/ FLORAL MORPHOLOGY IN : RELATION. TO ADAPTATION AND TAXONOMY IN, THE CARYOPHYLLACEAE BY . KATHERINE L NOVOSEL Thesis presented for the Degree of Doctor of Philosophy University of Edinburgh 1982 5.. ABSTRACT The floral morphology of several species in the 3 subfamilies of the family Caryophyllaceae has been investigated from different aspects. The reproductive biology of the family has been examined to determine if style number and stigmatic area are in any way related to ovule/seed number, and it has been found that there is no positive relationship. The genera in the family have also been placed in groups according to the distribution of the stigmatic papillae and the degree of style fusion. Species in 4 genera, Spergula, Spergularia, Stellaria, Myosoton have been examined in greater detail but again few correlations could be found. De-styling experiments on 3 5-styled species in the subfamily Dianthoideae have shown that pollen tubes readily cross between 'carpels' in these species and that if only 3 styles remain the number of seeds formed is the saine.as in 5-styled ovaries. The vascular tissue of the ovary and the position of the trans- mitting tissue has been studied in species in the subfamilies Dianthoideae and Paronychioideae. This has revealed that the trans- mitting tissue is part of the septal tissue and confirmed the views of other authors that the ovary in this family has not evolved from the traditional 'carpel' but that the ovary is composed of a sterile part and a fertile part. The taxonomy of the subfamily Paronychioideae has been investi- gated. A Key to the genera has been constructed and a description of each of the genera in this subfamily is given. CONTENTS Acknowledgements. List of Figures. List of Plates. List of.Tables. Chapter 1. Introduction ........•..•..•.••••. Chapter 2. 2.1 Introduction •...................................s.... 13 2.2 Materials and Methods ........................•..•... 18 2.3 Styles, stigmatic area and ovules in the family Caryophyllaceae 25 2.4 Comparisons between genera with three and five styled species. 38 2.5 De—styling experiments. 85 2.6 Discussion 95 Chapter 3. Ovary Morphology 3.1 Introductory Review . ... .. oo.0000. 97 3.2 Materials and Methods ooe00000000*4o*oo**e000eoe000000 116 3.3 Results ............................... 124 3.4 Discussion ..........•................................ 194 Chapter 4.. ParOnychioideae 4.1 Introduction 200 4.2 Floral Morphology 203 4.3 Classification 216 4.4 Description of Genera 234 Chapter 5. Conclusions. .. • 1 • • • • • . • . 31 Q Bibliography ....................................•.........•. 314 Appendix I Appendix II Index to Species in Chapters 2, 3, & 4. ACKNOWLEDGEMENTS. I am indebted to the Carnegie Trust for the Universities of Scotland who funded this research. I wish to thank my supervisors, Prof. P. H. Davis and Dr. P. N. Smith for all their help and encouragement during the period of research and for constructive criticism of this thesis. I would also like to thank all members of the. Botany Department who have helped me, especially those in the electron microscope laboratory for their technical assistance. To the Curator and staff of the Royal Botanic Gardens, Edinburgh, for their help and for allowing me to work in the herbarium for many months, I am grateful. I also acknowledge my gratitude to all the institutions which donated seeds or living material. I would also like to thank my husband and family for their continuous support through some very difficult periods. I also wish to thank Mrs. Marion Sharp for typing this thesis. LIST OF FIGURES Fig. 1. Agrostemma githago L. ........ • • • • . 130 Fig. 2. Agrosternrna githago L. 134 Fig. 3. Cucubalus baccifer L. 151 Fig. 4. Sj.lenecoeli-rosa(L.)Godron .... ......... ......... 10 1 Fig. 5. Silene conoadea L. 16 7 Fig. 6. Scieranthus annuus L. 184 Pig. 7. Drymaria arenarioides Huxnb., & Bonpi. 209 Fig. 8. Paronychia 23 8.1 P. americana (Mutt. ) Fenzl 8.2 P.canadensis (L.) Wood 8.3 P. setacea Torr. & Gray 8.4 P. herniarioides (Nichx.) Mutt. 8.5 P. argentea Lam. 8.6 P. kapela (Hacq.) Kermer 8.7 P. argyrocorna (Michx.) Mutt. 8.8 P. capitata (L.) Lam. Fig. 9. Herniaria ..•... .... ................... 251 9.1 H. glabra L. 9.2 H. polygama J. Gay 9.3 H. fontanesli J. Gay Fig. 10.lLochiabracteataBalf.f. .......................... 251 10.2GymnocarposdecanderForsk. ...-.................... 251 Fig. 11.1 Scierocephalus arabicus Boiss. ..................... 258 11.2lllecebrumverticillatumL. ..................,.... 258 11.3 Chaetonychia cyinosa (L.) Sweet 11.4 Cardionema congesta (Berth.) Nels. & Macbride 11.5 Corrigiola litoralis 11 C. propinqua Cl. Gay Fig. 12.1 Pollichiacainpestris Solander .................... 263 12.2 P. campestris Solander 12.3 Achyronychia cooperi Hemsi. 12.4 A. parryi Torrey & Gray 12.5 Scopuluphila rix±'ordii (Brand.) Munz & Johnston Fig. 13.1 Pteranthus . •........... .. 268 13,2 P. d.ichotomus Forskal 13.3/ Fig. 13.3 Dicheranthus 13.4 D. plocainoides Webb 13.5 Cometes 13.6 C. surranttensis L. Fig. 14. Polycarpaea 275 14.1 P. longiflora F. Muell. 14.2 P. staminodina F. Muell. 14.3 P. teneriffae Lam. Fig. 15. Dryrnaria ... .. ... ............ ........• 282 15.1 D. pachyphylla Wooton & Standi. 15.2 D. leptophylla (Chain. & Schlecht) Fenzl 15.3 D. arena.rioides Hunib. & Bonpi. ex Roem. 15.4 D. cordata (L.) Wilid. ex Roem. & Schult. Fig. 16.1LoeflingiahispanicaL. ........................... 289 1,.2 Microphyes litoralis Phil. 16.3 Polycarpon tetraphyllum (L.) L. 16.4 P. coloinense Porta 16.5 P. robbairea 0. Kuntze Fig. 17.1StipulicidasetaceaMichx. ........................ 2%. 17.2 Ortegia hispanica L. 17.3 Sphaerocoma hookeri'T. Anderson Fig. 18.lCerdiacongestiflora ............................. 296 18.2 Haya obovata Balf. f. Fig. 19.1 Spergu.la arvensa.s L. 301 19.2 Spergularia rubra (L.) J. & C. Presi. Fig. 20.lTelephiumirnperatiL. ............................. 301 20.2 Krauseola mosambicina (Moss) Pax & Hoffm. Fig. 21.1 Scieranthus annuus L. ...... ...... .... ...... 308 21.2 S. biflorous (Forst.) Hook. f. 21.3 Habrosia spinuliflora (Ser. ex DC.) Fenzl. LIST OF PLATES Plate l.a Nyosoton aq.uatici (L.) Noench .................. 67 l.b Spergula arvensis L. 1.c Spergularia rubra (L.) J. & C. Presi. l.d Stellaria media (L.) J. & C. Presi. 1,e Nyosoton aquatic nl (L.) Moench Plate 2 AgrostemmagithagoL.............................. 135 Plate 3 AgrostemmagithagQL. ........................... 145 Plate 4 AgrosteminagithagoL. ........................... 147 Plate 5 Lychnit coronaria (L.) Desv. ................... 156 Cucubalus baccifer L. Silen? coeli-rosa (La) Godron Plate 6 Silene coeli-rosa (L) Godron ................... 163 Plate Silene conoidea L. ............................. 169 Plate S SperguJ.aria media (L.) C. Presi. ............... 174 S. rubra (L.) J. & C. Presi. Plate 9 Spergalaria media (L.) C. Presi. ............... 175 Plate 10 Polycarpon tetraphyllum (L.) L. ................ 180 Drymaria cordata Wilid. ex Roem. & Schult. Plate 11 Sclera nthusannuusL. 188 Corrigiola litoralis L. Plate 12 HerniariaglabraL. ............................ 190 Plate 13 Illecebrum verticillatum L. .................... 193 Paronychia echinulata Chater. LIST OF TAMES Table 1 Genera listed according to style number .......... 26 Table 2 Percentage of genera with 2, 3, 4 or 5 styles .... 27 Table `3 Style categories 400-46 a•aa.. • . a • • a a • • a • a • a . • • • • 30 Table 4 Percentage of genera in each style category ....... 33 Table 5 Style number compared to style length and ovule number ...............................•...•• 33 Table6 Two fused styles ..............................••• 34 Table 7 Three fused styles ............................... 34 Table. 8 Two free styles ................................•. 35 Table 9 Three free styles . 35 Table 10 Four free styles .......*•se••••s•• 35 Table 11 Five free styles . a . • • • . 36 Table 12 Results for Stellaria/ media ...................... 40 Table 13 Ace, 30, Stellaria media •........................ 40 Table 14 Stellaria media, 4 styles ........................ 43 Table 15 Results for Stellaria graminea •.................. 44 Table 16 Results for Stellaria holostea ................... 48 Table 17 Stellaria holostea , 2 styles .................... 50 Table 18 Ace. 82,Nyosotonaquaticum •..................... 51 Table 19 Results for Myosoton aguaticum ................... 52 Table 20 6 styles ..............................a........s. 55 Table 21 7 styles •..••.•S•..S•S.....•••S.•..•.....•.S..e•• 55 Table 22 4 styles •.................................•...... 55 Table 23 Total results for Stellaria/yosotonN ............. 56 Table 24 Aces 321, Spergularia media ...................... 59 Table 25 Results for Spergularia media .................... Go Table 2 6.. Ace. 387, Spergularia marina, results. ........... 64 Table 27 Ace. 314, Spergularia rubra ...................... 68 Table 28 Results for Spergularia media .................... 6 9 Table 29 Ace. 20, Spergula arensis ...................... 72 Table 30 Results for Spergula arvensis ................... 74 Table 30A 4 styles •...................a.ss..•.s•t.s•••••a• 76 Table 31 Ace. 124, Spergula morisonhi ..................... 77 Table 32 Results for Spergula morisonii ................... 78 Table 33 4 styles •....a..•.....as.a••.•s....ae.eae•aes•et- 81 Table 34/ Table 34 Total results for Sperlaria/Sperla ........... 82 Table 35 Experiment 1 Silene coeli-rosa •................. 87 Table 36 Experiment S. coeli-rosa •..................... 88 Table 37 Experiment