FOLIAR ANATOMY OF IN ARASBARAN, NW.IRAN F. Zarrinkamar, A. Jalili, B. Hamzehee, Y. Asri, J. G. Hodgson, K. Thompson, S. Shaw Zarrinkamar, F., Jalili, A., Hamzeh'ee, B., Asri, Y. Hodgson, 1. G., Thompson, K. & Shaw, S. 2002 12 30: Foliar anatomy of some Carex species in Arasbaran, NW. Iran. -Iran. Journ. Bot. 9(2): 261-270. Tehran.

The foliar anatomy of genus Carex () from the Arasbaran protected area was studied under the light microscope. The following species have been analysed: C. distans, C. divisa, C. divulsa, C. halleriana, C. liparocarpos, C. nigra, C. pallescens.

Fatemeh Zarrinkamar, A del Jalili, Behnam Hamzeh'ee, Yunos Asri, Research Institute of Forests and Rangelands, P. 0. Box 13185-116. Tehran, Iran. - John G. Hodgson, Ken Thompson and Sue Shaw, Department of Animal and Sciences, the University, Sheffield S10 2 TN, UK.

Key word. Anatomy, Carex, Cyperaceae, Arasbaran, Iran.

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INTRODUCTION selected for analysis, and at each site, the data The Cyperaceae family are grass-like, recorded is an average of 15 samples of 3 herbaceous comprising about 70 genera plants from the same species. and 4000 species, they are commonly found in The species under anatomical study are as inundated or boggy conditions. Fox (2000) follows. defines this group of the as Carex distans L. -Arasbaran, Serahie Vayghan, being of particular significance as 1300 m, N, 10%, Harnzeh ee & Asri. environmental indicators for wetland habitats. Carex divisa Hudson. -Arasbaran, Toopkhaneh Cyperaceae leaves are long, alternate, and highland, 2250 m, N, 30% Harnzeh ee & Asri. mostly arise near the base, usually within a Carex divulsa Stokes. -Arasbaran, between closed sheathing base, parallel veined, and Abbasabad & Mahmoodabad highland, 2150 strap-shaped. The flowers of this family are m, SW, 50%, Hamzehee & Asri. bisexual or unisexual, and very minute (Judd Carex halleriana Asso. Arasbaran, Serahie & al. 2000).They reproduce by rhizomes and Vayghan, 1300 m, N, 10%, Harnzehee & seeds (Fox 2000). Asri. Carex is one of the major genera contained Carex liparocarpos Gaudin. -Arasbaran, within the Cyperaceae family; about 45 between Mahmoodabad & Makidi, 2150m, S, species of Carex are recorded in Iran, growing 60-70%, Harnzeh' ee & Asri. in diverse climatic conditions, but mostly in Carex nigra L. -Arasbaran, Abbasabad wetland or river basins. highland, 2426 m, N, 60%, Harnzehee & Asri. This work is part of a series of anatomical Carex pallescens L. -Arasbaran, Abbasabad studies of the main flora in the Arasbaran highland, 2426 m, N, 60%, Hamzehee & Asri. protected area. This area, in the northwest part Herbarium specimens are housed at the ofIran, latitude 38° 41 '-39° 7' N and longitude Research Institute of Forests and Rangelands, 46° 42'-46° 58' E has been studied previously and fixed materials are stored at the Laboratory (Assadi, 1987; Zarrinkamar, 200 I). Altitude of Vegetal Anatomy at this institute varies from 350 to 2840 m above sea level, mean annual temperatures are from 5 tol4° C, OBSERVATIONS and annual precipitation is between 316 to 686 Carex distans L. mm (Thompson & al. 2001). The following Leaf surface. Epidermis: short epidermal cells study presents the foliar anatomy of the genus with thin and sinuous walls on both surfaces Carex in this area. (Fig. 1, E, F). Stomata: absent on adaxial surface; paracytic subsidiary cells mostly MATERIAL AND METHODS dome-shaped sometimes triangular on abaxial Material was fixed in FAA, and sectioned with surface. The leaf had an average stomatal a sliding microtome. Sections were cleared density of 120 per rnnr' on the lower surface. with sodium hypochlorite, dehydrated, T. s. Lamina. Outline: V-shaped. Epidermis: coloured with methyl green and carmine-vest, adaxial larger than abaxial cells (Fig. 2, A). and mounted in gelatine. In order to study Bulliform cells: regular groups present above stomatal density, the diafanizaion technique midrib (Fig. 2, B). Stomata: superficial (Fig. 2, was employed (Stritmater, 1973), and the D). Mesophyll: chlorenchyma not radial, results observed using a light microscope. To though tending to radial around the vascular minimise misinterpretation, the middle of the bundles (Fig. 2, C, D). Air-cavities: well leaf lamina in relatively mature leaves was developed (Fig. 2, C). Vascular bundles: about IRAN. JOURN. BOT. 9 (2), 2002 Carex anatomy 263

double, in small vascular bundles; sometimes T S. Lamina. Outline: V-shaped, Epidermis: outer bundle sheaths interrupted abaxially, adaxial larger than abaxial cells (Fig. 3, B). larger vascular bundles incomplete adaxially Bulliform cells: well developed in midrib (Fig. and abaxially. Sclerenchyma: mostly small 3, C). Stomata: superficial (Fig. 3, D). vascular bundles not accompanied by Mesophyl/: chlorenchyma not radial, though sclerenchyma, but sometimes with abaxial tending to radial around the vascular bundles strands, large vascular bundles accompanied (Fig.3, D). Air-cavities: well developed in by girder adaxially and abaxially mesophyll (Fig. 3, B). Vascular bundles: about 7-8 in each half of lamina. Bundle-sheaths: Carex divisa Hudson double, outer bundle sheaths incomplete in Leaf surface. Epidermis: Both abaxial and larger vascular bundles (Fig. 3, D). adaxial surfaces comprise short cells, with thin Sclerenchyma: mostly small vascular bundles and sinuous walls (Fig. 1, G, and H). Prickle- without sclerenchyma, but sometimes hairs: at leaf margin (Fig. 1, G). Stomata only accompanied with adaxial strands; large on abaxial surface: with dome-shaped, vascu lar bundles with girders adaxially and sometimes triangular subsidiary cells, (Fig. 1, abaxially. H). The leaf had an average stomatal density of 126 per mm/ on the lower surface. Carex halleriana Asso. Leaf surface. Epidermis: short form cells, with T S. Lamina. Outline: V-shaped, Epidermis: thin and sinuous walls on both surfaces. adaxial larger than abaxial cells (Fig. 4, E). Prickle-hairs: at leaf margin. Silica-bodies: Bulliform cells: present in midrib (Fig. 4, F). conical, over the veins. Stomata: present only Stomata: superficial. Mesophyll: on abaxial surface, with dome-shaped, chlorenchyma not radial, sometimes sometimes triangular subsidiary cells. The leaf surrounding vascular bundles. Air-cavities: had an average stomatal density of 136 per generally well developed, decreasing in size mm2 on the lower surface. toward leaf margins and midrib (Fig. 4, E). T S. lamina. Outline: V-shaped. Epidermis: Vascular bundles: about 5-6 in each half of adaxial larger than abaxial cells (Fig. 4, A). lamina. Bundle sheaths: double, outer bundle Bul/iform cells: regular groups present above sheaths incomplete adaxially and abaxially, midrib (Fig. 4, B). Stomata: superficial (Fig. 4, and consisting of large vascular bundles. B). Mesophyl/: chlorenchyma not radial, well Sclerenchyma: small vascular bundles without developed around air-cavities and in leaf sclerenchyma, but large vascular bundles margins (Fig. 4, A). Air-cavities: rectangular, accompanied with girders adaxially and many containing remnants of thin-walled abaxially. translucent cells. Vascular bundles: about 7 in each half of lamina. Bundle-sheaths: outer Carex divulsa Stokes bundle sheaths interrupted adaxially and Leaf surface. Epidermis: short- cells, with thin abaxially, sometimes with extensions and sinuous outline walls on both adaxial and surrounding adaxial and abaxial sclerenchyma. abaxial surfaces (Fig. 1, A, B). Stomata: absent Sclerenchyma: central vascular bundles with on adaxial surface, but with parallel-sided wide T-shaped abaxial girders and a minute subsidiary cells on abaxial surface. The leaf adaxial girder, but other vascular bundles with had an average stomatal density of 143 per adaxial and abaxial girders. rnrrr' on the lower surface. 264 Zarrinkamar & al. IRAN. JOURN. BOT. 9 (2), 2002

Carex liparocarpos Gaudin bundle sheaths incomplete on either side. Leaf surface. Epidermis: short and rectangular Sclerenchyma: most vascular bundles with form with thin and sinuous walls on both wide girders at xylem and phloem poles. surfaces (Fig. 1, C, D). Prickle-hairs: angular prickle at leaf margin. Stomata: absent on Carex pallescens L adaxial surface, abaxial dome-shaped, cells Leaf surface. Epidermis: short and rectangular with variable triangular subsidiary cells. The form with thin and sinuous outline on both leaf average stomatal density is 152 per mm/ surfaces. Macro-hairs: short, rigid with on the lower surface. swollen and superficial bases between the T S. lamina. Epidermis: adaxial larger than veins only on abaxial surface. Prickle-hairs: at abaxial cells but not especially inflated (Fig. 3, leaf margin. Stomata: absent from adaxial A). Bulliform cells: regular groups only above epidermis, but present on abaxial, with midrib (Fig. 3, A). Stomata: superficial. triangular subsidiary cells. The leaf had an Mesophyll: chlorenchyma not radial (Fig. 3, average stomatal density of 149 per mm" on F). Air-cavities: variable in shape, many the lower surface. containing remnant of thin-walled translucent T S. lamina. Outline: V-shaped. Epidermis: cells (Fig. 3, E). Vascular bundles: about 5-6 adaxial larger than abaxial cells. Bulliform small and large vascular bundles in each half cells: only in midrib (Fig. 4, D). Stomata: of lamina. Bundle-sheaths: double, outer superficial. Mesophyll: chlorenchyma not bundle sheaths incomplete abaxially in smaller radial (Fig. 4, C). Air-cavities: well developed, vascular bundles and in larger vascular bundles mostly rectangular and variable in size (Fig. 4, interrupted adaxially and abaxially. C). Vascular bundles: about 9 vascular bundles Sclerenchyma: vascular bundles with abaxial in each half of lamina. Bundle-sheaths: outer girders near margins, and with adaxial and bundle-sheaths incomplete. Sclerenchyma: abaxial girders elsewhere. most vascular bundles with T-shaped girders at xylem poles, but in midrib with abaxial Carex nigra L. girders. Leaf surface. Epidermis: short and rectangular form with thin and slightly sinuous walls on DISCUSSION both surfaces. Prickle-hairs: angular prickle at As described in the observations, several leaf margin. Stomata: absent on adaxial common characteristics were observed epidermis, but on abaxial with triangular between the species studied. Here, some subsidiary cells. The leaf average stomatal common characters are highlighted. Study of density is 179 per mnr' on the lower surface. leaf anatomy demonstrates the presence of T S. lamina. Outline: V-shaped. Epidermis: stomata on abaxial surfaces only; stomata are adaxial larger than abaxial cells. Bulliform paracytic, with mostly dome-shaped, cells: well developed in midrib (Fig. 2, E). subsidiary cells sometimes triangular but Stomata: superficial, except above adaxial parallel sided in Cidivulsa. sclerenchyma and near leaf margins. The cuticles are of intermediate thickness Mesophyll: chlorenchyma not radial. A ir- (about 4-7.511); adaxial surface is often thicker cavities: variable in size and shape. Vascular than abaxial surface (Table 1). bundles: about 8 vascular bundles in each half of lamina. Bundle-sheaths: double, outer Table 1. Anatomical characters of Carex species. IRAN. JOURN. BOT. 9 (2), 2002 Carex anatomy 265

Superficial view T. S.

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Carex distans - 120 - 39.3 4 5 not radial + In midrib 2 Carex divisa - 126.33 - 33.05 4 4 not radial + In midrib 2 Carex divulsa - 142.76 - 35.88 7 7 not radial + In midrib 2 Carex halleriana - 136.26 - 33.63 7.5 5 not radial + In midrib 2 Carex Iiparoca1J2os - 152.07 - 32.86 6 5 not radial + In midrib 2 Carex nigra - 179.3 - 27.06 5 3 not radial + In midrib 2 Carex pallescens - 148.72 . 32 5 3 not radial + In midrib 2

In epidermis cells bubble shaped bulliform make confused their mesophyll arrangement; cells are found on the midrib. An important therefore, physiological study is needed to adaptation for many of the Cyperaceae recognize photosynthesis pathway. including Carex is the appearance of bull iform The mesophyll of these species contains air- cells as part of the epidermis allowing the cavities of thin-walled, translucent cells control of leaf rolling and unrolling at the without chloroplasts. This character makes the cellular level, by turgor pressure. When water internal atmosphere of the leaves more availability is limited, the bulliform cells lose efficient for gas exchange during water and become flaccid. Thereby the leaves photosynthesis. are less exposed to sunlight and transpiration is From an ecological view, the existence of reduced. these air-cavities in Carex leaves enables it to Most of the Cyperaceae util ise the C4 adapt to their variable habitats in wetland or photosynthetic pathway. According to Mauseth marginal aquatic conditions. (1988) the principal characteristic of Kranz anatomy is the radial arrangement of ACKNOWLEDGMENT mesophyll around the bundle sheath; however This project supported by Research Institute of a double bundle sheath is present in all Carex Forests and Rangelands (RIFR) and Darwin species and in the most of species radial Initiative for survival of species (DETR. mesophyll are not observed. The air-cavities London). We would like to express our 266 Zarrinkamar & at. IRAN. JOURN. BOT. 9 (2), 2002 appreciation to Azimi, Dehghan, Boostani, Mauseth, 1. D. 1988. Plant Anatomy. Kowsari and Siavash, laboratory technicians of -University of Texas, Austin. plant anatomy. Strittmater, C. G. R. 1973: Noeva tecnica de diafanization. -Bol. Soc- Arg. Bot, 15 (l): REFERENCES 126-129 Assadi, M. 1987. Plants of Arasbaran Thompson. K., lalili, A. Hodgson. J., Harnzehee, 8., Asri, Y., Shaw, S., Protected area NW of Iran. - Iran. Journ. Bot. 3 (2):131-175. Shirvany, A., Yazdany, Sh., Khoshnevis, Fox, R. 2000: An introduction to the anatomy M., Zarrinkamar, F., Ghahramany, M., of the Graminoid, Monocots. -Lander Safavi, R. 2001: Seed size, shape and University, South Carolina. persistence in the soil in an Iranian flora. Judd, W. S. Campbell, E. A. Kellogg, P. S. Seed Science Research: II: 345-355 Stevens. 2000: Families. (2001). -University of Hawaii, Botany Department. Zarrinkamar, F. 2001; foliar anatomy of the caryophyllaceae family in Arasbaran, NW. Iran. Iran, Journ. Bot. 9(1): 93-102. IRAN. JOURN. BOT. 9 (2), 2002 Carex anatomy 267

Fig. 1. Ovservations of abaxial epidermis in superficial view of the Cyperaceae. A, B. Carex divulsa. C, D. C. liparocarpos. E, F. C. distans. G, H. C. divisa. A, C, E, G (x 72); B, D, F, H (x 144). 268 Zarrinkamar & al, IRAN. JOURN. BOT. 9 (2), 2002

Fig. 2. Leaf of Carcx spp. in T. S. A-D. C distans; A. general aspect; B. midrib; C, D. detail of mesophyll. E. midrib ofC nigra. A (x 72); B, C, E (x 144); D (x 288). IRAN. JOURN. BOT. 9 (2), 2002 Carex anatomy 269

Fig. 3. Leaf of Carex spp. in T. S. A, E, F. C. liparocarpos; A. general aspect; E. midrib; F. detail of mesophyll. B-O. C. divulsa; B. general aspect; C. midrib; O. detail of mesophyll. A, B (x 29); C (x 72); O-F (x 144). 270 Zarrinkamar & al. IRAN. JOURN. BOT. 9 (2), 2002

Fig. 4. Leaf of Carex spp. in T. S. A, B. general aspect of C. helleriana. C, D. C. pallescens; C. detail of rnesophyll ; D. midrib. E, F. C. divisa ; E. detail of mesophyll ; F. midrib. A (x 29); B (> 72); C-F (x 144).