Introduction
The sedges (Cyperaceae) constitute a large, morphologically diverse, geographically widespread, and ecologically and economically important family. The family contains more than 5,000 species in 108 genera (Goetghebeur, 1998; Simpson et al., 2005). Despite the importance of sedges, understanding of phylogeny in the Cyperaceae is still in its infancy. Recently, several phylogenetic analyses based on molecular data have advanced knowledge of relationships within the family (Muasya et al., 2009 for this volume). However, phylogenetic analyses based on morphologic data remain scarce for sedges. Such analyses are daunting because of the number and severity of obstacles to using morphology for phylogenetic reconstruction in sedges.
Seven problems are often cited as formidable obstacles to inferring sedge phylogenies from morphology. First, identifying an adequate number of phylogenetically valuable characters is a problem, because sedges are relatively morphologically reduced, and many species are morphologically similar (Goetghebeur & Van den Borre, 1989; Gonzalez-Elizondo et al., 1997; Starr et al., 2004). Second, correctly determining homologies of morphologic structures, particularly those that are reduced or unique to the family, is a difficult endeavor with an often uncertain result (Reznicek, 1990; Bruhl, 1995; Muasya et al., 1998; Yen & Olmstead, 2000; Simpson et al., 2003). Incorrect assessment of homology could result in errors of character recognition and definition, as well as errors in polarization of character states. Such errors would be especially serious if few characters are available for phylogenetic inference. Third, existing infrageneric groups in large genera (e.g., Carex L., Cyperus L.) are poorly resolved and often not monophyletic (Crins, 1990; Reznicek, 1990; Muasya et al., 2002; Ford et al., 2006). Fourth, selecting appropriate outgroups for character state polarization is hampered by the lack of understanding of infrafamilial phylogenetic relationships (Thomas, 1984; Crins, 1990; Yen & Olmstead, 2000). Fifth, the fossil record is poor, rendering direct polarization of character states nearly impossible (Thomasson, 1983; Crepet & Feldman, 1991; Goetghebeur, 1998). Sixth, ontogeny of sedges is also relatively poorly known (Alexeev, 1988; Richards, 2002). Seventh, in some groups of sedges, hybridization is frequent, and hybrids are at least partially fertile (Standley, 1990; Cayouette and Catling, 1992). In such groups, hybridization must be recognized as a phenomenon that could obscure phyletic relationships.
Because of the problems perceived in using morphology to reconstruct phylogeny of sedges, few authors have attempted to reconstruct relationships using such characters. In all, only 11 publications contain phylogenetic analyses of members of Cyperaceae based on morphologic data. These few analyses vary greatly in methodology, taxonomic breadth, and size of ingroup. Thomas (1984) used the groundplan divergence method to reconstruct the phylogeny of 25 species and varieties of Rhynchospora Vahl section Dichromena (Michx.) Griseb. In this analysis, Thomas used outgroup comparison to determine the polarity of character states, in an attempt to avoid character state polarization, Crins and Ball (1988) and Crins (1990) used character compatibility analysis to infer phylogenies for Carex sects. Ceratocystis Dumort., Limosae (Heuff.) Meinsh., and Phyllostachyae Kuk. They conducted a separate analysis for each section, treating a total of 23 species and varieties. Parsimony was the algorithm for the remainder of the phylogenetic analyses, all of which employed outgroup comparison for polarization of character states: Seberg (1988) for 17 species and subspecies of Oreobolus R. Br. (generic revision); Simpson (1992) for 71 species of Mapania Aubl. (generic revision); Bruhl (1995) for 108 genera and generic segregates (tribal and subfamilial revision of Cyperaceae); Simpson (1995) for 12 genera representative of the family (assessment of Cyperales relationships); Gonzalez-Elizondo et al. (1997) for 29 species and varieties of Eleocharis R. Br. (Eleocharis sect. Pauciflorae Beauverd circumscription); Muasya et al. (2000) for 61 genera representative of the family (examination of tribal and subfamilial relationships); Naczi and Ford (2001) for three species of the Carex jamesii Schwein. complex (assessment of relationships within the complex); and Muasya and Simpson (2002) for 46 taxa of Isolepis and related genera (generic revision).
In order to evaluate the gravity of the off-cited obstacles to phylogenetic analysis of sedges, I used morphologic data to reconstruct phylogeny of sedge groups additional to those previously studied. To employ as many phylogenetically useful characters as possible, I attempted to discover novel characters when studying morphology in a broad sense (macromorphology, micromorphology, and anatomy). To permit comparisons and more robust conclusions, I inferred phylogeny for two disparate groups, one a member of Carex subgenus Carex, the other Carex subgenus Vignea (Lestib.) Peterm. In this paper, I report the results of these analyses, and discuss their implications for assessing relationships and refining classifications in the specific sedge groups. Furthermore, I compare the phylogenetic trees from these analyses to those from other groups of flowering plants, including sedges, in order to assess the effectiveness of morphologic data in reconstructing sedge phylogenies.