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 Torilidinae, with the former including representation of tribe Laserpitieae sensu Drude. These taxa are allied with the monophyletic Scandiceae subtribe Scandicinae whose members lack spiny fruits. The relationship among these three subtribes is equivocal when nuclear ribosomal DNA internal transcribed spacer 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 in order to study the evolution of these characters and to ascertain their utility for resolving relationships by comparison to the results of previous molecular analyses. Maximum parsimony analysis of 56 morphological characters resulted in a paraphyletic Torilidinae (Astrodaucus, Caucalis, Glochidotheca, Lisaea, Szovitsia, Torilis, Turgenia, and Yabea) from which a monophyletic Daucinae (Agrocharis, Ammodaucus, Cuminum, Daucus, Laserpitium, Orlaya, Pachyctenium, and Pseudorlaya) are derived. Scandicinae are maintained as monophyletic, sister to Daucinae plus Torilidinae. The genus Artedia, previously attributable to either Daucinae or Torilidinae, shows affinity with the former. The Daucinae plus Torilidinae clade is supported by three fruit synapomorphies—the regular distribution of appendages on both primary and secondary ridges, the presence of primary ridges and hairs on the face of the commissure, and the presence of vittae under the lateral ridges—but each subtribe cannot be circumscribed unambiguously on the basis of morphological data. Characters of the primary appendages exhibit less homoplasy than those of the secondary fruit appendages and support many clades identified in the molecular analyses. Parsimony analysis of combined morphological and ITS data, however, reaffirms the monophyly of subtribe Torilidinae and provides greater resolution of relationships within each of the subtribes than do either of the separate analyses.

Communicating Editor: Paul Wilson