Éfi1Ð A TAXONOMIC REVISION OF PRQITANTHERÀ Labill. SECTION KLANDERTA (F.v. Muel-l-. ) Benth. ( LABIATAE ) by Barry John Conn, iul.Sc., B.Sc.Ed. (MELB. ) Department of Botany, University of Adelaide Thesls presentèd for the Degree of Doctbr of Phllosophy at the University of Adelaide June, 1982 TABLE OF CONTENTS Ab-stract iv Decl.aration of OriginalitY vi Acknowl-edgements vii Introduction 1 Taxonomic HistorY 5 Methods, Materials and Presentation 8 Di-scussion of Select.ed Morphological Characters 11 Habit 11 Indumentum 12 Leaves i3 Infforescence 14 Prophylls 20 CaJ-yx 20 Coroll-a 21 Androecium 22 .l Disc and GYnoecium 24 !, Fruits and Seeds - 25 Pollinâti-on and Fl-oral- Biology Introduction 26 Field Observations 26 Pollinatlon mechanism i-n section Klanderia 27 Pollinat,ion mechanism in section Prostanthera 28 Fl-oral biology and ornithophily in section Klandenia 28 Breeding system in section Klanderia 31 i Seed Dispersal- and SeedÌing Establishment 33 Numerical Analysis Introduction 34 Sel-ection of Morphological Characters 36 Pre-numerical analYsis 37 Method used to seLect morphological- characters 38 Evaluation of character set 44 ' Numerical analyses of sPecimens 52 Numerical- anal-ysis of Prostanthera aspalathoides 66 Numerical- analysis of the Prostanthera P. mi-crophy l-]a-P. serpv]lifol-ia complex 70 Numerlcal analysis of the Prostanthera laricoides complex 76 Geographic Variation 83 Môrphological variati-on i-n Prostanthera aspalathoides 85 Morphologic al- variation in the Prostanthera P. microphvlla-P. serpyllifolia complex 97 Morphologi cal- variation in the Prosr-anthera laricoides complex 112 Conclusion fnom Geographic Variation studies 125 VoLatile 0i1s (Terpenoids) 127 Material-s and methods 127 Numerical analyses of volatil-e leaf-oiIs of Prostanthera aspalathoides 131 Evaluation of volatile leaf-oil character set 131 Numerical *^fy="" of sPecimens 131 11 Geographic variation of Pnostanthera aspalathoides based on volatile leaf-oils 143 Function of volatile oils in Prostanthera: A discussion 151 Volatil-e oils as a defence against animals and insects 151 Vol-atile oils and pollination 153 Systematic Treatment 154 a. Prostanthera sect. Prostanthera 154 b. Prostanthera Labil-l-. sect. Klanderia 156 Recognized taxa and their distribution 157 Key to Species 159 [slecies Descript,ions] 162 Nomen sedis incertae 257 Appendix Index to coll-ections used in the numerical analyses -' -- 258 (a) Names of taxa of Prostanthera section,Klanderia-- with their key ¡umbers 258 (b) Enumeration.of -col-l-ections 259 References - 265 Index to Scientific Names Index to plant names 286 Index to bind names 291 L1a ABSTRACT A taxonomic revision of Prostanthera sectjon KLanderia ís presented. Generat chapters on taxonomic history, morphology, pollination, and breeding syste$s precede the systematic treatrent. Fourteen species are recognized of which seven are descrjbed for the first time. The new species are P. fLorifena" P. íncuruata' P. Laricoides, P. monticoLa" P. patens, P. pedicelLata and P. sem¿- ter.es. rvo subspecies of P. sez'pyLldfoLia and two subspecies of P. serniteres are recognized. P. sem¿teres ssp. intrieata is des- cribed for the first time. Keys to the species and subspecies are provided, AIl recognized taxa are provided with fuII descriptions, distribution information (including maps), ecological and other relevant notes. AIl species are illustrated. I,torphological variation of P. aspaLathoides, the P. caLycina- P. mùcrophyLLa-P.serpyLlifoLia complex, and Ì.he P. Larícoides complex, plus the volatile leaf oil variation of P. aspaLatVnides, were analysed with the multivariate nunerical techniques: canonical variate analysis, principal components and principal factor analyses, princi- pal coordinates analysis, surface trend analysis (contour mapping) and differential systernatics. Manhattan metric distances were used as a measure of dissimilarity between.individuals and,/or groups; The univariate significance tests used were: t - test (often modified), F - test, and SÈudent - Newman - Kuels multiple-range test (SNK). patterns of variation appeared to be associated with envir- onmental and historical factors in P. a.spq,Lathoides and in the P. ca.LA- J-V eina-P. microphylLa-P. serpyLLifoLia compLex. The distinctness of the Kangaroo Island populations appears to reflect the relatively Iong separation of this island from the mainland.. v DECLARATION OF ORIGINAIITY I hereby declare that this thesis is my own work, excePt where specifically sÈated to the contrary. To my knowledge, it is not sr:bstantially the same as any other thesis which has been sutrnitted to any UniversitY. Barry iI. Conn vl- ACKNOWLEDGEMENTS I sincerely acknowtedge the following people for the technical assistance which they generously provided: for computing techniques - Dr D.T. Blackburn (then ADU), Messrs P.J. Lang, c.J. Merrick, K.P. Nicholson (a11 ADU) and Dr T. Whiffin (Department of Botany, Latrobe Univ., Victoria); and for gas-Iiquid-chromatographic techniques - Drs E.V. Lassak (Biological c Chemical Research Institute, Rydalmere, NeI^7 South V,IaIes), R.H. Prager (Department of Organic Chemistry, Univ. of Adelaide, South Australia) and T- Vlhiffin. I am indebted to the Directors and Curatorsof the many herbaria who made availabfe loans of herbarium material. I am extremely grateful to Dr J.p. Jessop for making availabfe the faciLities at the State Herbar- ium of South Australia (AD) - I gratefully acknowl-edge the support of the Nd.ional Parks and gilildlife Service of New South Wa1es, the Departrnent of Environment and planning (South Australia), the National Parks Authority of Victoria, and the Forests Commission of Victoria for freely granting permission to collect from areas u¡rder their control. t wish to sincerely thank Drs D.c. christophel (ADU) and J.P. Jessop for their continued interest in this study, the valuable discuss- ions which I had with them and the detailed comments which they made on all aspects of the manuscriPt- To Dr p.S. Short (MEL) I am thankful for his constructive criticisms of the manuscriPt. The plant illustrations were skilfully done by l"lrs H.M. Bennett (FiS. 53F), Ms A. Podwyszynski (MEL) (rigs 3 & 4) and Mr L. Dutkiewicz (AD) vrl_ (alt other plant illustrations). Mr & Mrs P. Al-thofer kindly offered their hospitality, field assistance and the facilíties of the Burrendong Arboretum (New South lfales) . !'lr G. Althofer generously .shared his extensive knowledge of the genus. I sincerely thank l"lr 6. l':rs G. Jackson for their hospitalíty and field assistance on Kangaroo Island (South Australia). I express my sincere appreciation to Misses C.E. Marcelline and T. Munro for carefully typing the manuscript. Finally, I wish torexpressrmy':deepest gratitude to Ïelen and my family for patiently enduring the rnany hours spent on this dissertation. vt-J-r 1 INTRODUCTION Prostanthera specÍes are evergreen sub-shrubs, shrubs or small trees (p. lasianthos) which are characterized by having decussate leaves, racemiform inf lorescences, two-J-irbed calyces, four fertil_e two-celled anthers in each flower, terminal styres, and fruits composed of four mericarps. Prostanthera with Eichlerago, Hemiandra , Hemigenia. Ivlicro- corys, vilestringia and wrixonia are grouped together in the subfamiry Prostantheroideae of the Labiatae (Briquet 1895; l"telchior L964¡ Carrick L976, 1-977). Carrick (1977) offered a diagnosis for prostantheroideae. His early paper (Carrick 1976) summarizes the key differences between aII the genera in the subfamily (with the exception of Eichleraqo which was not described at that tirne). The results from the work of Sharma ç singh (1982) on carper morphology (refer pp. 24 s,25) require.carrickrs diagmosis of the Prostantheroideae to be modified to: Stamens 4, fertile, or only 2 ( ad,axial or abaxial pair sterile); style terminal (may superficially appear subgynobasic); fruit of 4 separate mericarps or entíre, dry and indehiscent (Eichlerago). Comment on generic delimitations within the Prostantheroideae must await critical evaluation which is beyond the scope of this present study. The Prostantheroideae are endemic to Australia with Èhe most dis- junctdistribution being record,ed by Jacobs e Pickard (198L) for l¡estringia fruticosa which they list as occurring on Lord Howe Island. Furthermore, this subfami-ly is the only one (of the Labiatae) in which any genus is endemic to Australia (Jessop 1980). The Prostantheroideae appear to be a distinct taxon of Èhe La-l¡iatae. This is indirectly verified by 2 the fact that the Prostantheroideae, as circumscribed by Bentham [as 'Tribus VII. Prostanthereae.' (Bentham 1834, p. 4471¡ and Bentham & Hooker j-BZ6l, has been maintained almost unaltered by subsequent authors. Cunningham recognized the homogeneity and distinctness of the group of genera which are now classified within this subfamily as early as 1825 (Cunningham, in Field L825). However, prior to 1834 the genera which are now regarded as belonging to the Prostantheroideae were frequently ptaced in separate infrafamilial groups (e.g. Reichenbach 1828; Bentham, in Lindley 1-829-1-83O; Bartting 1830). 'l{ithout evaluating the relation- ship between the subfamities of the Labiatae, workers have usually regarded the Prostantheroideae as most closely related to the Ajugoideae (e.g. Briquet 1895; Hillson l-959) - The prostantheroideae, together with the Ajugoideae and Rosmarin- oideae, are regarded as transitional subfamilies between the remaining Labiatae and Verbenaceae (Cronquist 1981). The relationship'between