The evolution of fruit in Scandiceae subtribe Scandicinae (Apiaceae) Krzysztof Spalik, Aneta Wojewódzka, and Stephen R. Downie Abstract: Evolutionary relationships among 66 representatives of the family Apiaceae, including 37 species of tribe Scandiceae subtribe Scandicinae, were inferred from separate and combined analyses of fruit morphology and anatomy and nuclear ribosomal DNA internal transcribed spacer (ITS) sequences. Phylogenetic trees inferred from analysis of 35 fruit characters were not congruent to those derived from molecular data and, overall, had much lower bootstrap sup- port values than the latter. Contrary to molecular data, fruit characters did not support the monophyly of subtribe Scandicinae. Fruit data do, however, corroborate the monophyly of nearly every genus within Scandicinae, the affinity of members of the “crown” clade—Anthriscus, Kozlovia (including Krasnovia and Neoconopodium), Geocaryum, Myrrhis, and Osmorhiza—that had been identified in previous molecular analyses, and the sister group relationship be- tween the “crown” clade and the genus Scandix. Phylogenies derived from the analysis of combined ITS and fruit char- acters were congruent to those inferred from molecular data alone. Reconstructions of ancestral character states using the results of the combined analysis suggest that among extant Scandicinae, the fruits of Athamanta have retained the most plesiomorphic characters. Evolutionary tendencies that have occurred in the fruits of Scandicinae include the broadening of the vascular bundles and vittae, the thickening of the cuticle and epidermal cell wall, the origin of bris- tles from hairs, the appearance of a pedicel-like appendage, the development of a long beak, and lateral wings. These changes are interpreted as adaptations to fruit dispersal and seed defense. Key words: Apiaceae, Scandiceae subtribe Scandicinae, ITS, fruit morphology. Résumé : Parmi 66 représentants de la famille des Apiaceae, incluant 37 espèces de la tribu des Scandiceae sous-tribu Scandicinae, les auteurs ont déduit les relations évolutives, en utilisant des analyses séparées et combinées de la mor- phologie et de l’anatomie des fruits, ainsi que des séquences de l’espaceur interne transcrit (ITS) de l’ADN ribosomal nucléique. Les dendrogrammes phylogénétiques, obtenus de l’analyse de 35 caractères des fruits, sont incongrus par rapport à ceux dérivés des données moléculaires et, en général, montrent des valeurs de support en lacet beaucoup plus faibles que ceux-ci. Contrairement aux données moléculaires, les caractéristiques des fruits ne supportent pas la mono- phylie de la sous-tribu Scandicinae. Cependant, les données sur les fruits corroborent la monophylie d’à peu près tous les genres appartenant aux Scandicinae, l’affinité des membres du clade “crown”—Anthriscus, Kozlovia (incluant Kras- novia et Neoconopodium), Geocaryum, Myrrhis et Osmorhiza—qui ont été identifiés dans des analyses moléculaires précédentes, ainsi que la relation de sororité entre le clade “crown” et le genre Scandix. Les phylogénies dérivées de l’analyse combinée des caractères des ITS et des fruits sont congruentes avec celles obtenues à partir des données mo- léculaires prises isolément. Les reconstructions des caractères ancestraux, basées sur les résultats d’analyses combinées, suggèrent que parmi les Scandicinae actuelles, les fruits de l’Athamanta ont retenu les caractères les plus plésiomor- phes. Les tendances évolutives qui se sont manifestées chez les fruits des Scandicinae incluent l’élargissement des fais- ceaux vasculaires et des vittae, l’épaississement de la cuticule et de la paroi des cellules épidermiques, l’origine des soies à partir des poils, l’apparence de l’appendice pédicelloïde, le développement d’un long bec, et les ailes latérales. On inter- prète ces changements comme des adaptations pour la dispersion des fruits et la protection des graines. Mots clés : Apiaceae, Scandiceae sous-tribu Scandicinae, ITS, morphologie des fruits. [Traduit par la Rédaction] Spalik et al. 1374 Introduction Ever since Morison’s (1672) Plantarum umbelliferarum, fruit morphology and anatomy have been regarded as essen- tial to the taxonomy of Apiaceae (Umbelliferae; Drude 1898; Constance 1971). Despite their general similarity, umbellifer fruits vary with respect to their external and inter- nal features, and nearly all classification systems of the fam- ily are based on these characters. Koch (1824), for instance, divided the family into two principal groups, Multiiugatae and Pauciiugatae, on the basis of the number of ribs. He sub- sequently segregated these groups into 15 tribes upon con- sideration of fruit shape and compression and characteristics of the ribs and vittae. de Candolle (1830) stressed the impor- tance of endosperm shape, arranging the umbellifers into three groups: Orthospermae, Campylospermae, and Coelospermae. Bentham (1867) gave priority to the number Can. J. Bot 79: 1358–1374 (2001) © 2001 NRC Canada 1358 DOI: 10.1139/cjb-79-11-1358 Received March 7, 2001. Published on the NRC Research Press Web site at http://canjbot.nrc.ca on November 16, 2001. K. Spalik 1 and A. Wojewódzka. Department of Plant Systematics and Geography, Warsaw University, Aleje Ujazdowskie 4, 00-478 Warsaw, Poland. S.R. Downie. Department of Plant Biology, University of Illinois at Urbana-Champaign, Urbana, IL 61801, U.S.A. 1 Corresponding author (e-mail: spalik@bot.uw.edu.pl). Can. J. Bot. Downloaded from www.nrcresearchpress.com by Canadian Science Publishing on 06/08/15 For personal use only.