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Comparative Leaf Anatomy and Systematics in Dendrobium, Sections Aporum and Rhizobium (Orchidaceae) Barbara S

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Comparative Leaf Anatomy and Systematics in Dendrobium, Sections Aporum and Rhizobium (Orchidaceae) Barbara S Eastern Illinois University The Keep Faculty Research & Creative Activity Biological Sciences January 1997 Comparative leaf anatomy and systematics in Dendrobium, sections Aporum and Rhizobium (Orchidaceae) Barbara S. Carlsward Eastern Illinois University, [email protected] William Lois Stern University of Florida Walter S. Judd University of Florida Terence W. Lucansky University of Florida Follow this and additional works at: http://thekeep.eiu.edu/bio_fac Part of the Biology Commons Recommended Citation Carlsward, Barbara S.; Stern, William Lois; Judd, Walter S.; and Lucansky, Terence W., "Comparative leaf anatomy and systematics in Dendrobium, sections Aporum and Rhizobium (Orchidaceae)" (1997). Faculty Research & Creative Activity. 260. http://thekeep.eiu.edu/bio_fac/260 This Article is brought to you for free and open access by the Biological Sciences at The Keep. It has been accepted for inclusion in Faculty Research & Creative Activity by an authorized administrator of The Keep. For more information, please contact [email protected]. Int. J. Plant Sci. 158(3):332-342. 1997. C 1997 by The Universityof Chicago. All rightsreserved. 1058-5893/97/5803-0012$03.00 COMPARATIVELEAF ANATOMY AND SYSTEMATICS IN DENDROBIUM, SECTIONSAPORUM AND RHIZOBIUM (ORCHIDACEAE) BARBARA SUE CARLSWARD,* WILLIAM LOUIS STERN,I't WALTER S. JUDD,t AND TERENCE W. LUCANSKYt *Departmentof Ecology, Evolution and Marine Biology, Universityof California,Santa Barbara, California93106-9610; and tDepartmentof Botany,University of Florida, Gainesville, Florida 32611-8526 The specialized leaf anatomyfor species of Dendrobiumwithin section Aporum is similarto thatof species in section Rhizobium.In both sectionsleaves are characterizedby a unifacialor nearlyunifacial surface where the exposed surface is abaxial. However, leaves in section Rhizobiumalso featurea lacuna submergedin the mesophylland surroundedby an adaxial epidermis.In contrast,leaves in sectionAporum merely present an internalsuture that divides the leaf into bilateralhalves. These two sections of Dendrobium are hypothesizedto be sistertaxa because of synapomorphiesin theirfoliar anatomy.A cladistic analysis performedwith various anatomical charactersof the leaf demonstratesthat both groups are monophyletic.Little resolutionwas found withinthe anatomicallyand morphologicallydistinctive Aporumclade. Introduction Indonesian islands, Papua New Guinea, Andaman Is- lands, Thailand,Burma, Laos, Vietnam,and regionsof Dendrobium Swartz, a genus withintribe Dendro- northeasternIndia and southernChina. Plants of Apo- bieae, subtribeDendrobiinae (Orchidaceae), comprises rum are characterizedby shortrhizomes that bear fas- about 900 species (Bechtel et al. 1992; Mabberley cicles of uprightor pendentslender stems,sometimes 1993) that exhibit a wide range of vegetative mor- obscured by overlapping leaves. Leaves are coria- phology and anatomy. Dendrobium is distributed ceous, equitant,and laterally flattened;they vary in throughouttropical Asia, some Pacific islands, and outlinefrom scalpelliform (fig. 1), lanceolate, and fal- Australasia (Dressler 1993). Plants may be epiphytic, cate (fig. 2), to triangular.The laminae arise from a lithophytic,or terrestrialwith slender to pseudobul- sheaththat is obliquely insertedon the stem (Kranzlin bous stems composed of one to several nodes. Leaves [1910] 1957). may be equitantor more or less widely separateddis- Leaf anatomy of members of section Aporum has tichouslyalong the stem, conduplicate,laterally flat- ex- tened,or cylindrical. not been studiedextensively and was only briefly Schlechter ([1912] 1982) is the most widely cited amined by Morris et al. (1996) in theirtreatment of authorityin the classificationof Dendrobiumbased on subtribeDendrobiinae and by Solereder and Meyer his work in the orchid floraof German New Guinea. (1930) in their survey of the vegetative anatomy of We used Kranzlin's ([1910] 1957) classification,even Orchidaceae. Solereder and Meyer reported on the thoughit is not favoredby currentworkers, because it anatomy of Orchidaceae but made no attemptto use contains a more comprehensivetreatment of section this informationsystematically. They mentionedonly Aporum Blume. Kranzlin treatsforty-nine species of one species of Aporum, Dendrobium aloifolium,the which we had ten for studyin additionto Dendrobium leaves of which were characterizedas having: (1) eq- brevimentum,more recentlydescribed by Seidenfaden uitant arrangement,unifacial orientation,and an ex- (1985). posed abaxial epidermis; (2) glandular,sunken, mul- Kranzlin divided section Aporum (as a subgenus) ticellular hairs; (3) peripheral "bast" fiber bundles intotwo subgroups,Hemiphylla and Holophylla,based below the subepidermal cell layer ("nur periphere upon the distributionof leaves along the stem. Plants Bastfaserbtindel,zumeist unter der subepidermalen in sectionHemiphylla possess leaves thatonly partially Zellschicht"; Solereder and Meyer 1930, p. 118); (4) cover the stem, whereas leaves in section Holophylla vascular bundles arranged in two symmetricalrows completelycover the stem. within the mesophyll; and (5) stegmata containing Species of sectionAporum are centeredaround Bur- spherical silica bodies. In addition,they cited, as gen- ma, but they also occur in Malaysia, throughoutthe eral characteristicsof the Dendrobium leaf, calcium- oxalate raphidesand druse-likestructures and "antho- 'Authorfor correspondenceand reprints. cyanin pigments" (anthocyanartige Farbstoffe) in Manuscript received June 1996; revised manuscriptreceived De- solutionwithin the cell vacuoles. They describedfiber cember 1996. bundles, variouslythickened walls of cells comprising Figs.1-5 Leaves of Dendrobium section Aporum. Bars = 10 ,um.Figs. 1 and 2 show leafy stems. Fig. 1, D. leonis with scalpelliform, equitant,closely overlappingleaves; x 0.40. Fig. 2, D. acinaciformewith falcate,equitant, widely separatedleaves; x 0.5. Figs. 3-5 show cross sections of leaves. Fig. 3, D. acinaciformewith reniformsuberin-like bodies (arrow) in adaxial epidermalcells; X 1360. Fig. 4, D. distichumunder polarized light showing thickenedadaxial epidermalcell walls (arrow) impregnatedwith a suberin-likesubstance; X 1360. Fig. 5, D. acinaciformewith heterogeneousfiber bundles; x 2200. 332 This content downloaded from 139.67.65.234 on Wed, 14 May 2014 15:29:04 PM All use subject to JSTOR Terms and Conditions CARLSWARD ET AL.-LEAF ANATOMY DENDROBIUM SECTION APORUM 333 __ _~~M 07e This content downloaded from 139.67.65.234 on Wed, 14 May 2014 15:29:04 PM All use subject to JSTOR Terms and Conditions 334 INTERNATIONAL JOURNAL OF PLANT SCIENCES Table1 Table2 SOURCES OF SPECIMENS CHARACTERS USED IN CLADISTIC ANALYSIS OF SECTIONS APORUM AND RHIzOBIuM Taxon Sourcea 1. Cells surroundinghair base: thin- and thick-walled(0), thin- D. acinaciformeRoxb ...................... WLS hort.FLAS walled only (1). D. aloifolium(Bi.) Rchb. f ........... ..... K s.n. 2. Stomatal apparatus:paracytic (0), tetracyticand cyclocytic(1), D. anceps Sw............................... WLS hort.FLAS mainlyparacytic with some anomocytic(2) [unordered]. D. brevimentumSeidenf .................... WLS hort.FLAS 3. Cells surroundingsubstomatal chamber: thin-walled(0), thin- D. distichum(Presl) Rchb. f............... SEL 1987-0062A and thick-walled(1). D. indivisum . K (Bi.) Miq .................... 1984-3270 4. Abaxial cuticle: thin,less than 3 pm (0); thick,more than 3.5 D. leonis (Lindl.) Rchb. f.................. WLS hort.FLAS PAm(1). D. linguiformeSw ........................... WLS hort. 5. Epidermal cell shape (x-s): rounded, width:length ratio less D. mannii Ridl............................... K 1986-1031 than 46 (0); elongate,width: length ratio greaterthan 54 (1). D. nathanieleRchb. f....................... K 1986-5692 6. Fiber bundles: absent (0), presentwith heterogeneous cell com- D. rigidumR. Br. ........................... WLS hort.FLAS position (1). D. rosellumRidl ............................. SEL 1978-0200A 7. Water-storagecells: absent (0), presentwith banded thickenings D. salaccense (Blume) Lindl.i............. WLS hort.FLAS (1), presentwith pleats (2) [unordered]. D. toressae (F M. Bailey) Dockrill....... WLS hort. 8. Distributionof stegmata: presentalong vascular bundles only (0), absent (1), presentalong vascular bundles and fiberbundles a = = WLS William Louis Stern,private collection; K Royal Bo- (2), presentalong fiberbundles only (3) [unordered]. tanic Gardens, Kew; SEL = The Marie Selby Botanical Gardens. 9. Leaf orientation:bifacial (0), primarilyabaxial epidermis ex- Abbreviationsof herbariaare according to Holmgrenet al. (1990), posed with lacuna or incipientlacuna (adaxial groove) present and those of authorsfollow Brummittand Powell (1992). (1), only abaxial epidermisexposed and no lacuna (2) [unor- dered]. 10. Hypodermis: absent (0), composed of thick- and thin-walled hypodermal tissue, sunken trichomes, and guard cells banded cells (1), composed of thin-walledbanded and unbanded with small lumens and thickened walls with protruding cells (2), composed of thin-walledunbanded cells (3) [unorder- outer ledges. ed]. Plants in section Rhizobium (Dendrobium) possess 11. Leaf width:length ratio: narrow,0-4 (0); wide, 43-70 (1); in- termediate,10-38 (2) [unordered]. some vegetative characteristics similar to those of Apo- 12. Mesophyll canal: absent (0), present(1). rum, such as the unifacial leaf, and it has been sug- 13. Basal leaf sheath: present(0), absent (1). gested that section Rhizobium is the sister taxon of 14. Suturewithin
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