Proc. lndian Acad. Sci. (Plant Sci.), Vol. 97. No. 5, October 1987, pp. 385-389. Printed in India.

Morpho-taxonomy of Anthoceros crispulus (Mont.) Douin from south India*

S C SRIVASTAVA and A K ASTHANA Department of Botany, University of Lucknow, Lucknow 226 007, India MS received 3 July 1986; revised 5 January 1987 A~traet. Anthoceros crispulus (Mont.) Douin, with ah extended distribution from plains of UP to montane areas in the Palni and Nilgid hills in peninsular India, is characterized by radially lamellate (erispate) dorsal surface and lobed rosettes of thalli with stomata 44 x 33- 56 x 44/~m, and inner lining layer cells of capsule wall with irregular thickening bands. Scanning eleetron microscopie studies of the spore provide additional taxonomic character as both proximal and distal lates are distinct in the sporoderm arehitecture. Keywords. Bryophyta; Anthocerotales; Anthocerotaceae; Anthoceros.

1. lntroduction

Anthoceros crispulus (Mont.) Douin is a widely distributed species, though restricted in some regions of the world (Bharadwaj 1950). It is known from Europe: Gallia (Stephani 1912), Great B¡ (Macvicar 1926) and USA (Evans 1924; Schuster 1953). In India, the species was first reported by Pande and Ahmad (1944) from the neighbourhood of Lucknow where it was very common (near Chinhat). However, in recent years due to fast increasing urbanization and biotic disturbances in the microhabitat, this species now appears to have depleted from its original sites. Repeated collections from different localities in the Himalaya also did not reveal the occurrence of this taxon in these territories. In a recent trip to south India (Tamil Nadu) some fertile plants of this taxon growing on soil covered rocks forming small but distinctive patches (rosettes) on hill slopes in Palni (Kodaikanal) as well as in Nilgiri hills (Ootacamund), were collected. The discovery of A. crispulus from south India extends the range of distribution of this taxon from Indo-gangetic plains of UP to mountainous ranges in peninsular India. Detailed investigations revealed some interesting details, not provided earlier for Indian plants. Earlier contributions on A. crispulus include the details of thallus structure, development of sex organs and the sporophyte based on local plants from Lucknow (Bharadwaj 1950). Of the 4 evolutionary tendencies recognized by Bharadwaj (1960) in the genus Anthoceros on the basis of sporoderm characteristics, A. crispulus falls under A. punctatus group (with A. husnoti and A. neesii) which is characterised by lamellate dorsal surface of thallus, monoecious sexuality (antheridia not seen in our plants) and brown to dark reticulate spores with reticulations in contact with triradiate rays on the proximal face.

* Contribution from Botany Department: New Sedes Bryophyta No 221. 385 386 S C Srivastava and A K Asthana

2. Taxonomic description

Plants monoecious (?), thalli spongy, radially oriented in form of rosette, up to 1.5 cm in diameter, dorsal surface covered with radially oriented lamellate outgrowths; involucres narrow and long up to 5 mm; epidermal layer of capsule wall stomati- ferous, stomata 44 x 33-56 x 44 #m with 5-11 stomata/mm2; inner lining layer cells of capsule wall with occasional irregular thickening bands; spores dark-brown to black, 42--44 #m in diameter with distinct proximal and distal faces, proximal face with reticulate sculpturing (pitted in appearance) in contact with triradiate rays, distal face prominently spinulate (pits not clearly seen as compared to proximal face), spines acute to subacute with adhering tendency; elaters light-brown, up to 4-celled, 110-150 #m long, some times branched.

2.1 Specimens examined

LWU 7167-E/83, 7317/83, 7453/83, 7460/83, 7497/83, 7498/83, 7544/83, 7704/83, Loo.: Kodaikanal, Oct., 1983, Leg.: R Udar and party, Det.: RU, SCS and AKA; LWU 7554/83, 7568/83, Loc.: Shembaganur, Oct. 1983, Leg.: R Udar and party, Det.: RU, SCS and AKA; LWU 7167-A/83, Loo.: Naduvattam; LWU 7167-C/83, Loe.: Emerald; LWU 6941/83, 7014/83, 7069/83, 7167-B/83, 7167-(3/83, Loo.: Ootacamund, Oct. 1983, Leg.: R Udar and party, Det; RU, SCS and AKA. Habitat: Terricolous.

3, Results and discussion

A. cri~pulus was synonymized under A. punctatus L. by Proskauer (1948). According to hito A. punctatus and A. crispulus cannot be sharply demarcated from each other as they resemble in almost all the features except the densely c¡ (lamellate) dorsal surface of thallus in the latter. He gave no importance to this character as he considered it to be ah ecological va¡ He, however, suggested 8 different conditions which contribute to this c¡ appearance, of which one is the deve- lopment of lamellate outgrowths (see Bharadwaj 1950). Meijer (1950) provided ah independent status to A. crispulus, for the plants of Java having crispate dorsal surface of thallus. Bharadwaj (1950) reinstated A. crispulus as distinct species on the basis of lamellate (crispate) nature of thallus surface and slightly larger spores and elaters. He also emphasized the divisions of the zygote by transverse wall as an important feature of this taxon. Apart from typical lamellate thallus (figure 1), 2-4 celled, often branched elaters (figures 8-11) and stomata (figure 2) ranging from 44x33-56x44/am, our investigations on south Indian plants, revealed some hitherto unknown features like stomatal frequency (5-11/mm 2) and occasional presence of dark thickening bands (figure 3) on the tangential waU of the cells of the inner lining layer of capsule wall. Of particular significance is the sporoderm architecture under LM (figures 4-7) and SEM (figures 12-15). The spores under SEM study revealed a basically reticulate sporoderm architec- ture which is rather pitted on the proximal faces (figures 13, 15). Fimb¡ bordered triradiate mark (laesura) is in contact with sculptured surface (figures 12, Morpho-taxonomy of A. crispulus from South India 387

4 ._. 6

5

i 2mm 2--3 ,o$mm 4--11 905 mm

Figures 1-11. Anthoceros crtspulus (Mont.) Douin. 1. Female thallus showing lamellate surface. 2. Epidermal layer of capsule wall. 3. Inner lining layer of eapsule wall showing band like thickenings. 4 and 5, Spores (proximal face). 6 and 7. Spores (distal late). 8-11, Elaters.

13, 15). Muri or ridges enclosing the pits (or lumina) are more or less devoid of spines (figure 15). Distal face is densely spinulate with spines adhering at base (figures 12, 13). The dense impregnation of spines together with their adherant bases make the pits inconspicuous (figure 14). Spines are acute to subacute and even blunt at some places with rounded" apices (figure 14), In the light of investigations made by Bharadwaj (1950, 1960) together with our observations on stomata, inner lining layer of capsule wall and SEM study of spores, A. crispulus maintains an independent status forming a natural group with A. punctatus, A. husnoti and A. neesii as already reeognized by Bharadwaj (1960). 388 S C Srivastava and A K Asthana

Figures 12-15. Anthoceros crispulus (Mont.) Douin. 12 anfl 13. Spores (distal and proximal facesx 307, x 660, respectively). 14. Spore (distal facex 843). 15. Spore (proximal face x 884).

Acknowledgements

The authors are indebted to the late Prof. Ram Udar, for the valuable suggestions regarding the identity of A. crispulus. Thanks are also due to the authorities of the Indian Institute of Technology, Kanpur for providing SEM facilities and to the University Grants Commission, New Delhi. for financial assistance. Morpho-taxonomy of A. crispulus from South India 389

References

Bharadwaj D C 1950 Studies in lndian Anthocerotaceae I~The morphology of Anthoceros crispulus (Mont.) Douin; J. Indian Bot. Soc. 29 145-163 Bharadwaj D C 1960 Studies in Indian Anthocerotaceae III~The morphology of Anthoceros erectus Kash. and some other species; J. Indian Bot. Soc. 39 568-592 *Evans A W 1924 Hepaticae of Nebraska; Bryologist 27 52 Macvicar S M 1926 Students Handbook ofBritish Hepatics (London: V V Sumfield) Meijer W 1950 Notes on some Malayan species of Anthoceros L. (Hepaticae)-II; J. Hattori Bot. Lab. 18 1-13 *Pande S K and Ahmad S 1944 Liverworts of Lucknow and its neighbourhood; Proc. Indian Sci. Cong. Delhi 3 80 Proskauer J 1948 Studies on the morphology of Anthoceros L.; Ann. Bot. New Ser. 12 237-265 Schuster R M 1953 Boreal Hepaticae: manual of the liverworts of Minnesota and adjacent regions; Aro. Midl. Nat. 49 291-297 Stephani F 1912 Species Hepaticarum V (Geneve: Georg & C ie)

*Not seen in original.