Relationships within the spiny-fruited umbellifers (Scandiceae subtribes Daucinae and Torilidinae) as assessed by phylogenetic analysis of morphological characters

Previous molecular systematic studies have indicated that the spiny-fruited umbellifers (Apiaceae tribe Caucalideae sensu Heywood) comprise two major lineages, recently delimited as Scandiceae subtribes Daucinae and Torilidin ae, with the former including representation of tribe Laserpitieae sensu Dr ude. These taxa are allied with the monophyletic Scandiceae subtribe Scandi cinae whose members lack spiny fruits, The relationship among these three s ubtribes is equivocal when nuclear ribosomal DNA internal transcribed space r sequences are compared. Evidence from plastid DNA, however, suggests that Daucinae and Torilidinae are sister taxa. Herein, we provide results of a phylogenetic study of these spiny-fruited umbellifers based on morphology i n order to study the evolution of these characters and to ascertain their u tility for resolving relationships by comparison to the results of previous molecular analyses. Maximum parsimony analysis of 56 morphological charact ers resulted in a paraphyletic Toriliclinae (Astrodaucus, Caucalis, Glochid otheca, Lisaea, Szovitsia, Torilis, Turgenia, and Yabea) from which a monop hyletic Daucinae (Agrocharis, Ammodaucus, Cuminum, Daucus, Laserpitium, Orl mia, Pachyctenium, and Pseudorlaya) are derived. Scandicinae are maintained as monophyletic, sister to Daucinae plus Torilidinae. The genus Artedia, p reviously attributable to either Daucinae or Torilidinae, shows affinity wi th the former. The Daucinae plus Torilidinae clade is supported by three fr uit synapomorphies-the regular distribution of appendages on both primary a nd secondary ridges, the presence of primary ridges and hairs on the face o f the commissure, and the presence of vittae under the lateral ridges-but e ach subtribe cannot be circumscribed unambiguously on the basis of morpholo gical data. Characters of the primary appendages exhibit less homoplasy tha n those of the secondary fruit appendages and Support many clades identifie d in the molecular analyses. Parsimony analysis of combined morphological a nd ITS data, however, reaffirms the monophyly of subtribe Torilidinae and p rovides greater resolution of relationships within each of the subtribes th an do either of the separate analyses.