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Taxonomy Oe the Notholaena 5Inhata Compeex The taxonomy of the Notholaena sinuata complex Item Type text; Thesis-Reproduction (electronic) Authors Hevly, Richard H. Publisher The University of Arizona. Rights Copyright © is held by the author. Digital access to this material is made possible by the University Libraries, University of Arizona. Further transmission, reproduction or presentation (such as public display or performance) of protected items is prohibited except with permission of the author. Download date 06/10/2021 11:59:11 Link to Item http://hdl.handle.net/10150/551396 ; ' THE; TAXONOMY OE THE NOTHOLAENA 5INHATA COMPEEX ; V by; ; iRichard Holme s Hevly v ■; A The s is Submitted to the Faculty of the ' . DEPARTMENT OF BOTANY In Partial Fulfillment of the Re<|UirementS 'A;- .i 1 T For .the Pegree of : 1; ' M AST ER O F .SCIENCE:';- ' ’l; V the: Graduate Colle ge ; . • % : UNIVERSITY OF ARIZONA A a 1959 . • .y/y/ , , 959 f : ::: 9 5 ^ y - ST.ATEMEN'T by a u t h o r . : . ■ This thesis has been submitted in partial fulfillment of require- . ments for an advanced degree at the University of Arizona .and is . deposited in the "University Library to be made available to borrowers under rules of the Library. ' . .. ' y , ' : ;: y ' 'Brief quotations, frofn this thesis are allowable .without special permission, provided that accurate acknowledgment of source is made. Requests for permission for extended quotation from or reproduction of this manuscript in whole or in part may be granted by the head of the major depd.rtment or the Dean of the Graduate College when iL .their judgment the proposed use of the material is in the interests of scholar ship> In all other instances, however, permission must "be Obtained frOm the author, y ' y ;' y. y y. y' ■ ■' . -y; : ay-' " SIGNED J ? A / A b - A J APPROVAL "BY- THESIS DIRECTOR This thesis has been approved on the date shown below: OoL-O-k Vl/l o ^ -11, .rfsTj . ■V ' CHARLES TLMiSObr, Jr." % ate 1 Associate Professor of Botany ii : : .'i ■ ■ ACKN:qWDEDGME^^N ' : I wish to express .my. deep appreciation to Dr. Charles TV 4. Mas oil, Jr. , for his inspiration: and, guidance, without which, this Study would never have been initiated and completed. I am much, indebted to Dr. Phillips, Dr. Kurtz, and Dr. Harris K-' as well as Dr. Mason, for criticism and helpful suggestions in the pre t paration.of this'thesis. / : : ..J.'•: ;vp;; : :V ; «- In. addition, thanks is due Dr. Harris and Albert T. Ellis of the Department of Botany for translation of critical papers in German' arid • -French, respectively, and to Dr. .Ayers of the Depaftment of Glassies 1 •for‘the Datin diagnoses. > .'■ . ’ . ■ . ' . ... ; : ‘ : ;:• Gratitude is due to all the members of the 'Department of Botany for their encouragement and assistance throughout the cdiirse of the - i' inyestigations, and to the 'Depaftment of Bdfany and the University Of' ;' Arizona for the use of equipment and facilities necessary to undertake - this work: . ■- . , . ■ . ;* A special yote:of,tisanks is also due to M rsyLutfetla Birea^eale Hamiitori for kindly p reparihg the drawings of the mature Spdrophytes. I would also like to thank the following institutipris for the use of herbarium material: Gray Herbarium at Harvard XJniversity, University Of California Herbari'um, Dudley Herbarium at Stanford Uriiyersity, . Califorriia Academy of Science, Rancho Santa :Ana Botanic Garden, . ■••Pomoha Gbllege, the University of Arizona, and the Uriiyersity of Wash- /: ington Herbaria. • :;'/.• ;' p::: .•.■•'•' : ; 'g : , ' ■■ TABLE QE CONTENTS; P a g e List of Tables v List of Figures« , . ».. O O O Q O p © o » © » .» © o o © » » » vi CHAPTER . ' ' ' . \y'-/;' I INTROb.UCTlON.:lFlil'V%L^'Llll',La'iaL^ .. 1 II MATERIALS AND METHODS. ...................... 4 Ill COMPARATIVE MORPHOLOGY AND ANATOMY. 7 ' , ' _ : ; ' ' .1 . /, ■ • M a teria ls and. iviecinoSs ^ o #» »@ @ * * * * * @ @ @ @ #»*»@ 9 Gross structure and anatomy of the roots. ,A. .. 11 Gross structure aAd anatomy of the rhizom es.... 13 Gross structure and anatomy of the fronds... ... 15' Spores.......«......«....«»............- #.... * * 24 - ' Prothallia. i...... .. & e ©■ » © a © o 9 30 - Young spordphytes. = . O O O O O O » O O O O > » . O 0,0 35 IV PHYSIOLOGY, 39 V CYTOLOGY. ; o o©oooos?ooooaaooooooo© 54 VI THE NOTHOLAENA SIN DATA COMPLEX AND ITS INTERPRETATION. ... ..... 59 LITERATURE CITED;. ... .. .. 89 V-v 1 ■ - . ■ List ABLss; ■ Table Page i :■ ' : - :: :1 ; ■ Light requirement for germination and light ' intensity requirement for normal development ' b : 3 r ■ of' the prothalins • • «« * v * ® a* • •• ■* • > © • • ©■'« w, © • © « © ' The effect of photoperiod on spore germination©. ' ' ■•■■■ :'V2 ; ; 'Effect■ of light color on germination. ■ v ; 4-;:: The effect of dark and light on spore imbibition • . and ge rm in at ton,.*.1. .... ® . * . @.. *. .. >■. © .... 5° : , ' : . , V ' ; ' ' . " V; . Summary of the factors which c ans e the .anthOr . to believe that siimata and cochisenBis are dig- / tinet .apec.ie.S o o' »• • • > •• * * & * @ «_ <>:« « ® >•« » > » > * «.• ■* • *• •. *• * c » © , LIST OF FIGURES F ig u re • ' P ag e ■ ' ■ . ■ • ■ . , ' ■ Io - Transverse section of N* cocliisenSis root in the region of maturation (ca» 100x)« 0 <> o » » .. .. 12 2. Transverse sections of rhizomes (ca» 4'0x)o o o „ o » »* 15 ■ : 3q’ . Indument of the rhizome (ca.» 30x).»«, . * . *«.» « » o» o « « 16 4a Transverse sections of the Stipe (cao 50x)o » 6 » »»o 18 5o Transverse sectiohs of the pinnae (c.a, 100x)» » » » , • 20 ‘ 6 Pinnae venation, soral arrangement, and sporangia© © «< <> » © » © © © e » o »© © o ® © © ® © » © © »© © © » ©.© © © © © » © 21 7a Indument of the lamina® ©; © © © :© ® ® « o 23 ■ 1 - - ' ■ ' - ; ■ ■ ' . " V*:. Indument of the lamina.* , . ■ 2 5"' ' V 9 < Spores, » ©■ e ©o© o © a o © : o ® a o o o o e © » © © © © ® < 27 10. Prothallia. 32 11. Young sporophyte de-velopment. .............. 34 12. The germination response of spores of varying ages to pH. ..... o-o o d o o © o « d d O' O O O • O 9 9 42 The germination response of spores to pH and c emp e r atur e..... ./. >. ..... ........... .. ; 44 14. Cytology. .... 9 0 0 9 0 0 0 0 0 0 57 15. N» cochisensis x 1/2. 65 v i :. MST ‘ OF FIGURES!' (cont 'd) F ig u re P a g e 16. Distribution map of the principal varieties of Notholaena coob.isen.sis.. * 6 . *. * ...... .... ... 66 17. N. sinuata var. sinuata x 1 /2.................. 74 18. Distribution map of the principal varieties of ” Notholaena sinuata. .......... .. .». ...... 75 19. : N.;: sihuata var. ' rnadriensis x 1/2......... .. 82 20. bio infegerrima .x 1/2. .... ... ..... ... .. 85 ; 21. .Distribution map of the principal varieties of Notholaena integerrima.. .. .. .... ... ... .. 86 \'V' -'Vt. ,, » - ' ; ■ ‘ • : CHAPTER. I vB^-ftbEtrcTioiir: ■ Although relatively inconspicuoass one of the common elements of the vegetation of the rocky, arid foothills of southwestern United ; • ;States: and Mexico:iis an assemblage of plants which are .frequently 'l. referred to as xerophytic rock ferns. Among the more numerous are . those with linear, .pinnate-pinnatifid,.paleaceous fronds presently known :: kyyi- ' - ;.-V yV'f'ktkk--, •, • ■ as Notholaena sinuata (Lagasca ex Swartz) Kaulfuss. , k y . y : . - 1 : ■ k. ; y f ;;Motholaena sinuata. Was originally described from Pernyby .Mariano y -y. ' Lagasca in Swartz's. -Synopsis Filicurn. (Ig'bbj - as Acrostichum sinuatum. : . .yiy ; I . According to R. Tryon (1956) Kaulfuss transferred this complex to the genus Nothola,ena in 1824, where it has been retained except by P resl •" ■ ' y and Mettenius who included it in the. genus Gymnogramma and Domin who • :' considefed it part of the genus Chstlanthes. y k- '■ /. .f i .; k. -' 1' . : ;y; The very. polymorphic nature of this complex has presented tax~ek y -'i ' onomiC difficulties which have resulted in the publication of .nume rous ■ >a\y. 'k^yy-y y k : yy e .■■ - y.,.^- 'yy;y: y y yy . k. .yk '. ; ;.:.ke.y .k ....- f^/kv:..-, specific and varietal names. Some of these were based, unfortunately, ,, '• y yi. vyii-ik^/d 'yk% y'-y y on juyenile, depauperate, dr otherwise peculiar morphological abherations y - y . 2 as cited by R„ Try on. (1956). W eatherby, who considered this complex to be the truest example of the genus Notholaena in the Americas (1941), attempted to resolve some of the confusion surrounding three of the more conspicuous elements in the northern portion of the range in 1943. He recognized only one species, reducing Notholaena cochisensis Goodding to the rank of variety and maintaining,besides the typical form,Notholaena sinuata,. var. integer rim a Hooker. Weather by left undecided the treat- ' ment of another element variously known as Notholaena c rassifolia Moore and Houlst.or Notholaena prulnosa Fee, but suggested that it might be reduced to a variety of Notholaena sinuata. Without modification Tryon (1956) extrapolated Weather by’S treatment to all of the various forms of the N. sinuata complex. ,, / " The complex has an extensive distribution which ranges from northwestern Argentina along the Andes to the eastern Cordillera and the island of Hispanola, through Central America and Mexico to the southwestern United States. Throughout its range it is found growing in rock crevices and on rocky hillsides, some of the forms of the com­ plex being restricted to particular kinds of rock. Since N. sinuata and its allies may be found growing from near sea level to several thousand feet above, it is not surprising to find them occupying such diverse habitats as thorn scrub desert and oak-pine communities.
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