, 10). They extend
beyond the anthers.

3.2. Soymida febrifuga

The floral anatomy of S. febrifuga follows the same pattern as described by Narayana
(1959). Sepals and petals are single traced (figures 11-13). Aestivation of sepals
is quinquncial and that of petals is twisted. Sepal and petal bundles show profuse
branching. No ridges are found in between the petals in the herbarium material.
I0 traces supply the staminal tube (figure 14). No carpellary dorsals are found.
After the supply of ovules carpellary ventrals move towards the septum and adja-
cent carpellary ventrals fuse to form five fused inverted carpellary ventrals situated
on septal radii and continue throughout the style where they may show branching.
Ovary is pentalocular at the base but it becomes unilocular above and ovules appear
on parietal placentae (figures 16--18). Disc is adnate to the base of the ovary (figure
15). Stigma is umbrella shaped and splits into five lobes at higher level (figures 18, 19).
The staminal tube immediately after its formation splits into 10 bits containing
concentric vascular bundles (figures 15, 16). From each bit a stamen separates
towards inside and the rest of the part breaks up into two non-vascular teeth like
structures lying at the back of each anther (figures 16, 17).

3.3. Chukrassia valutina

The pedicel in a transverse section shows a ring of vascular tissue (figure 20). The
vascular supply to calyx tube, petals, staminal tube and disc resembles that of
 Floral anatomy in Meliaceae 57

Figures 1-29. 1-10. Swietenia mahogonL Serial transverse sections of flower from
base upwards. 11-19. Soymidafebrifuga. Serial transverse sections of flower from
base upwards. 20-29. Chukrassia valutina. Serial transverse sections of flower from
base upwards. ScaleA: figures 1-I0, 20-29. Scale B:figures 11-19.

Swietenia mahogoni (figures 21-24). The five lobed calyx tube is longer and extends
up to the level of ovule attachment (figures 23-26).
After the departure of the compound strands which supply the staminal tube, disc
and ovary wall, the central vascular tissue organises into four carpellary dorsals that
move outward from the receptacle and four fused carpellary ventrals situated on

l'. 03)---4
 58 Y S Murty and Sushma Gupta

same radii as the carpellary dorsals (figures 24-26). Each carpellary ventral divides
into two and supplies the ovules oa axile placentae (figure 26). The adjacent carpel-
lary ventral traces move towards the septum and fuse to form fused inverted car-
pellary ventrals situated on septal radii (figure 27). At the top of the ovary car-
pellary dorsals, ovary wall bundles and branches of the carpellary ventrals fuse and
continue into style and end in the stigma (figures 28, 29). The inconspicuous disc
is adnate to the base of the ovary.
The staminal tube gets differentiated into ten stamens. At few places very small
non-vascular teeth-like structures situated in between anthers are found at the top
of the staminal tube (figures 28, 29).

3.4. Dysoxylum binectariferum

In a transverse section pedicel shows a ring of 20-25 vascular bundles of unequal size.
Calyx tube is supplied by a whorl of 8 traces representing alternately sepal medians
and sepal laterals which arise from the receptacle (figure 30). All of them show pro-
fuse branching at the periphery of the receptacle. Numerous bundles are found
arranged in a ring on the inner side of the calyx tube which splits into four, short,
thick, valvate lobes at a higher level.
Each petal is single traced (figure 31). The petals are slightly connate, thick,
hairy and show valvate aestivation with 6-14 bundles in each. Stamens are supplied
by two rings of four traces each of which the antepetalous ones arise at a lower level
from the central vascular tissue. The staminal tube contains all the 8 stamen traces
(figures 32, 33).
Above the level of stamen supply the remaining vascular tissue in the centre orga-
nises into 4 secondary marginals, 4 carpellary dorsals, 8 carpellary ventrals and some
ovary wall traces. Ovary wall traces and secondary marginals show some branch-
ing. At about this level 4 locules appear in the ovary showing the dorsals at the
back of the locules and 4 fused inverted carpellary ventrals on alternate radii with the
carpellary dorsals (figure 32). The fused ventrals supply the ovules arranged on the
projecting placentae. Ovary is observed to be pentalocular in one of the flowers.
However, the ovary becomes unllocular in all the cases higher up (figure 32). The
carpellary dorsals and the ovary wall traces extend only for some distance in the style.
Carpellary ventrals and secondary marginals continue throughout style and fuse at a
higher level and end in stigma (figures 33-35). A non-vascular disc is present in
between the staminal tube and ovary (figures 32, 33). It separates and extends up
to the level of base of the anther.
The staminal tube containing the eight concentric stamen bundles becomes
gradually dilated. At a higher level 8 dithecous anthers appear on inner side of the
staminal tube. The anthers are syngenesious with interlocking teeth connecting the
adjacent anthers and forming a tube like structure (figures 34, 35). The non-vascular
staminal tube as well as the tube formed by syngenesious anthers continue after the
separation of the filaments from the staminal tube (figures 34, 35). The non-vascular
staminal tube beyond the level of anthers splits into 8 unequal lobes one situated at
the back of each anther (figure 36).
 Floral anatomy in Meliaceae 59

Figures 30-53. 30-36. Dysoxylum binectariferum. Serial transverse sections of

flower from base upwards. 37-43. dphanamixisrohituka. Serial transverse sections
from base upwards. 44-53. Heynea trijuga. Serial transverse sections of flower
from base upwards. Scale A: figures 30-36. Scale B: figures 37-43. Scale C:
figures 44-53.

3.5. Aphanamixis rohituka

A ring of vascular tissue is found in the pedicel. Sepals are supplied by two whorls
of five traces each which arise from the receptacle (figure 37). Within the sepals all
these bundles branch at higher levels to form 6--12 or more bundles in a sepal. Sepals
show quinquncial aestivation.
60 Y S Murty and Sushma Gupta

A whorl of 5 petal traces arise from the receptacle on alternate radii of sepal medians
(figure 38). They enter 3 structures. 2 of these are larger and receive double supply
which branches into 6-14 bundles at higher level (figure 39). Petals show imbricate
aestivation.
6 or 7 stamen-ovary compound traces move outward from the receptacle (figures
39, 40). Each of these compound traces branches tangentially. The outer portion
supplies the staminal tube. The inner portions alternately constitute carpellary
dorsal and secondary marginal. Some of them may branch and supply the ovary
wall (figure 40). The central vascular tissue forms 3 carpellary ventrals. Disc is
inconspicuous and adnate to the base of ovary. Ovary is trilocular at the base and
becomes unilocular above (figures 40, 41). Placentation is axile (figure 40). The
carpellary dorsals and the carpellary ventrals are arranged on the same radius. Each
fused carpellary ventral divides into two which supply the ovules. Above the level
of ovule supply, adjacent carpellary ventrals move towards the septum and fuse to
form fused inverted carpellary ventrals (figure 41). Ovary wall traces, carpellary
dorsals, secondary marginals and carpellary ventrals extend up to different levels in
the style (figure 42).
The anthers are sessile and the stamen bundles enter directly the base of the anther
from the staminal tube (figures 42, 43). The non-vascular staminal tube extends up
to some distance only (figure 43). The anthers appear to be arranged in one whorl.
Hairs (non-glandular unicellular and needle like and glandular unicellular and
branched multicellular) are present especially on pedicel, sepals, petals and staminal
tube of all members studied. Tannin like substance and oil are also found in scattered
cells of the different floral parts of all the members studied.

3.6. tleynea trijuga

In a transverse section the pedicel shows a ring of 6--8 conjoint, collateral vascular
bundles of unequal size. The number increases due to splitting at the base of the
receptacle. 10 compound strands arise from these for the supply of sepals, petals,
stamens, disc and ovary wall (figures 44, 45). Out of these 5 strands arise a little
earlier than the other 5 which are situated on alternate radii. The 5 strands in the
outer ring divide each into one outer sepal median and an inner conjoint stamen-
disc-ovary wall strand (figure 45). The inner ring of 5 compound strands divide
each into an outer conjoint sepal traces and an inner conjoint petal-stamen-disc-
ovary wail strand (figure 46). The conjoint sepal trace branches into two to supply
the adjacent sepal mergins. At higher leveIs the valvate sepals show up to 5 bundles
(figure 48).
Each petal receives a single trace from the conjoint petal-stamen-disc-ovary wall
strand (figures 46, 47). This divides within the imbricately arranged petals which
show up to 5 bundles at higher level (figures 49-53). The staminal tube is supplied
by 10 traces derived from the 10 conjoint stamen-disc-ovary wall strands (figure 48).
The central vascular tissue constitutes the gynoecium supply, The 10 disc-ovary wall
strands supply the disc and ovary wall. The ovary wall bundles include carpellary
dorsals situated at the back of each locule (figure 48). 2 secondary marginals move
out-ward from the central vascular tissue and the remaining vascular tissue organises
into four carpellary ventrals, arranged on alternate radii with secondary marginals
(figures 47, 48). The ovary is bilocular at the base while it becomes unilocular
 Floral anatomy in Meliaceae 61

above (figures 47--49). Ovules are supplied by carpellary ventrals (figure 48). The
ovules are found attached to the placentae in both bilocular and uniloeular condi-
tions (figures 47, 48). The carpellary ventrals are consumed in supplying the ovules
at higher levels. Ovary wall bundles including carpellary dorsals fuse with the secon-
dary marginals and continue into style (figures 49, 50). They expand and show
branching at the base of the stigma (figures 51, 52). Disc is adnate to the base of the
ovary and separates immediately above the separation of the staminal tube (figure 50).
It is supplied by disc traces.
The staminal tube which is uniformly circular at lower levels becomes lobed and
wavy containing ten concentric stamen bundles. This splits at the level of stigma
into ten boat shaped structures each containing a bundle. 5 of these are outer and
opposite the petals (figure 51). At a slightly higher level an anther appears attached
to each lobe. The vascular bundle moves into the connective through the short
filament. With the departure of the vascular bundle the non-vascular part of the boat-
shaped structure splits into 2 lobes situated at the back of each anther (figures 52, 53).
Thus 20 non-vascular lobes are formed. The anthers project beyond these lobes.

4. Discussion

Floral anatomy of Swletenia mahogoni, Soymida febrifuga and Heynea trijuga

(Narayana 1958b, 1959) and Swietenia mahogoni, Aphanamixis rohituka and
Dysoxylum binectariferum (Nair 1962, 1963) has been worked out earlier. The
vascular supply to calyx, petals, staminal tube, disc and to some extent to ovary is
similar to the description given by Narayana (1958b) for Swietenia mahogoni and
Soymida febrifuga and by Nair (1962, 1963) for Swietenia mahogoni. The present
observations agree with those of Nair (1962, 1963) in the vascular supply to the
petals, stamens and ovary in Dysoxylum binectariferum and that of ovary in Aphana-
mixis rohituka. The vascular supply to calyx tube, petals, staminal tube, and disc
in Chukrassia valutina resembles that of Swietenia mahogoni. Slight variations are
observed in the vascular supply to the ovary.
While Nair (1962, 1963) has observed that in Dysoxylum binectariferum and
Aphanamixis rohituka each sepal receives single trace which divides into three, in the
present study, the calyx tube in Dysoxylum binectariferum is found supplied by 4
sepal medians and 4 conjoint sepal laterals arranged in one whorl. In Aphanamixis
rohituka the sepals are supplied by 2 whorls of 5 traces each. The outer 5 are sepal
medians and the inner 5 the conjoint sepal laterals. Narayana (1959) has observed
in Heynea trijuga that perianth is supplied by 2 whorls of 5 traces each. However,
in the present study the sepals and petals are distinct and they are supplied by 10
compound strands; 5 of these are conjoint sepal median-stamen-disc-ovary wall
strands and the other alternating 5 are the conjoint sepal lateral-petal-stamen-disc-
ovary wall strands..
Nair (1962, 1963) has observed only 3 petals in Aphanamixis rohituka each receiv-
ing a single trace. This species is generally described as showing 3 petals (Hooker
1875, etc.). In the present material also, only 3 structures are observed of which 2
are larger than the third one. The larger ones are found to be supplied by 2 petal
traces each. Hence they are to be considered as double and the petal members to
be actually 5 and not 3 as was reported by others.
 62 Y SMurty and Sushma Gupta

The 10 stamen traces in Heynea trijuga are considered by Narayana (1959) as

arising from the central vascular tissue and supplying the staminal tube but in the
present material as mentioned above the staminal tube is supplied by stamen traces
which are branched out of the compound strands. In Dysoxylum binectariferum,
the anthers are found to be syngenesious due to interlocking teeth connecting the
adjacent anthers and forming a tube like structures. Nair (1962, 1963) has not
reported such a type of condition of anthers.
Narayana (1958b, 1959) and Nair (1963) have observed only continuation of car-
pellary ventrals into the style in the members of Meliaceae studied by them. The
present observations reveal that the fused product of ovary wall traces and carpellary
ventrals continue into the style in Swietenia mahogoni and fused product of ovary
wall traces, secondary marginals and carpellary dorsals in Heynea trijuga. The
carpellary ventrals are found to be consumed in supplying the ovules in the present
material. The vascular supply to the ovary in Chukrassia valutina differs from that
of Swietenia mahogoni and Soymidafebrifuga. The ovary is supplied by 4 carpellary
ventrals, 4 carpellary dorsals which arise from the vascular tissue in the centre and
ovary wall bundles. A similar vascular supply of ovary is observed in Azadirachta
indica by Nair (1956). Narayana (1958b) also reported in Melia azedarach and
Cedrela toona the formation of carpellary dorsals from the central vascular tissue.
The fused product of ovary wall bundles, carpellary dorsals and branches of car-
pellary ventrals continue throughout the style in Chukrassia valutina.
The ovary which is bilocular (Heynea trijuga), triloeular (Aphanamixis rohituka),
tetralocular (Dysoxylum binectariferum and Chukrassia valutina) and pentalocular
(Soymida febrifuga and Swietenia mahogoni) below becomes unllocnlar above except
in Chukrassia valutina where it continues to be tetralocular. The attachment of
ovules is observed at the level of unilocular condition in Dysoxylum binectarifentm,
Soymidafebrifuga, Swietenia mahogoni and Heynea trijuga and when the ovary is bi-,
tri or tetralocular respectively in Heynea trijuga, Aphanamixis rohituka and Chukrassia
valutina. The ventral bundles are situated on septal radii in Swietenia mahogoni
and Dysoxylum binectariferum and on inner side of locules in Soymida febrifuga,
Heynea trijuga, Chukrassia valutina and Aphanamixis rohituka. The placentation is
considered as parietal in Swietenia mahogoni, Soymidafebrifuga and Heynea trijuga
by Narayana (1958b, 1959) and in Dysoxylum binectariferum and Swietenia mahogoni
by Nair (1963) and axile in Aphanamixis rohituka (Nair 1963). However, only parietal
placentation in Dysoxylum binectariferum, Swietenia mahogoni and Soymidafebrifuga
and axile in Aphanamixis rohituka and Chukrassia valutina and axile at the base and
parietal above in Heynea trijuga (see Puri 1952) are observed in the present investi-
gations.
Disc is free in Swietenia mahogoni, Dysoxylum binectariferum and adnate at the
base in Soymida febrifuga, Aphanamixis rohituka and Chukrassia valutina. It is
inconspicuous in Aphanamixis rohituka and Chukrassia valutina. In Heynea trijuga
the disc immediately separates from the ovaryabove the level of supply to the staminal
tube. The disc is supplied by compound strands in Heynea trijuga, Swieteniamahogoni,
Aphanamixis rohituka and Chukrassia valutina and it is without vasculature in
Dysoxylum binectariferum and Soymida febrifuga. The disc is considered recepta-
cular in all the members of Meliaceae studied in the present investigations (Murty
and Gupta 1976, 1978a, b, c).
Narayana (1958b, 1959) has observed extension of staminal tube as 10 teeth-like
 Floral anatomy in Meliaceae 63

structures arranged on alternate radii with anthers in Swietenia mahogoni and 20

teeth-like structures situated at the back of anthers in Soymida fabrifuga. Similar
teeth-like non-vascular structures are observed in the material studied. They are
10 in Swietenia mahogoni and Chukrassia valutina arranged on alternate radii with
that of anthers; 20 in Heynea trijuga and Soymida febrifuga arranged in pairs on the
back of each anther; 8 in Dysoxylum binectariferum one situated at the back of each
anther and a non-vascular tube-like structure in Aphanamixis rohituka which may
finally break into small teeth-like structures at a higher level.
Teeth-like structures are mentioned by Hooker (1875), Kanjilal (1901) and Duthie
(1903). Although Rendle (1938) did not mention about the teeth-like structures,
his figure of the flower of Swietenia mahogoni dearly showed ten teeth-like structures
of staminal tube situated on alternate radii with anthers. The non-vascular lobes/
teeth-like structures may be considered as interpetiolar stipules as has been inter-
preted by Murty and Gupta (1978b). The structures in Swietenia mahogoni and
Chukrassia valutina are in alternate position with anthers. These may be compared
with fused interpetiolar stipules. The condition in Dysoxylum binectariferum may
be considered as fused stipules. Where there are 20 teeth as in Soymida febrifuga
and Heynea trijuga they may be interpreted as due to lobation of the fused stipules.
The condition in Aphanamixis rohituka may be interpreted as a continuous stipular
tube.

Acknowledgements

We wish to express our thanks to Prof. V Puri for his interest and encouragement,
and also to Messrs K C Sahni, K M Vaid and Naresh Bahadur, F R I, Dehra Dun for
collection of the materials.

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