BLUMEA 24 (1978) 101-117

Epidermalhairs of

Khwaja+J. Ahmad

Plant Anatomy Laboratory, National Botanic Gardens, Lucknow-226001, India.

Summary

two belonging to Structure and distribution of the foliar epidermal hairs of 109 species and varieties 39 hairs have Acanthaceae have been studied. Both and non-glandularepidermal genera ofthe family glandular hairs The sub- been recorded in the investigated taxa. The glandular may be subsessile or long-stalked. Glandular head of Glandular head 2-celled, and ii) sessile glandular hairs are two types: i) panduriform, —8- more-celled. Subfamilies and are character- globular or disc-shaped, 2 or hairs i«*H hv thfi nanduriform hairs, while Mendoncioideae and have glandular with a globular also only in nine species. Non-glandular hairs are widely head. Long-stalked glandular hairs are present

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

the hairs are of at species uniseriate; rarely they are branched. Though non-glandular diagnosticimportance The like , and Äphelandra, are quite characteristic. present level only, in some genera ,, they delimitation of the family Acanthaceae, involving the transfer study does not support Bremekamp's (1965) and the of his subfamilies Thunber- Nelsonioideae to raising of Lindau's (1895) subfamily Scrophulariaceae, rank of families. the retention of Nelsonioideae, gioideae and Mendoncioideae to the independent Instead, Acanthaceae is favoured. Thunbergioideae, Mendoncioideae, and Acanthoideae within the family

Introduction

The taxonomic and phylogenetic significance of trichomes has long been recognised

number ofworkers Solereder, 2 Cowan, by a (Bachmann, 1886; 1908; Cooper, 193 ! 195°! According to Carlquist Metcalfe & Chalk, 1950; Goodspeed, 1954; and Sporne, 1956).

their accessibility, the most important (1961), trichomes are, because of easy perhaps The of the tricho- anatomical features which could be used for taxonomic purposes. study and of by (1954) showed that mes ofRhododendron by Cowan (1950) Nicotiana Goodspeed

excellent characters for distinctions and levels. they are making out at subgeneric generic of the most The epidermal hairs of Solanum, according to Roe (1971), provide some of the features for as hair forms are characteristic important diagnostic purposes many studies by Inamdar in for a species or a section. Recent in Jasminum (1967), have Compositae by Ramayya (1972), and in Loganiaceae by Bendre (1973) similarly of these demonstrated the significance of trichomes in the systematic studies taxa of

flowering .

about of with and over The Acanthaceae are a large family flowering plants 250 genera „. Though the family has, for been treated 2500 species spread over the tropics. many years, and has been the of contro- as a distinct taxon, its delimitation subdivision subject great

and and views have been expressed regarding the systematic versy divergent conflicting into foursubfamilies: Nelsonioi- position of various taxa. Lindau (1895) divided Acanthaceae

deae, Mendoncioideae, Thunbergioideae, and Acanthoideae on the basis of types of fruits, the absence of Nees number of and the or retinacula and their shape. (1847) ovules, presence Acanthaceae: Anechmatacantheae (without retinacula) had earlier recognised two groups in 102 BLUMEA VOL. 24, No. I, 1978

and Echmatacantheae (with retinacula). While Anechmatacantheaehas two tribes, Thunbergi- and which include Lindau's eae Nelsonieae, together Thunbergioideae, Nelsonioideae, and Echmatacantheae has Mendoncioideae, the second group nine tribes comprising Lindau's

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 Lindau's Acanthoideae raised Tieghem comprised only. Bremekamp (1953, 1955, 1965) Thunbergioideae and Mendoncioideae to family rank and transferred Nelsonioideae to the Thu the Acanthaceae family Scrophulariaceae. s, family 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 hairs of this epidermal family. Among relatively early reports of the studies of foliar of trichomes Acanthaceae may be mentioned the works of Lindau (in Engler &

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

investigated the epidermal features, including trichomes of six species of , and and and acaulis campestris. Singh Jain (1975) studied the structure and of different of trichomes of ontogeny types present on the floral appendages of 41 taxa Acanthaceae. The which deals with the and present investigation, structure, distribution, taxonomic of the foliar hairs of of Acantha- significance epidermal 39 genera(109 species) has been carried with better of the ceae, out a view to help in a understanding of this large and complicated family of flowering plants.

MATERIAL AND METHODS

The material of the species investigated consisted ofmature and healthy leaves collected herbarium from the Botanical of India from locally, or procured as specimens Survey or botanical gardens and herbaria of Sri Lanka, Singapore, Malaysia, Indonesia, and . The of the names species (and varieties) investigated are listed in the TableI at the end of cuticles this paper. Epidermal hairs were examined from separated from leaves by mechani-

cal peeling (scraping with a safety razor blade) or by macerationwith 10%—30% nitric acid. The cuticles with were washed water, stained with aqueous safranin, mounted in and the with Canadabalsam. While trichome their glycerine cover slip was ringed types, and structure, distribution, range of variation in the family are described under 'Obser- trichome characters of individual listed vations', species are in the TableI.

OBSERVATIONS

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

(a) GLANDULAR HAIRS

recorded all They have been in investigated species of the family and can be distin- guished into three 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- hairs with celled, globular or disc-shaped head, and (iii) long-stalked I- to several-celled 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-celledhead is roughly dumb-bellshaped 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 the hairs have been recorded all layer (Fig. 36). In present study, panduriform glandular in of the investigated species the genera (Fig. 1),Elytraria (Fig. 2), Nelsonia (Fig. 3), and of while Thunbergia (Figs. 4—7). The shape the glandular head basically remaining often in the panduriform, may vary same species, e.g. Thunbergia laurifolia (Figs. 5—7). the of the There is comparatively less variation in size glandular head among different species (Table I).

Subsessile hairs with head (ii) (short-stalked) globular or disc-shaped (Figs. 8—35

These hairs in all and 37, 38). are recorded the investigated genera except Staurogyne, and the The Elytraria, Nelsonia, Thunbergia, in except species H. serpyllum. head of cells. The glandular is globular or disc-shaped and is formed 2—8 4-celled con- dition is the In most common (Figs. 8, 21). genera like , , andRuellia glandular hairs with 8-celled head are more common (Figs. 9—11). In

R. with tweediana, lorentziana (Fig. 12), and Ruttya speciosa (Fig. 13) glandular hairs a more than 8-celled head also Glandular hairs with 2-celled head are common. are present in maderaspatensis (Fig. 14), nilotica (Fig. 15), and C. infundibuliformis.

hairs with in Glandular 2—4-celled head are common tetragona (Fig. 16), latifolium (Fig. 17), tinctoria (Fig. 18), purpuricaulis

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

In surface view, the outline of the glandular head is circular but 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). of head The diameter the globular varies only slightly in a species. Among species of the variationis same the diameter does not vary more thanI to 1.5. Thus the 23 —34,11m of Ruellia and of Strobilanthes. with in seven species 22—34 pm in nine species Species typically small glandular heads are: barlerioides (19 pm, Fig. 30), Hygrophila 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 pictum (38 pm), Justicia procumbens (35 pm, Fig. 23), and Barleria courtallica (35 pm, Fig. 24).

hairs These hairs of (iii) Long-stalked glandular (Figs. 39—53). consist a 1—5-celled uniseriate stalk terminatedby a I—several-celled globular or hemispherical head. The foot cell arise from the middleof cell and be may an epidermal (Figs. 41 —44, 48, 49, 51) or may surrounded to several cells Such hairs by 2 epidermal (Figs. 39—40, 45—47, 50). occur intermixed with the short-stalked glandular hairs in several species. In Hygrophila serpyl- 104 BLUMEA VOL. 24, No. 1, 1978

hairs — hairs with Glandular (Figs. 1—53). Figs. 1 —7. Sub-sessile glandular panduriform head. 1. Staurogyne

Nelsonia — — longifolia ; 2. Elytraria acaulis var. lyrata; 3. campestris;; 4. Thunbergia erecta; 5 7. T. laurifolia. hairs Figs. 8—35. Sub-sessile glandular with globularhead. 8. nobilis; 9. Strobilanthes ixiocephalus;

10.Pseuderanlhemum Ruellia lorentziana; malaccense; 11—12. 13. Ruttya speciosa; 14. Blepharis maderaspatensis; Crossandra 15. nilotica; 16. Aphelandratetragona; 17. Gymnostachyum latifolium; 18. Peristrophe tinctoria; 19—20.

L. Lepidagathispurpuricaulis; 21. trinervis; 22.Justicia vahlii; 23.J. procumbens; 24. Barleria courtallica; 25.Asysta- sia dalzelliana; 26. Fittonia verschaffeltii var. argyroneura; 27—28. Hemigraphis colorata; 29. Hygrophila poly-

Dianthera spermy 30. Petalidium barlerioides; 31. nodosa; 32—33. var. sparatteria; 34—35. M. coccinea. in — Figs. 36—38. Glandular hairs sectional view. 36. Thunbergia grandiflora; 37. Hemigraphis

hirta; 38. Petalidium barlerioides. — Figs. 39—51. Long-stalked glandular hairs. 39—43. Hygrophila serpyllum;

44. erecta ; 45—48. Dyschoriste vagans; 49—51. Lepidagathis cuspidata. — Fig. 52. Sectional view of

hair of — A branched trichome of long-stalked glandular roxburghiana. Fig. 53. Dyschoriste vagans

with glandularhead on onebranch. — Scale 100 µm for Figs. 39—51; scale 50 µm for Figs. 1 —38, 52; scale

250 µm for Fig. 53. K. Ahmad: Hairs Acanthaceae J. of 105

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

in and occur densely Hygrophila serpyllum (Figs. 39—43) Dyschoriste vagans (Figs. 45—48);

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 rosea, Strobilanthes heyneanus they are sparse and generally restricted to the veins and the margin.

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

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

to 1062 in and up /im length. In Hygrophila serpyllum (Figs. 41 —43) Lepidagathis cuspidata

these hairs arise from cell instead of (Figs. 49, 51), commonly a single epidermal each, hair-base. the others having a multi-celled Branched hairs with one branch glandular and

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

(b) NON-GLANDULAR HAIRS

Distribution Acantha- and frequency. Non-glandular hairs are of wide occurrence in of ceae. In a majority species, they are common (+ + Table I), in thirteen species they are , found to be dense (+ + + Table I), and in 21 (+, TableI). Non-glandular , species sparse

hairs are the intercostal generally more numerous on the veins and the margin than in in confined the areas, and some species they are to veins and the margin. In many species, those and especially of the genera Strobilanthes, , Hygrophila, the non-glandular the the hairs on lower epidermis are confined 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, nitidum, and O.

strictum.

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

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

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

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

Eranthemum and Thunbergia grandiflora (Fig. 56, 73 /im), wattii(Fig. 63, 57 /mi), Hygrophila polysperma (55 /Mn).

The uniseriate in acaulis hairs are all the investigated species except Elytraria var. lyrata which hairs in they are sometimes biseriate near the base (Fig. 94). Branched are rarely found in Pseuderanthemum bicolor Dyschoriste vagans (Fig. 85) and (Fig. 93). both Unicellular hairs Generally, unicellular and multicellular hairs occur in a species. are, however, predominant in Staurogyne longifolia (Figs. 54 —55), Thunbergia grandiflora

(Fig. 56), T. laurifolia (Fig. 57), T. kirkii, Mendoncia coccinea (Fig. 58), Dyschoriste dalzellii

D. Pseude- (Fig. 59), erecta (Figs. 60—62), Eranthemum wattii (Fig. 63), E. capense (Fig. 64), ranthemum grandiflorum (Fig. 65), Hygrophila serpyllum (Fig. 70), Lepidagathis trinervis

L. (Fig. 72),. cuspidata (Figs. 73—74), L. purpuricaulis (Figs. 75—76), L. incurva (Figs. 77—78), Barleria cuspidata (Fig. 79), B. lupulina, B. prionitis, andGymnostachyum 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 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.

E. 66. D. erecta; 63. Eranthemum wattii; 64. capense; 65. Pseuderanthemum grandiflorum; Asystasia dalzelliana;

67. Justicia betonica; 68. J. vahlii; 69. Dianthera nodosa; 70. Hygrophila serpyllum; 71. Rhinacanthus nasuta;

L. L. 72. Lepidagathis trinervis; 73—74. cuspidata; 75—76. L. purpuricaulis; 77—78. incurva; 79. Barleria cuspi-

— Multicellular data. Figs. 80—98. non-glandular hairs. 80. Fittonia verschaffeltii var. argyroneura; 81. F. verschaffeltii var. pearcei; 82. Gymnostachym febrifugum; 83. echioides; 84. Hemigraphis hirta; 85.

86. 8 Dyschoriste vagans (branched hair); Ruellia formosa; 87. Justicia procumbens; 88. Hygrophila serpyllum; 9. Adhatoda Pseuderanthemum Asystasia chelonoides; 90. vasica; 91. Aphelandra tetragona; 92—93. bicolor; 94. Ely- Mendoncia traria acaulis var. lyrata; 95. velloziana var. sparatteria; 96. M. velloziana var. sparatteria (stellate hair-base); 97. Ruttya speciosa;98. Lepidagathis cuspidata. — Figs. 99—101. Portions of non-glandular hairs

Eranthemum magnified, to show wall ornamentation. 99. purpurascens; 100. Ruttya speciosa; 101. Dyschoriste vagans. — Scale 100µm 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 deal and be small vary a great in shape size. They may slender conical or small and or papillose (Figs. 54—56, 59 —63, 65, 66, 68, 71), stout, and needle-shaped (Figs. 70, 73, 74), or comparatively large, stout, erect (Figs. 72, 78, 79). and They are typically thick-walled lanceolate in Barleria cuspidata (Fig. 79), B. lupulina, hair-base but B. prionitis. Generally, the hairs have a several-celled in some cases they

from the middle of an cell arise epidermal (Figs. 54, 62, 64, 70 —71, 73 —77).

The 80 —6-celled. multicellular hairs (Figs. —95, 97—98) are generally 2 be 12-celled Pseuderanthemum bicolor and They may up to in (Figs. 92, 93) up to 15-celled the of the in Elytraria acaulis var. lyrata (Fig. 94). As in the case of unicellular hairs, shape

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

slender (Fig. 92), or long and stout (Figs. 82—83, 86—89, 95)- The non-glandular hairs of some species are characteristic. In Aphelandra the hair cells tetragona (Fig. 91) is built oftwo or more small basal surmounted by a much

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

here lies than the basal cells. In at right angles to the basal part and has a thicker wall thick-walled hairs Adhatoda vasica (Fig. 90) the hairs are sometimes T-shaped. Lanceolate

occur in several species of Barleria (Fig. 79). Thick-walled uniseriate hairs with bulgings

at the nodes are recorded Mendoncia in velloziana var. sparatteria (Fig. 95). The hair-base is generally multicelled, with polygonal, isodiametric, straight- walled cells. and However, small and slender, or more rarely long stout non-glandular hairs from the middle of cell often spring an epidermal (Figs. 84, 90, 98). Hygrophila

from a serpyllum has both these types: 1—2-celled, small, slender hairs emerging single cell hair- epidermal (Fig. 70) and 2—4-celled, longer, and stouter hairs with a multicelled the cells less base (Fig. 88). In species with sinuous-walled epidermal cells, hair-base are

sinuous or straight-walled. In Mendoncia coccinea and M. velloziana var. sparatteria, non- of have —several glandular hairs the upper epidermis a typically stellate hair-base with 2

arms (Fig. 96). Barleria the Thenon-glandular hairs are moderately thick-walled. In species of thickness The wall of the wall is considerable and the hairs consequently have narrow lumina. tubercles ornamentationgenerally consists ofround, oval, elliptic, or linear (Figs. 60—62, the wall be used for 99—101). Sometimes, ornamentation can diagnostic purposes.

DISCUSSION

The of of of Acanthaceae reveals study the foliar epidermal hairs 39 genera (109 species) that various features of the epidermal hairs, like their shape, size, the number of cells comprising them, thickening and ornamentationof the wall, type of hair-base, etc. play hairs of a very useful role in the systematic considerationof various taxa. The glandular mentioned earlier intermixed While the three types occur with the non-glandular ones. the tribe the glandular hairs are more important at the higher level (subfamily, etc.), 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 ofthe Acanthaceae. Thunbergioideae, whilst the disc-shaped glands occur in the rest

The shows that the hairs are present investigation, however, panduriform glandular

of these two sub- uniformly present in Nelsonioideae as well, and they are characteristic families of the Acanthaceae. Kumar andPaliwal (1975), who studied the foliar trichomes of six species of Thunbergia (subfamily Thunbergioideae)) and Elytraria acaulis and Nelsonia campestris (subfamily Nelsonioideae) have recorded long-stalked glandular hairs in Nelsonia the campestris. However, they have not reported presence ofsubsessile panduriform glandu- lar hairs in the species investigated by them. BLUMEA VOL. 24, No. 1, 1978 108

of the Acanthoideae and Mendoncioideae (of Lindau, 1895) In the genera subfamily the hairs head. This presently investigated, glandular have only a globular or disc-shaped Mendoncia is in conformity with the observations of Solereder (1908). In (sub-family hairs Mendoncioideae), while the heads of glandular are basically globular or disc-shaped, ,, be and in surface view. they may frequently modified appear quadrangular or triangular hairs it While the diameter of the head of glandular does not vary appreciably, can the Thunbergia sometimesbe used for distinguishing one species ofa genus from other, e.g. 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

used taxonomic character. In some the glandular head is quite important and can be as a All the of the sub- cases it is constant for a sub-family or a genus. investigated genera The nine families Thunbergioideae and Nelsonioideae have only 2-celled glandular heads. —8-celled heads. species of Strobilanthes presently studied uniformly show 4 glandular heads. All the Similarly, the seven species of.Barleria have 4—8 (mostly 4)-celled glandular

six of Pseuderanthemum have —8 heads. On the other species 4 (mostly 8)-celled glandular within be distinguished from the others by the hand, a genus, some species can easily difference in the number of cells comprising their glandular heads. According to Santapau and minute and relate (1951), the differences between Dyschoriste dalzellii D. vagans are the shows that they only to the size of the anther spurs. However, present investigation hairs. D. dalzellii has can easily be distinguished from each other by their epidermal the head is D. subsessile glandular hairs with a 4-celled head, but in D. vagans 8-celled; hairs in D. also has of which are lacking vagans a dense covering long-stalked glandular dalzellii. subfamily The long-stalked glandular hairs have been recorded only in nine species of Kumar and Paliwal Acanthoideae and not in the other subfamilies. However, according to also. While their are Nelsonia (subfamily Nelsonioideae) ( T 975)> they present in campestris

taxonomic level be are present in widely significance at higher appears to limited, as they different from of view species taxa, they are nevertheless important diagnostic point jit D. level. For example, such hairs occur in Hygrophila serpyllum, Dyschoriste erecta, vagans, and Strobilanthes heyneanus, Dipteracanthus patulus, Lepidagathis cuspidata, Justicia tranque- of the bariensis, but are absent in the other investigated species genera. hairs role in the taxonomic consideration of Non-glandular play a very significant thicken- various show considerable variationin their size, wall taxa as they range of shape, unlike glandular hairs, the ing, and hair-base structure. However, as pointed out earlier, also useful in distin- hairs useful they are non-glandular are more at species level; rarely others. guishing some genera from the unicellular The ofBarleria hairs which are species have very characteristic non-glandular lumen canal-like having a (rarely I- to several-celled) with thick walls and a narrow Solereder, spherical enlargement at the base of the hair (Fig. 79). According to Hobein(see

these of the tribe Barlerieae investigated by him. 1908) hairs are present in most genera various Metcalfe and Chalk (1950) have also characterised various tribes of Acanthaceae by

f hairs. the shows that in Lepidaga- types o non-glandular However, present investigation

which with unthickened walls occur. The this, belongs to the same tribe, uniseriate hairs same view is expressed by Solereder (1908).

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

hair-base of cells forming a rosette several-celled, stout, with a several-celled polygonal around the foot basal distribution also characteristic as they or cell. Their pattern is quite are generally confined to costal and marginal areas. hairs. These A number of species are characterised by their typical non-glandular K. J. Ahmad: Hairs of Acanthaceae 109

include: conical unicellular hairs on the upper epidermis of Dyschoriste dalzellii (Fig. 59),

hairs of branched D. vagans (Fig. 85) and Pseuderanthemum bicolor (Fig. 93), conical unicel- lular hairs ofLepidagathis 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 of hair-base is also an important diagnostic character. The two species Mendoncia Rizzini are characterised by typically stellate hair-bases on the upper epidermis (Fig. 97). divided of the hair- (1948) Mendoncia into four subgenera on the basis of the characters

base. The two species investigated here have non-glandular hairs with a conspicuously stellate hair-base the the lower the hair-base on upper epidermis; on epidermis cells, though basically indistinguishable from the other epidermal cells, frequently form

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 is characteristic from the middle ofan epidermal cell and this feature very for those species. the hairs often within The density of non-glandular varies considerably 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 from of these genus can be distinguished each other on the basis characters (Table I). hairs often within but where Non-glandular show a wide range of variation a species,

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

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

in there is and in the division that no sharp distinction between the two groups, any case related illustrates the separates obvioulsy very closely types'. Dyschoriste vagans strikingly

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 Where the hairs and hairs of this species. are branched, one branch may be non-glandular

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

The delimitationof Acanthaceae by Nees (1847), Bentham & Hooker (1876), Lindau

(1895), Van Tieghem (1908) and Bremekamp (1953, 1955, 1965) has been explained

earlier (see 'Introduction'). The present study shows thatLindau'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) Nelsonioideae the latter transfer of to Scrophulariaceae as the glandular hairs in are not

this Nelsonioideae close to panduriform (Metcalfe & Chalk, 1950). In respect is more 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 Thun- Nelsonioideae to Scrophulariaceae and raising of his subfamilies Mendoncioideae and in bergioideae to family rank as suggested by Bremekamp (1953, 1955, 1965) his delimita- tion of Acanthaceae is not justified. This view is also supported by evidence from the stomata & and (Ahmad, 1974, 1974a; Paliwal, 1966, 1967; Kumar Paliwal, 1975) 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 Dr. R. for his valuable in this work and owe my deep gratitude to V. Sitholey guidance to my colleague D. B. in assistance. I thank Mr. Shukla for helping me the local collection and laboratory wish also to the

individuals and for for following organisations supplying me the bulk of the material required this study:

Dr. A. N. Rao, University of Singapore; the Curator, Lembaga Biologi Nasional, Bogor (Indonesia); Dr.

K. C. Cheang, Waterfalls Garden, Penang (Malaysia); the Superintendent, Royal Botanic Gardens, Perade- niya (Sri Lanka); Dr. Margaret Emmerich, Curator of the Herbarium Museu Nacional, (Brazil); and the Director, Botanical Survey of India, Calcutta, India.

Botanic I wish especially to record my gratefulthanks to Dr. T. N. Khoshoo, Director, National Gardens,

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

REFERENCES

K. in Bot. Linn. Soc. AHMAD, J. 1974. Cuticular studies 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.

BACHMANN, O. 1886. Untersuchungen iiber die systematische Bedeutung der Schildhaare. Flora 69: 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 in BENTHAM, G., & J. HOOKER, 1876. Plantarum. 1862—1883; Acanthaceae 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 Anatomy. Holt, Rinehart & Winston.

D. The Amer. Bot. COOPER, C. 1932. development of the peltate hairs of Shepherdia canadensis. J. 19:

423 —428.

M. A COWAN, J. 1950. The Rhododendron leaf: study of the epidermal appendages. Oxford & Boyd,

London.

T. H. Botanica GOODSPEED, 1954. The genus Nicotiana. The Chronica Co.,Waltham.

A. Proc. Indian INAMDAR, J. 1967. Studies on the trichomes of some Oleaceae, structure and ontogeny.

Acad. Sci. Sect. B, 66 (4): 164—167.

B. & H. of JOHRI, M., SINGH, 1959. The morphology, embryology and systematic position Elytraria

acaulis (Linn, f.) Lindau. Botanisk. Notiser. 112: 227—251.

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

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

METCALFE, C. R., & L. CHALK, 1950. Anatomy ofthe Dicotyledons. Oxford, Clarendon Press.

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

Regni Vegetabilis 11: 46—519. Paris.

G. S. Structure in 16 PALIWAL, 1966. and ontogeny of stomata some Acanthaceae. Phytomorphology (4):

527—532.

1967. Stomatal ontogeny in the family Acanthaceae and the systematic position ofthe genus Elytraria.

Proc. Indian Sci. Congr. 54, Part III, Abst., Sect. VI. Botany: 306—307.

RAMAYYA, N. 1972. Classification and phylogeny of the trichomes of angiosperms. In Research Trends in Plant Anatomy. Ed. Ghouse & Yunus, Aligarh, India. Acanthacearum Brasiliensia. do RIZZINI, C. T. 1948. Disquisito Circa aliquot genera Arquivos Jardim Rio Botanico, de Janeiro8: 295 —372.

K. The ofhairs the Solanum. Taxon ROE, F. 1971. terminology in genus 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.

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

K. R. SPORNE, 1956. The phylogenetic classification ofthe angiosperms. Biol. Rev. 31: 1 —29. Cuticular studies aid Bull. Hist. STACE, C. A. 1965. as an to plant taxonomy. Br. Mus. nat. 4 (1): 1—78. 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 Acanthaceae of 111

TABLE I. EPIDERMAL HAIRS OF ACANTHACEAE

— D: Explanation of abbreviations. average diameter of glandularhead; Glob: globularhead of subsessile and and glandular hair; L: length oflong-stalked glandular non-glandularhairs (minimum, average, maxi- in mum that order); Low: trichomes on lower epidermis; Ls: long-stalked glandularhairs; Pand: panduri- form head of sub-sessile rhomboid glandular hairs; Rhomb: or quadrangular head of glandular hairs; Ss: subsessile St: stalk of the U: trichomes glandular hairs; long-stalked glandular hairs; on upper epidermis.

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

taken is The source of the material from which the voucher specimens are indicated in abbreviated form and is given within brackets. The followinginternationally accepted abbreviations have been used: BLAT, Blatter Herbarium, St. Xavier's College, Bombay (India); BOG, Lembaga Biologi Nasional, Bogor

(Indonesia); BSI-North, Botanical Survey of India, Northern Circle, Dehra Dun (India); BSI-South, Botanical Survey of India, Southern Circle, Coimbatore (India); BSI-Wcst, Botanical Survey of India,

Western Circle, Poona (India); CHAMOL, Parsari Farm, Chamoli, U.P. (India); CNH, Central National Herbarium, Botanical Survey of India, Calcutta (India); DE, Dr. Anima De, Bose Research Institute,

Calcutta (India); DUN, Dehra Dun, U.P. (India);LUCK, Lucknow, U.P. (India); LWG, National Botani- 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. VOL. No. 112 BLUMEA 24, 1, 1978

2- fim) fim.) 2- 944944 only. 2- 427/im)427/1111) 743) 187/im) U.U. 115115 187 (rarely ii) stellate. 71m) & 59—190,0m) (L: Low. 3-celled,3-celled, I038jum)103871m) 59— fim) 1570^m) 1486/im)148672m) 1014/im)ioi4,um) I334yum)I334yum) 165—554—743) fim) 1570 slender, 140,um);14071m);Low.Low. 944 upper, rare) 335,11111) unicellular 335 +, biseriate. (L: margin,margin, on (L: (L: 79— (L:(L: (L: (very (L: 4-4-4-,+ 71m) short, toto + 4-celled,4-celled, short, 825825 often ii) +, i)-celled. 45—155— , (L: i)-celled. i)-celled. + (L: hair-base +, + (L: hair-base i)-celled. + 7)-celled. (L: (L: (L: i)-cclled. i)-celled. (L: (L:(L: confined + fim) confined + hairshairs (L: + U. stout, 1298/im)129871m) to i) i) stout, jum) 826,um)826 137/zm);13771111); i) 15-celled, & 3(mostly 6-celled. 2(up 3-celled. 519 to 73—40— 1—3(mostly types; conical, (L: types: long, 1—2(up17— 1—6-celled; 80—-191— 1—3(mostly191— 554— 1—3-(mostly42—89—165—42— 1—3(mostly 2—6-celled. Low. 2—9-celled.476— 106—476—106— 82—590—590— 1—6-celled. 212—424— 1—3(mostly 73—40— 1— 637—2—6-celled.637— 176— I—20—S5—20—55— 1— 654—236—2—2—6-celled., 720—2—6-celled.720— 106— 2- 79—1—30— 2—2— 1—17—70—349—1—3-celled.up I—3-celled.45— 155— I—70— 349— 47—330— 1—3(mostly 1—3-(mostly 1—3-celled. +, 47—330— 270—20— + two only.20—270— two 578—224—224— ++, +, Non-glandularNon-glandular +, +, +, 59—204— +, +, +, +, +, +, +, +, + +4-4-4-, celled), celled, + + + + + + +, fim) + + 4-4-,+ Of + + Of + + (L: + +, +, (L: U. + (L:

2671m)26yum) /jm)fim) 32,um) 32,um) 34/

SING)SING)LUCK) SING) LUCK) LUCK) LUCK) LUCK) LUCK) LUCK) LUCK) MALAY) MALAY) LUCK) LUCK) RIO) BSI- BSI- (ex(ex (ex (ex(ex (ex RIO) (ex (ex (ex (ex (ex (ex (ex (ex (ex (ex (ex (ex (ex (ex specimen (ex 64561 6461464614 64546 64593 64606 64607 64609 64589 6459564595 64590 64563 64562 64615 64617 42757 114.908114.908 38.323 BSI-South) BSI-South) 76626 Voucher LWG LWG LWG LWG LWG LWG LWGLWG LWG LWG BSD North) West) LWG LWG LWG R R LWG LWG LWG LWG (ex (ex BSI

Ktze. Bremek. Nees O. var. Anders.

Lindau Br. Nees (Schumach.) T. (Nees) (Nees) Moore Nees R. Veil. i I Nees S. Sims Anders. Hook./Hook./. Anders. s (Buch.-Ham.) (L./.) (Roxb.) (Buch.-Ham.) lyrata (Mart.) ex Roxb. T.T. (Vahl) longifolia coccinea (Mart.) affinis Anders.Anders. Roxb. Lindl. auriculataauriculata T.T. var. campestriscampestris Hook./Hook./. nobilis acaulis BojerBojcr T. velloziana grandiflora laurifolia mysorensis polysperma quadrivalvis salicifoliasalicifoUa serpyllumserpyllum Staurogyne Elytraria acaulis Nelsonia Mendoncia sparatteriasparalleria Thunbergia alata erecta T.T.fragransfragrans kirkii Sanchezia Hygrophila Heine H. E. M. T. T. T. T.T. T. T. H. H. H.

5- Species I.1. 2. 3- 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) /im)ftm) 354/im)354/mi)fim) yum)fim) only. 490490 637,um) 1191/jm) 1298/im)1298/im)57/mi) unicellular, 295 46046o/im) 1191 3S7ftm) rarely 400400 3 550,um)55O/tm) Low. /im)fim) yum)fim) 168—168— 2i2//m)212/zm) short, 800^m)800 6016oi,uin) 100100 4)-celled,4)-celled, 205205 4-celled,4-celled, ++, /im)fim) (L: (L: (L: (L: (L: (L: (L: (L: 849 ii) 849 (L: (L: 4)-celled.4)-celled. 4)-celled. 4)-celled. ij-celled. /xm) (L: (L: 2 fim) ++■++, (L: (L: hairs 920yum);92Oyum); o(moscly9(mostly (L: several-celled. 684y«m) inifim)1121/xm) 2 262/xm) 262 i) several-celled. several-celled. several-celled. several-celled. several-celled. several-celled. only. to several-celled. several-celled. 4-celled. (L: 5-celled. to 8(mostly8(mostly 8(mostly8(mostly types: up to to 1-to to to to 1—1— U.141—531—141—531— 2—2— 2—5(often50—204— 283— 94—118—94— 200—94—200— 55— 635—195—to932—195— 272— 613—236—155— 90—227— 175—340—175—340— 8(mostly8(mostly 635—932— to to 283— 118— 1- 42—42— 1—5-celled.106— 62—85—85— 1—4-celled.12—192— 2—5-celled.no—no— 2—5-celled.276—167—94—167— 1- 1- 1- I-1- I-1- 224—495—684 1—192—12— 276—2—94— +,!- I—2—2— 1— 2— 1—153—365— 147—205—: +, 291—94—291— 2— 1— two 1- 1- 495—224— I— 205—147— +, +, I— +, +, + +, +, +, +, ++,+ +, +, +, +, , , Non-glandular branched. , conical, + + +, + Of (L: + + + + + + + + + + + + + + (L: (L:(L: (L:

/xm) /xm) /xm) 8- /xm)fim) fim)fim) /xm) /xm) fim);/xm); um); io62^m)25/im) 24/jm) 25,um) 27^) 26/xm) 2S/xm)25,um) 2S/xm)22/xm) 34,um)34jxm) 27,um) than 32/im) Glob., Glob., i 24 25 27 26^01) 4—8- 27 27 32 8)-8)- Glob., Low. 2.6fim); 25 22 23 26 Ls., 4— 27/xm)27,um) (D. (D. (D. (D. Ss., fim)Ss., 5-celled, (D. (D. (D. ßm) (D. (D. (D. (D. (D. (D. (D.(D. (D. +, more (D. 117 , 2—5-celled, Glob., 7}Oftm) +, + 750 4-4-, U. 2-celled,2-ceiled, U. ii) + + Ss., fim) 8-celled.8-celled. 8-celled.8-celled. (L: 8-celled. 8-celled. 8-celled. 8(mostly & & St. 8-celled. 8-celled. 8-celled. 8-celled. 8-celled.8-celled. 8-celled. 8-celled.8-celled. 8(often8(often 26ßm) 2—8(mostly /xm) 4-celled.4-celled. i) i) 4—8-celled. 4—8-celled. 4—8-celled. 4—8-celled. 4—8-celled. i) fim); 4—8-celled. 4—8-celled. 4—8-celled. 26 4—8-celled. 19/im) (L: hairs 19 St. Ls., Low. Low. 27 (D. types: types: U. types: (D. Ls., +, (D. (L: &&U. + 384—106— 4— 4— 4— 4— 4—4— 22,um) Glob., Glob., Glob., Glob.,4— 22/xm) 4— Glob., Glob., i— 1— Glob., Glob., 4—Glob., Glob., Glob., 4—Glob., Glob.,Glob., Glob., Glob.,Glob., 4—2— 77—106— 2— Glob., 4— 4— 3—4-celled. 4— 4— 4— 4— 384—106— 4— 4— two two 77— 106—two + +, 4— Glandular Ss., celled. Ss., 4-celled,4-celled, only. 8-celled, Low. Ss., Ss., Ss., Ss., Ss., celled. Ss., Ss., Ss., Ss., Ss., 8)-cclled.8)-celled. Ss., Ss., Of ii) Of ii) Ss., Ss., (D. Of St. Ss.,

MALAY) LUCK) LUCK) LUCK) LUCK) SING) BSI-West) BOG) LUCK) BLAT) BSI-West)BSI-West) LUCK) BLAT) DE) LUCK) DE) BSI- BSI- BSI-West) BSI-West)BSI-West) BLAT) (ex (ex (ex (ex (ex (ex (ex (ex (ex (ex (ex (ex (ex (ex (ex (ex(ex specimen (ex (ex (ex 64588 64586 490 64567 54354 6455964559 64580 64613 6454864548 64549 64583 CHAMOL) 64588 64586 3 64558 66489 66604 BSl3857(exBSI-West)3857 79955 9685 74982 Voucher BSI66489 LWG LWG BLAT LWG LWG BSI West) BSI West) BSI LWG LWG LWG (ex BSI BSI LWG LWG BLAT BSI LWG BLAT LWG LWG LWG

Nees

Anders. ex Ktze. Nees T. O. Wall, z. Greenman Benth. Andr. Ktze.Ktze. Blume ClarkeClarke Dalz.Dal Griseb. Ktze. alternataaltemata Anders. alatus Nees barlerioides dalzellii Hall./. NeesNees Nees O. (L.) Nees O T. Nees ata s formosa „._,. lorentziana malacosperma erectaerecta vagans color hirta/ii'rfa barbatus callosus crispus heyneanus ixiocephaluspulneyensispulneyensis scaber scrobiculatusscrobiculatus lorentziana Petalidium Dyschoriste Hemigraphis StrobilanthesStrobilanlhes RuelliaformosaRuellia D. D. H. H. S. S. S.S. S. S.5. S. S. S. R.i?. R.

Species 21. 22. 23.23- 24. 25.25-_., 26. 27-27. 28. 20.29. 30. 31.31- 32. 33-33. 34.34- 35- 36. 37-37. 38.38. 39-39. 114 BLUMEA VOL. 24, No. 1, 1978

/im) /im)731/im) 743743 731 /im) 260/im)260/im) /im)/im) 10031003/im) /im) 195/im) 450/im) 170/im) 235/im) 162/im)162/1111) 195 127/cm) 450/im) 735/Mn)735/im) 170 —235/1111) 127 400/im)400/tm) 512/im)512/101) 175/tm)175/1111) 4)-celled.4)-celled. (L: (L: 4)-celled.4)-celled. 4)-celled.4)-celled. (L: /im) /im) (L: (L: (L:(L: (L: (L: (L: hairs 650/im) 225/im) (L: 425 225 4-celled.4-celled. (L: (L: (L: several-celled. thick-walled. 8(mostly8(mostly 337—650/Hn) i-celled. 8(mostly8(mostly 8(mostly8(mostly I—4-celled. 1—4-celled.1—4-celled. to410—410— 188—188— 247—483—483—247— 82— 164—1— 440—212— 1—4-celled. l-celled.i-celled. i-celled. i-celled. I—82— 164— 440—212— 70—27—1—4-celled. 87—53— 1—4-celled. 57—22— 1—4-celled. 1—4-celled.167—15—167—15— 205—50—205— 1- 70—27— 1—4-celled. I—4-celled.22—22— 1—4-celled.87—53—100—100 1—4-celled.57— 102— 272—272— 117— 22— 142—142— 58— 119—119— 1- 2—2—1—1—137—245—I—2—2— 1— 142—337— 2— 1— +.+, +, i-celled, 142— I—2— (L:i48—(L:I48—166—166— +, + +, +, +, +, +, +, +, +, +, + +, +, Non-glandularNon-glandular , + ++, + + + +4-+, + + +, + (L: +, (L: ++, + + ++ + + +4- +

8)- /im) /im) +, +, than 27/im) +++, 27/rm)27/im) 24/im) than + 2323 Ls. 24 /im) /im) /im) /im) /rm)/im) 4)-celled. 4)-celled. ii) 4)-celled. ii) 220/tm)220/im) 25/im) 26/im) 34/tm)34/im) 26/tm)25/rm)25/im) 24/im)24 ii) 26/tm)26/im) (D. +, (D. more Glob., (D. 25 26 26 Glob., 23 (D. (D. (D. /im); (L: more-celled. (D. (D. (D. (D. (D. /im); (L: (D. Ss., 27/im); Ss., 22/im); 8-cclled. 8-celled; 8(often8(often 27 8-celled. 22 4-celled.4-celled. 8-(mostly8-(mostly 8(mostly8(mostly 8(mostly 8-celled. i) i) i) 4—8-celled. 4—8-celled; 4—8-celled. 34/im) (D. 3-celled.3-celled. 4—8-celled. or 4-celled. 4-celled.4-celled. 4-celled. 4-celled.4-celled. 4-celled. 4-celled.4-celled. (D. 5-celled.5-celled. 330/tm)330/im) 4-celled. 4-celled 4—8(mostly hairs types: 8- types;types: 4— 4—4—(D.(D. 2—2— 4—4— 4— 4—8-celled. 30/im) 35yUm) 4—4— 26/tm)26/im) 34/tm)34i"m) 30/tm) 4—8-celled. 35/tm) Glob., Glob.,4— Glob., Glob., Glob.,Glob., Glob.,Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glob., 2—2— Glob., 26/tm)26/im) 4— St. Glob., 4—Glob., Glob., two St. Glob., Glob., two GlandularGlandular two 4—8-celled. 4—8-celled. 2—2— 27—135— Ss., Ss., (D. Ss., Ss., celled. Of Ls., Ss., Ss., (D. Ss., Ss., Ss., Ss., Ss., Ss.,Of Ls., Ss., Ss., Ss., Ss., (D. Ss., (D. Ss., (D.

SING)SING) LUCK) LUCK) LUCK) POND) CNH) LUCK) LUCK) SING) DUN) BSI-Dun) LUCK) BSI- BSI-West)BSI-West) BSI-West) BSI-West) BSI-West) BSI- (ex (ex(ex (ex (ex (ex (ex (ex (ex (ex (ex (ex (ex (ex specimen (ex (ex (ex (ex (ex (ex 64560 64596 64604 64605 64602 64603 80113801 64608 64610 64547 64612 40247 64598 12502 POND) 13 POND) 72973 79530 71408 86000 12502 Voucher LWG BSI72973 LWG LWG LWG LWG LWG LWG (ex MH South) LWG LWG (ex LWG BSI LWG BSI BSD BSI BSI LWG MH South)

Nees Mart. (Jacq.) Br. Nees Nees Nees Br. Willd. Nees R. ex Nees ex ex patulus Nees R. Nees Griseb. (Poir.) albomarginata cristatacrislata Wall, Don. Heyne Linn.Linn. Linn. (Vahl) Wall, Wall, Linn.Linn. Hemsl. (Vahl) Stapf. D. courtallicacourtallka Linn. tweedianatweediana prostratus nervosum purpurascens cuspidata purpuricaulis cuspidata repens rosea tuberosa Dipteracanthus Eranthemum capense roseum wattii Lepidagathis incurvaincurva trinervis cristatacrislata R. R. R. D. Barleria R. E. E. E.E. E. E.E. L.L. L. L. L. B. B.

Species 40 41. 42. 43. 44. 45. 46. 47-47. 48. 49. 50. 51. 52. 53. 54-54. 55-55. 56. 57. 58-58. 59.jo. K.J. Ahmad: Hairs of Acanthaceae 115

/tin)ftm) 32m)/im) /im) 4)-4)- 731731 625625 59059032m) 2-celled). 32m)ftm) /im) /im) (rarely 531/cm) 755/tm) 125/Mn)12532m) 401401 25032m) 531 —755/im) upper. 250 (mostly 150/tin)150/im) thick-walled.thick-walled. thick-walled. 77 on 4)-celled. (L: (L: (L: /im)/on) 4)-celled.4)-celled. 82—365182—369— 4)-celled.4)-celled. 4)-celled.4)-celled. (L: unicellular 4)-celled.4)-celled. 2—4)-celled. upper, 1203-1203 ftm)fim)(L: (L: /im) (L: 9)-celled (L:(L: fim) (L:(L: hairs 1416/im)141632111) 483/im)48332m) (L: thick-walled. thick-walled. 287/an)28732m) 15532m) to 436/im)43632m) (mostly 507/rm) several-celled,several-celled, thick-walled. several-celled, 660660 778 several-celled. to conical, 50/im)5032m) 507 7(mostly7(mostly 8(mostly8(mostly 6(mostly 77 3(up3(up in—155/im) 7(mostly7(mostly to to HI— to 362—212— 389—389— 177—177— 448—531—531— 1—6(mostly 212—362— 1—6-celled. I—4-celled.1—4-celled. confined2—2— (L:1— 267—267— 1—6-celled.118— 98—98—37—37— short, , 3—6-celled.236—413— 1- I—3-ceIled.74—74—27— 1—3-celled. 1- 3—6-celled.236—413— i- 70—377—7783^1) 55—55—116—116— 1- 2— 1— 2—2 — + 2—3-celled. +■ 2—+,2—1—1— i-celled, i-celled, 3—3—1—1— 3—3—1—70—377— 81—424— 2—2—1—1—94—224— 47—224— 118—330—2—3-celled. 6—590—590—10106— 61—280— 52—142— 75— 224—94— 330—118— +, 61—280— 52—142— +, +, 26—12—12—+, + + Non-glandular +, +, +, +, +, +, +, (L:75— +, +, +, + + +, +, + + + + celled. + + +, + (L: +, (L: + + (L: + + (L: + + (L: + (L:(L: + + (L: (L:(L: (L: +

2732m)/im) 4)-cclled.4)-celled. 4)-celled. 4)-celled. 4)-celled. /im)3i^m) 2sμm)25/im) 24/4111)24/im) 32/im) 27 8)-celled. 8)-celled. 4)-celled. 4)-celled. 4)-celled. 4)-celled. 4)-celled.4)-celIed. 4)-celled. 38/im)3832m) 2932m)/im) 8)-celled.8)-cclled. 31 (D. 29

(D. (D. (D. (D. (D. (D. 8(mostly8(mostly 8(mostly 8(mostly8(mostly 8(mostly 4-celled.4-celled. 4—8(mostly4—8(mostIy 6(mostly6(mostly 4(mostly4(mostly 8(mostly4—8(mostly 8(mostly8(mostly 8(mostly4—8(mostly 8(mostly8(mostly 4—8(mostly8(mostly hairs 4—8(mostly 4—8(mostly 2-celled. 4-celled. 2-cellcd. 2-celled. 4—8(mostly 8-celled. 4-celled. /im). 4— /im) 2—2— /im) 4—4— /im) 4— ftm) 4— [im) 4— 4—4— 27//m)27/im) 4— 4—4— 35/itn) 3i/«n) Glob., 2732m) Glob., 2432m)24/im) 2—2— 23/tm) 4—4— 33/im)3332m) 25/im) 2432m).24/Lim). Glob., 4— 29/tm)29/im) 30/tm)30/im) 31/tm) Glob.,4— Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glandular 32 31 27 23 31 25 Ss., (D. Ss., (D. Ss., (D. Ss., (D. Ss., Ss., Ss., Ss., Ss., Ss., (D. Ss., (D. Ss., (D.(D. Ss., (D.(D. Ss., (D. Ss., (D. Ss., (D. Ss., (D.(D. Ss., Ss., Ss.,(D.

LUCK) LUCK) LUCK) LUCK) LUCK) DE)LUCK) LUCK) POND) LUCK) MALAY) LUCK) POND) BSI-West)BSI-West) BSI-West) BSI-West)BSI-West) BSI- BSI- BSI- x (ex (ex (ex (ex (ex (ex(® (ex (ex (ex (ex (ex (ex (ex (ex (ex specimen (ex(ex (ex (ex (ex (ex 6460164601 64599 6460064600 64591 64611 64544 64570 64572 6457664576 64569 i93i7(exBSI-17 64542 64571 64564 7477674776 97409 1273512735 2031320313 193 LUCK) Voucher BSI76416BSI76416 LWG646n(exLUCK) Voucher LWG LWG LWG LWG LWG LWG LWG LWG LWG LWG BSI BSI MH South) MH South) MH South) LWG (ex LWG LWG

Nees Roth Nees J.inn.) (Linn.) ex Benth.Bench. terminalis (Linn.) Anders. var. Anders. terminalis Anders. Nees Wall, Griff. Nees Nees Nees T. T. T. atropurpureum var. febrifugumfebrifugum SantapauSantapau pictum infundibuliformisinfundibuliformis tetragona echioides (Burm.) (Dalz.) Clarke (Linn.) coccinea Anders. Lindl. Linn. Willd. maderaspatensis montanus Oliver chelonoides BlumeBlume T.T. ClarkeClarke -angularis Bailey lupulina nilotica paniculatapankulata latifolium dalzelliana gangeticagangetka intrusa lawii lupulina prionitis strigosa BlepharisBkpharis Crossandra Aphelandra Andrographis Gymnostachyum Asystasia quadrangularis GraptophyllumPachystachys Pseuderanthemum (Nees) C. A. quadt A. A. A. (Bull.) ies B.B. B. B. B. A. G. ' A. SpecSpecies 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

rarelyrarely apical cells, hairs hairs 74m)ftm) slender,slender, pm) basal fim) /urn) 10731073 ftm) upperupper upperupper /urn) 332 235yum) 357/im)35774111) 37774m) 2807(01) 26074m) thick-walled 377 82674m) i)-celled. i)-celled. 280 119—23574m) 260 small 826 conical. i2)-celled, 354jUm)35474m) 59—174—33271111) 4)-celled.4)-celled. 74m)4)-celled,4)-celled, 59—174— 3)-celled.3)-celled. several 4)-celled;4)-celled; 442/4111) to elongated 8o2yum)80274m) (mostly (L: (L: (L: (L: (L: ftm) 200—442 (L: i)-celled, hairs (L: (L:(L: an 475yum) (L: several(mostlyseveral(mostly I30yum)13074m) iisfim)11574m) 354pm)354 I42;um)14274m) several(up several by 2— 1—io(mostly (L: 5(mostlys(mostly io(mostlyio(mostly io(mostly io(mostlyio(mostlyconical.conical. to to 1—3-celled. 1—4-celled. 1—5-celled. (L: 5(mostly5(mostly to 822—424— 1—7-celled. 3—7-celled,several 1—1— 2— 1— I—2—2— 1— 1-1- 67— 1—3-celled.424—152—822— 1—7-celled. 42— 1—4-celled.119—32— 130—130— 75— 1—3-celled.142— 7j—150—47574m) 42—42— 162—236—59—236—59—1—5-celled. 228—228— 118—118— 1- 67— 1—3-celled.152— 75— 1—3-celled.142—2—I—2—1—3J—155—35—155— 2—2—1—1—75—150— 1—1— (L: I-1- 578—389— 50—200— , 27— 37—64— 578—389— +, 62—62—27— +, +■+, +, +, 82—55— +, +, + 82—55— Non-glandular +, +, +, +, branched. surmounted ++, conical.conical. typicallytypically , +, + (L: +, + + +4-+, +, + cell. + + +, ++, (L: + (L: + (L:(L: + (L: + (L:

1 fim 1 6574m) 3374m)33//m) 65 2774m)27/im) 2774m)27/im) 74m) 8)-celled.8)-celled. 8)-celled.8)-celled. 8)-celled. 74m) 74m)ftm) ++, 4)-celled. 4)-celled. 4)-celled. 8)-celled.8)-ceUed. 8)-celled. 35/im)5 2874m)28,um) 3574m)35/im) 25/Mm) 3274m)32//m) 3174m)3i,um) (D.(D. (D. (D. 25 3 25 Glob., ii) (D. (D. (D. (D. (D. (D. more-celled.more-celled. (D. (D. Ss.,Ss., 8(mostly 8(mostly 8(mostly 8(mostly 8(mostly 4-celIed.4-celled. (L: 8-celled. 8-celled. 8(mostly8(mostly 8(mostly8(mostly 8(mostly i) 2674m); 4—8(mostly 4—8(mostly 4—8(mostly 4—8(mostly 4—8(mostly 4-celled.4-celIed. 4-celIed.4-celled. 4-celled.4-celIed. 4—8-celled. 4—8-celled. 8-celled.8-celled. 4-celled.4-celled. or 4—8(mostly 4-celled. 4-celled. hairs 8- types: (D. 2-celled. 4— 74m) 4— 4— 51—42—51— 4— 4— 4— 4—4— 4— Glob., Glob.,Glob., Glob., 2—2— Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glob., 28,um) 3274m)32/

DE) SING) MALAY) LUCK) SING) LUCK) MALAY)LUCK) DUN)DUN) DE) DE) CNH) PDA) 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 64553 64554 LUCK)LUCK) 64555 64585 64556 64584 64557 64565 64577 64551 64552 64594 64578 64579 64592 64575 64574 64587 64566 Voucher LWG LWG (ex LWG LWG LWG LWG LWG LWG LWG LWG LWG LWG LWG LWG LWG LWG LWG LWG LWG

Kurz. Andre Engl. var. var. Nees Ktze. Alston ex Nees Nees Radlk. Radlk. O. (Linn.) LindenLinden Domin Br.)Br.) Nees (Forsk.) (Hochst.)(Höchst.) Coemans Coemans Linn.Linn. LindenLinden (Retz.) Ktze.Ktze. nasuta Hort. Bailey acuminata nitidum nasuta Regel (Schrank) Bailey (R. Nees roxburghiana O. pectinata 1 viride speciosa gigantea Regel betonica bicolorbicohr grandiflorumgrandiflomm kewense malacceme variabile bicalyculata linctoria strictum verschaffeltiiverschajfeltii verschaffeltii calycotricha PeristrophePeristrophe RungiaDiclipteraDicliptera Odontonema Rhinacanthus Ruttya Fittonia argyroneura O.O. F. F.F. pearcei JusticiaJusticia P. P. P. P. P. P.P. P. "' J. Species 80. 81. 82. 83. 84. 85. 86. 87.87. 88. 89. 90. 91. 92. 93. 94. 95. 96. 97-97. 98. 99. K.J. Ahmad: Hairs of Acanthaceae 117

/im) /im) /im) conical. 20062006 /im) 362362/(in) /im) /im) 483/im)554/im)554/im) 342//m) 287287 287/im)287/im) 300300 152/im)152/mi) 200/im)200/im) 342 i-celled), 250250 35—239— 52— (L: (L: (L: (mostly (L: (L: (L: (L: (L: fun) (L: hairs (L: celled (L: 5-celled. 5-celled.5-celled. 5-celled. 290290/(111) 4-celled. 4-celled. I—57—57—189—1—5-celled.189— 247—1—5-celled.129—893—306— 1—1— 45—45—1—5-celled.161—1— I—55 1— 1—4-celled.142—3—6-celled.267—142—267— 157—91—157—50—91—I—4-celled.1—2-celled. 1—4-celled.I—4-celled.155—197—239—35—I—4-celled.35—266—266—1—35— 2—4-celled.1—42— 105—105— +,+, +, +, +, +, 4-+, + 92—42—92— +, +.+1 +, + +, Non-glandularNoo-glaodular + + + + +, + + + (L: + + + +, + + +

Uμ™.)34/im) 29/im)/im) /im) 27/im)/im) 30/im) /im) Ls. 29 /im) 31 /im) 27

26/im) 5 +, 25/4m) 27/im) 26/im) (D. 3 (D. 25 (D. 29 (D. (D.

Glob., ii) (D. (D. (D. (D. (D. (D. Ss., 4-celled. 4-celled.4-celled. 4—8-celled. i) 26/iin);26/im); 2—4-celled. 4—8-celled. 4—8-celled. hairs 4-celled. 4-celled. 4-celled. 4-celled. 4-celled. 4-celled. types: (D. /im) 2— 2—2— Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glob., Glob., GlandularGlaodular two 4-celled. 123 Ss., Ss., Ss., Of (L: Ss., Ss., Ss., Ss., Ss., Ss., Ss., Ss.,

LUCK) LUCK) DE) LUCK) LUCK) MALAY)LUCK) MALAY) BSI- BSI- BSI- BSI- (ex (ex (ex (ex (ex (ex (ex (ex specimeospecimen (ex (ex (ex (ex 50 64597 64582 64550 20431 MHi73i9(exBSI-17319 12639 16622 64568 64545 64573 64581 64543 645 Voucher MH South) LWG MH South) MH South) MH South) LWG LWG LWG LWG LWG LWG LWG

Hook./.

& Beoth.Benth. Hook./Hook./.Nees Lino.Linn. & Nees & Nichols. Burm./. Lino.Linn. amherstiae LiodleyLindley vasica candicans Beoth.Benth. amherstiae BraodegeeBrandegee Willd. Roth carneacornea gendarussagendarussaprocumbens tranquebariensis nodosa guttata oblongata diffusa vahlii Adhatoda Dianthera Beloperone JacobiniaJacobinia J. J.J. J. D. B. B. J.

Species ioo.100. IOI.101. 102.102. 103. 104 105. 106. 107. 108. 109. no.110. in.