Dfd79afeec92bb782b4474bb2e8

Dfd79afeec92bb782b4474bb2e8

Korean J. Pl. Taxon. pISSN 1225-8318 eISSN 2466-1546 46(2): 199−212 (2016) Korean Journal of http://dx.doi.org/10.11110/kjpt.2016.46.2.199 Plant Taxonomy Taxonomic significance of the leaf micromorphology in the tribe Sorbarieae (Spiraeoideae: Rosaceae) Jun-Ho Song and Suk-Pyo Hong* Laboratory of Plant Systematics, Department of Biology, Kyung Hee University, 26, Kyungheedae-ro, Dongdaemun-gu, Seoul, 02447, Korea (Received 17 March 2016; Revised 28 April 2016; Accepted 2 June 2016) áƴ\μ&λྐྵ\)7+,˔ů7/,ʻᄡሞ ůͬጏലሽ̿,ጏԽʀ,̫ɢሽ̿,ɀƹͥ ɺȉΉԂΞǶƭ( ǻংnݯʎ ŚȺnݯ ʓÚݯȺ ABSTRACT: A comparative study of leaf epidermal microstructures in the tribe Sorbarieae (Adenostoma: 3 spp., Chamaebatiaria: 1 sp., Sorbaria: 11 spp., Spiraeanthus: 1 sp.) including related genera Gillenia (2 spp.) and Lyonothamnus (2 spp.) was carried out using scanning electron microscopy (SEM) in order to evaluate their significance in taxonomy. The leaves of Adenostoma, Chamaebatiaria, and Spiraeanthus were amphistomatic, whereas Gillenia, Lyonothamnus, and Sorbaria were hypostomatic. The size range of the guard cells is 7.84−48.7 × 5.86−38.6 µm; the smallest one was found in Sorbaria tomentosa var. tomentosa (7.84−11.8 × 6.84−10.5 µm), while the largest measured example was Adenostoma fasciculatum var. obtusifolium (30.3−48.7 × 18.8−38.6 µm). Anomocytic stomata complex were the most frequent type (rarely cyclocytic), with usually both anomocytic and actinocytic types occurring in one leaf. On the surfaces, both the adaxial and abaxial anticlinal walls of the subsidiary cells vary (e.g., straight/curved, undulate, sinuate). Four types (unicellular non-glandular trichome, stellate, glandular trichome, pustular glandular trichome) of trichomes are found in the leaves. The epicuticular wax can be divided two types: membraneous platelets (Lyonothamnus) and platelets (Sorbaria arborea var. arborea, S. arborea var. subtomentosa, S. kirilowii, S. tomentosa var. tomentosa, Spiraeanthus schrenkianus). The trichome diversity (in particular, stellate, gland) and the existence of epicuticular wax may have taxonomic significance, although the leaf epidermal micromorphological characteristics do not provide synapomorphy in this tribe. These leaf micromorphological features are most likely better understood in the Sorbarieae when used in conjunction with external morphological characters. Keywords: leaf epidermal cell, stomata complex, stellate, glandular trichome, epicuticular wax ੘ࢼ: :ʳǮÒɗ(Adenostoma: 3 spp., Chamaebatiaria: 1 sp., Sorbaria: 11 spp., Spiraeanthus: 1 sp.)Ⱥ ξȾ ̢ ݒ ƖˮʪɖΞ nݲٲ ݾݲ ̛ 20̢˞ʮľ Ŵ؂ ʮŞ Gillenia̓ 2̢˞ʮ, Lyonothamnus̓ 2̢˞ʮĪ˞̢ ,ݯǧ Ć˧Ī ƾζݮȞŶ ˢɢǪŶࠚƖǻ(SEM)Ī Śݮκ ȾȮݮȞ ͮƒݮώR. Adenostoma̓˞ Chamaebatiaria̓, Spiraeanthus̓Ξ˒ ͮȳʓݿʲ(stomatal complex) ޻ΓȺ ǎΓ FƾΞ ͮȳŚ ɚƒ ݮ ͮ͟ȳϋ(amphistomatic type)ŚΖȞ, Gillenia̓, Lyonothamnus̓Ⱥ Sorbaria̓Ξ˒ ǎΓ(abaxial *Author for correspondence: [email protected] http://e-kjpt.org, © 2016 the Korean Society of Plant Taxonomists. This is an open-access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/4.0) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited. !"" !"" !"#$%&'(&#)'*#+'(",$-.&'%&#) ľ ʂͮ 7.84−48.7 × 5.86−؂side)Ξ͒ ɚƒݮ Śͮȳϋ(hypostomatic type)ĢǢ Ǯ˖ȂR. ȳɞˮ 38.6 µmĢǢ ̓Ⱥ ɫ͎R R͂ ȦŚ̂ ʒŚÞ, Sorbaria tomentosa var. tomentosa (7.84−11.8 × 6.84− Ǣ Ǯ˖ȂȞ, Adenostoma fasciculatum var. obtusifolium (30.3−48.7 × 18.8−؂µm)ľ ľƋ ŷĦ ȳɞˮ 10.5 µm)Ξ˒ ľƋ ʂNJ Ǯ˖ȂR. ͮȳʓݿʲľ ࠫ˲ n̢̞ ̦̚oࠫ(anomocytic)Ś ˮݮ, ̦̚oࠫ 38.6 Ⱥ LJɢࠫ(actinocytic)Ś Fƾ Ǯ˖ǮƮǮ ɊÚNJ Rˢࠫ(cyclocytic)Ś Ǯ˖Ǯ ̢˞ʮŚ ࡫ŞƆΖR. ̞ˮ ; (ľ ώKɛ(anticlinal wall)Ħ Χ˖ࠫȺ ȟ˖ࠫŚ ŇªΞ ȾȮ(straight/curved) ƆƮǮ Ԣɷࠫ(undulate؂ -VF(unicel؂ʲȟࠫ(sinuate)Ś ȾȮƆΖR. ξʪƊ ̢˞ʮΞ˒ Ǯ˖Ǯ Fľ ɫ˞ ʂNJ 4ɫ˞Ǣ Vˮ -F(pustular glandular tri˖ ˧؂lular non-glandular trichome), ˧ɷF(stellate), ˖F(glandular trichome), ˣ Ξ˒ ͏˧ Ԧ(membraneous platelets), Ԧ؂ݒˮٲ chome)ľ ࡫ŞƆΖR. ξʪƊ ̢˞ʮ ľÞ Ţ̞ ̢˞ʮľ ݒ Ɩˮࠫ˲ݯǧ ࠫή Ȋ ʖ ɗľ ȳٲ ݒɷǿή(epicuticular wax)Ś ȾȮƆΖR. Ϣǣ Ŵٲ F͟(platelets)ľ ݒɷǿή ĆÒľ ̢˞ݯǧ Ć˧Īٲ ĆԢʓࠫήĪ ȼĪ ώ ΔΖΦ͒, Fľ R͟˧(ӏঞ, ˧ɷF, ˖F)Ⱥ ݮÞ ʒRފݒ Ȋ R͟ݲ Ɩˮࠫ˲ݯǧ ࠫήĦ ^̞ࠫ˲ݯǧ ࠫήȺ Â̦΂ ʖ ɗĪ Śٲ ࡫ŞݮώR. Ŵ .Ćݲ ǻʒ̂ Ȃȳݶ ǁŚR ݒɷǿήٲ ,ȳʓݿʲ, ˧ɷF, ˖Fͮ ,؂ݒˮٲ ୱࢼ߂: Ŵ О ׻ dž ˞ʚɪӆƞ Ԃgł̾ β Lyonothamnus A.Gray˿Ӿڛ Ԙ ᇦ዆ ᎟ݒ͚ *ዦ ͺɦΦ ˞ʚዣ֋ӆƞ ̦к ዦ ͷዪӎ ዾϞӆβ, ̐ӊ ዆Ԛgɪ dž͚ʎ ˞ʚዣ֋, ׺ႧƪˎǶ dž͚ʎ ֟Ꮯዦ ׻ dž ˞ʚዣ֋ Ҏ௢ʾ ৈ݊ ƾ༿ዣ֋ ҪгʣӾ źܧł͒(Wilkinson, 1979; Stace, ዢǚ ԒΦ Žӆƞ ƪ๊ƾǚ, Rydberg (1908), Hutchinson 1984; Zhou et al., 1994; Yang and Lin, 2005; Zoric et al., (1964), Schulze-Menz (1964), Takhtajan (1997)͚ Ӣዾ ΊǶӢ Anil Kumar and Murugan, 2013; Deng et al., 2014). ;̦ʾ ĐΦ ၃ݟӆƞ ï˿ņ Adenostomateae Rydb.׻ӆƞ ;2009 ǯӢ ᎟๦, Ԃgł͒ Adenostoma˿Ӿ Ϟɯƪ׻͚ ᅶዲł̾, ԯ̪ ,ڛ ၃ᔒ, Ԙ ᇦ዆͚ʎ Ǻ঺ዪ Ί ԒΦ ᎟ݒ ݊݊ ŒǶ dž ʺ ˞ʚɪȎǶΦ ɦȞłΦ ï˿ņ ׻Ԏӎ ڛ ΢î ũ гӢ ؏ʚϊ ˞ᅶ țӊ ̛ዦ ˞ʚɪ͚ʎ Оዦ ᎟ݒƞ ͺɦł͒ӆβ(Ma et al., 2004; Moon et al., ዢΊ. ̺ዦ, Hutchinson (1964), Kalkman (2004)͚ Ӣዾ Chamaebatiaria˿, Sorbaria˿, Spiraeanthus˿Ƕ ዲϒ ,̪ڎ ,Adebowale et al., 2014; Cho et al., 2014), Ԙ ;2009 ˶Ƕn ᇦζ͚ ƪ๊ƪΦ ᇦ዆ȳƻݒ(epicuticular waxes) ͷ Gillenieae Maxim.׻ӆƞ Ԃgņ Gillenia Moench˿ӊ ૦ .f ˞ʚዣ֋ӆƞ ú௢ ԒΦ ᎟ݒƞʎ ͺɦł͒ ȳ׻(Supertribe)Ԃ Pyrodae C.S. Campb., R.C. Evans, D.R (Barthlott et al., 1998; Wissemann, 2000; Ensikat et al., 2006; Morgan & T.A. Dickinson͚ Ҏ௢ዢΊ. Tomaszewski and Zielinski, 2014). ӾĮዦ ᎟ݒӊ ̛ዦ Ϟɯƪ׻(Sorbarieae) sensu Potter et al. (2007)͚ ˿ዢ ,͚ʎ ֋гł̾ ׻ dž ǯҪ႖ɏ᎟ݒ Φ 4˿Ӣ Ԙ, ˾·Ӣ Ђ˚᎟๦Φ লӾʾ ɎӾΦڊ׻ Ί ڛ ʚƾ̃˞ ๚ȻӾƪ ˿þ, ؏þ gȞ᎟ݒ ႖ˏӎ Ҏዾ fîł͒ Adenostoma˿Ӣ Ʒк, ȳƟʣԂ ·ӊ ʒ᎟ӆƞ ௲·Ƕ (Husain et al., 1990; Taia, 2004; De S-Haiad et al., 2009; ҪȞዦ ᎟๦ʾ ƪ๊džǚ(Jepson, 1993), Chamaebatiaria˿ Moon et al., 2009; Heo et al., 2013). ӊ ȳƟʣӢ 2−3ᐖ кȳɏ·ӆƞ ̛௢gӢ ᅶԚ·Ƕ Ϟɯƪ׻(Sorbarieae Rydb.)ӊ ԯŒǶ(Rosaceae Juss.) ҪȞዦ ᎟๦ʾ ƪ๊NJ(Kalkman, 2004). Sorbaria˿ӊ Ԇ džӢ ׺ႧƪˎǶ(Spiraeoideae Arn.)͚ ˿ዢΦ ˞ʚɪӆ Ų֋Ԃ кȳɏ·Ӣ ᎟๦ʾ ɎӾβ(Lee, 2007), Spiraeanthus ƞ, ૦˶ ˞Ԛƾ༿ዣ֋ ͺɦʾ ̪Ųӆƞ ዦ ԯŒǶ ˞ʚਾ ˿ӊ ؋ӊ ʒ᎟Ӣ кȳɏ·ӆƞ 20−35ĖӢ ˾·ӆƞ ɦ ƾ͚ʎΦ Adenostoma Hook. & Arn., Chamaebatiaria Maxim., ʣłΦ, Chamaebatiaria˿Ƕ ҪȞዢƆ ֎ਾ֋ӆƞ ̛ Sorbaria (Ser.) A.Braun, Spiraeanthus Maxim. 4Ė ˿ӎ ᅶዲ ௢·Ƕ ΞAዦ ᎟๦ʾ ɎԂ(Komarov, 1939). ͺǺ ˞ʚ ዢǚ Ԓ(Potter et al., 2007). Adenostoma˿ӊ ˛Œ ʎ˚͚ ɪӆƞ Lyonothamnus˿Ӣ Ʒк ԘӢ Ӿ᎟Ꮮʾ ɎӾΦ, ̈́ Chamaebatiaria˿ӊ 1؏Ӿ Œɧ ʎ˚͚ ˞ᅶዢǚ, ֎ͺӢ · ᎟๦Ԃ L. floribundus subsp. floribundusϊ ,؏2−3 .ˣˎfˎϊ ăˎfˎ͚ Ųਗ਼ Ɔ кȳӆƞ Ă¾ׂ ɎӾΦ ᎟๦Ԃ L. floribundus subspڛ Sorbaria˿ӊ 4−9Ͷ؏Ӿ ;Ԛɏዢβ, Spiraeanthus˿ӊ ௾Ԛᓚ.๎ ǚҪ˿ӆƞ 1؏Ӿ aspleniifoliusӆƞ ɦ˞łβ(Raven, 1963; Jepson, 1993 ˞ᅶዦ(Jepson, 1993; Gu and Alexander, 2003; Kalkman, Kalkman, 2004), Gillenia˿ӊ 3ଦ·ӎ ݊Ϟ(Jepson, Ϟɯƪ׻ӊ ਢӖ Rydberg (1908)͚ Ӣዾ ӖӢ ʪ 1993). ૦˶͚Φ ԘӢ Ђ˚᎟๦ዣ֋ ᎟ݒ লӾ͚ʎ ႖ɏ .(2004 Chamaebatiaria, Schizonotus Lindl., Porteranthus Britton) łΦ dž˚ɦ׺Ӣ লӾ ႖ˏ ũ ׻ dž ǯҪ႖ɏ᎟ݒ ๚Ȼ)˿ ӎ ᅶዲዢΦ ׻ӆƞ Ԃgņ ӾÚƞ ̐ӊ ዣԚȎ͚ Ӣዾ ӎ Ҏዾ, ԘӢ ዾ˚ዣ֋ ᎟ݒ͚ *ዦ ͺɦú Ίጓł͒ ׻Ӣ ዦƾ ʖ֟Ӿ ȚƷł̾ Ϟ. ၃ᔒ, Potter et al. (2007) (Song and Hong, 2014). ዢ݊̎, ԘӢ Œʪ᎟๦ዣ֋ ᎟ݒ Ӣ ˞ʚਾƾ͚ ɚʶζ, Schulze-Menz (1964)͚ Ӣዾ Ϟɯƪ ͚ *ዦ ͺɦΦ ˎ݋ ֎ዢ. !"#$%&'(")#&%*'"+',*%&-'.%/"&"01'2"*3'45'6"3'7'879:5; /0*1'2345&2&567&8&).'3#'970'953:0'(&5:*530*0 !"# ɚ¾ʎ, ɒ ͺɦӢ ֋ӊ Ϟɯƪ׻ 4˿Ƕ ͺǺ ˞ʚ Ӣ ̲Ӿ(L), ȚΞ(W), ΢î(F) þӢ Ǻƾʾ ƪ๊dž̪ Ҏዾ Ȟ֎ԚᎎŒ ዆̾2 ȳǺǺƾ(Pearson correlation)ʾ ӾгዢͶ ˞ʏዢچ ɪ 2˿(Gillenia, Lyonothamnus)ӎ *ȳӆƞ Ʒ(SEM)ӎ ӾгዢͶ, ԘӢ ᇦ዆͚ʎ ᏟԂዪ Ί ԒΦ ᇦ዆ Ί(IBM SPSS Statistics for Windows, Version 19.0., .(ʪᅶ, ̪ǯ, г, ᇦ዆ȳƻݒ țӢ Ԙ ᇦ዆ Œʪɦ׺ʾ ȳ Released 2010, IBM Corp., Armonk, NY, USA ОʣǶ Ԙ ᇦ዆ Œʪ᎟ݒ͚ Ǻዦ г̾Φ Roe (1971), Wilkinsonڛ ʪᔒ Ǻ঺ዢǚ ̪Զዢǚ, Ӿʾ ༿ዾ Œʪ᎟๦Ӣ ƞ ɚÒǚ, ̪ǯӢ ΢îӢ Ʒкچ ׻ dž ǯҪ႖ɏ᎟ݒ ๚Ȼ ũ ˿þ, ؏þ gȞ᎟ݒƞʎӢ (1979), Stace (1984)ʾ ú௢ʾ ႖ˏዢΦ Ԓ. Croxdale (2000)Ӣ ƃǯӎ ূǚዢΊ. µŵ Ð êŖ I1 Ϟɯƪ׻ 20˞ʚɪӢ ԆŲ֋Ԃ ݎ̾ͅ ɒ ͺɦ͚ Ȟгņ Ԙ ԶȎΦ Adenostoma˿ 2؏ 1Ț؏, ɒ ͺɦƪǶ Chamaebatiaria˿ 1؏, Gillenia 2؏, Lyonothamnus˿ 1؏, 1 Ԙ ᇦ዆Ӣ Œʪ᎟๦ዣ֋ ၃ݟȎӊ Table 1͚ ֟ˮዢΊӆ .Sorbaria˿ 6؏, 4Ț؏, 1ሒ؏, Spiraeanthus˿ 1؏, ધ β, Figs. 1−4͚ ֦fዢΊ ,؏ˎ 20˞ʚɪ(36Ė ᇦɒ)ӆƞ ɧdžĹЂᇦɒǺ(A, C, GH, K, KH, KHUS, MO, SNUA, abbreviations according to Thiers, ƭʍȑĽ 2015)ӆƞ˚຺ *Ͷዦ ʏ·ᇦɒ͚ʎ ᏟɎዢΊǚ, Ԇ˚ ˞ ᇦ዆ʪᅶӢ ᎟๦Φ țƃ᎟(isometric, Fig. 1D), ˎțƃ ʚɪȎӊ ݋֛ ̆Ђ ৎݛዢͶ FAA (40% formalin: 40% ᎟(sub-isometric, Fig. 1I), ˢ˖௣᎟(irregular, Fig. 1C)ӆ :glacial acetic acid: 70% ethyl alcohol) Ꮖዳ˫͚ ǚ֟ዦ ᑎ, ƞ Ǻ঺ł͒ӆβ, *˚˞Ӣ ˞ʚɪ͚ʎ ጯଟζ(AD 70% ˖ೞζ͚ ԯ̪ ɎǺዢζʎ Ǻ঺ዢΊ. adaxial side)Ƕ Ɗଟζ(AB: abaxial side)͚ʎ ź ҪȞዢ ʣ΋ዦ Ԙӎ ʒȞዢǚ, гǶ ʒ֚ӎ Ǻ঺ዢ̪ Ҏዾ ዾ Ɔ ƪ๊ƾ(Table 1). ̎, Gillenia˿Ƕ Sorbaria˿͚ʎ ˚ᎎŒƷ(SM, SteREO Discovery. V8, Carl Zeiss Microscopy Φ ጯଟζǶ ƊଟζӢ ᇦ዆ʪᅶ ᎟๦ú ˾ ʶƆ Ǻ Ȟ֎ԚᎎŒƷ ঺ł͒. ˚ʪᅶӢ Ίிȗ(AW: anticlinal wall)ӊ ݋ʒچ ,GmbH, Jena, Germany)ӎ ȞгዢΊǚ .SEM)ӎ ӾгዢͶ Ԙ ᇦ዆ Œʪ᎟ݒȎӎ Ǻ঺ዢΊ. Ӿ ᎟Ƕ Ǜʒ᎟Ӿ ăf͚ Ǻ঺(straight/curved) łŪƪ(Fig) Œ Ů׺ņ ᇦɒӾƪ, *ଦᇦɒӆƞ˚຺Ӣ ԶȎΦ ̇ 90oC 1J), ႖ȳ᎟(undulate) ̺Φ ɮǛ᎟(sinuate)Ӿ ƪ๊ƾ .(͚ʎ 3˞þ ťӾŪƪ ͺᏞ֦(Agepon, Agfa Gevaert, (Fig.

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