419 Jabaily & al. • Systematics of Goodeniaceae TAXON 61 (2) • April 2012: 419–436 INTRODUCTION The Goodeniaceae, with ca. 420 recognized species, are a conspicuous part of the Australian and Pacific Island floras. This family, the 9th-largest in Australia with over 400 spe- cies and seven of the twelve recognized genera endemic to the continent (Table 1; Fig. 1), contributes greatly to the spring and summer wildflower displays particularly in the Southwest Aus- tralian Floristic Region (SWAFR, George & al., 1979; Lamont & al., 1984; Hopper & Gioia, 2004), as well as being of signifi- cant horticultural value. Goodeniaceae herbs, subshrubs and shrubs are found in most vegetation types and biogeographic zones, from the northern monsoon tropics, to the arid Eremaean interior to the seasonally dry southwest and the cool temperate areas of Victoria and Tasmania. Three genera ( Goodenia Sm., Dampiera R. Br., Scaevola L.) comprise the vast majority of species. Scaevola spans the greatest geographical space and is the only genus with a significant diversity outside of Aus- tralia (ca. 35 species), including eight endemic species in New Caledonia (Müller, 1990) and ten in Hawaii (Patterson, 1990; Wagner, 1996). Scaevola taccada and S. plumieri are the most widespread species found in coastal habitats throughout the tropics and subtropics. The principal morphological synapomorphy for the Goode- niaceae is the stylar indusium, a cup-like or bilabiate append- age at the style apex that collects and packs pollen as it pushes past the stamens during development in bud (Fig. 2A). At an- thesis, the stamens are withered and the pollen mass placed within the indusium is presented to pollinators. The stigmatic lobes subsequently emerge from within the indusium. Darwin was fascinated by the unusual pollination mechanisms of the Goodeniaceae but he was somewhat frustrated in his experi- ments with the enigmatic genus Lechenaultia R. Br., reflected in a letter to Hooker “… the Goodeniaceae have weighed like an incubus for years on my soul” (Darwin, 1860). It took Darwin many years to determine that the stigmatic region in Lech- enaultia is located adjacent to its unique two-lipped indusium (Fig. 2B), a mechanism that promoted outcrossing (Darwin, Systematics of the Austral-Pacific family Goodeniaceae: Establishing a taxonomic and evolutionary framework Rachel S. Jabaily, 1,6 Kelly A. Shepherd,2 Mats H.G. Gustafsson,3 Leigh W. Sage,2 Siegy L. Krauss, 4 Dianella G. Howarth5 & Timothy J. Motley1 1 Department of Biology, Old Dominion University, Norfolk, Virginia, U.S.A. 2 Department of Environment and Conservation, Kensington, Western Australia; and School of Plant Biology, University of Western Australia, Nedlands, Western Australia, Australia 3 Department of Biological Sciences, Ecoinformatics and Biodiversity, Aarhus University, Aarhus, Denmark 4 Kings Park and Botanic Garden, Botanic Gardens and Parks Authority, Perth, Western Australia, Australia; and School of Plant Biology, University of Western Australia, Nedlands, Western Australia, Australia 5 Department of Biological Sciences, St. John’s University, Queens, New York, U.S.A. 6 Department of Biology, Rhodes College, Memphis, Tennessee, U.S.A. Author for correspondence: Rachel Jabaily, jabailyr@gmail.com Abstract The Goodeniaceae, close relatives of the Asteraceae, are a conspicuous part of the flora of Australia and many islands in the Pacific. A comprehensive molecular phylogenetic analysis of the family using cpDNA regions trnL-F and matK is pre- sented, including representatives of all genera and nearly half the species. The family resolves into the two large clades: ‘LAD’ (Lechenaultia, Anthotium, Dampiera) and ‘Core Goodeniaceae’ (Brunonia, Scaevola, Diaspasis, Coopernookia, Goodenia, Selliera, Velleia, Verreauxia, Pentaptilon), which are also supported by morphological characters. The former Brunoniaceae, comprising the single species Brunonia australis, is clearly placed within the Goodeniaceae as sister to the remainder of Core Goodeniaceae while possessing many autapomorphic characteristics. Current subgeneric taxonomic groups are partially sup- ported as monophyletic within Goodenia and Dampiera, but are generally non-monophyletic within Lechenaultia and Scae- vola. Fruit types and floral traits involved in pollination and adaptations to aridity have evolved in a highly convergent fashion within several major clades. Rates of molecular evolution and number of extant taxa differ at several points between sister clades, particularly between Brunonia and the rest of Core Goodeniaceae, and between Scaevola s.l. and Goodenia s.l. Future taxonomic changes will involve the synonymization of Selliera and monotypic Diaspasis and, pending more comprehensive taxon and molecular sampling, the large, paraphyletic genus Goodenia may be split into at least three genera. Keywords Asterales; Australia; Goodeniaceae; Hawaii; molecular systematics; Pacific Islands; SWAFR; taxonomy Supplementary Material The Appendix (in the Electronic Supplement) and the alignment files are available in the Supplementary Data section of the online version of this article (http://www.ingentaconnect.com/content/iapt/tax).