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The Dicot Order, Caryophyllales

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The Dicot Order, Caryophyllales lnterfamilia/Relationships of Cactaceae within the Dicot Order, Caryophyllales by James E. Oliver A Thesis Submitted to the Faculty of the Charles E. Schmidt College of Science in Partial Fulfillment of the Requirements for the Degree of Master of Science Florida Atlantic University Boca Raton , Florida December 1998 lnterfamilial Relationships of Cactaceae ~vithin the Dicot Order, Caryophyllales by Jarne.s E. Oliver Tt·ds thesis was prepamd ur.dor the direction of the candidate's thesis advisor, Dr. David M. Binninger, Department of Bioiogic:a: SGi ence, and has t-,efm c..pproved by members of his supervisoiy comm ittAG lt wa.s submitted to th0 f;:~cu!ty of t;·,e Cr,ar ies E. Schmidt College of Sc1ence and was accepmd as ~~ ailial fulfillment of the requirements for the dsg:-ee of Master of Sc;snce. S UPERVIS08'r' COMiVliTTc~ : 1-Jd.-70 ---'---~ - · -.. - ~--~. · --·- ·· ·- -· - J O::;, t ~ il Acknowledgments I would like to thank my advisors, Dr. David Binninger, Dr. Daniel Austin and Dr. Ralph Adams for their patience and guidance in enabling me to complete this project. I also want to give special thanks to Chris Doughtery for his work in generating the genetic sequence data that made my research possible. ill Abstract Author: James E. Oliver Title: lnterfamilial Relationships of Cactaceae within the Dicot Order, Caryophyllales Institution: Florida Atlantic University Thesis Advisor: Dr. Daniel F. Austin and Dr. David M. Binninger Degree: Master of Science Year: 1998 The position of the cactus family, Cactaceae, within the order Caryophyllales was examined by outgroup analysis of chloroplast rbcL gene sequence data. Comparative data came from 28 outgroup species in 15 families and six ingroup species. Phylogenetic Analysis with P AUP 3.1 produced twelve equally parsimonious trees; these were used to generate a strict consensus tree. MacClade 3.06 was used to refine the polytomies in the consensus tree. Analysis indicates that the Cactaceae is a monophyletic clade; moreover, these results support the work of other analyses that the sister family to Cactaceae is the Portulacaceae. Other relationships within the order are not completely consistent with previous studies. For example Molluginaceae and Caryophyllaceae occupy a central position in the cladogram; Achatocarpaceae aligns with the Amaranthaceae and Chenopodiaceae; and Basellaceae is presented as the basal family within the lV order. Lastly, the Jamaican cactus Opuntia spinosissima was derived from a common ancestor with the Florida Keys endemic, Opuntia corallicola. Key Words: Cactaceae, Caryophyllales, chloroplast rbcl gene, polytomies, monophyletic v Table of Contents Page List of Tables .................................................................................................................. vii L1st. ofF' 1gures ............................................................................................................... Vlll... Introduction ...................................................................................................................... ! Hypothesis ........................................................................................................................ 8 Materials and Methods ................................................................................................... 9 Results ............................................................................................................................. lO Discussion ....................................................................................................................... 18 Literature Cited ............................................................................................................. 23 Vl List of Tables Page Table 1. Taxa, Sources and Accession Numbers for rbcL Data ......................... 3 Vll List of Figures Page Figure 1. Cronquist's 1981 Arrangement of Families Within the Caryophyllales ............................................................................................. 6 Figure 2. Strict Consensus of 12 Equally Parsimonious Trees Generated with PAUP 3.1 ....................................................................... 12 Figure 3. MacClade 3.06 Soft Polytomy Resolution of Strict Consensus Tree .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ... 14 Figm·e 4. MacClade 3.06 Hard Polytomy Resolution of Strict Consensus Tree ......................................................................................... 16 viii To my family and friends, whose faith in me and support made this possible. lX Introduction Morphological, chemical, and molecular characters indicate that the dicot order, Caryophyllales, comprised of 12 families (Table 1) and ca. 10,000 species, is monophyletic (Cronquist 1988; Downey and Palmer 1994; Downey et al. 1997). Past investigations have distinguished the order on the basis of specific embryological features (Cronquist 1988); the presence of unique sieve tube plastid elements; and the replacement of anthocyanin pigments with the class of pigments called betalains (Gibson and Nobel1986). Another widespread but unique characteristic of the order, though not present in Cactaceae, Portulacaceae, or Didieraceae, is anomalous secondary thickening of successive cambia (Gibson and Nobel1986). Nevertheless, the position of Cactaceae and its sister group is still debated (Downey and Palmer, 1994; Hershkovitz and Zimmer, 1997). Cronquist's (1981) arrangement of the order placed Caryophyllaceae and Molluginaceae basally with the remaining 11 families derived from Phytolacaceae (Figure 1). Cronquist (1988) grouped Cactaceae with Aizoaceae and Didieraceae as a trichotomy and aligned Portulacaceae with Basellaceae (Figure 1). Some evidence suggests that the sister taxon to the Cactaceae is either the Portulacaceae (Gibson and Nobel1986; Downey and Palmer 1994; Hershkovitz and Zimmer 1997), Aizoaceae (Rodman 1984) or 1 Didiereaceae (Gibson and Nobel1986). Other studies put Cactaceae and Didiereaceae in an alliance of Aizoaceae, Portulacaceae and Basellaceae (Rodman 1984). Many of the uncertainties regarding the interfamilial relationships within Caryophyllales may be attributed to the various methods employed in arranging the families (Downey and Palmer 1994). This variation was demonstrated by Dougherty (1996) in which chloroplast rbcL gene sequence data were used to construct phylogenies to delineate the species status of an endangered cactus. The rbcL gene is useful because it contains >1400 base pairs for comparative information and its rate of evolution is appropriate for phylogenies at the subfamiliallevel and higher (A vise 1994; Bayer et al. 1996). However, certain representative species of Caryophyllales failed to align with those of Cactaceae according to traditional taxonomic arrangements (Dougherty 1996). The apparent grouping of cacti away from other species of the order suggests that cacti represent a separate clade (Dougherty 1996). 2 Table 1. Taxa, Sources, and Accession Numbers for rbcl Data Used in the Study. 3 Taxa Accession 1 Outgroupllngroup Source Number Family Genus Species 1 Achatocarpaceae I Phaulothamnus spinescens GenBank M97887 Outgroup 2 Aizoaceae I Lithops ~ GenBank M97889 Outgroup 3 Aizoaceae I Mesembryan- crystallinum GenBank M98515 Outgroup themum 4 Aizoaceae Trianthema portulacastrum GenBank M62572 Outgroup 5 Amaranthaceae Amaranthus hypochondriacus GenBank X51964 Outgroup 6 Amaranthaceae Amaranthus tricolor GenBank X53980 Ou!grOUIJ 7 Basellaceae Base II a alba GenBank M62564 Outgroup *"" 8 CC!rYopl}yllaceae Arenaria drummondii GenBank M83541 Outgroup 9 Caryophyllaceae Cerastium glomeratum GenBank M83542 Ou!group 10 Caryophyllaceae Silene gallica GenBank M83544 Outgroup 11 I Caryophyllaceae Dianthus caryopl"lyll us GenBank M77699 Ou!_g_roup 12 I Chenopodiaceae A triplex petula GenBank M33795 Outgroup 13 I Cheno_Qodiaceae AtrlQiex rose a GenBank M33794 Outgroup 14 I Chenopodiaceae Spinacia oleracea GenBank J01443 Outgroup V00168 15 Didiereaceae Alluaudia pro cera GenBank I M62563 Ou!grQI.!P 16 Moll~naceae Mollugo verticillata GenBank I M62566 Outgroup 17 Nyctaginaceae Bougainvillea _g!abra GenBank I M88340 Outgroup 18 Nyctaginaceae Mirabilis jalapa ! GenBank I M62565 Outgroup Taxa Source Accession Outgroupllngroup Family Genus Species Number 19 Ph_ytolaccaceae Gisekia pharnacioides GenBank M97890 Outgroup 20 Phytolaccaceae Phytolacca americana GenBank M62567 Outgroup 21 Phytolaccaceae Rivina humilis GenBank M62569 Outgroup 22 Ph_ytolaccaceae Stegnosperma halimifolium GenBank M62571 Outgroup 23 Plumbaginaceae Plumbago capensis GenBank M77701 Outgroup 24 Polygonaceae Rheum xcultorum GenBank M77702 Outgroup 25 Portulacaceae Portulaca grandiflora GenBank M62568 Outgroup 01 26 Sarraceniaceae Darlingtonia californica GenBank L42211 Outgroup 27 Sarraceniaceae Heliamphora nutans GenBank L02433 Outgroup 28 Sarraceniaceae Sarrracenia flava GenBank L01952 Outgroup 29 Cactaceae Cereus pentagonus Daugherty FAU OP-4 lngroup 30 Cactaceae Opuntia corallicola Daugherty FAU OP-9 lngroup 31 Cactaceae O_Q_untia s_pinosissima Daughe_rty FAU OP-8 lngroup 32 Cactaceae Opuntia spinosissima Daugherty FAU OP-23 lngroup 33 Cactaceae Pereskia aculeata Genbank M97888 lngroup 34 Cactaceae Schlu'!lbe_r-_ger'!_ truncata Genbank M83543 L__ lngroup ·-·· _ - ---· - - --· - ·-- - - -- Figure 1. Cronquist's 1981 arrangement of families within the Caryophyllales 6 r--------~~----~-----~~-----~--~--~, I I : s. Dldiereaceae CARYOPHYLLALES I I I I I I I I 4. Alzoaceae 8. Cactaceae J I I I I ~ ;{ 7. Cbonopodlac.
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