BLUMEA 24 (1978) 101-117

Epidermal hairs of Acanthaceae

Khwaja+J. Ahmad

Plant National Botanic Gardens,

Anatomy Laboratory,
Lucknow-226001, India.

Summary

Structure and distribution of the foliar epidermal hairs of 109 species and two varieties belonging to
39

hairs have
genera of the family Acanthaceae have been studied. Both glandular and non-glandularepidermal
The sub-
been recorded in the investigated taxa. The glandular hairs
may be subsessile or long-stalked.
and Glandular head
sessile glandular hairs are of two
types: i) Glandular head panduriform, 2-celled, ii)

—8- more-celled. Subfamilies Nelsonioideae and Thunbergioideae are character-

globular or disc-shaped, 2 or

i«*H hv thfi nanduriform hairs, while Mendoncioideae and Acanthoideae have glandular hairs with a globular
hairs also widely
head. Long-stalked glandular hairs are present only in nine species. Non-glandular are

multicellular
distributed in the family; they are in all but ten
species. They may be unicellular, or
present

uniseriate; rarely they are branched. Though the non-glandular hairs are of diagnostic importance at species
The
level only, in some genera
like Barleria, Ruttya,, and Äphelandra, they are quite characteristic. present

delimitation of the family Acanthaceae, involving the transfer

study does not support Bremekamp's (1965)
and the of his subfamilies Thunber-
of Lindau's (1895) subfamily Nelsonioideae to Scrophulariaceae ,
raising
of Nelsonioideae,
and Mendoncioideae to the rank of independent families. Instead, the retention
gioideae
and Acanthoideae within the family Acanthaceae is favoured.
Thunbergioideae, Mendoncioideae,

Introduction

The taxonomic and phylogenetic significance of trichomes has long been recognised

by a
number of workers (Bachmann, 1886; Solereder, 1908; Cooper, 193 2 ! Cowan, 195°!

Metcalfe & Chalk, 1950; Goodspeed, 1954; and Sporne, 1956). According to Carlquist

(1961), trichomes are, because of their easy

accessibility, perhaps the most important
the tricho-
anatomical features which could be used for taxonomic purposes. The study of

mes of Rhododendron by Cowan (1950) and of Nicotiana by Goodspeed (1954) showed that

they are
excellent characters for making out distinctions at subgeneric and generic levels.

The hairs of Solanum, according to Roe (1971), provide some of the most
epidermal
for of the hair forms characteristic
important features diagnostic purposes
as many are

by Inamdar
for a species or a section. Recent studies in Jasminum (1967), in

have
Compositae by Ramayya (1972), and in Loganiaceae by Bendre (1973) similarly
of these of
demonstrated the significance of trichomes in the systematic studies taxa

flowering plants.

with about and

The Acanthaceae „.
are a large family of flowering plants 250 genera over

the tropics. Though the family has, for been treated

2500 species spread over many years,

as a distinct taxon, its delimitation and subdivision has been the subject of great contro-

and views have been expressed regarding the systematic

versy
and divergent conflicting
(1895) divided Acanthaceae into four subfamilies: Nelsonioi-
position of various taxa. Lindau

deae, Mendoncioideae, Thunbergioideae, and Acanthoideae on the basis of types of fruits, the

number of ovules, and the presence or

absence of retinacula and their shape. Nees (1847)

had earlier recognised two groups in

Acanthaceae: Anechmatacantheae (without retinacula)
102 BLUMEA VOL. 24, No. I, 1978

and Echmatacantheae (with retinacula). While Anechmatacantheaehas two tribes, Thunbergi-

eae
and Nelsonieae, which together include Lindau's Thunbergioideae, Nelsonioideae, and

Mendoncioideae, the second Echmatacantheae has nine tribes comprising Lindau's

group

Acanthoideae. Bentham and Hooker (1876) divided Acanthaceae into five distinct tribes:

Thunbergieae, Nelsonieae, Ruellieae, Acantheae, and Justicieae. Lindau's subfamilies Thun-

bergioideae and Mendoncioideae together constitute Bentham and Hooker's Thunbergieae

while his Nelsonieae is equivalent to the latter's subfamily Nelsonioideae. The remaining

three tribes of Bentham and Hooker together form Lindau's subfamily Acanthoideae.

Van Tieghem (1908) constituted the three subfamilies Nelsonioideae, Mendoncioideae, and

Thunbergioideae of Lindau into a new family Thunbergiaceae, and Acanthaceae sensu Van

Tieghem comprised Lindau's Acanthoideae only. Bremekamp (1953, 1955, 1965) raised
and Mendoncioideae family rank and transferred Nelsonioideae the
Thunbergioideae to to

Thu the
family Scrophulariaceae. s, family Acanthaceae sensu Bremekamp consists only of

Lindau's subfamily Acanthoideae.

In contrast to
many reports of palynological, embryological, and cyto-taxonomic

studies of Acanthaceae, no
detailed and systematic investigation has been carried out on

the foliar epidermal hairs of this family. Among relatively early reports of the studies of

foliar trichomes of Acanthaceae be mentioned the works of Lindau (in Engler &
may

Prantl, 1895), Solereder (1908), and Metcalfe & Chalk (1950). Kumar and Paliwal (1975)

investigated the epidermal features, including trichomes of six species of Thunbergia, and

Elytraria acaulis and Nelsonia and

campestris. Singh Jain (1975) studied the structure and

ontogeny of different types

of trichomes present on the floral appendages of 41 taxa of

Acanthaceae. The
present investigation, which deals with the structure, distribution, and
taxonomic significance of the foliar epidermal hairs of 39 species) of Acantha-
genera (109

ceae, has been carried out with a view to help in a better understanding of the taxonomy of

this large and complicated family of flowering plants.

MATERIAL AND METHODS

The material of the species investigated consisted of mature and leaves collected
healthy
herbarium
locally, or procured as specimens from the Botanical Survey of India or
from

botanical gardens and herbaria of Sri Lanka, Singapore, Malaysia, Indonesia, and Brazil.
The names
of the species (and varieties) investigated are listed in the Table I at the end of

this paper. Epidermal hairs were examined from cuticles separated from leaves by mechani-

cal peeling (scraping with a safety razor blade) or by maceration with 10%—30% nitric

acid. The cuticles were washed with water, stained with safranin, mounted in
aqueous

glycerine and the cover slip was ringed with Canada balsam. While trichome types,
their

structure, distribution, and of variation in the family are described under 'Obser-
range

vations', trichome characters of individual species are

listed in the Table I.

OBSERVATIONS

Basically two types

of hairs are found on the leaf of Acanthaceae: (a) Glandular and
(b) Non-glandular.

(a) GLANDULAR HAIRS

They have been recorded in all investigated species of the family and can be distin-

into three
guished major categories: (i) subsessile (short-stalked) hairs, with two-celled,
 K.J. Ahmad: Hairs of Acanthaceae 103

panduriform, glandular head; (ii) subsessile (short-stalked) hairs, with two- or more-

celled, globular disc-shaped head, and (iii) long-stalked hairs with I- to several-celled
or

stalk and one- or more-celled, globular or hemispherical, glandular head.

(i) Panduriform hairs (Figs. I—7 and 36). This term was applied to the glandular

hairs of Thunbergioideae by Hobein (see Solereder, 1908). In this type the two-celled head

is roughly dumb-bell shaped or oblong with a slight narrowing in the middle. In side view

the hair shows a short one-celled stalk with the foot cell embedded in the epidermal

layer (Fig. 36). In the present study, panduriform glandular hairs have been recorded in all

the investigated species of the genera Staurogyne (Fig. 1), Elytraria (Fig. 2), Nelsonia (Fig. 3),
and Thunbergia (Figs. The shape of the glandular head while basically remaining
4—7).

panduriform, may
often
vary
in the same species, e.g. Thunbergia laurifolia (Figs. 5—7).
There is comparatively less variation in the size of the glandular head among different

species (Table I).

(ii) Subsessile (short-stalked) hairs with globular or disc-shaped head (Figs. 8—35
and 37, 38). These hairs are recorded in all the investigated genera except Staurogyne,
Elytraria, Nelsonia, Thunbergia, and in Hygrophila except the H. serpyllum. The
species
glandular head is globular or disc-shaped and is formed of 2—8
cells. The 4-celled con-

dition is the most common (Figs. 8, 21). In like Strobilanthes, Pseuderanthemum,

genera

and Ruellia glandular hairs with 8-celled head are more common (Figs. 9—11). In Ruellia

tweediana, R. lorentziana (Fig. 12), and Ruttya speciosa (Fig. 13) glandular hairs with a more

than 8-celled head are also common. Glandular hairs with 2-celled head are present in

Blepharis maderaspatensis (Fig. 14), Crossandra nilotica (Fig. 15), and C. infundibuliformis.
Glandular hairs with 2—4-celled head are common
in Aphelandra tetragona (Fig. 16),

Gymnostachyum latifolium (Fig. 17), Peristrophe tinctoria (Fig. 18), Lepidagathis purpuricaulis

(Figs. 19, 20 ), Justicia vahlii (Fig. 22), and Beloperone guttata. In some species, e.g. Asystasia
dalzelliana (Fig. 25), Fittonia verschaffeltii var.
argyroneura (Fig. 26), Hemigraphis colorata
and Dianthera nodosa have variable number of
(Figs. 27, 28), (Fig. 31), glandular hairs a

cells.

In surface view, the outline of the glandular head is circular but Mendoncia coccinea and

M. velloziana often have glandular heads which are triangular, quadrangular, or rhomboid

in shape (Figs. 32—35).

Like the panduriform hairs, these hairs also have a
short unicellular stalk with the foot

cell embedded in the epidermal layer (Figs. 37, 38).

The diameter of the globular head varies only slightly in a species. Among species of the

same genus the diameter does not vary more than I to 1.5. Thus the variation is 23 —34,11m
of Ruellia and of Strobilanthes.
in seven species 22 —34 pm in nine species Species with
typically small glandular heads are:
Petalidium barlerioides (19 Fig. 30), Hygrophila
pm,

and
polysperma (21 pm, Fig. 29), Hemigraphis colorata (22 pm, Figs. 27, 28), Lepidagathis

cuspidata and Strobilanthes scaber (both 22,1ml). Species with conspicuously large glandular
heads are Graptophyllum pictum (38 pm), Justicia procumbens (35 pm, Fig. 23), and Barleria
courtallica (35 pm, Fig. 24).

(iii) Long-stalked glandular hairs (Figs. These hairs consist of

39—53). a 1 —5-celled
uniseriate stalk terminated by a I—several-celled globular or hemispherical head. The foot

cell may arise from the middleof an epidermal cell (Figs. 41 —44, 48, 49, and 51) or
be
may

surrounded by to several epidermal cells 50). Such hairs

2
(Figs. 39 —40, 45 —47, occur

intermixed with the short-stalked glandular hairs in several species. In Hygrophila serpyl-
 104 BLUMEA VOL. 24, No. 1, 1978

Glandular hairs glandularhairs with

(Figs. 1—53). Figs. 1 —7. Sub-sessile panduriform head. 1. Staurogyne
—

longifolia ; 2.
Elytraria acaulis var. lyrata; 3. Nelsonia campestris;
; 4. Thunbergia erecta; 5 —
7. T. laurifolia. —

hairs with
Figs. 8—35. Sub-sessile glandular globular head. 8. Sanchezia nobilis; 9. Strobilanthes ixiocephalus;
10. Pseuderanlhemum Ruellia lorentziana;
malaccense; 11—12. 13. Ruttya speciosa; 14. Blepharis maderaspatensis;
Crossandra nilotica; 16. Aphelandra tetragona; Gymnostachyum latifolium; 18. Peristrophe tinctoria;
15. 17. 19—20.

Lepidagathispurpuricaulis; L. trinervis; Justicia vahlii; 24. Barleria

21. 22.
23.J. procumbens; courtallica; 25.Asysta-
sia dalzelliana; Fittonia Hemigraphis colorata;
26. verschaffeltii var. argyroneura; 27—28. 29. Hygrophila poly-
spermy 30. Petalidium barlerioides; 31. Dianthera nodosa; 32—33. Mendoncia velloziana var.
sparatteria; 34—35.
M. coccinea. hairs in sectional
—
Figs. 36—38. Glandular view. 36. Thunbergia grandiflora; 37. Hemigraphis
hirta; 38. Petalidium barlerioides. —

Figs. 39—51. Long-stalked glandular hairs. 39—43. Hygrophila serpyllum;

44. Dyschoriste erecta ; 45—48. Dyschoriste vagans; 49—51. Lepidagathis cuspidata. —

Fig. 52. Sectional view of

long-stalked glandular hair of Dicliptera roxburghiana. Fig. A branched trichome of Dyschoriste

53.
—

vagans
with glandular head branch. Scale 100 µm for Figs. scale 50 µm for 52; scale
on one —
39—51; Figs. 1
—38,
µm for Fig. 53.
250
 K. Ahmad: Hairs
J. of Acanthaceae 105

lum, however, short-stalked glandular hairs are absent. The long-stalked glandular hairs

occur densely in Hygrophila serpyllum (Figs. 39 —43)

and Dyschoriste (Figs. 45—48);
vagans

they are
quite common in Lepidagathis cuspidata (Figs. 49—51), Dicliptera roxburghiana
(Fig. 52), and Dipteracanthus patulus. In Dyschoriste erecta (Fig. 44), Justicia tranquebariensis,
Ruellia and Strobilanthes heyneanus they are and generally restricted to the
rosea, sparse

veins and the margin.

The length of the long-stalked hairs varies from than
glandular 25 pm to more a

millimeter. They are the longest in Dyschoriste vagans (Figs. 45 —48) where they measure

up
to 1062
/im
in length. In Hygrophila serpyllum (Figs. 41 —43) and Lepidagathis cuspidata
51), these hairs
(Figs. 49, commonly arise from a single epidermal cell each, instead of
having a multi-celled hair-base. Branched hairs with one branch glandular and the others

without the glandular head are rarely found in Dyschoriste vagans (Fig. 53).

(b) NON-GLANDULAR HAIRS

Distribution and frequency. Non-glandular hairs are of wide occurrence in Acantha-

of Table I), in thirteen species they

ceae. In a
majority species, they are common (+ +
,
are

found to be dense (+ + + Table I), and in (+, Table I). Non-glandular

,
21 species sparse

hairs are generally more numerous on

the veins and the margin than in the intercostal

areas, and in some species they are confined to

the veins and the
margin. In many species,

especially those of the genera Strobilanthes, Eranthemum, and Hygrophila, the non-glandular
hairs the lower confined the
on epidermis are to margin and veins but on
upper epidermis

they are
also common
in the intercostal areas. Non-glandular hairs are
absent in Thun-

bergia affinis, T. mysorensis, Ruellia tweediana, Eranthemum albo-marginata, Barleria courtallica,

Pseuderanthemum atropurpureum, P. kewense, P. variabile, Odontonema nitidum, and O.

strictum.

Shape and size. The non-glandular hairs in shape, size, number of cells, thickness
vary

and ornamentation of the wall, and in the structure of the hair-base. The size variation
within be considerable, (12 —800 /<m), Barleria lawii
a
species may e.g. Dyschoriste erecta

(106 —1416/(m), and Asystasia chelonoides (81 —1203 /im). But some species have, on an

average, longer hairs than the others. Conspicuously elongated hairs are found in Stro-

bilanthes heyneanus (average length 932 pm), Justicia procumbens (Fig. 87, 893 pm), Peristrophe
bicalyculata (822 pm), Hygrophila salicifolia (720 /<m), H. quadrivalvis (654 /<m), H. auriculata

(637 pm), Barleria lawii (590 /im), and Mendoncia velloziana var. sparatteria (590 /an). Short

hairs occur
in Pseuderanthemum malaccense (64 /im), P. grandiflorum (Fig. 65, 62 ,um),

Thunbergia grandiflora (Fig. 56, 73 /im), Eranthemum wattii (Fig. 63, 57 /mi), and Hygrophila
polysperma (55 /Mn).
The hairs are
uniseriate in all the investigated species except Elytraria acaulis var. lyrata
in which they are sometimes biseriate near the base (Fig. 94). Branched hairs are rarely
found in Pseuderanthemum bicolor
Dyschoriste vagans (Fig. 85) and (Fig. 93).
Generally, both unicellular and multicellular hairs occur in a species. Unicellular hairs

are, however, predominant in Staurogyne longifolia (Figs. 54 —55), Thunbergia grandiflora

(Fig. 56), T. dalzellii
laurifolia (Fig. 57), T. kirkii, Mendoncia coccinea (Fig. 58), Dyschoriste
(Fig. 59), D. erecta (Figs. 60—62), Eranthemum wattii (Fig. 63), E. capense (Fig. 64), Pseude-
ranthemum trinervis
grandiflorum (Fig. 65), Hygrophila serpyllum (Fig. 70), Lepidagathis
(Fig. 72),.L. cuspidata (Figs. 73—74), L. purpuricaulis (Figs. 75— 76), L. incurva (Figs. 77 —78),

Barleria cuspidata (Fig. 79), B. lupulina, B. prionitis, and Gymnostachyum latifolium. Unicellular
hairs are also quite common in Asystasia dalzelliana (Fig. 66), Justicia betonica (Fig. 67),
J. vahlii (Fig. 68), Dianthera nodosa (Fig. 69), and Rhinacanthus nasuta (Fig. 71).
 BLUMEA VOL. No.
24, 1, 1978
106

Non-glandular hairs (Figs. 54—101).

—

Figs. 54—79. Unicellular non-glandular hairs. 54—55. Staurogyne

longifolia; 5 6. Thunbergia grandiflora; 57. T. laurifolia; 58. Mendoncia coccinea; 59. Dyschoriste dalzellii; 60—62.
D. erecta; 63. Eranthemum wattii; 64. E. capense; 65. Pseuderanthemum grandiflorum; 66. Asystasia dalzelliana;
Dianthera 71. Rhinacanthus
67. Justicia betonica; 68. J. vahlii; 69. nodosa; 70. Hygrophila serpyllum; nasuta;
L. L. L.
72. Lepidagathis trinervis; 73—74. cuspidata; 75—76. purpuricaulis; 77—78. incurva; 79. Barleria cuspi-
data. —

Figs. 80—98. Multicellular non-glandular hairs. 80. Fittonia verschaffeltii var.

argyroneura;
81. F.

verschaffeltii 82. Gymnostachym febrifugum; 83. Andrographis echioides; 84. Hemigraphis hirta; 85.
var.
pearcei;
Dyschoriste vagans (branched hair); 86. Ruellia formosa; 87. Justicia procumbens; 88. Hygrophila serpyllum; 89 .

Asystasia chelonoides; 90. Adhatoda vasica; 91. Aphelandra tetragona ; 92—93.

Pseuderanthemum bicolor; 94. Ely-
traria acaulis lyrata; Mendoncia velloziana M. velloziana sparatteria (stellate
var. 95. var.
sparatteria; 96. var.

hair-base); 97. Ruttya speciosa;98. Lepidagathis cuspidata. —

Figs. 99—101. Portions of non-glandular hairs

magnified, to show wall ornamentation. Eranthemum 100. Ruttya speciosa; 101. Dyschoriste
99. purpurascens;

vagans. —
Scale 100
µm for Figs. 54—84, 86—98; scale 62 µm for Fig. 85; scale 50 µm for Figs. 99—101.
 K.J. Ahmad: Hairs of Acanthaceae 107

The unicellular hairs vary a great

deal in shape and size. They may be small

conical small and stout, slender

or papillose (Figs. 54—56, 59 —63, 65, 66, 68, 71),
or

needle-shaped (Figs. 70, 73, 74), or comparatively large, stout,

and erect (Figs. 72, 78, 79).
They are typically thick-walled lanceolate in Barleria cuspidata (Fig. 79), B. lupulina, and
B. prionitis. Generally, the hairs have a several-celled hair-base but in some cases they
arise from the middle of an epidermal cell (Figs. 54, 62, 64, 70 —71, 73 —77).
The multicellular hairs (Figs. 80 —98) are generally 2 —6-celled.
—95, 97

They be to 12-celled in Pseuderanthemum bicolor (Figs. 92, 93) and to 15-celled

may up up

in Elytraria acaulis var. lyrata (Fig. 94). As in the case of unicellular hairs, the shape of the

multicellular hairs varies greatly. They be short and conical (Figs. 80—81), long and
may

slender (Fig. 92), or long and stout (Figs. 82 —83, 86—89, 95)-
The non-glandular hairs of some species are
characteristic. In Aphelandra

tetragona (Fig. 91) the hair is built of two or more small basal cells surmounted by a much

elongated apical cell. The hairs of Ruttya speciosa (Fig. 97) are similar but the apical cell

here lies at right angles to the basal part and has a thicker wall than the basal cells. In

hairs
Adhatoda vasica (Fig. 90) the hairs are sometimes T-shaped. Lanceolate thick-walled
occur in several species of Barleria (Fig. 79). Thick-walled uniseriate hairs with bulgings
at the nodes are recorded in Mendoncia velloziana var. sparatteria (Fig. 95).
The hair-base is generally multicelled, with polygonal, isodiametric, straight-
walled cells. However, small and slender, or more rarely long and stout non-glandular
hairs often spring from the middle of an epidermal cell (Figs. 84, 98). Hygrophila
90,

serpyllum has both these types: 1 —2-celled, small, slender hairs emerging from a single

epidermal cell (Fig. 70) and 2 —4-celled, longer, and stouter hairs with a multicelled hair-

base (Fig. 88). In species with sinuous-walled epidermal cells, the hair-base cells are
less

sinuous or straight-walled. In Mendoncia coccinea and M. velloziana var. sparatteria, non-

glandular hairs of the epidermis have a typically stellate hair-base with 2

—several
upper

arms (Fig. 96).

The non-glandular hairs are moderately thick-walled. In species ofBarleria the thickness

of the wall is considerable and the hairs consequently have narrow

lumina. The wall

ornamentationgenerally consists of round, oval, elliptic, or

linear tubercles (Figs. 60—62,

99—101). Sometimes, the wall ornamentation can be used for diagnostic purposes.

DISCUSSION

The study of the foliar epidermal hairs of (109 species) of Acanthaceae reveals
39 genera

that various features of the epidermal hairs, like their shape, size, the number of cells

comprising them, thickening and ornamentation of the wall, type of hair-base, etc. play
useful role in the consideration of various The glandular hairs of
a
very systematic taxa.

the three types mentioned earlier occur

intermixed with the non-glandular ones. While

the glandular hairs are more important at the higher level (subfamily, tribe etc.), the non-

glandular hairs are of diagnostic value at lower levels such as genus, species, and variety.
Hobein (see Solereder, 1908) observes that the panduriform glands are characteristic
of the of the Acanthaceae.
Thunbergioideae, whilst the disc-shaped glands occur in the rest

The however, shows that the panduriform hairs

present investigation, glandular are

uniformly present in Nelsonioideae as well, and they are characteristic of these two sub-

families of the Acanthaceae. Kumar and Paliwal (1975), who studied the foliar trichomes of

six species of Thunbergia (subfamily Thunbergioideae)) and Elytraria acaulis and Nelsonia
hairs in Nelsonia
campestris (subfamily Nelsonioideae) have recorded long-stalked glandular
campestris. However, they have not reported the presence ofsubsessile panduriform glandu-
lar hairs in the species investigated by them.
 BLUMEA VOL. 24, No. 1, 1978
108

In the genera
of the subfamily Acanthoideae and Mendoncioideae (of Lindau, 1895)
the hairs have only globular disc-shaped head. This
presently investigated, glandular a or

with the observations of Solereder Mendoncia (sub-family

is in conformity (1908). In

Mendoncioideae), while the heads of glandular hairs are basically globular or disc-shaped,
,,

they frequently be modified and quadrangular or triangular in surface view.

may appear

While the diameter of the head of glandular hairs does not

vary appreciably, it can

sometimes be used for distinguishing one species ofa genus from the other, e.g. Thunbergia
S. scrobiculatus
erecta (42 /im) and T. kirkii (31 /mi); Strobilanthes scaber (22 /tm) and

(34 /im); Justicia diffusa (26 /mi) and J. procumbens (35 /«n). The number of cells comprising
the glandular head is quite important and can be used as a taxonomic character. In some

for All the investigated of the sub-

cases it is constant a sub-family or a
genus. genera

The nine
families Thunbergioideae and Nelsonioideae have only 2-celled glandular heads.

species of Strobilanthes presently studied uniformly show 4

—8-celled glandular heads.
heads. All the
Similarly, the seven species of.Barleria have 4—8 (mostly 4)-celled glandular
six species of Pseuderanthemum have
4
—8 (mostly 8)-celled glandular heads. On the other

hand, within easily be distinguished from the others by the

a genus, some species can

difference in the number of cells comprising their glandular heads. According to Santapau

dalzellii and D. minute and relate

(1951), the differences between Dyschoriste vagans are

that they
only to the size of the anther spurs. However, the present investigation shows
D. dalzellii has
can easily be distinguished from each other by their epidermal hairs.
subsessile glandular hairs with the head is 8-celled; D.
a 4-celled head, but in D. vagans

also has of hairs which lacking in D.

vagans a dense covering long-stalked glandular are

dalzellii.

The long-stalked glandular hairs have been recorded only in nine species of subfamily

Acanthoideae and Kumar and Paliwal

not in the other subfamilies. However, according to

in Nelsonia campestris (subfamily Nelsonioideae) also. While their

( T 975)> they are
present
taxonomic significance at higher level to be limited, as they are present in widely
appears
view
different taxa, they are nevertheless important from diagnostic point of jit species
level. For example, such hairs occur in Hygrophila serpyllum, Dyschoriste erecta, D. vagans,

and
Strobilanthes heyneanus, Dipteracanthus patulus, Lepidagathis cuspidata, Justicia tranque-

bariensis, but are absent in the other investigated species of the genera.

hairs role in the taxonomic consideration of

Non-glandular play a
very significant
thicken-
various taxa as they show considerable
range of variation in their shape, size, wall
the
ing, and hair-base structure. However, as pointed out earlier, unlike glandular hairs,
also useful in distin-
non-glandular hairs are more
useful at species level; rarely they are

guishing some genera from the others.

which unicellular
The species of Barleria have
very characteristic non-glandular hairs are

walls and canal-like lumen having

(rarely I- to several-celled) with thick a narrow a

spherical enlargement at the base of the hair (Fig. 79). According to Hobein (see Solereder,

of the tribe Barlerieae investigated by him.

1908) these hairs are
present in most
genera
various
Metcalfe and Chalk (1950) have also characterised various tribes of Acanthaceae by

types o
f
non-glandular hairs. However, the investigation shows that in Lepidaga-
present
The
this, which belongs to the same tribe, uniseriate hairs with unthickened walls occur.

same view is expressed by Solereder (1908).

hairs which to
The
genus Strobilanthes is quite uniform in its non-glandular are 1-

several-celled hair-base of cells forming rosette

several-celled, stout, with a polygonal a

around the foot or basal cell. Their distribution pattern is also quite characteristic as they

are generally confined to costal and marginal areas.

A number of species characterised their typical non-glandular hairs. These

are by
 K. J. Ahmad: Hairs Acanthaceae
of 109

include: conical unicellular hairs on the upper epidermis of Dyschoriste dalzellii (Fig. 59),
branched hairs of D. vagans (Fig. 85) and Pseuderanthemum bicolor (Fig. 93), conical unicel-
lular hairs of Lepidagathis trinervis (Fig. 72), and typical hairs of Adhatoda vasica (Fig. 90),

Aphelandra tetragona (Fig. 91), and Ruttya speciosa (Fig. 97). These have been described
under 'Observations'.
The hair-base is also The of Mendoncia
an important diagnostic character. two species
are characterised by typically stellate hair-bases on the epidermis (Fig. 97). Rizzini
upper

(1948) divided Mendoncia into four subgenera on the basis of the characters of the hair-

base. The two species investigated here have non-glandular hairs with a conspicuously
stellate hair-base on the upper epidermis; on the lower epidermis the hair-base cells,

though basically indistinguishable from the other cells, frequently form

epidermal
rosette or stellate pattern.

In many species, e.g. Hygrophila serpyllum (Fig. 70), Dipteracanthus patulus, Lepidagathis
cuspidata (Figs. 73—74), L. purpuricaulis (Fig. 75 —76), non-glandular hairs frequently
arise from the middle of an epidermal cell and this feature is characteristic for those
very

species.
The density of the non-glandular hairs often varies considerably within a
genus and

is not an absolutely reliable character for diagnostic purposes. However, the size of the

hair and the number of cells comprising it, are

important features and several species of a

genus can be distinguished from each other on the basis of these characters (Table I).

Non-glandular hairs often show wide of variation within but where

a
range a
species,
they are of a characteristic form they can serve as a means of distinction among the

species. Stace (1965, p. 52) remarks: 'The most usual division, between glandular and
non-glandular types (of trichomes), suffers from the same disadvantages as other systems

in that there is no sharp distinction between the two and in case

the division
groups, any

closely related types'. Dyschoriste illustrates strikingly the

separates obvioulsy very vagans

transition between glandular and non-glandular hairs. Except for the terminal gland,

there is little difference between the long-stalked glandular hairs and the non-glandular
hairs of this species. Where the hairs are branched, one branch be non-glandular and
may

the other terminated by a gland (Fig. 53).

The delimitation of Acanthaceae Hooker
by Nees (1847), Bentham & (1876), Lindau
(1895), Van Tieghem (1908) and Bremekamp (1953, 1955, 1965) has been explained
earlier (see 'Introduction'). The present study shows that Lindau's (1895) four subfamilies

namely: Nelsonioideae, Mendoncioideae, Thunbergioideae, and Acanthoideae, have broadly

similar non-glandular hairs. While Nelsonioideae and Thunbergioideae have glandular

hairs with panduriform head, those of the Acanthoideae and Mendoncioideae have globular
or disc-shaped heads. This character, however, does not justify Bremekamp's (1965)
transfer of Nelsonioideae to Scrophulariaceae as the glandular hairs in the latter are not

panduriform (Metcalfe & Chalk, 1950). In this respect Nelsonioideae is more close to

Thunbergioideae. Also, members of the Mendoncioideae have glandular and non-glandular

hairs basically similar to those of the Acanthoideae.

In the light of the present investigation, therefore, the transfer of Lindau's subfamily
Nelsonioideae Thun-
to Scrophulariaceae and raising of his subfamilies Mendoncioideae and
bergioideae to family rank as suggested by Bremekamp (1953, 1965) in his delimita-
1955,

tion of Acanthaceae is not justified. This view is also supported by evidence from the

stomata (Ahmad, 1974, 1974a; Paliwal, 1966, 1967; Kumar & Paliwal, 1975) and embry-

ology (Johri & Singh, 1951). The present study, therefore, lends support to the treatment

of Nelsonioideae, Mendoncioideae, Thunbergioideae, and Acanthoideae, either as subfamilies,

or as tribes under die family Acanthaceae.

 BLUMEA VOL. 24, No. 1, 1978
110

ACKNOWLEDGEMENTS

I owe my deep gratitude to Dr. R. V. Sitholey for his valuable guidance in this work and to
my colleague
Mr. D. B. Shukla for helping me
in the local collection and laboratory assistance. I wish also to
thank the

following individuals and for the bulk of the material for this study:
organisations supplying me required
Dr. A. N. Rao, University of Singapore; the Curator, Lembaga Biologi Nasional, Bogor (Indonesia); Dr.

K. C. Waterfalls Garden, Penang (Malaysia); the Botanic Gardens, Perade-

Cheang, Superintendent, Royal
of the Herbarium Museu
niya (Sri Lanka); Dr. Margaret Emmerich, Curator Nacional, Rio De Janeiro

(Brazil); and the Director, Botanical Survey of India, Calcutta, India.

I wish record grateful thanks Khoshoo, Director, National Botanic Gardens,

especially to my to Dr. T. N.

Lucknow, for the facilities provided to me and for taking keen interest in this work.

REFERENCES

K. studies in Bot. Linn. Soc.

AHMAD, J. 1974. Cuticular some Nelsonioideae (Acanthaceae). J. 68: 73—80

1974a. Cuticular studies in some species of Mendoncia and Thunbergia (Acanthaceae). Bot. J. Linn.

Soc.
69: 53 —63.

O. 1886. Untersuchungen iiber die systematische Bedeutung der Schildhaare. Flora 69:
BACHMANN, 387 —

400; 403 —415; 428—448.

BENDRE, A. M. 1973. Studies in the family Loganiaceae. I. Trichomes. Jour. Indian bot. Soc. 52: 225—234.

D. Genera Plantarum. Acanthaceae in

BENTHAM, G., & J. HOOKER, 1876. 1862—1883; 2: 1060—1122.

BREMEKAMP, C. E. B. 1953. The delimitation of the Acanthaceae. Proc. Koninkl. Nederl. Akad. Wet. Amst.

Ser. C, 56: 533—546-

1955. The Acanthaceae of the Malesian area.
I. General considerations. Proc. Koninkl. Nederl. Akad.

Wet. Amst. Ser. C, 58: 162—171.

1965. Delimitation and subdivision of the Acanthaceae. Bull. Bot. Surv. India 7 (1—4): 21—30.

CARLQUIST, S. 1961. Comparative Plant Anatomy. Holt, Rinehart & Winston.

COOPER, D. C. The development of the peltate hairs of canadensis. Amer. J. Bot.

1932. Shepherdia 19:

423 —428.

COWAN, J. M. The Rhododendron leaf: A of the Oxford Boyd,

1950. study epidermal appendages. &

London.

T. H. Botanica
GOODSPEED, 1954. The genus Nicotiana. The Chronica Co.,Waltham.
A. 1967. Studies the trichomes of and Proc. Indian
INAMDAR, J. on some Oleaceae, structure
ontogeny.

Acad. Sci. Sect. B, 66 (4): 164—167.

B. & H. The and of
JOHRI, M., SINGH, 1959. morphology, embryology systematic position Elytraria
acaulis (Linn, f.) Lindau. Botanisk. Notiser. 112: 227—251.

KUMAR, S., &G. S. PALIWAL, 1975. Foliar anatomy of the family Acanthaceae. Acta Bot. Indica 3:121—131.

G.
LINDAU, 1895. Acanthaceae. In A. Engler & K. Prantl, Die natiirlichen Pflanzenfamilien 4 (3B): 274—354.

Leipzig.
METCALFE, C. R., & L. CHALK, 1950. Anatomy of the Dicotyledons. Oxford, Clarendon Press.

NEES VON ESENBECK, C. G. 1847. Acanthaceae. In A. P. De Candolle, Prodromus Systematis Naturalis

Paris.
Regni Vegetabilis 11:
46—519.
PALIWAL, G. S. Structure and of stomata in Acanthaceae. 16 (4):
1966. ontogeny some Phytomorphology
527—532.

Stomatal ontogeny in the family Acanthaceae and the systematic position ofthe genus Elytraria.
1967.
Proc. Indian Sci. Congr. 54, Part III, Abst., Sect. VI. Botany: 306 —307.
N. 1972. Classification and phylogeny of the trichomes of In Research Trends in
RAMAYYA, angiosperms.
Plant Anatomy. Ed. Ghouse & Yunus, India.
Aligarh,
C. T. Circa Acanthacearum Brasiliensia. do Jardim
RIZZINI, 1948. Disquisito aliquot genera Arquivos
Botanico, Rio de Janeiro 8: 295 —372.

ROE, K. F. 1971. The terminology of hairs in the genus Solanum. Taxon 20: 501—508.

H. Bot. Memoirs
SANTAPAU, 1951. The Acanthaceae of Bombay. 2. Bombay Univ., Bombay.

SINGH, V., & D. K. JAIN, 1975. Trichomes in Acanthaceae. I. General structure. J. Indian bot. Soc. 54:

116—127.

H. & F. E.
SOLEREDER, 1908. Systematic anatomy of the dicotyledons. Transl. L. A. Boodle Fritsch; Revis.

D. H. Oxford.
Scott,
SPORNE, K. R. 1956. The phylogenetic classification of the angiosperms. Biol. Rev. 1
31: —29.

STACE, C. A. 1965. Cuticular studies as an aid to plant taxonomy.

Bull. Br. Mus. nat. Hist. 4 (1): —78.
1

VAN TIEGHEM, PH. 1908. Structure du pistil et de 1'ovule, du fruit et de la graine des Acanthacees. Dedouble-

ment de cette famille. Ann. Sci. nat. Ser. 9, Bot. 7: 1—24.

 K.J. Ahmad: Hairs of Acanthaceae 111

TABLE I. EPIDERMAL HAIRS OF ACANTHACEAE

of abbreviations. D:
Explanation average diameter of glandularhead; Glob: globular head of subsessile
—

glandular hair; L: length of long-stalked and non-glandularhairs (minimum, and maxi-

glandular average,
in that Low: trichomes lower Ls: Pand:
mum order); on epidermis; long-stalked glandular hairs; panduri-
form head of sub-sessile glandular hairs; Rhomb: rhomboid or quadrangular head of glandular hairs; Ss:

subsessile glandular hairs; St: stalk of the long-stalked glandular hairs; U: trichomes on epidermis.
upper

+ : sparse; + -f: common; + + +: dense.

The source of the material from which the voucher specimens are
taken is indicated in abbreviated form

and is within brackets. The following internationally accepted abbreviations have been used:
given BLAT,
Blatter Herbarium, St. Xavier's College, Bombay (India); BOG, Lembaga Biologi Nasional, Bogor

(Indonesia); BSI-North, Botanical of India, Northern Circle, Dehra Dun (India); BSI-South,
Survey
Botanical Survey of India, Southern Circle, Coimbatore (India); BSI-Wcst, Botanical Survey of India,
Western Circle, (India); CHAMOL, Central National
Poona Parsari Farm, Chamoli, U.P. (India); CNH,

Herbarium, Botanical Survey of India, Calcutta (India); DE, Dr. Anima De, Bose Research Institute,
Calcutta (India); DUN, Dehra U.P. Lucknow, U.P. National Botani-
Dun, (India); LUCK, (India); LWG,
cal Gardens, Lucknow (India); MALAY, Waterfalls Garden, Penang (Malaysia); POND, Medical College,
Pondicherry (India); PDA, Royal Botanical Gardens, Peradeniya (Ceylon); RIO, Herbarium Museu

Nacional, Rio De Janeiro (Brazil); SING, Department of Botany, University of Singapore, Singapore.
 BLUMEA VOL. No.
112 24, 1, 1978

2-
2-
fim) fim.)
427/4i2m7/1) 944944
743) 187/im) (rarely
only. 2-

59—190,m)
U.
U.
stelate. 115115 187 ii)

Low. 3-cel e3-cdle, I038jum1)03871m)

165— 4 73) 71m) &

59— fim) (L:

(L:
101io4,u/mi) I3 4yIu3m4yu)m 1 5 7 0 ^ m ) 1481648/7i2m) slender, 140,u1m407)1Low.;m
uniceluar margin,

1I—1—77—0032473(94u9p—1—-celd. 24—79-6celd.1047 1—6-celd;8529—0 16-celd. 8-01—9 3(1mo—stly1—3(42mostly 1—(mostly734—0 513(mostly842965—- 13(mosIt—ly3-4ce5ld.15—6237-c—eld.16— I2—0S5 127—0— 62534—-celd. 72—60-celd. 16— 2-791—30 2— 57284—
fim) 1570
944 up er, rare) 35,1 1)
biseriate. (L:
335 +,
79—
(L:
(L: on (L:
(very (L: (L:
(L:
4-4-4-, 71m) +
short, 4-cel e4-cdl ,
i)-celed. i)-c led. i)-celed. i)-celed. i)-celed. i)-celed. 45—15 —
short,
to
to +

often 825825
7)-celed. hair-base fim) ,

confined
ii)
(L:
+,

12981/2im9871)m
+,
+
+
+ (L:
(L: (L: (L: (L: (L:
(L:
haihairsrs
15-cel d, jum) 882626,um) 137/z1m371); 6-celed.
+
+

to
(L:
+
+ U.
ststout,
3(mostly
i) i)
i)

3-celd.
&
&

1—3-(mostly 1—3-cel d. 2—6-cel d. types; conical,

519
2(up types: Low. long,
47—3 0—
(L:
(L:

Non-glandular
to

up

59—204—
,

+,
two only. ++, + ttwo
wo
+, +, +,
+
+ + +
+, +,
+
(L:
+,
+ +
+,
+,
+,
+
fim)
+,
+ +,
+,
+ +,
+,
4-4-,
+ Of cel ed), (L: U.
U. +
+
+ +,
+ Of celled, 4-4-4-, (L: +

26y267u1m)m)
/jm)
fim)
32,um) 32,um) 34/,<umm)) 5-cel ed. 5-celed. 38,1401)fim)40/«n) 42,um) 38/3871jm) 35/3571im) 3i3171m)/^m) 387,u1m) 36/36j7m1m) )4)-celed. 8)-celed. 4)-celed. 8)-celed.
39 32
3—5-cel d. 3—5-cel d. 40 (D.

2— 4— 4— 4— 4— 1—25—102—
1—3-3c-ceeldl. d2. 40f2t4m071)m
3— 3—
(D. (D. (D. (D. (D. (D. (D. (D. (D. (D. (D. (D.

8(mos8t(lmyostl 8(mostly 8(mos8(tmloyst 8(mostly 4—8-cel8-celdd..

4—8(mostly
hairs
2-cel d. 2-celcd. 2-celed. 2-cel d. Rhomb., Rhomb., 2-cel d. 2-cel d. 2-cel d. 2-cel d. 2-cel d. 2-celed. 2-cel ed. 2-cel ed. 4—8(mostly St.
St.

Ls.,
fim) 71m)
fim) fim)
Pand., Pand., Pand., Pand., Glob., 33/«imn)) Glob., 30,um) Pand., Pand., Pand., Pand., Pand., Pand., Pand., PaPand.nd., Glob., 27 1m) Glob., Glob., 2171m) Glob., 27f27 1im) Glob.,
Glandular
+,

Ss., Ss., Ss., Ss., Ss., (D. Ss., (D. Ss., Ss.,
Ss., Ss., Ss.,
Ss., Ss., Ss., Ss., Ss.,
Ss., Ss., (D. Ss., (D. Ss., (D.
27 25 21

Ss., (D. Ss.,

Ss.,
4-4-4-,
+

+ (L:

SING) LUCK) SING) LUCK) LUCK) LUCK) LUCK) LUCK) LUCK) LUCK) MALAY) MALAY) LUCK) LUCK) BSI-
RIO) RIO) BSI-
specimen (ex (ex (ex
(ex (ex (ex
(ex
(ex
(ex (ex (ex (ex (ex (ex (ex (ex (ex (ex
(ex
(ex

64561 64614 64546 64593

114.908 38.323
64606 64607 64609 64589 64595 64590 64563 64562 64615
64615 64617 BSI- outh) BSI- outh) 4427572757 76626
Voucher LWG LWG LWG LWG R
R R
R
LWG LWG LWG LWG LWG LWG LWG LWG LWG LWG (ex (ex BSD North) BSI
BSI West)

Ktze.
Bremek. Anders.
Nees

(Schumach.)
O. var.

(Nees) Lindau (Nees) Nees Moore

(Buch.-Ham.)
Br. T.
T.
Nees
R. Veil. Nees Sims Anders. Hook./ Anders.
(Roxb.)
i I

(Mart.)
S.
(L./.) lyrratata
Anders. Roxb. Roxb.
Hook./ nobilis auriculat
s

longifolia ex
Lindl. (Vahl)
campestris co cinea (Mart.) affinis
T.
T. T.
T.

acaulis var. Bojecr

velozian
T.

grandiflorakirkii laurifolia mysorensi polysperma quadrival is salicifoUlia serpylum

StaurogyneElytraria acaulis E.
Nelsonia Mendo ncia M.
spar tleria Thunbergia alata erecta T.fragrans
T. T. T. T.
T. T. T. T.
Sanchezia Hygrophila Heine H. H. H.
H. H.
H.

SpSpecies I.
1. 2. 3- 4- 5-
4- 5- 6. 7- 8. 9-
9.
10. II.
II. 12. 13- 14- 15.
15- 16. 17-
17. 18. 19-
iy. 20.
 K. J. Ahmad: Hairs of Acanthaceae 113

pm)
fim)
only. uniceluar, 490490
637,um) ftm) 354/3i5m4/ )i 46o/im) 1 91/jm)
/im) fim)
yum)
fim) 1291829/im57/3) S7mftmi)) 5 0,um5O/t)
295 1191
rarely 460
400400 3

Low. 168—168—
2i2/2m1 /z)m /im) 80 ^m) yum)
fim)
6oi,un)
short, fim)

42— 1056-—celd. 682154——53 1—4-29celd. 2— 2941— 2n—o5-celd.21—96745-celd. 2—50(o4ften28328—3—19148 2094— 5—613952— 7 62135— 2907— I—2175—340 1—4295— 2—1 2—1 205—147
4-cel4-ced, ++,
800
10010 4)-cele4)-dc,led fim)/im) 205205
601

(L: (L:

849849
(L: (L: (L:
(L: (L: (L:
4)-cele4)d-c.led 4)-celed.
ii)

ij-celed.
(L: (L:
4)-celed.
sevral-ce d. sevral-ce d. sevral-ce d. sevral-ce d. sevral-ce d. sevral-ce d. sevral-ce d. sevral-ce d. sevral-ce d. 8(mos8t(lmyostl
/xm)
684y«m) in1if2m1/x)m) 262/xm)
fim)
hairs
(L: ++■
++,
9209y2Oum); (L: (L:
9o(mosctly (L:
(L: 2

262

8(mos8(mtolsy
i)

only.
4-celd. 5-celed.
24—95 684I— 153—65I8——(mos8(mtolsy 147—205—:
to

types: (L: to
1-to

Non-gladu r
up to to to to to
U. to

1- 1- +,!- 1- I-
1-
to
I-
1- 1- to

branched.
+,
two ++,
conical,
1- 1-
+, +, + +, +, +, +, + +, +, +, +, +, +,
, ,
,

+ Of (L: +
+ + + + + + + +
+ + + + + + + + (L: + (L: +, (L:
(L:

Glob., fi/xm);
m);Low. Glob., um);
2.6fim); io62^m)25/im) /xm) /xm) /xm)
24/jm) 25,um) 27^) 26^/x0m1) 25,2S/xumm)) 4—8-
8- fim) fim) fim)
/xm) /xm)
2S/xm)2 /xm) 34,34juxm)m)27,um) than
/xm)
32/im)
27,/uxm)
8)-8)- i
24 25 27 27 25 22 27 32

5-celed, Ls.,
2—5-cel d,
23 26
fim)Ss.,

42— i— 177——101606—— 2— 3834841——06106 4— 4— 4— 4— 4— 4— 3—4-celd. 4—4— 4— 4—4—4— 4— 4—

4— (D. (D. (D. (D. (D. (D. ßm) (D. (D. (D. (D. (D.
Ss.,
Ss.,
(D.
(D. (D. Glob., +, 7}Oftm) more (D.
(D.
(D.
+, 2-cele2-cdile, 117
+
,

750
8(mostly /xm) 4-4-, U. 8-celed8-c.eld 8-cele8-cdl. 8-celd. 8-celd. 8-celd. 8-cel d8-celd. 8-celd. 4—8-cel8-celdd.. 8-cel d8-c.eld 4—8-cel8-celdd.. 8(ofte8(onfte fim)26ßm) 4—8-cel8-celdd..
ii)

4—8-cel d. 4—8-cel8-celdd..
U.

2—8(mostly 19/im)4-celed. 4—8-cel d. 4—8-cel d. 4—8-cel d. Ss.,

4—8-cel d.
+ +
& St. (L:
i) i)
&
i) fim); 26
hairss Ls., (L:
types:Low.Low. types:Low.
19 St. 27
(D.
U. types: (D.
(D. +,
&U.
Glob., Glob., 22,/uxm) Glob., Glob., Glob., Glob., Glob., two
Ls.,
Glob., Glob., (L: &
Glob., Glob., Glob., Glob., Glob., Glob., Glob.,
Glandular 8)-celed.
+
two two
Ss., cel ed. Ss., Of
4-celed, +,

ii)
only. Of 8-celed, +

ii)
Low. Ss., Ss., (D. Ss., Ss., Ss., Ss., Ss., Of cel ed. St. Ss., Ss., Ss., Ss., Ss., Ss., Ss.,

LUCK) SING) MALAY) LUCK) BSI-West) BSI-West) BOG) LUCK) L L

BLAT) BSI-West) UCK) BLAT) DE) UCK)
BSI- BSI- BSI-West) DE)

specimen
BSl3857(exI-Wst)
(ex (ex (ex (ex (ex (ex (ex (ex (ex (ex
(ex (ex (ex

CHAMOL) 79 5
(ex (ex (ex
(ex (ex
(ex
64613 64548 64549 64583
64583 64588 64586 490 64567 354 64559 64580
64580 64558
54

66489 66604 3857 9685 74982

74982 3

Voucher LWG BSI6 489 West)

BSI BSI
West) BSI LWG LWG LWG (ex BSI
BSI BSI LWG LWG BLBLATAT BSIBSI LWG BLAT LWG LWG LWG

Nees
Ktze.
Anders. ex

Nees
O.
T.
T.
Wall,
BBeenth. Clarke DalDalz. Andr. Griseb.
z.
Gre nman
altermnat a Hall./. Anders. alatus NeesNees Blume Nees
barleiodes dalzeli Ktze.
KtKtzze.e.
Nees
Nees formosa
lorentzian mal cosperma
O. (L.)

i x o c e p h a l u s p u l n e y e
n s i s c r o b i c
scabberer Rueliaform sau l a t s
O
O T.
T.
ata
h e y n e a n u s
s

Petalidum Dyschoriste ererecta

D. D.
vagans Hemigraphis
„._,.
colorr
H.
/ihii'rtafa
H.
H.
Strobilanthes calosus crispus
S.
b a r
S. S. S.
b
S. S.
S.
a t u s
S.
5. S. S.
S. S. S.
S.
Ruel ia i?.
R. R.
R.

Species 21.
21. 22. 23.
23- 24. 25._., 26.
25- 27. 28. 29.
27- 20. 30. 31-
31. 32. 33. 34-
33- 37.
34. 35- 36. 37- 38.
38. 39.
39-
114 BLUMEA VOL. 24, No. 1, 1978

/im) /im)
743743 731
731/im)
/im)
260/2i6m0/i ) 10 3/im) /im)
170/im) 2—32355//1im1)) 162/16i2m/1 ) 195/im)127/cm) 40 /40im/t )450/4i5m0/i )
/im) /im)
735/7M35/nim)

4 1 0 — 8 2 4 7 — 3
1003 195
170 127
512/5i1m2/)01 175/1t7m5/1)

I 8
—2 1 6 —
4 421 0
— 4 - c e l d . 7 0
2 — 1 4 - c e l d . I — 4 -2c e l d . 1
8 —
5743-
0 c e l d . 1 5
4 -7c —
e l d . 1 05
2—7 4 - c e l d . 1 —6
54 -
7 c e l d . 250—
4)-cele4)-dc.led 4)-cele4)-dc.led 4)-cele4)-dc.led
(L:

72 — 1 7
(L:

2—1— I—2 1— 2I—1(—L:i4I8—16— 2 — 1 4 58 — 1 9 —

(L:

sevral-ce d. thick-waled.
/im) /im)
650/im) 2 5/im)
(L: (L: (L: (L: (L: (L: (L:
(L:
hairs
8(mos8t(lmyostl
425
37—650/Hn)
8(most8l(myostly 8(mos8t(lmyostl
225
4-cele4-cdl .
I—4-cel d. 1—4-cel de.
(L: (L:

i-celed.
(L:

137—245— 11442—23—7— il-celed. i-celed. i-cel d.

Non-glandular
to

+.
+, +,
1-
1- i-celed,
+, +
+ +,
+, +, +, +, +,
+, +,
+, +,
+, +, +, + +, +,
+ (L: +, (L: +,
(L: +
,

+
+ +
+ +
+
++,
+ + 4- + + +
+ + + 4-+,
+ + +
+
+

+
+ +,

8)-
/im) /im)
24/im)
+,
27/im) +,
++, 27/rim)
26/tim) 4)-cel ed.
than
23
23 Ls. 24
+
+

2 0/tim) /im) /im) /im)

25/im) 26/im) 34/tim) 26/tm)25/rim) 24/im) /im) +
/rm)
/im)
4)-celed. 4)-celed.

4— 4— 4— 4— 4—8-celd.2— 4—
ii) ii)
ii)
momorre Glob., (D.

more-cled.
24
(D.
Glob., (D.

4—8-celd.2— 2— 4— 4— 4—
+, 25 26 26 23
(D.
/im); (L: (D. (D. (D. (D. (D. (D. /im); (L: (D.

8-cled. 8-celd; 8-celd. 8(ofte8(noften 27/im); 8-celd. Ss., 22/im); 4-cel4-ce d. 8-(mo8s-(mtloys 8(mos8(mtolsy 4—8(mo8s(mtloystly
(D.

3-cele3-cdle. 4—8-cel d.
Ss.,

4—8-cel d. 4—8-cel d; 4—8-cel d. 27 22

hairs
34/im)
i)

(D.
or
4-celed. 4-celed. 4-celed. 4-celed. 4-celed. 4-celed. i)
i)

(D. 5-cele5-cdle. 3 0/tim) 4-celed. 4-celed

types:
8-
types:pes;
(D.
(D.
Glob., Glob., 30/tim) Glob., Glob., Glob., Glob., Glob., Glob., two
Glandular
Glob., Glob., 26/tim) Glob., Glob., two
Ss., Ss., (D. Ss., Ss., Ss., cel ed. Of
4—8-cel d. St.

Ss., (D. Ss.,

Ls., Ss., Ss., Ss., Ss., Ss., Ss., Ss., Of
Ss.,
4—8-cel d. 27—135— Glob., Glob., Glob., Glob., 35/35ytUmm)) Glob., 26/tim) Glob., 34i34/"tm)
St.

Ls., Ss., Ss., Ss., Ss., (D. Ss., (D. Ss., (D.

SING) LUCK) LUCK) LUCK) POND) CNH)

BSI-
LUCK) LUCK) SING) BSI-West) DUN) BSI-West) BSI-Dun) BSI-West) BSI-West) LUCK)
BSI-
specimen (ex (ex (ex (ex
(ex (ex (ex
(ex
(ex
(ex (ex (ex
(ex
(ex
(ex
(ex
(ex (ex
(ex
(ex

64560 64596 64604 64605 64602 64603

64603 13801 64608 64610 POND) 64547 72973 64612
801
79530 40247 71408 860 0
64598
64598
POND) 12502
LWG BSI72973 LWG
13

Voucher LWG LWG LWG LWG LWG LWG (ex MH South) LWG LWG (ex BSI BSI BSD BSI BSI LWG MH South)

Nees
Mart.
(Jacq.) Nees
Nees
Wil d. Nees
Br.
Br.
Nees Br.
Br. Nees

(Poir.) albomarginat
R.
R. ex
ex ex
Nees ex

Griseb. patulus
ex
Nees R. Nees
cristlata Wall, Don. Heyne
Linn. (Vahl) Wall,
Linn. Hemsl. Linn. (Vahl) Stapf. D. courtalikca Linn.

repens rosea tuberosa twe diana Dipteracnthus prostrausEranthemu capense nervosum pur ascen roseum watti Lepidag this cuspidat pur icaulis
incurva trinervis cristlata cuspidat
R.
R.
R. R.
R. R.
R. D. E.
E. E. E. E.
E. E. E. E.
E. L.
L. L.
L. L. L.
Barleria B.
B. B.

Species 40 41. 42. 43. 44. 45. 46. 47-

47. 48. 49. 50. 51. 52. 53.
48. 55- 56.
54- 55.
54. 56. 57. 58-
58. 59.
jo.
 K.J. Ahmad: Hairs of Acanthaceae
115

/tin)
ftm) /im)
32m) /im)
731731 625625 590
59032m) 2-celed). 4)-4)-

ftm)
32m) /im)
7—575/t/mim)) /im)
(rarely 531/cm)
125/M125n32)m up er. 25025032m)

365—9016—89—17 45381— 6218—0 3—1—52—142I7——34-2celd.1—33—-celd1. 2316— 267—1-8celd. 2— 9387— 2— 84112—— 2—1—294— 2—1— 2—13—01—85— 13624—-celd.
(mostly 150/imt n) 531
thick-waled. thick-waled.
401401

77
42)—-c4e)-lceeld.d. 4)-cele4)-dc.led
(L: (L:
8822——3366591— 4)-cel4)-ced. 4)-cele4)d-c.led (L:
on
uniceluar /on)
/im)
4)-cel4)-ecdl .
-1203
1203
9)-celed up er,
sevral-ce d, 14161/4i6m321)thick-waled. thick-waled. sevral-ce d, sevral-ce d. 436/im4362m)(mostly 550707/rm) 7(mos7t(lmostly 483/4i8m32m)
(L:
(L:
fim)(L:
ftm) (L:
/im) (L:
(L:
fim) (L:
(L:
hairs 287/28a73n2m) 15 32m) conical, 50/5i0m32m) (L:

I—41-c—el4-cde.l d8.(mo8s(mtolsy 1—66((mmosotlsyt7l0y—730—3778—3^1) in—HI15—/im) 1—6-cel d.

to
6606 0 778
7(mos7(mtloys
to

26—12—
77

confined 94—2 4— 47—2 4— 1 8—3 0 2—3-cel d.

3(up3(up

Non-glandulra
to
to to
to
to

i-
1- i-celed, i-celed, 1- 1-
1-
short, (L: ,

+■ +,
75— +,
2 +

+,
+
10

(L: +, +,
+,
+
+ (L:
+, +,
+
+ +
+ (L: +
(L: +
+, +,
+ (L: +
+
+, +,
+
+
+,
+
+
(L:75—
(L: +
+
+,
+
+ (L:
(L:
+,
+,
+
+
cel ed. +
+,
(L:
+
+
+
+ (L:
(L:
+
+

+ (L: +,
+,
+

/im)
2732m)
4)-celed. 4)-celed. 4)-celed. 4)-celed. /im)
3i^m) 25/2sμimm)) 24/24/41im1) 32/im) (D.
27
8)-celed. 8)-celed. 4)-celed. 4)-celed. 4)-celed. 4)-celed. 4)-celIed. 4)-celed. 2932m) 8)-celed.
38/383i2m) /im)

4— 4— 4— 4— 2— 4— 2— 4— 4— 4— 4— 4—
31 29

(D. (D. (D. (D. (D. (D.

8(mo8s(mtolsy 84—(m8(omsotsltyly 8(mos8t(lmyostl 84—(m8(omsotsltyly 4-cel4-ced. 4—8(mostIyl 6(mos6t(lmyostl 4(mos4t(lmyostl 84—(m8(omsotsltyly 8(mos8(tmloyst 84(—m8o(msotlsytly 8(mos8t(lmyostl 8(mostly
hairs
2-celed. 4-celed. 2-celcd. 2-celed. 4—8(mostly 8-celed. 4-celed. 4—8(mostly
/im). /im) [im) /im)
Glob., 27//im) Glob., Glob., 35/itn) Glob., 3i/«n) Glob., Glob., Glob., Glob., Glob., Glob., 30/tim) Glob., 2732m) Glob., 24/243i2m) Glob., 23/tm) Glob., 31/tm) Glob., 33/3 3i2m) Glob., 25/im) Glob., 243/L2im). Glob., Glob., Glob., 29/tim)
/im) ftm)
Glandular 32

Ss., (D. Ss., (D. Ss., (D. Ss., (D. Ss., Ss.,
31

Ss., Ss., Ss., Ss., (D. Ss., (D. Ss., (D.

(D. Ss., (D.
27

(D. Ss., (D. Ss., (D. Ss., (D. Ss., (D.

23

Ss., Ss., Ss., (D.

(D. Ss.,
31 25

BSI-West) LUCK) LUCK) LUCK) LUCK) LUCK) DE)

LUCK) LUCK) POND) LUCK) BSI-West) BSI-West) MALAY) LUCK) POND)
BSI- BSI- BSI-

LWG64n(exLUCK)
BSI-
specimen i937(exBSI-
x

(ex (ex (ex (ex (ex (® (ex

(ex (ex (ex
(ex (ex (ex (ex (ex
(ex (ex
(ex (ex
(ex
(ex (ex (ex

64601 64599 64600 64591 64611 64544 64570

64544 64570 64572 64576 64569 64542
64542
LUCK) 64571 64564
17

74776 97409 12735 20313

BSI76416
193

Voucher LWG LWG LWG LWG LWG LWG LWG LWG LWG LWG BSI BSI MH South) MH
MH South) South) MHMH South) LWG (ex LWG LWG
Nees
Roth Nees
(LJ.inn.) ex Bencth.
terminalis (Linn.)
Anders. Anders. var.
Anders.
Nees Nees Wall,
febrifugum
Nees
Grif . Nees
atropu reum
maderspatensi montaus infudbilformis Oliver
T.
T. T.
T.
var. T.
T.
Santapau
teragona echiodes (Burm.) (Dalz.) Clarke pictum
coc inea
Anders. Lindl. Linn. Wil d. chelon ides (Linn.) Blume
Clarke
GraptohylumPachystachys Pseudranthemu (Bull.)
Bailey
Aphelandra Androgaphis pankiculat a Gymnostachyum -angularis dalzelian gangetikca
T.
T.

lupulina prionit s strigosa nilotica latifolium quadrngularis intrusa

lawii
B.
B. B.
B. B.
B. B.
(Nees) Blkepharis Acanthus Cros andra C. A.
A. G.
Asy tasia quadt A. A.
A. A.
ies '

Species
Spec 60. 61. 62. 63. 64. 65. 66. 67. 68. 69. 70. 71.
71- 72. 73.
73- 74- 75- 76. 77.
77- 78. 79-
79.
 BLUMEA VOL. 24, No. 1, 1978
116

rarely apical
cells, hairs hairs
74m)
ftm)
slender, pm)
/urn) 10731073 332 235yum) 35735/7im41 ) ftm)
basal
thick-waled uupperp er uupperp er fim)37 4m) /urn)
2807( 1)
59—174321 ) 19—23574m) 26026074m)
82674m)

2 — 1 i o ( m s t l y
377

3 — 7 - c es l
v r a d , 1 —
i2)-cel d, 354jU35m47)mi)-celed. i)-celed.

28—1 8— 627— 67— 185432-ce—ld. 17-celd. 42— 1 -3c2e9—ld.130— 75— 123I4-—ce2l—d.1— 57389— 2—17j—1—5047m) 250—I—2 421— 65239—15-celd.18—52—
2 — 1
280 small 826
conical.
59—174— 3)-cele3)d-c.led several elongated 8o2yu80m274m)
4)-cele4)-dc.led 4)-cel 4)-cdel; 74m)4 2/41 1)4)-cele4)-dc,led
to
to
(mostly 20—42 i)-celed,
hairs
sev ral(up sevral(mostlyI30yum130)74m) several i s1fi5m74)m)
(L:
(L: (L: (L: (L:
(L:
(L:
(L:

5(mos(mtolsy
ftm)
354pm)
354
an
475yum)
(L:

io(mosi(mtloys io(mostly io(mosit(lmyostconical. 5(mo5s(mtolsy

(L:
(L:
I42;um1)4274m)
1—3-cel d. 1—4-cel d. 1—5-cel d.
by
(L:

Non-glandulra
to
to
to

37—64— 35J—15 — 75—150— (L:

surmounted
to
to
1-
1-

branched.
1-
27— (L:

typical y
I-
1-

++, (L: +, conical.

,

+, +,
+, +, +, +, +, +■
+, +, +, +,
+

+, +
+ (L: + +, + + + + 4-+, +, +
+ (L:
(L: +
cell. + + +
+ +, +
+, (L:
(L:
,

+ (L: +
1

fim
1

333/74/m) 65
6574m) 27/27 4im) 27/27 4im)
8)-celed. 8)-celed. 8)-celed. 8)-ceUl ed. 8)-celed. 74m) ftm)
74m)
35/im) 287,u4m) ++, 335/57i4m)74m)
25/Mm) 4)-celed. 4)-celed. 4)-celed. 32/327/4m) 3i31,7u4m)m)

4— 4— 4— 4— 4— 2— 5142— 4— 4— 4— 4— 4—
25 (D.
(D. (D. (D.
Glob.,
5
25

more-cled.
3

ii)
(D. (D.
(D. (D. (D. (D. (D. (D.

8(mo8(smotly 8(mos8t(lmyostl 8(mostly

4—8(m8(omsosttllyy 4—8(m8(omsosttllyy 4—8(mo8s(mtolystly 4—8(mo8s(mtolystly 4—8(mo8s(mtolystly 4-celIe4-cdle. 8-celd. 8-celd.
4-celIed. 4-celIed. 4-celIed. 2674m); 4—8-cel d. 4—8-cel d. 8-cel ed. 4-celed. 4—8(mostly
Ss.,
Ss., (L:

hairs
i)
or 4-celed. 4-celed.
typpes:es: 2-cel ed.
8-
8-
(D.

Glob., 74m)
28,um) Glob., 327/<4m)m) Glob., 28/2874im) Glob., 32/3274im) Glob., 27,u4m) Glob., Glob., Glob., Glob., two Glob., Glob., Glob., Glob., Glob., i3O0f7im4m) ) Glob., 3i31/7*4m) Glob., 26,267u4m)m) Glob., 29,714m) Glob., Glob.,
Glandular 28

Ss., (D. Ss., (D. Ss.,

Ss., (D. Ss., (D. Ss., (D. Ss., Ss., Ss.,
Ss., Ss.,
Ss., Of
St.
4-celed. Ls.,
Ls., Ss., Ss., Ss., Ss., Ss., (D. Ss., (D. Ss., (D. Ss., (D.
(D. Ss., Ss.,

DE) SING) MALAY) LUCK) SING) LUCK) MALAY)LUCK) DUN) DE) DE) CNH) PDA) CNH) LUCK) LUCK) LUCK) LUCK) PDA) LUCK)
specimen (ex (ex (ex (ex (ex (ex (ex (ex (ex
(ex (ex (ex (ex (ex (ex (ex (ex (ex
(ex (ex (ex (ex (ex

64553 64554 64555 64585 64556 64584 64557

64557 64565 64577
64577 64551 64552 64594 64578 64579 64592 64575 64574 64587 64566
LUCK)
Voucher LWG LWG (ex
LWG LWG LWG LWG LWG LWG LWG LWG LWG LWG LWG LWG LWG LWG LWG LWG LWG

Kurz. Andre
Ktze. Alston Engl. var. var.
Nees ex

Radlk. Radlk. Nees Nees O. (Linn.)

Domin (Retz.) Nees
Ktze. (Forsk.) nasutaa
(Höochst.) Linden Coemans Coemans LiLinnn.n. Hort.
(Schrank) Bailey
Linden Br.)
Br.)
(R.
acuminat Nees roxburghian nitdum Regel

"
pectinat
O.
viride gigantea
grandiflormu bicalyculat a linctoria
speciosa
verschafjelti argyoneura verschafelti Regelbetonica calycotricha
1

bbiicohlor kewense mal c em variabile Peristrophe Odont ema strictum Ecbolium Rhinac thus
RungiaDiclpitera Ruttya Fit onia pearcei Justicia
'

P.
P. P. P. P. P. P.
P. P.
P. O.
O. F. F.
F.
J.

Species 80. 81. 82. 83.

83. 84. 85. 86. 87.
87. 88. 89. 90. 91. 92. 93. 94. 95. 96. 97-
97. 98. 99.
98.
 Ahmad: Hairs of Acanthaceae
K.J. 117

/im)
/im)
/im) conical.
362/(in) /im) 200620 6
/im)
483/im)5 4/im5 4)/im)20 /2im0 /i)m
362
342/ m) 287287
287/im287)/im 300300 152/1im52//im)
m)i 250250
i-celed),

I—57189 — 247—1582-c3e90l6d—. 1 45— 16— I— 9412—— 1432—6-7celd.195—70 I42-celd. 145-9c2e—73ld. 51—62 —4-2celd.105—
342

35—239—
(L: (L: (L: (mostly 52—
(L: (L:
(L: (L: (L: (L:
hairs fun)
(L:
celled 290/(1 ) (L:

1—5-5cel-cd.el d. 5-cele5-cdle. 5-celed.

1—5-cel d. 55
290
I—4-4c-ceelledd..I—4-4c-ceelledd..
Non-gladuolar +
+,
+,

+ +,
+,
+

+
+,
+

+
+
+,

+ (L:
+, +.
+ +
+1 +,
+ + +,
+,
+

+ + +
4-+, +,

/im) /im) /im)

Uμ™.34/im) /im) Ls. 29/im)
29
/im) 31
/im)
27/im) 30/im)
27
26/im) (D.
5 +,
(D.
25/4m) 27/im) (D. (D. (D.
26/im)

2— 2—
29

(D. (D.
3

Glob., ii)
25

(D. (D. (D. (D.

4—8-cel d. 4-celed. 26/imn); 24—-c4-ecleledd.. 4-celed. 4-celed. 4—8-cel d. 4-celed. 4-cele4-cdle. 4—8-cel d. 4-celed.
Ss.,

hairs 4-celed. i)

types: (D. /im)

Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glob.,
Glanodular Ss., Ss., Ss., Of
two
4-celed. 123

(L: Ss., Ss., Ss., Ss., Ss., Ss., Ss., Ss.,

LUCK) LUCK) DE) LUCK) LUCK) MALAY)LUCK) MALAY)

BSI- BSI- BSI- BSI-
specimeno (ex (ex (ex (ex (ex (ex (ex (ex

MHi739(exBSI-
(ex (ex (ex (ex
50
64597 17319 12639
16 2 64568 64545 64573 64581 64543 64582
12639 64582 64550
645
20431
Voucher MH South) LWG MH South) MH South) MH South) LWG LWG LWG LWG LWG LWG LWG

Hook./
&
&

Benoth.
Hook./.Nees
Linn.o. &
&
Nichols.
Burm./ Linn.o. Nees
amherstiae Branodeg e Liondley
Wil d. vasica candicans Benoth. coarnea

dif usa gendarus a procumbens tranquebarins Roth

vahli
nodosa
Adhatoda Dianthera Beloperone gut ata Jacobina
oblongat
J. J. J. J. D. B. B.
J.

Species io100.o. I101.

OI. 102.
102. 103. 104 105. 106. 107. 108. 109. no.110. in.