Ninety-three taxa comprising thirty-two genera (plus four outgroups from Lobeliaceae) were used to estimate a phylogeny of the Campanulaceae based on ITS sequences of nuclear ribosomal DNA. From 2629 most parsimonious trees, a strict consensus tree with bootstrap values was constructed, in addition to a phylogram showing branch lengths. The topologies of these two trees are discussed in relation to the pollen and capsule morphology within the family, in addition to chromosome number and geographical distribution. The results show that there is a major dichotomy between the colpate/colporate pollen alliance (platycodonoid taxa) and the porate pollen alliance (wahlenbergioid and campanuloid taxa). Both these major alliances are monophyletic. Within the porate alliance there are two major clades, the wahlenbergioids and the campanuloids. The campanuloid clade is further subdivided into two major clades representing the Rapunculus and the Campanula s. str. groups of taxa, plus three smaller clades that are considered as ''transitional'' taxa. It is argued that the family originated in a fragmenting West Gondwanaland and that tectonic processes are responsible for the major dichotomy in the family. The colpate/colporate platycodonoids subsequently remained relatively relictual in Asia, whereas the porate taxa spread over much of the Northern and Southern Hemispheres. The campanuloid lineage spread over the Northern Hemisphere from a major evolutionary center in the Mediterranean region and is represented in North America only by the Rapunculus group. The wahlenbergioid lineage is widely dispersed across the southern continents and oceanic islands but has a major secondary center of diversification in southern Africa. The use of ITS provides insights for future investigations and a phylogenetic framework that can be tested with other data sets. Its limitations for phylogeny reconstruction are briefly discussed. More extensive taxon sampling and additional data sets are required to refine these results and for a new classification of the Campanulaceae to be proposed.
Cainpanulaceae herein are treated in a narrow niodate \lci,ir:a (IV Candolle. 1859). Schoiiland circumscription (without bobeliaceae) as a mono-( 1889) also divided the tribe Campanuloideae phyletic group with a distinct geographical disln-(Cainpanulaceae s. str. here) into three groups, hulion and with well-delined morphological char-based on mode of dehiscence and man position, acters. Campanulaceae s. str.. despite their size and hut these three group-dill, red in composition from importance in temperate floras, remain unrevised. those of l)e Candolle. These two classifications be-This family, with about 50 genera and 800 species came the basis (or all future treatments (Table I). distributed worldwide, is the largest and most prim-Although the current systems differ greatly from the (Lammers, 1992; Takhtajan, 1997). Although rep-been added during the last century, it is easy to resentatives of the family occur on all continents understand what classification each particular auexcept Antarctica, the vast majority of genera and thor is following. Schonland's treatment has been species are found in temperate regions of the Old used often and remains a currently useful refer-World. Haven and Axelrod (1971) considered the ence. family to have a baurasian-African origin. The cen-Fedorov (1957). on the contrary, followed in genlers ol dislrihiitiou and diveisilv include the Med-eral De Candol|e"s position and published a deiterranean, Kasl Asia, and South Africa (Shulkina. tailed classih, .iti.ui I,., ( .ampaiiulaceae growing in 1978; Kolakovsky. 1995; Hong. 1995; Kddie. the former Soviet Union (FSU). Fedorov proposed l ' ,< '' •â-8 tribes ((> new) based on ca|)sule dehiscence, colie's (1850) comprehensive monograph rolla shape and pre>en, . and sha|ie ..I appendages provided a solid basis for between the calyx lobes. Kolakovsky (1995) proall subsequent works. Me divided the family into posed a new system with 4 subfamilies and 22 two tribes and later added a third tribe to accom-tribes based on internal fruit structure. He 1 We are indebted to Ihsan M-Shehhaz. I Vie, Stevens, anil IVler Much f„r ..-..(ling the text and providing useful remarks. Main thank-I,. \ i<-|.,.ia IMI.mell I,., caiel.il editing. T. <|„dkina lliank-llie people who helped her ohlain Cainpanulaceae seeds from different farts ol the world: J. !'. \1. Ureiian. \. Crimquist. \. Dolukhanov. K Oagmd/e K. Kamelin, A. kolakoxskv. \. Moiin. II. I!, lis, â-roll. ami |> Wendelbo. '\li-sonri liolam.al ( .aid.n. I'.O K,,x 2<>>>. <\. I,,,,,-. \1,-„,iii tU I (,<>( rJ09. I >. \. t , lh ana.-lmlkma<« iih.I ... 'Present address: I Sl)\. \KS-\I'\KL I'.O. Pox l(,;. Sidm-v. Montana 59270, U.S.A.' ' Section of Integrative liiolog). Universitv ol lexa-at \uslm. Texa-7i!7ll>. I S. V. IVe-enl address: Office of Lifelong learning, University ol Kdml.urgh. I I liuccleiicli I'laee. Kdinhurgh. Scotland. I k. weddie I («staffmail.ed.ac..ik.
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