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Anatomical Adaptations in the Leaves of Selected Fynbos Species

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Anatomical Adaptations in the Leaves of Selected Fynbos Species S.Afr.J.Bot., 1994, 60(2): 99 - 107 99 Anatomical adaptations in the leaves of selected fynbos species Al ison M. van der Merwe (nee Summerfield),· J.J.A. van der Walt and Elizabeth M. Marais Department of Botany, University of Stellenbosch, Stellenbosch, 7600 Republic of South Africa Received 23 August 1993; revised 6 December 1993 Fynbos plants experience very harsh conditions during the hot and dry summer months and their leaves are adapt­ ed to reduce the loss of water due to transpiration. The leaves of 46 selected fynbos species of 24 families were examined to determine which anatomical adaptations contribute to the reduced rate of transpiration and subse­ quent reduced water loss. Without exception, all species examined show leaf adaptations typical of xerophytic species. Four typical leaf types are recognized and proposed as models of leaf adaptation: 1. Myrsine type - dorsi ventral or isobilateral leaves; more palisade parenchyma present than spongy parenchyma; tissues contain large amounts of phenolic substances. 2. Meta/asia type - small dorsiventral leaves with involute margins and a single groove in the adaxial surface; mesophyll is usually inverted. 3. Retzia type - dorsi ventral or isobilateral leaves with revolute margins and one or two grooves in the abaxial surface; spongy parenchyma is the main component of the mesophyll. 4. Spatalla type - small centric or near-centric leaves; little or no spongy parenchy­ ma tissue. Fynbos plante ondervind uiterste toestande tydens die warm, droa somermaande, en hulle blare is aangepas om waterverlies tydens transpirasie te beperk. Blare van geselekteerde fynbos-spesies uit 24 families is ondersoek am die bydrae van die verskillende anatomiese aanpassings tot verminderde transpirasietempo en gevolglike water­ verlies, vas te stel. AI die spesies wat ondersoek is, vertoon sonder uitsondering, tipiese xerofitiese blaaraanpas­ sings. Vier verskillende blaartipes word beskryf en voorgestel as modelle vir blaaraanpassings: 1. Myrsine-tipe - dorsiventrale- of isobilaterale blare; meer palisadeparenchiem as sponsparenchiem is aanwesig; weefsels bevat groot hoeveelhede fenoliese verbindings. 2. Metalasia-tipe - klein dorsiventrale blare met blaarrande involuut en 'n enkele adaksiale groef; mesofil meestal omgekeerd. 3. Retzia-tipe - dorsiventrale- of isobilaterale blare met blaarrande revoluut en een of twee abaksiale groewe; mesofil bestaan hoofsaaklik uit sponsparenchiem. 4. Spatal­ la-tipe - klein sentriese of byna sentriese blare; min of geen sponsparenchiem . Keywords: Anatomical adaptations, fynbos, leaf types. * To whom correspondence should be addressed Introduction Material and Methods Fynbos vegetation is unique to the Cape Floristic region and is Twenty-four families occurring in the south-western Cape, readily recognized by the sclerophyllous to microphyllous including all shrubby endemic families, were chosen as represen­ nature of almost all the woody taxa (Kruger 1979). tative of fynbos vegetation. From these families 46 perennial The fynbos biome extends from the Nieuwoudtville escarp­ species were randomly selected and collected in their natural ment to the Gifberg Massif and the Nardouw-Pakhuis-Cedar­ habitat (one species from four localities, two from three, 23 from berg mountains (32°S to 34°S). The inland border runs along two and 20 from one locality, Table 1). The additional species collected from single localities were also included in the study as the base of the sandstone mountain belt of the Witteberg, ' they were valuable for comparisons. Voucher specimens of each Swartberg, Baviaanskloof and Great Winterhoek mountains. species are kept at the Stellenbosch University Herbarium These ranges form an almost continuous chain running west to (STEU). At each locality, ten mature sun leaves were collected east (l8°E to 24°E) and terminating at the Indian Ocean near from more than one plant of each species. Leaf material was fixed Port Elizabeth (Bond & Goldblatt 1984). in FAA, dehydrated and infiltrated and finally embedded in The climate of the fynbos region is mostly mediterranean or Paraplast, following the tertiary Buthanol method. Transverse semi-mediterranean with a rainfall range of 200 - 3000 mm sections of 5 - 10 fl-m were made through the mid-section of the per annum. The mean annual temperatures throughout the laminae, using a rotary microtome and stained with Alcian Green region are close to 17°C except in the immediate vicinity of Safranin (Joel 1983). the coast where the mean annual temperature range is less than 8°C. Fynbos usually occurs on coarse-grained, infertile, acidic, Results and Discussion sandy soils derived from quartzites and sandstone of the Cape Thick cuticle, waxes and trichomes Supergroup (Cowling & Holmes 1992). In the leaves of all the species examined at least one, if not all, Although fynbos is such a diverse and interesting vegetation, of the above protective coverings were present. very little is known about the anatomy of the leaves of fynbos Although cuticle thickness is not considered to be a very species. The aim of this investigation was to determine which reliable character, in most of the leaves examined the cuticles anatomical adaptations contribute to the reduced rate of trans­ were relatively thick, when compared with the cuticle of a piration and subsequent reduced water loss which enables normal mesophytic leaf. Where a thick cuticle is not present, fynbos plants to survive the hot, dry summer months. Leaves the outer periclinal walls of the epidermal cells are extremely were selected for study as they are the most exposed to aerial thick, as in Roella incurva var. incurva, Aspalarhus cephalores environmental conditions and therefore serve as good indica­ subsp. violacea, Penaea mucronara, Salrera sarcocolla, tors of any anatomical and morphological changes in the plant Thesium ericaefolium, Freylinia lanceolaraand Campylo­ (Mauseth 1988). srachys cernua. A thick cuticle, together with waxes, prevents 100 S .-Afr .Tydslcr .Plantk., 1994, 60(2) Table 1 List of selected species with collector, collection number and locality Family/Species Collector Locality Aizoaceae Galenia africana L. Summerfield 53 Clanwilliam Summerfield 87 Porterville Asteraceae Brachylaena neriifolia (L.)R.Br. Summerfield 28 Eersterivier Summerfield 79 Jonkershoek, Stellenbosch Eriocephalus africanus L. Summerfield 25 Eersterivier Summerfield 45 Clanwilliam Helichrysum patulum (L.) D.Don Summerfield 50 Clanwilliam Summerfield 61 Jonkershoek, Stellenbosch Heterolepis aliena (L.F.) Druce Summerfield 10 Kogelberg, Betty's Bay Summerfield 86 Gydouw Metalasia densa (Lam.) Karis Summerfield 09 Jan Marais, Stellen bosch Summerfield 23 Kogelberg, Betty's Bay Summerfield 29 Hermanus Metalasia dregeana DC. Summerfield 52 Clanwilliam Metalasia seriphiifolia DC. Summerfield 30 Hermanus Bruniaceae Berzelia lanuginosa (L.) Brongn. Summerfield 63 Jonkershoek, Stellenbosch Brunia albif10ra E. Phillips Summerfield 12 Kogelberg, Betty's Bay Brunia alopecuroides Thunb. Summerfield 18 Kogelberg, Betty's Bay Campanulaceae Roella incurva DC. var. incurva Summerfield 16 Kogelberg, Betty'S Bay Summerfield 82 Jonkershoek, Stellenbosch Ericaceae Erica longifolia Ail. Summerfield 22 Kogelberg, Betty'S Bay Erica plukenetti L. Summerfield 62 Jonkershoek, Stellenbosch Coetzenberg, Stellenbosch Fabaceae Aspalathus cephalotes Thunb. subsp. violacea Dahl gr. Summerfield 57 Jonkershoek, Stellenbosch Summerfield 74 Coetzenberg, Stellenbosch Grubbiaceae Grubbia tomentosa (Thunb.) Harms Summerfield 06 Kogelberg, Betty's Bay Summerfield 31 Hermanus Lobeliaceae Lobelia pinifolia L. Summerfield 32 Hermanus Myricaceae Myrica cordifolia L. Van der Walt 1602 Y zerfontein Myrsinaceae Myrsine africana L. Summerfield 07 Jan Marais, Stellenbosch Summerfield 26 Eersterivier Summerfield 37 Hermanus Summerfield 43 Clanwilliam Myrtaceae Metrosideros angustifolia (L.) Smith Summerfield 27 Eersterivier Summerfield 67 Jonkershoek, Stellenbosch Penaeaceae Penaea mucronata L. Summerfield 14 Kogelberg, Betty'S Bay Summerfield 59 Jonkershoek, Stellenbosch Saltera sarcocolla (L.) Bullock Summerfield 33 Hermanus Polygalaceae Muraltia heisteria (L.) DC. Summerfield 38 Hermanus Summerfield 56 Jonkershoek, Stellenbosch Proteaceae Diastellafraterna Rourke Summerfield 15 Kogelberg, Betty's Bay Protea repens (L.) L. Summerfield 08 Jan Marais, Stellenbosch Summerfield 72 Coetzenberg, Stell en bosch Serruria kraussii Meisn. Summerfield 68 Jonkershoek, Stellenbosch Summerfield 92 Porterville Spatalla mollis R. Bf. Summerfield 17 Kogelberg, Betty's Bay S.Afr.J.Bot., 1994, 60(2) 101 Table 1 Continued Family/Species Collector Locality Retziaceae Relzia capensis Thunb. Summerfield 11 Kogelberg, Betty's Bay Rhamnaceae Phylica buxifolia L. Summerfield 34 Hermanus Summerfield 90 Citrusdal Phylica cryptandroides Sond. Summerfield 46 Oanwilliam Phylica lasiocarpa Sond. Summerfield 21 Kogelberg, Betty' s Bay Phylica spicala L.f. Summerfield 55 Jonkershoek, Stellenbosch Summerfield 91 Citrusdal Phylica slipularis L. Summerfield 58 Jonkershoek, Stellenbosch Summerfield 89 Gydouw Roridulaceae Roridula gorgonias Planch. Summerfield 13 Kogelberg, Betty's Bay Rosaceae Clifforlia ruscifolia L. Summerfield 02 Jan Marais, Stellenbosch Summerfield 44 Oanwilliam Summerfield 66 Jonkershoek, Stellenbosch Rutaceae Agalhosma bifida (Jacq.) Bartl. & Wendl. Summerfield 41 Hermanus Diosma hirsula L. Summerfield 20 Kogelberg, Betty's Bay Summerfield 95 Jonkershoek, Stellenbosch Diosma subulata Wendl. Summerfield 42 Hermanus Sapindaceae
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