Systematic Botany (2009), 34(1): pp. 182–197 © Copyright 2009 by the American Society of Plant Taxonomists

Phylogenetic Classification of Subtribe Castillejinae (Orobanchaceae)

David C. Tank,1,3, 4 J. Mark Egger, 2 and Richard G. Olmstead1,2 1 Department of Biology, University of Washington, Box 355325, Seattle, Washington 98195 U.S.A. 2 Herbarium, Burke Museum of Natural History, University of Washington, Box 355325, Seattle, Washington 98195 U.S.A. 3 Present Address: Department of Forest Resources and Stillinger Herbarium, University of Idaho, PO Box 441133, Moscow, Idaho 83844-1133 U.S.A. 4 Author for Correspondence ( [email protected] )

Communicating Editor: Lena Struwe

Abstract—Recent molecular systematic research has indicated the need for a revised circumscription of generic boundaries in subtribe Castillejinae (tribe Pedicularideae, Orobanchaceae). Based on a well-resolved and well-supported phylogenetic hypothesis, we present a for- mal reclassification of the major lineages comprising the Castillejinae. Prior to this treatment, subtribe Castillejinae included Castilleja (ca. 190 spp.), Cordylanthus (18 spp.), Orthocarpus (9 spp.), Triphysaria (5 spp.), and the monotypic genera Clevelandia and Ophiocephalus. In the clas- sification presented here, Orthocarpus and Triphysaria retain their current circumscriptions, Castilleja is expanded to include Clevelandia and Ophiocephalus , and Cordylanthus is split into three genera; a key to the genera as they are recognized here is provided. Two new combinations, Castilleja beldingii and Castilleja ophiocephala , are proposed within the expanded Castilleja . The concept of Cordylanthus is restricted to the 13 species formerly recognized as subg. Cordylanthus , while subg. Dicranostegia and subg. Hemistegia are elevated to genus level (Dicranostegia and Chloropyron, respectively). We resurrect the generic name Chloropyron for the halophytes previously recognized as subg. Hemistegia . Five new combinations are proposed for Chloropyron ( Chloropyron maritimum subsp. canescens, Chloropyron maritimum subsp. palustre, Chloropyron molle subsp. hispidum, Chloropyron palmatum, and Chloropyron tecopense ). In addition to the formal classification, we provide phylogenetic clade definitions for Castillejinae, each of the genera, and two additional clades that are not assigned formal ranks. Morphological characteristics used to recognize traditional groups are evaluated, and synapomorphies are discussed. Finally, the current infrageneric classifications for Castilleja and Cordylanthus are evaluated in light of the recent molecular phylogenetic analyses. Keywords— former Scrophulariaceae , hemiparasites , Lamiales , Lamiidae , revision , western North America.

With the explosion of molecular phylogenetic studies in the monotypic genus Gentrya did not differ enough from Castilleja last two decades, systematists have been better able to evaluate to warrant generic status, and it was reassigned to Castilleja traditional classifications in the context of an explicit phylo- as subg. Gentrya. Therefore, Castillejinae (sensu Chuang and genetic hypothesis. A molecular systematic study investigat- Heckard 1991) included six genera: Castilleja (ca. 190 mostly ing phylogenetic relationships within subtribe Castillejinae perennial species), Clevelandia (1 annual species), Cordylanthus G. Don (tribe Pedicularideae Duby, Oroban chaceae Vent.), (18 annual species), Ophiocephalus (1 annual species), Orth- based on data from chloroplast (cp) and nuclear ribosomal ocarpus (9 annual species), and Triphysaria (5 annual species). (nr) DNA regions, indicates the need for a revision of the clas- Traditional classifications place subtribe Castillejinae in sification of this group ( Tank and Olmstead 2008 ). Scrophulariaceae Juss. (e.g. Bentham 1846 , 1876 ; Wettstein As it is currently circumscribed, Castillejinae comprises 1891; Cronquist 1981; Takhtajan 1997). However, numer- ca. 220 hemiparasitic species in six genera distributed pri- ous molecular systematic studies have demonstrated that marily in western North America. The group was originally Scrophulariaceae, as it is traditionally recognized, represents proposed to include Castilleja Mutis ex L.f., Cordylanthus an unnatural assemblage of plants comprising more than seven Nutt. ex Benth., and Orthocarpus Nutt. ( Bentham 1846 ). Three distinct lineages of Lamiales (reviewed in Tank et al. 2006 ). monotypic Mexican genera, Clevelandia Greene, Gentrya One of these lineages, now recognized as Orobanchaceae s.l. Breedlove & Heckard, and Ophiocephalus Wiggins, were (APG II 2003), contains the obligately parasitic Orobanchaceae subsequently added to Castillejinae (Greene 1885; Wiggins and all of the hemiparasitic Scrophulariaceae traditionally 1933; Breedlove and Heckard 1970). With the exception of placed in tribes Buchnereae Benth. and Pedicularideae Duby Ophiocephalus , all of these taxa possess anthers with unequal of subfamily Rhinanthoideae Link (sensu Wettstein 1891 ). anther sacs that are unequally attached (i.e. the larger anther Furthermore, molecular systematic studies (e.g. Young et al. sac attached to the filament medially, and the smaller by its 1999 ; Wolfe et al. 2005 ; Bennett and Mathews 2006 ) place sub- apex), and this character was used by Bentham (1846) to tribe Castillejinae within a clade of New World taxa distributed define the group. Throughout the history of the subtribe, in North and South America. defining generic boundaries has been problematic, especially The evolutionary relationships of the major lineages com- between Castilleja and Orthocarpus , where numerous spe- prising subtribe Castillejinae was the focus of a recent molec- cies have been shuffled between the two genera (e.g. Gray ular phylogenetic investigation (Tank and Olmstead 2008). 1862 ; Watson 1871 ; Eastwood 1909 ; Jepson 1925 ; Keck 1927 ; This study explicitly addressed evolutionary hypotheses Chuang and Heckard 1991, 1992 ). The most recent treatment presented by Chuang and Heckard’s (1991) realignment of of Castillejinae ( Chuang and Heckard 1991 ), based on exten- Castillejinae, based on sequences from two cpDNA regions sive morphological and cytological study, departed from the (trnL/F and the rps16 intron) and two nrDNA regions (ITS traditional generic boundaries of Castilleja and Orthocarpus . In and the 3¢ end of ETS; Tank and Olmstead 2008 ). Sequences of this treatment, Orthocarpus subg. Triphysaria was restored to the four DNA regions were obtained from a broad sampling its original status as a genus and 12 other Orthocarpus spe- of all six genera comprising subtribe Castillejinae sensu cies were moved to Castilleja , forming sect. Oncorhynchus of Chuang and Heckard (1991), including 46 species of Castilleja , subg. Colacus (Chuang and Heckard 1991, 1992 ), and reducing complete sampling of Orthocarpus and Triphysaria , and 14 of Orthocarpus from 25 species (sensu Keck 1927) to only nine. the 18 species of Cordylanthus (Appendix 1), as well as three In addition, Chuang and Heckard (1991) concluded that the outgroups. In this study, Orthocarpus and Triphysaria each 182 2009] TANK ET AL.: CLASSIFICATION OF CASTILLEJINAE 183 formed well-supported monophyletic groups, representing the two monotypic genera Clevelandia and Ophiocephalus two separate evolutionary lineages within Castillejinae (Fig. 1). were derived from within Castilleja ( Fig. 1 ). The major- Chuang and Heckard (1991) were correct in their interpreta- ity of Castilleja species are perennial (ca. 170 species), and tion that Castilleja subg. Colacus sect. Oncorhynchus , which, these taxa, along with Clevelandia and Ophiocephalus , formed like Triphysaria , comprises species previously recognized a well-supported clade derived from a grade of annual as Orthocarpus , belonged with Castilleja ; however, these lineages belonging to subg. Colacus sect. Oncorhynchus . annuals did not represent a distinct clade within Castilleja This clade also included the enigmatic species Castilleja rac- (Fig. 1). Although all three of these groups were previ- emosa, which, prior to Chuang and Heckard’s (1991) revision, ously included in Orthocarpus (sensu Keck 1927), they were was isolated in the monotypic genus Gentrya . The final genus confirmed to be more closely related to other members of Castillejinae, Cordylanthus, was polyphyletic in all molec- of the subtribe than they were to each other, as suggested ular analyses; two of the subgenera (subg. Cordylanthus and by Chuang and Heckard (1991). Separate and combined subg. Hemistegia ) were each monophyletic, and the mono- analyses of the cpDNA and nrDNA regions also indicated typic subg. Dicranostegia formed the sister group to subg. that the large genus Castilleja was not monophyletic, because Hemistegia ( Fig. 1 ). In the present study we develop a revised classification of Castillejinae and reevaluate the morphological charac- ters traditionally used to recognize genera. The congruence of the independent data sets from both the chloroplast and nuclear genomes strongly supports the revised classification of Castillejinae proposed here. In an explicitly phylogenetic classification, only monophyletic taxa are recognized (Hennig 1966; Wiley 1979; de Queiroz 1988; de Queiroz and Gauthier 1990, 1992 , 1994 ); however, under the traditional Linnaean system, as governed by the International Code of Botanical Nomenclature (ICBN), taxa are ultimately defined by types and the ranks to which they are assigned, with no explicit ref- erence to evolutionary relatedness. In contrast, phylogenetic nomenclature provides clade definitions of taxon names that refer directly to inferred phylogenetic relationships among taxa (de Queiroz and Gauthier 1990; de Queiroz 1992; de Queiroz and Gauthier 1992, 1994 ). Most importantly, clas- sifications based on monophyly provide an evolutionary framework for the communication, storage, and retrieval of taxonomic information ( Farris 1979 ; Wiley 1979 ). A phyloge- netic classification of Castillejinae, including phylogenetic definitions for each of the major lineages, will convey the cur- rently available information regarding evolutionary relation- ships within the group.

Materials and Methods The best estimate of phylogeny, and thus, primary basis for classification of Castillejinae, is provided by the results of molecular phylogenetic anal- yses focused on relationships of the major lineages comprising subtribe Castillejinae ( Tank and Olmstead 2008 ). In this study, Tank and Olmstead (2008) developed a well-supported phylogenetic hypothesis for sub- tribe Castillejinae to explore patterns of phenotypic evolution, focusing primarily on the evolution of life history variation, but also considering morphological characteristics that have been important historically for cir- cumscribing taxonomic groups. A summary of the current evolutionary hypothesis for Castillejinae, based on the aforementioned molecular sys- tematic study, is shown in Fig. 1 and photographs of each of the genera as Fig. 1.A summary of the best estimate of phylogenetic relation- circumscribed here are shown in Fig. 2 . A synoptical overview to species ships for Castillejinae based on molecular data from the chloroplast and of the phylogenetic classification proposed here and a comparison to the nuclear genomes and analyzed with a range of analytical methods most recent taxonomic treatment of Castillejinae ( Chuang and Heckard ( Tank and Olmstead 2008 ). Numbers associated with branches repre- 1986, 1991 , 1992 ) and the major lineages comprising the subtribe is given sent Bayesian posterior probabilities (PP), maximum likelihood bootstrap in Appendix 1. percentages (ML), and parsimony bootstrap percentages (BS) for In addition to the traditional classification given here, as governed the major clades as reported by Tank and Olmstead (2008) . The height by the ICBN, we also give clade definitions for subtribe Castillejinae, of the clades with solid boundaries correspond approximately to the each of the genera comprising the subtribe, and two additional clades numbers of species comprising those clades. The height of the clade with not formally named or assigned ranks in the traditional classification the dashed boundary corresponds to approximately 35% of the species (indicated by a slash mark preceding the clade name, e.g. /Rhynchapetala ). found in that clade. The classification of Castillejinae sensu Chuang and The unranked clades are discussed following the formal taxonomic Heckard (1986, 1991 , 1992 ) is indicated in the left column. The phyloge- treatment. Phylogenetic definitions follow the draft PhyloCode netic classification proposed in this paper is indicated in the right column. (Cantino and de Queiroz 2007) and are given following the discussions Compare to Appendix 1 for a synoptical classification to species accord- of individual groups. Node-based definitions are given for the majority ing to both systems. Two rankless clade names are indicated on the tree of groups, when taxon sampling and/or confidence in the given (see text). clades permits. Stem-based definitions are given for those clades where 184 SYSTEMATIC BOTANY [Volume 34 uncertainty (due to incomplete taxon sampling) in the exact circumscrip- Corolla strongly gamopetalous, bilabiate, the upper lip folded tion exists. into a beak (open at tip) or galeate (closed at tip), the lower lip either simple-saccate, 3-saccate, or reduced and rudimen- Taxonomic Treatment tary 3-toothed. Stamens 4 (2), epipetalous, didynamous, the anther sacs 2 (1), unequal in size and placement. Gynoecium We present a revised classification of subtribe Castillejinae of 2 fused carpels, forming a bilocular, superior ovary with ( Fig. 1 , Appendix 1). Of the six genera (Castilleja , Clevelandia , axile placentation. Ovules few (4) to many, anatropous or Cordylanthus , Ophiocephalus , Orthocarpus , and Triphysaria ) campylotropous. Style slender with a capitate, bilobed, or included in Castillejinae (sensu Chuang and Heckard 1991), unexpanded stigma. Seeds with a terminal or lateral hilum Orthocarpus and Triphysaria retain their previous circumscrip- and loose or tight-fitting seed coat. Chromosome numbers, n tions. New nomenclatural combinations are proposed in = 10, 11, 12, 13, 14, 15, 16, 21, 24, 36, 48, 60, 72. Castilleja, where generic boundaries have been expanded to include the monotypic genera Clevelandia and Ophiocephalus . Included genera: Castilleja Mutis ex L.f., Chloropyron Behr, Cordylanthus is restricted to include only those species tradition- Cordylanthus Nutt. ex Benth., Dicranostegia (A. Gray) ally recognized as subg. Cordylanthus, while the two remain- Pennell, Orthocarpus Nutt., Triphysaria Fisch. & C.A. Mey. ing subgenera, Dicranostegia and Hemistegia , are reassigned to Family-wide molecular phylogenetic analyses place the rank of genus (Dicranostegia and Chloropyron , respectively). Castillejinae in a New World clade including the hemiparasitic Thus, Castillejinae will now include the six genera Castilleja ( Fig. genera Agalinis , Seymeria , and Lamourouxia ( Young et al. 1999 ; 2F -I), Chloropyron ( Fig. 2C ), Cordylanthus ( Fig. 2B ), Dicranostegia Wolfe et al. 2005 ; Bennett and Mathews 2006 ). Chloroplast and ( Fig. 2D ), Orthocarpus ( Fig. 2A ), and Triphysaria ( Fig. 2E ). nuclear DNA sequences strongly support the monophyly of Castillejinae G. Don, Gen. Hist. 4: 614, 1838.—TYPE: Castillejinae ( Tank and Olmstead 2008 ), and, with the exception Castilleja Mutis ex L.f. of Ophiocephalus and those species that have undergone a reduc- Plants annual or perennial hemiparasitic herbs. Leaves tion to only one anther-sac per stamen (e.g. Triphysaria and some entire to pinnately or palmately dissected. Inflorescence usu- species of Cordylanthus ), the group is characterized by having ally a short or elongate spike, raceme, single-flowered flo- anther sacs that are unequal in size and unequally attached. rescence (Cordylanthus ), or capitate or spicate clusters of We provide the following node-based clade defini- single-flowered florescences (synflorescence; Cordylanthus ). tion: Castillejinae are the least inclusive clade that con- Bracts subtending individual flowers gradually differing tains Cordylanthus rigidus (Benth.) Jeps. (Cordylanthus ), from the leaves upwards, either green throughout or becom- Orthocarpus luteus Nutt. (Orthocarpus ), and Castilleja fissifolia ing colored apically. Calyx tubular to spathe-like, 2- to 4-cleft. L.f. ( Castilleja ). Key to the Genera of Subtribe Castillejinae 1. Corolla bilabiate, sometimes only slightly 2-lipped, upper corolla lip folded longitudinally forming a beak (apex of the 2-fused corolla lobes open and the opening directed forward), or slightly bilobed and scarcely beaked, or upper corolla lip 2-lobed and not beaked; stigma expanded, capitate, or bilobed; hilum of seed terminal, seed coat loose or tight-fitting ...... 2 2. Stamens each with 2 anther-sacs; corolla throat not indented (except in Castilleja campestris ); seed coat predominately loose-fitting ...... 1. Castilleja 2. Stamens each with a single anther-sac; corolla throat abruptly indented forming a fold below the lower lip (except in Triphysaria pusilla with unusually small, 4–6 mm long corolla); seed coat tight-fitting (except in Triphysaria micrantha and some Triphysaria eriantha ) ...... 6. Triphysaria 1. Corolla bilabiate, upper corolla lip folded longitudinally to form a distinct galea (apex of the 2-fused corolla lobes closed and the opening directed downward forming a hood); stigma unexpanded; hilum of seed lateral, seed coat tight-fitting ...... 3 3. Calyx tubular or short campanulate, unequally 4-cleft with the deepest cleft in back (adaxial); lower corolla lip minutely 3-toothed; inflorescence a spike or spike-like raceme ...... 5. Orthocarpus 3. Calyx spathe-like, completely or nearly completely cleft to the base abaxially; lower corolla lip not toothed; inflorescence a spike, or single-flowered florescence, or spike-like or capitate clusters of single-flowered florescences (synflorescence) ...... 4 4. Leaves entire, linear or filiform, or 3-parted, or subpalmately 5- to 7-parted into narrow segments; inflorescence a short dense spike, less than 2 cm in length (except in Cordylanthus kingii rarely to 5 cm), or of scattered single-flowered florescences, or of short spicate or capitate synflorescences; fertile stamens 4 (except in Cordylanthus capitatus ); middle lobe of lower corolla lip tightly revolute, the tip distinctly folded inside-out ...... 3. Cordylanthus 4. Leaves entire, oblong to lanceolate, or pinnately dissected; inflorescence an elongate spike, usually 3 cm or more in length; fertile stamens 2 (except in Chloropyron maritimum ), the sterile pair inserted lower in corolla tube reduced to rudimentary filaments or bearing sterile, hyaline, villous appendages; middle lobe of lower corolla lip erect, not revolute ...... 5 5. Leaves entire; calyx equal to or slightly shorter than corolla, shallowly cleft at apex (ca. 1 mm deep); anther connective not elongated and anther-sacs remaining approximate; halophytes ...... 2. Chloropyron 5. Leaves pinnately dissected; calyx ca. 1/2 the length of the corolla, cleft at apex greater than 1/2 its length; anther connective conspicuously elongated, separating the two anther-sacs; not halophytic ...... 4. Dicranostegia 2009] TANK ET AL.: CLASSIFICATION OF CASTILLEJINAE 185

Fig. 2. A. Orthocarpus tenuifolius (Pursh) Benth. B. Cordylanthus kingii S. Watson subsp. helleri T. I. Chuang & Heckard C. Chloropyron maritimum (Nutt. ex Benth.) A. Heller subsp. canescens (A. Gray) Tank & J. M. Egger D. Dicranostegia orcuttiana (A. Gray) Pennell E. Triphysaria versicolor Fisch. & C. A. Mey. F. Castilleja exserta (A. Heller) T. I. Chuang & Heckard var. venusta (A. Heller) J. M. Egger G. Castilleja ophiocephala (Wiggins) Tank & J. M. Egger H. Castilleja chrymactis Pennell I. Castilleja talamancensis N. H. Holmgren. Photographs by J. M. Egger.

Circumscription of Genera in Subtribe Castillejinae Ophiocephalus Wiggins, Contr. Dudley Herb. (Stanford Univ.) 1: 175. 1933.—TYPE: Ophiocephalus angustifolius 1. Castilleja Mutis ex L.f., Suppl. Syst. Veg. 47. 1781.—TYPE: Wiggins. Castilleja fissifolia L.f. Gentrya Breedlove & Heckard, Brittonia 22: 20. 1970.—TYPE: Clevelandia Greene, Bull. Calif. Acad. 1: 182. 1886.—TYPE: Gentrya racemosa Breedlove & Heckard. Clevelandia beldingii (Greene) Greene. Orthocarpus beldin- Oncorhynchus Lehm., Index Sem. Hort. Hamb. 1833.—TYPE: gii Greene. Oncorhynchus pinnatifidus Lehm. Castilleja laciniata. 186 SYSTEMATIC BOTANY [Volume 34

Euchroma Nutt., Gen. Am. 2: 54. 1818.—TYPE: Euchroma coc- in part), as in Castilleja ( Wiggins 1933 ). Castilleja ophiocephala cinea (L.) Nutt. Bartsia coccinea L. is also unique in having strongly-exserted stamens that are Perennial or annual herbs, (5–) 10–100 (–125) cm tall with inserted equally at the top of the corolla tube, each possessing few to many ascending-spreading or decumbent branches, two subequal anther sacs. It is interesting that Castilleja rac- hemiparasitic; herbage usually green, sometimes suffused emosa, also a narrow endemic formerly recognized as a mono- with deep red or purple anthocyanins, glabrous to tomen- typic genus in Castillejinae, was originally excluded from tose; roots forming haustoria along finer branches. Leaves Castilleja based primarily on its small (1–2 cm), yellow corolla entire to pinnately dissected. Inflorescence a short or elongate with a beak that is curved nearly at a right angle to the corolla spike, often becoming racemose below, or rarely entirely rac- tube (Breedlove and Heckard 1970). However, with their emose. Floral bract one per flower, gradually differing from expansion of Castilleja to include all members of Castillejinae the leaves upwards, usually becoming more incised and often with a basic chromosome number of x = 12 and seeds colored apically. Calyx tubular, 4-cleft, the lobes subequal, with a terminal hilum, Chuang and Heckard (1991) decided or in many perennial species the lateral lobes partly to com- that the corolla morphology alone was not sufficient for pletely connate. Corolla bilabiate, variable, mostly elongate maintaining C. racemosa as a distinct genus. Despite this deci- and narrow, ventral pouch (of throat and lower lip) 3-saccate sion, Clevelandia and Ophiocephalus, both with n = 12 chro- to lacking; upper lip folded longitudinally forming a beak mosomes and terminally attached seeds, were maintained with the apex open and the opening directed forward, or only as distinct genera ( Chuang and Heckard 1991 ). To the credit scarcely beaked and slightly bilobed (C. ophiocephala ), or not of Chuang and Heckard, the corolla of Castilleja racemosa beaked and distinctly 2-lobed (C. beldingii ); lower lip mostly is much more similar to the remainder of Castilleja than are highly reduced and rudimentary 3-toothed, or less commonly the highly modified corollas of either C. beldingii or C. oph- the lobes varying up to as long as the beak, or lobes triangular, iocephala. Given the phylogenetic relationships inferred from spreading and slightly plicate (C. beldingii ). Style slender with molecular data ( Tank and Olmstead 2008 ), the floral morphol- a capitate to bilobed stigma. Stamens 4, mostly didynamous, ogy of all three of these narrowly endemic annual Castilleja included; anther sacs 2, unequal in size and placement (but species has been modified from the more or less straight- in C. ophiocephala stamens inserted equally at summit of beaked corolla commonly found in the closely related peren- corolla-tube, long-exserted; anther sacs subequal in size and nial species. The unique floral morphologies of Castilleja medially placed). Seeds numerous per capsule with a termi- beldingii , C. ophiocephala, and C. racemosa are likely the result nal hilum and reticulate and loose or tight-fitting seed coat. of increased specialization to pollinators, however, pollina- Chromosome numbers of n = 10, 11, 12, 24, 36 48, 60, 72. tor observations have not been previously reported for any This is by far the largest genus of the subtribe, with ca. 190 of these species. Field observations of Castilleja ophiocephala mostly perennial species distributed primarily in western by Egger are supportive of this hypothesis. A small species North America, but also found in eastern North America (four of Bombus (Hymenoptera) was observed collecting pollen species), northern Asia (ca. five species), Central America selectively and exclusively from the strongly-exserted sta- and Andean South America (ca. 20 species). The genus was mens of Castilleja ophiocephala for extended periods of time named in honor of Professor Domingo Castillejo (1744–1793), in a meadow rich in other flowering plant species. The new an instructor of botany at Cadiz, Spain. nomenclatural combinations necessary for these two taxa are Based on the phylogenetic relationships inferred from included below, as well as the lectotypification of Castilleja separate and combined analyses of cp and nrDNA sequence beldingii . Although there is no unequivocal morphological data ( Tank and Olmstead 2008 ), the traditional concept of synapomorphy for the Castilleja clade that excludes Triphysaria Castilleja did not represent a monophyletic group, because the (see discussion of /Rhynchapetala below), the genus is cyto- two formerly recognized monotypic genera Clevelandia and logically distinct from the remainder of the subtribe with a Ophiocephalus were derived from within Castilleja ( Fig. 1 ). We basic haploid chromosome number of x = 12. present here an expanded treatment of Castilleja that includes We provide the following stem-based clade definition: the two species comprising Clevelandia (= Castilleja beldingii ) Castilleja are the most inclusive clade that contains C. ambigua and Ophiocephalus (= Castilleja ophiocephala ). These two species Hook. & Arn. and C. fissifolia L.f., but not Triphysaria versicolor were historically maintained as distinct genera in Castillejinae, Fisch. & C.A. Meyer (Triphysaria ). despite their numerous similarities to Castilleja , including chro- mosome number (n = 12), vegetative morphology, attachment 1. Castilleja beldingii (Greene) Tank & J.M. Egger, comb. of the ovule and seed, and the ability to make fertile hybrids in nov. Orthocarpus beldingii Greene, Bull. Calif. Acad. Ser. experimental crosses (only C. ophiocephala was tested for cross 1, 3: 123. 1885. Clevelandia beldingii (Greene) Greene, compatibility; Chuang and Heckard 1991 ). Both of these spe- Bull. Calif. Acad. 1: 182. 1886.—TYPE: MEXICO. Baja cies possess unique corolla morphologies that formed the pri- California: “on Victoria Mountains, Lower California,” mary justification of their status as separate genera (Chuang Jun 1883, E. L. Belding s.n. (holotype: CAS, apparently and Heckard 1991). Castilleja beldingii, a narrow endemic of destroyed by fire in 1906; lectotype: here designated , GH the southeastern end of the Baja California peninsula, has a sheet no. 00078143!). Note: Although Greene did not spec- small (1–2 cm), rose-colored corolla in which the lobes of the ify the location of the holotype in his original publication upper corolla lip are not beaked, and the lower corolla lip has of Orthocarpus beldingii , we assume that it was located at three triangular, spreading lobes that are wider than those of CAS where many of the holotypes from species that he the upper corolla lip, forming a slightly zygomorphic corolla published in the Bulletin of the California Academy of ( Greene 1885 , 1886 ). Castilleja ophiocephala, narrowly restricted Sciences during that time are housed. Furthermore, the to the Sierra San Pedro Mártir in northern Baja California, has GH specimen that we designate here as the lectotype a purple, distinctly inflated corolla that is only slightly bilabi- was labeled with a “California Academy of Sciences ate. However, the upper corolla lip is fused into a beak (at least Botanical Section” label, and annotated in Greene’s hand 2009] TANK ET AL.: CLASSIFICATION OF CASTILLEJINAE 187

as an isotype. Unfortunately, this sheet only contains one Bentham’s sections, and thus, the majority of Castilleja spe- depauparate plant from the type collection, and we are cies, in their much broader view of subg. Castilleja comprising unaware of the existence of any other isotypes. species with flowers modified for hummingbird pollination 2. Castilleja ophiocephala Tank & J.M. Egger, nom. nov. (i.e. with floral bracts and calyx tips colored red, a well-devel- Ophiocephalus angustifolius Wiggins, Contr. Dudley Herb. oped corolla tube, and a beaked upper corolla lip that is much (Stanford Univ.) 1: 176. 1933., not Castilleja angustifolia longer than the reduced lower corolla lip). However, they (Nutt.) G. Don—TYPE: MEXICO. Baja California: Sierra were unwilling to commit to a sectional classification until San Pedro Mártir, in granitic soil under pines about the additional research could provide the information neces- margins of the main meadow at La Encantada, 2,200 m, sary to formulate satisfactory groupings. Their subg. Colacus 18 Sep 1930, Wiggins & Demarée 4888 (holotype: DS!; iso- included the majority of Castilleja species that have flowers types: DS!, GH!, NY!, PH!, RSA-POM [3]!, UC!, US!). modified for insect pollination (i.e. with floral bracts and Note: This epithet was selected to provide a connection calyx tips colored other than red and a well-developed lower to the previous generic concept and its descriptive value corolla lip with some degree of pouch development). Chuang with reference to the shape of the swollen corolla (ophio and Heckard (1991) further divided subg. Colacus , based on = snake; cephalus = head). There is an earlier name, characteristics of the calyx incision, into two sections of peren- Castilleja angustifolia (Nutt.) G. Don, preventing the use nials that had been historically shuffled between Orthocarpus of this specific epithet. and Castilleja (sects. Pilosae and Pallescentes) and one section comprising the annual Castilleja species (sect. Oncorhynchus ) Infrageneric Classification—Castilleja is a morphologically previously assigned to Orthocarpus by Keck (1927) . Subgenus complex group that historically has had numerous infrage- Gentrya was viewed as an isolated lineage in Castilleja , and neric assemblages and intraspecific taxa, in which the taxo- was restricted to the single species C. racemosa (formerly nomic complexity is extremely challenging. The complexity Gentrya racemosa ). in Castilleja stems from complex, overlapping morphological Phylogenetic analysis of sequence data from cpDNA and variation often attributed to the formation of polyploid com- nrDNA regions did not support the monophyly of either plexes (Heckard 1968; Holmgren 1970; Heckard and Chuang Castilleja subg. Castilleja or subg. Colacus ( Fig. 1 ; Tank and 1977 ), and the resulting difficulty in circumscribing distinct Olmstead 2008 ). In addition, these analyses placed Castilleja taxa, because of overlapping variation in many characters racemosa (subg. Gentrya ) within a group of Mexican and traditionally used to delimit infrageneric groups and even Central American perennial Castilleja species and not as an species ( Holmgren 1984 ). As a result, infrageneric classifica- isolated lineage of Castilleja . The molecular data recovered a tion in Castilleja has been difficult, and most efforts to clas- well-supported clade, including all of the perennial Castilleja sify this group (e.g. Bentham 1846 ; Gray 1862 , 1886 ; Wettstein species from both subgenera, that is derived from a grade of 1891 ; Fernald 1898 ; Eastwood 1909 ; Rydberg 1917 ; Pennell annual species from subg. Colacus sect. Oncorhynchus ( Fig. 1 ). 1935 , 1951 ; Ownbey 1959 ; Holmgren 1970 , 1971 , 1973 , 1976 , Furthermore, none of the three sections of subg. Colacus des- 1978 , 1984 ; Nesom 1992a , 1992b , 1992c , 1994 ) have been prob- ignated by Chuang and Heckard (1991) (sects. Oncorhynchus , lematic due to their failure to delimit cohesive units within Pallescentes , and Pilosae ) were found to be monophyletic ( Tank Castilleja , their narrow use of small or monotypic infrage- and Olmstead 2008 ). A rapid and recent diversification fol- neric groups, and the treatment of species groups for regional lowed by the retention of ancestral morphological poly- floras (e.g. Rydberg 1917; Pennell 1951; Ownbey 1959) with- morphism may explain, in part, the conspicuous amount of out regard to intraspecific variation outside of the region overlapping morphological variation between taxa that has under consideration. hindered infrageneric classification in Castilleja ( Tank and Bentham (1846) was the first to propose an infrageneric Olmstead 2008 ). classification for the 34 species of Castilleja known at the time, Thus, the current infrageneric classification of Castilleja fails separating the genus into four sections, Epichroma Benth., to delineate natural groups within the complex genus and Euchroma (Nutt.) Benth., Callichroma Benth., and Hemichroma sorely needs attention. However, our understanding of phy- Benth. (= sect. Castilleja ), based primarily on the nature of logenetic relationships within Castilleja , especially among the calyx incision, density of the inflorescence, and the color and perennial species, is still too incomplete to propose satisfac- shape of the calyx and associated floral bract. As more species tory infrageneric groups at this time. However, the complex were added to Castilleja , Bentham’s four sections were recog- history of perennial Castilleja species is the focus of an ongo- nized to be unsatisfactory in their delimitation of cohesive ing research program of the authors, and will be addressed in infrageneric groups and were thus combined in various ways future publications. (e.g. Gray 1862 ; Wettstein 1891 ), further divided (e.g. Pennell 2. Chloropyron Behr, Proc. Calif. Acad. Sci. 1: 62. 1855. 1935 ), or ignored completely (e.g. Rydberg 1917 ; Pennell 1951 ; Cordylanthus sect. Hemistegia A. Gray, Proc. Amer. Acad. Ownbey 1959). The taxonomic history of infrageneric group- Arts 7: 383. 1868. Adenostegia sect. Chloropyron (Behr) ings in Castilleja is reviewed in detail by Chuang and Heckard Ferris, Bull. Torrey Bot. Club 45: 418. 1918. Cordylanthus (1991) . subg. Hemistegia (A. Gray) Jeps., Manual Fl. Pl. Calif. 945. In their generic realignment of subtribe Castillejinae, 1925.—TYPE: Chloropyron maritimum (Nutt. ex Benth.) A. Chuang and Heckard (1991) proposed a new infrageneric Heller. classification for Castilleja including the three subgenera, Castilleja , Colacus , and Gentrya (Appendix 1). Prior to this Halophytic annual herbs of saline and saline-alkali soils, revision, the only use of subgenera in Castilleja was the desig- 10–60 cm tall with few to many ascending-spreading or nation of subg. Castilleja by Holmgren (1970 , 1976 ), in which decumbent branches, hemiparasitic; herbage green, or gray- he included Bentham’s sects. Epichroma and Hemichroma . In ish-green to glaucous, often with deep red to purple anthocya- contrast, Chuang and Heckard (1991) included all four of nins; roots yellowish, forming haustoria along finer branches. 188 SYSTEMATIC BOTANY [Volume 34

Leaves entire, sessile, lanceolate to oblong, 5–35 mm long. Morphologically, Chloropyron shares a number of character- Inflorescence a loose to dense spike, often elongate. Floral istics with Dicranostegia that can be used to distinguish them bract one per flower, entire or pinnately lobed. Calyx spathe- from Cordylanthus (see /Pseudocordylanthus below). A detailed like, cleft completely to the base abaxially, partially surround- taxonomic treatment has been published for the four species ing corolla tube laterally, entire to slightly bifid at apex. Corolla comprising Chloropyron (as Cordylanthus subg. Hemistegia ; bilabiate, club-shaped, tubular below, expanded laterally near Chuang and Heckard 1973 ). The necessary nomenclatural or above midpoint of corolla; upper lip folded longitudinally changes, including new combinations, are included below. to form a distinct galea, apex rounded and closed, the open- We provide the following node-based clade definition: ing directed downward forming a hood enclosing the anthers Chloropyron are the least inclusive clade that contains C. mar- and style; lower lip shorter than or about as long as upper lip, itimum (Nutt. ex Benth.) A. Heller, C. tecopense (Munz & Roos) obscurely 3-lobed. Fertile stamens 2 (4 in C. maritimum ), adax- Tank & J.M. Egger, and C. palmatum (Ferris) Tank & J.M. ial pair (attached lower in corolla tube) reduced to rudimen- Egger. tary filaments (except in C. maritimum with a single anther sac); anther sacs, tufted-hairy on lower end, dissimilar in size 1. Chloropyron maritimum (Nutt. ex Benth.) A. Heller, and placement. Gynoecium 2-carpellate; ovary bilocular, gla- Muhlenbergia 3: 133. 1907. Cordylanthus maritimus Nutt. brous, placentation axile; ovules numerous and campylotro- ex Benth. in DC. Prodr. 10: 598. 1846. Adenostegia maritima pous; style elongate, slightly enlarged towards apex and bent (Nutt. ex Benth.) Greene, Pittonia 2: 181. 1891.—TYPE : downwards with the small stigma barely exserted from apex UNITED STATES. California: San Diego, Nuttall s.n. of galea. Fruit a loculicidal capsule; seeds with a lateral hilum (holotype: K; isotypes: GH!, PH!). and reticulate tight-fitting seed coat. Chromosome numbers 1a. Chloropyron maritimum subsp. maritimum n = 14, 15, 21. 1b. Chloropyron maritimum subsp. canescens (A. Gray) Tank The disintegration of Cordylanthus , as traditionally recog- & J.M. Egger, comb. nov. Cordylanthus canescens A. Gray, nized, was one of the most surprising results of the molec- Proc. Amer. Acad. Arts 7: 383. 1868. Adenostegia cane- ular phylogenetic analyses (Tank and Olmstead 2008). scens (A. Gray) Greene, Pittonia 2: 181. 1891. Chloropyron Historically, Cordylanthus was viewed as one of the most canescens (A. Gray) A. Heller, Muhlenbergia 3: 134. 1907. distinct and recognizable members of subtribe Castillejinae Cordylanthus maritimus var. canescens (A. Gray) Jeps., (Chuang and Heckard 1991), based primarily on the char- Manual Fl. Pl. Calif. 947. 1925. Cordylanthus maritimus acteristic calyx that is abaxially cleft nearly or completely to subsp. canescens (A. Gray) Chuang & Heckard, Brittonia the base and fused adaxially to form a one-piece, spathe-like 25: 149. 1973.—TYPE : UNITED STATES. Nevada: near calyx. Nearly all previous workers on the group have fol- Carson City, C. L. Anderson 207 (hololectotype: GH!; lowed this general circumscription of the genus (e.g. Bentham isolectotype: US!). 1836 , 1846 ; Gray 1867 , 1883 , 1886 ; Wettstein 1891 ; Ferris 1918 ), Cordylanthus parryi S. Watson ex Parry, Amer. Naturalist 9: and in the most recent treatment of the genus, Chuang and 346. 1875. Cordylanthus canescens var. parryi (S. Watson ex Heckard (1986) recognized the three subgenera, Hemistegia (4 Parry) A. Gray, Synopt. Fl. North Amer. 2(1): 304. 1878. species), Dicranostegia (1 species), and Cordylanthus (13 spe- Adenostegia parryi (S. Watson ex Parry) Greene, Pittonia cies comprising 4 sections). However, as it is traditionally 2: 181. 1891. Chloropyron parryi (S. Watson ex Parry) A. circumscribed, Cordylanthus was not monophyletic in any Heller, Mulenbergia 3: 134. 1907. Cordylanthus maritimus of the molecular analyses, but rather, was biphyletic ( Tank var. parryi (S. Watson ex Parry) Jeps., Manual Fl. Pl. Calif. and Olmstead 2008); the two main subgenera, Cordylanthus 947. 1925.—TYPE : UNITED STATES. Utah: Washington and Hemistegia, were each monophyletic, and the monotypic Co., “Valley of the Virgin,” Jun 1874, C. C. Parry 155 subg. Dicranostegia was sister to subg. Hemistegia, forming the (holotype: GH!; isotypes: DS, F, GH!, K, MO [2]!, NY!, /Pseudocordylanthus clade (Fig. 1). Following the molecular PH!, US!). phylogenetic results, we return to a previous generic concept 1c. Chloropyron maritimum subsp. palustre (Behr) Tank & for subg. Hemistegia , resurrecting the genus Chloropyron to J.M. Egger, comb. nov. Chloropyron palustre Behr, Proc. accommodate this monophyletic group. Calif. Acad. Sci. 1: 62. 1855.—TYPE : UNITED STATES. The four species comprising Chloropyron are distinguished California: San Francisco, Russ Garden, Behr s.n. (holo- both morphologically and ecologically from Cordylanthus as type: CAS apparently destroyed by fire in 1906). UNITED defined here ( Fig. 1 ; see below). Even when included within STATES. California: San Francisco, 1866, A. Kellogg s.n. the traditional Cordylanthus , these species were distinguished (neotype, Chuang and Heckard, Brittonia 25: 148: MO; from the remainder of the genus, either as a section (e.g. Gray isoneotypes: GH!, NY). 1867 ; Ferris 1918 ) or a subgenus (e.g. Chuang and Heckard 2. Chloropyron molle (A. Gray) A. Heller, Muhlenbergia 3: 1973). Behr (1855), in his description of a new species from 134. 1907. Cordylanthus mollis A. Gray, Proc. Amer. Acad. California, created the genus Chloropyron , noting its inter- Arts 7: 384. 1868. Adenostegia mollis (Gray) Greene, Pittonia mediate morphology between Orthocarpus and Triphysaria . 2: 181. 1891.—TYPE : UNITED STATES. California: Solano However, in his description, Behr (1855) did not note any affin- Co., “Mare Island, Bay of San Francisco,” Nov 1855, ity to Cordylanthus , which had already been described. Given Charles Wright s.n. (holotype: GH!, NY!; isotype: K, NY!). the similar one-piece, spathe-like calyx and club-shaped corolla 2 a. Chloropyron molle subsp. molle of both Chloropyron and Cordylanthus, it is likely that Behr was 2 b. Chloropyron molle subsp. hispidum (Pennell) Tank unaware of Bentham’s (1836, 1846) work on Cordylanthus . The & J.M. Egger, comb. and stat. nov. Cordylanthus hispi- most striking difference between Chloropyron and the other dus Pennell, Proc. Acad. Nat. Sci. Philadelphia 99: 192. lineages of the traditional Cordylanthus is the halophytic 1947.—TYPE : UNITED STATES. California: Merced Co., nature of Chloropyron , occurring in both coastal and inte- Volta, 14 Jul 1937, R. F. Hoover 2605 (holotype: DS; iso- rior saline and alkaline habitats of western North America. types: CAS, JEPS!, MO, NY [2]!, PH [2]!, UC!, US [2]!). 2009] TANK ET AL.: CLASSIFICATION OF CASTILLEJINAE 189

Cordylanthus mollis var. viridis Jeps. ex H.L. Bauer, Ecology strongly supports the monophyly of this restricted circum- 11: 271. 1930.—TYPE : UNITED STATES. California: Kern scription of Cordylanthus ( Fig. 1 ). Cordylanthus can be recog- Co., 15 miles SE of Bakersfield, 22 Oct 1927, H. L. Bauer 8 nized by the following morphological synapomorphies: 1) (holotype: JEPS!). the architecture of the inflorescence has undergone an evolu- 3. Chloropyron palmatum (Ferris) Tank & J.M. Egger, comb. tionary reduction from the basic spike common throughout nov. Adenostegia palmata Ferris, Bull. Torrey Bot. Club the subtribe (and the majority of Orobanchaceae) to single- 45: 420. 1918. Cordylanthus palmatus (Ferris) J. F. Macbr., flowered florescences and the subsequent clustering of these Contr. Gray Herb. 59: 38. 1919.—TYPE : UNITED STATES. into capitate or spike-like synflorescences, and 2) the mid- California: Colusa Co., in alkaline soil, overflowed lands dle lobe of the lower corolla lip is tightly revolute, where the at Tule near College City, 17 Jun 1916, R. Stinchfield 284 tip of the middle lobe is curled inside-out. The characteris- (holotype: DS; isotypes: PH, UC, US!). tic, spathe-like calyx, viewed as an evolutionary reduction of Cordylanthus carnulosus Pennell, Proc. Acad. Nat. Sci. the 4-cleft calyx found throughout Castillejinae and used to Philadelphia 99: 191. 1947. Cordylanthus palmatus subsp. recognize the traditional Cordylanthus , has evolved indepen- carnulosus (Pennell) Munz, Aliso 4: 98. 1958.— TYPE : dently in the two lineages comprising the former group (i.e. UNITED STATES. California: Fresno Co., 6 miles S of Cordylanthus and / Pseudocordylanthus), and therefore, is not a Kerman, 29 Jul 1937, R. F. Hoover 2636 (holotype: DS; iso- diagnostic feature of Cordylanthus as recognized here. The flo- types: CAS, JEPS!, PH [2]!, UC!). ral bract found in Cordylanthus also differs from that seen in 4. Chloropyron tecopense (Munz & J.C. Roos) Tank & J.M. the / Pseudocordylanthus clade; in Cordylanthus, the floral bract Egger, comb. nov. Cordylanthus tecopensis Munz & J.C. is similar to the calyx in shape, size, and texture. Because of Roos, Aliso 2: 233. 1950.—TYPE : UNTIED STATES. their similarity, the floral bract was commonly misinterpreted California: Inyo Co., below Tecopa Hot Springs, 1400 ft, as part of the calyx by previous workers (e.g. Gray 1867, 1886 ; 9 Oct 1949, Munz & Campbell 14358 (holotype: RSA; iso- Wettstein 1891 ; Ferris 1918 ), where species belonging to the types: CAS, DS, GH!, NY!, PH!, RM, SD!, UC!). Cordylanthus clade were described as having a “diphyllous” 3. Cordylanthus Nutt. ex Benth. in DC., Prodr. 10: 597. 1846 calyx and the members of the /Pseudocordylanthus clade were (nom. cons.). Adenostegia Benth. in Lindley, Nat. Syst. described as having a “monophyllous” calyx ( Chuang and Bot., second ed. 445. 1836 (nom. rej.). Cordylanthus subg. Heckard 1986). Part of this confusion was due to the presence Adenostegia (Benth.) Jeps., Manual Fl. Pl. Calif. 945. 1925 of the outer bract, which usually subtends the basic spike (nom. inval.).—TYPE: Cordylanthus filifolius Nutt. ex inflorescence, but as a result of the reduction of the basic inflo- Benth. rescence to the single-flowered florescences, each flower has Annual herbs, (5–) 10–120 (–150) cm tall, hemiparasitic; multiple bracts associated with it. When the inflorescence is herbage gray or yellow-green, often turning red to pur- reduced to a single-flowered florescence, the outer bracts that ple; roots mostly yellowish, forming haustoria along finer typically subtend the basic inflorescence are found immedi- branches. Leaves entire and linear to filiform, or 3-parted, or ately subtending the single-flowered florescence, creating the subpalmately 5- or 7-parted into narrow divisions. Flowers appearance of a “diphyllous” calyx and an associated floral arranged in a short spike (mostly < 2 cm), or scattered single- bract. With the subsequent clustering of the single-flowered flowered florescences, or spicate or capitate clusters of sin- florescences into capitate or spike-like clusters of these flores- gle-flowered florescences. Outer bracts entire, 3- to 5-lobed, cences, it becomes even more difficult to interpret the homol- subpalmately 5- or 7-parted; floral bracts mostly navicu- ogy of inflorescence parts. Nine of the 13 species comprising lar, entire or with 1–4 pairs of pinnate lobes, rarely with the the genus were included in the molecular phylogenetic study apex shallowly 3- to 7-lobed. Calyx spathe-like, navicular, of Castillejinae ( Tank and Olmstead 2008 ). However, the four cleft abaxially to the base, or nearly so and leaving a short species missing from the analyses all possess the revolute tubular base, entire to slightly bifid at apex, rarely divided middle lobe of the lower corolla lip and the modified inflo- to 1/3 its length (C. capitatus ). Corolla bilabiate, club-shaped rescence architecture characteristic of Cordylanthus . A detailed or fusiform, tubular below, expanded laterally near or above taxonomic treatment has already been published for the 13 midpoint of corolla; upper lip folded longitudinally to form a species of Cordylanthus (as Cordylanthus subg. Cordylanthus ; distinct galea, apex rounded and closed, the opening directed Chuang and Heckard 1986), including numerous infraspe- downward forming a hood enclosing the anthers and style; cific taxa. lower lip shorter than or about as long as upper lip, obscurely We provide the following stem-based definition for the 3-lobed, the middle-lobe tightly revolute outwardly (the tip clade: Cordylanthus are the most inclusive clade that contains of the lobe curled inside-out). Stamens 4 (2 in C. capitatus ), C. rigidus (Benth.) Jeps. and C. capitatus Nutt. ex Benth, but didynamous; anthers sacs 2 (rarely 1), densely hairy at both not Orthocarpus luteus Nutt. (Orthocarpus ) or Chloropyron mar- ends and ciliate along the line of dehiscence, dissimilar in itimum (Nutt. ex Benth.) A. Heller (Chloropyron ). size and placement. Gynoecium 2-carpellate; ovary bilocular, Infrageneric Classification—The most recent infrageneric glabrous, placentation axile; ovules numerous and campy- treatment of Cordylanthus is that of Chuang and Heckard lotropous; style elongate, slightly enlarged towards apex and (1986 ; as subg. Cordylanthus ), where they recognized sects. bent downwards with the small stigma barely exserted from Anisocheila , Cordylanthus , and Ramosi , based primarily on apex of galea. Fruit a loculicidal capsule; seeds with a lateral the architecture of the inflorescence, morphology of the seed hilum, seed coat tight-fitting, reticulate or irregularly striate. coat, and chromosome number ( Chuang and Heckard 1972 , Chromosome numbers n = 12, 13, 14. 1976 , 1986 ). The hypothesized relationships expressed by We present here a considerably narrowed circumscrip- their treatment are similar to the four sections of Ferris (1918) , tion of the genus to include the 13 species comprising subg. but differ considerably from the eight subsections delim- Cordylanthus sensu Chuang and Heckard (1986; Appendix 1). ited by Pennell (1951). Pennell (1951) and Ferris (1918) both Molecular phylogenetic analysis of cp and nrDNA regions relied heavily on characteristics of the inflorescence and 190 SYSTEMATIC BOTANY [Volume 34 morphology of the calyx and corolla in their treatments, florescences, and in some species of sect. Cordylanthus the however, they overlooked the morphology of the seed coat. loose inflorescence has been reaggregated into capitate clus- Chuang and Heckard (1976 , 1986 ) defined section Cordylanthus ters of these single-flowered florescences. The inflorescence primarily by the presence of an irregularly striate seed of Cordylanthus ramosus (sect. Ramosi) is unique in that sin- coat that is not found in any other group of Castillejinae. gle-flowered florescences are clustered into a short spike, The remainder of the subtribe possess reticulate seed coats mimicking the ancestral condition ( Fig. 3 ). However, within that vary in the depth of the reticulations, but, nonethe- Cordylanthus, because these complex inflorescences are found less, are all conspicuously reticulate ( Chuang and Heckard in all three sections, the evolutionary reduction and subse- 1991). Chloroplast DNA and nrDNA sequence data from quent modification of the inflorescence has occurred multiple four of the six species of sect. Cordylanthus confirmed the times independently ( Fig. 3 ). Furthermore, even within a spe- monophyly of this group (Fig. 3), and, based on the pres- cies the diversity of inflorescence types and degree of reduc- ence of this unique seed coat morphology, C. nevinii and tion of the spike can vary significantly (Chuang and Heckard C. nidularius (two species that have not been included in phylo- 1986 ). Therefore, caution should be used when using charac- genetic analyses) should be part of this clade. Species belong- teristics of the inflorescence to assess evolutionary relation- ing to section Anisocheila and the monotypic section Ramosi ships. Chuang and Heckard (1976 , 1986 ) recognized this in are both reported to have shallowly reticulate seed coats, in their treatment of the group and used a combination of char- contrast to the deeply reticulate seed coat found through- acters to define their sections. However, previous workers out the /Pseudocordylanthus clade ( Chuang and Heckard (e.g. Ferris 1918 ; Pennell 1951 ) relied heavily on the diversity 1972, 1976 ). However, as employed by Chuang and Heckard of inflorescence types when determining interspecific rela- (1976) , the depth of reticulations is a subjective measure and tionships, which, consequently, led to significant confusion some species are difficult to categorize, thus, this is not a defining natural groups within Cordylanthus , including the clear distinction for these groups. Since the reticulate seed delimitation of species boundaries. coat is the ancestral condition for Cordylanthus , being pres- Chuang and Heckard (1986) relied heavily on chromosome ent throughout Castillejinae, it is not surprising that this number to make decisions of relationships in Cordylanthus . characteristic fails to delimit a monophyletic group within Each of Chuang and Heckard’s (1986) three sections are char- Cordylanthus ( Fig. 3 ). acterized by distinct chromosome numbers; the monophyl- Throughout Orobanchaceae, the architecture of the inflores- etic sect. Cordylanthus all have n = 14 chromosomes, sect. cence is consistent and is most often either a spike or raceme Anisocheila is characterized by a reduction to n = 13, and the ( Judd et al. 2007 ). In Cordylanthus , however, the diversity of monotypic sect. Ramosi has been reduced further to n = 12 inflorescence types is surprising and often pushes the limits ( Fig. 3 ). However, sect. Anisocheila was not monophyletic, of the standard terminology used to describe basic inflores- with one species, Cordylanthus capitatus, sister to the rest of cences ( Chuang and Heckard 1976 , 1986 ). This has resulted in the genus. The remainder of sect. Anisocheila that have been the misinterpretation of what constitutes homologous char- included in phylogenetic analyses were found to be mono- acteristics of the inflorescence, leading to erroneous decisions phyletic, and the monotypic sect. Ramosi was the sister lineage regarding infrageneric classification in the group (e.g. Ferris to this clade ( Fig. 3 ). Cordylanthus laxiflorus and C. parviflorus 1918; Pennell 1951). The evolutionary reduction of an elon- are both enigmatic in their morphological affinities to the gate spike to single-flowered florescences and the clustering rest of sect. Anisocheila and are thought to be closely related of these into capitate or spike-like synflorescences is a synapo- to each other (Chuang and Heckard 1986). However, neither morphy for the clade. Both sects. Anisocheila and Cordylanthus have been included in phylogenetic analyses, so it is unclear have some species with cymosely arranged single-flowered whether they will add to the basal grade of n = 13 lineages

Fig. 3 . Majority rule consensus tree for Cordylanthus , modified from the partitioned Bayesian analysis of chloroplast DNA and nuclear ribosomal DNA data of Tank and Olmstead (2008 ). The mean branch lengths shown are proportional to the number of substitutions per site as measured by the scale bar. Numbers above the branches represent Bayesian posterior probabilities (PP), and numbers below the branches represent maximum likelihood (ML) followed by par- simony bootstrap percentages (BS), as reported by Tank and Olmstead (2008) . The dashed lines indicate species that were not sampled by Tank and Olmstead (2008) and are, therefore, left unplaced (see text). The columns indicate the taxonomic section following Chuang and Heckard (1986) , haploid chromosome number (ch. #; Chuang and Heckard 1986), seed coat patterning (seed; Chuang and Heckard 1972), and the architecture of the inflorescence (inflor.; Chuang and Heckard 1976 ). The inflorescence types include: a reduced spike < 2 cm tall (short spike), loosely arranged single-flowered florescences (1-flwd. flors.), capitate clusters of single-flowered florescences (capitate clust.), and spicate clusters of single-flowered florescences (spicate clust.). 2009] TANK ET AL.: CLASSIFICATION OF CASTILLEJINAE 191 in Cordylanthus or join the clade comprising the remainder of hilum, seed coat reticulate and tight-fitting. Chromosome sect. Anisocheila and C. ramosus ( Fig. 3 ). number n = 16. In addition to the irregularly striate seed coat, species Dicranostegia is the second lineage that we have removed belonging to sect. Cordylanthus differ from the rest of the from Cordylanthus as it was defined by Chuang and Heckard genus both geographically and ecologically. The six species (1975 , 1986 ; Fig. 1 ), and it is morphologically and cytologi- comprising sect. Cordylanthus are restricted to the open oak- cally distinct from the other groups comprised of species pine woodlands and serpentine outcrops found west of the belonging to the traditional Cordylanthus (i.e. Chloropyron Cascade-Sierra Nevada crest from the Siskiyou Mountains in and Cordylanthus ). When Gray (1883) described Cordylanthus southern Oregon to the northern portion of Baja California orcuttianus, he also erected the monotypic sect. Dicranostegia (Chuang and Heckard 1986). In contrast the remaining seven to accommodate this narrow endemic from northern Baja species are distributed widely throughout the intermoun- California and adjacent California. Pennell (1947) raised the tain west in the Great Basin, southwestern deserts, and the taxon to genus (Dicranostegia ), based primarily on the small, adjacent arid regions of the Cascades, Sierra Nevadas, and two-lobed calyx unique to this species. In their detailed mono- Rocky Mountains, where they occur mostly in juniper-pin- graphic work, Chuang and Heckard (1975, 1986 ) reassigned yon woodlands and sagebrush scrub habitats ( Chuang and Dicranostegia to the monotypic subg. Dicranostegia within Heckard 1986). All of these species flourish during the hottest the traditional Cordylanthus ( Fig. 1 ). Their decision was based time of the year in these mostly arid environments, flower- primarily on their view that the calyx of Dicranostegia merely ing mostly from July to September, well after the majority of represents an earlier stage in the evolution of the spathe- desert annuals. like calyx (Chuang and Heckard 1975). Based on the distri- Although it is clear that sect. Cordylanthus is monophyletic bution of different calyx morphologies in Cordylanthus and / and is morphologically and geographically distinct from the Pseudocordylanthus (Fig. 1), this interpretation is likely correct, remainder of the genus, because of the numerous infraspe- but the evolution of the spathe-like calyx has occurred inde- cific taxa and the uncertainty in placement of the species not pendently in the two lineages ( Tank and Olmstead 2008 ). sampled in the molecular phylogenetic studies (Fig. 3), we Chloroplast DNA and nrDNA sequence data indicate a sis- do not think it appropriate to propose a revised infrageneric ter group relationship between Dicranostegia orcuttiana and classification at this time. Future molecular systematic work Chloropyron, the other lineage removed from Cordylanthus focusing on the wide array of morphological diversity not yet (Fig. 1); together, these two genera form the /Pseudocordylanthus represented in any molecular phylogenetic study will likely clade. We have chosen to return to Pennell’s (1947) treatment provide insight into the evolution and diversification of the of this single species as the monotypic genus Dicranostegia , clade, as well as provide the information necessary to develop rather than including it in Chloropyron , primarily because a meaningful infrageneric classification of Cordylanthus . Chloropyron comprises an ecologically distinct group of species adapted to saline and alkaline habitats of western 4. Dicranostegia (A. Gray) Pennell, Proc. Acad. Nat. Sci. North America. However, so the sister group relationship of Philadelphia 99: 189. 1947. Cordylanthus sect. Dicranostegia Chloropyron and Dicranostegia is not lost in the limitations of A. Gray, Proc. Amer. Acad. Arts 19: 95. 1883. Adenostegia the Linnaean classification proposed here, we also propose sect. Dicranostegia (A. Gray) Ferris, Bull, Torrey Bot. Club the unranked group /Pseudocordylanthus (see below). 45: 417. 1918. Cordylanthus subg. Dicranostegia (A. Gray) T.I. Chuang & Heckard, Madroño 23: 93. 1975.—TYPE: 1. Dicranostegia orcuttiana (A. Gray) Pennell, Proc. Acad. Dicranostegia orcuttiana (A. Gray) Pennell. Nat. Sci. Philadelphia 99: 190. 1947.Cordylanthus orcuttianus A. Gray, Proc. Amer. Acad. Arts 19: 95. 1883. Adenostegia Annual hemiparasitic herbs, (10–) 20–35 (–50) cm tall; herb- orcuttiana (A. Gray) Greene, Pittonia 2: 181. 1891.— TYPE : age green, often with dark red markings, densely to sparsely MEXICO. Baja California: “Tia Juana Lower California” hirsute; roots mostly yellowish, forming haustoria along finer [=Tijuana], H. C. Orcutt & Son s.n. (holotype: GH!; iso- branches. Leaves pinnately dissected with 4 or 5 pairs of lin- type: K). ear lobes, 3–6 cm long. Flowers arranged in a dense, indeter- 5. Orthocarpus Nutt., Gen. N. Amer. Plant. 2: 56. 1818. minate spike, 2–10 cm long, subtended by outer bracts. Floral Orthocarpus sect. Monosaccus D.D. Keck, Proc. Calif. Acad. bracts oblong-lanceolate with 1–3 pairs of pinnate lobes. Sci. Ser. 4, 16: 520. 1927.—TYPE: Orthocarpus luteus Nutt. Calyx short, less than one half as long as corolla, cleft abaxi- ally to the base, surrounding the base of the corolla laterally, Annual herbs, 10–50 cm tall, hemiparasitic; herbage green; divided over one half its length adaxially into two attenuate roots forming haustoria along finer branches. Leaves linear to lobes. Corolla bilabiate, club-shaped, tubular below, expanded linear-lanceolate, entire or upper leaves 3-parted, the lobes lin- laterally above midpoint of corolla; upper lip folded longi- ear-lanceolate to filiform. Flowers arranged in a short or elon- tudinally to form a distinct galea, apex rounded and closed, gate spike. Floral bracts lanceolate to ovate, entire, 3-parted the opening directed downward forming a hood enclos- to pinnately 5-parted, usually colored at tips, gradually or ± ing the anthers and style; lower lip shorter than upper lip, abruptly differing from the upper leaves. Calyx tubular or obscurely 3-lobed. Stamens 4, only 2 of which are fertile, didy- short campanulate, unequally 4-cleft with the deepest cleft in namous; the fertile pair of stamens with 2 anther sacs, dis- back (adaxial). Corolla bilabiate, club-shaped, tubular below, similar in size, separated by an elongate connective with one expanded and distinctly saccate above; upper lip folded lon- anther sac at either end; the sterile pair of stamens with ves- gitudinally to form a distinct galea, apex rounded and closed, tigial anther-sacs. Gynoecium 2-carpellate; ovary bilocular, the opening directed downward forming a hood enclosing the glabrous, placentation axile, ovules numerous and campy- anthers and style; lower lip subequal to the upper lip, pouch lotropous; style elongate, slightly enlarged towards apex and inconspicuously to distinctly 3- or 4-lobed with 2 longitudi- bent downwards with the small stigma barely exserted from nal grooves and a V-shaped indentation near the tip of the apex of galea. Fruit a loculicidal capsule; seeds with a lateral central lobe, terminating in 3 narrow-triangular or rounded 192 SYSTEMATIC BOTANY [Volume 34 teeth. Stamens 4, didynamous; anther sacs 2, dissimilar in 4-cleft. Corolla bilabiate, club-shaped and dolabriform in side size and placement. Gynoecium 2-carpellate; ovary bilocular, view (except T. pusilla ), tubular below, expanded and dis- glabrous, placentation axile; ovules few to many and campy- tinctly saccate above, the throat abruptly indented forming lotropous; style slender with an unexpanded stigma. Fruit a a fold below lower lip; upper lip folded longitudinally form- loculicidal capsule; seeds with a lateral hilum, seed coat retic- ing a beak, the apex open and the opening directed forward ulate and tight-fitting. Chromosome numbers n = 14, 15. with the stigma protruding; lower lip subequal to the upper The circumscription of Orthocarpus has changed signifi- lip, much inflated, deeply 3-saccate, minutely 3-toothed at cantly since it was first proposed by Nuttall (1818) with the apex (corolla in T. pusilla : throat lacking indentation, lower lip description of the most widespread species, O. luteus . The shorter than the upper lip, shallowly 3-saccate, conspicuously genus is distributed throughout western North America 3-lobed). Stamens 4, didynamous, enclosed in or exserted from California to northwestern Minnesota and adjacent from the beak of the upper lip, with a single anther-sac. Canada, and Orthocarpus luteus occupies the entire range Gynoecium 2-carpellate; ovary bilocular, glabrous, placen- ( Chuang and Heckard 1992 ). Bentham (1835 ; 1839 ; 1846 ) tation axile; ovules numerous and anatropous; style slender divided the genus into the three sections Oncorhynchus , with a capitate stigma. Fruit a loculicidal capsule; seeds with Orthocarpus , and Triphysaria , based on the degree of pouch- a terminal hilum, seed coat reticulate and loose or tight-fit- ing of the lower corolla lip and the number of anther sacs per ting. Chromosome number n = 11. stamen. Throughout the history of the genus, there has been Historically, Triphysaria has been included within considerable difficulty defining generic boundaries between Orthocarpus s.l., either as a section (e.g. Bentham 1846 ) or sub- Castilleja and Orthocarpus, and numerous species have been genus (e.g. Keck 1927). Based on chromosome number, seed shifted between the two genera (e.g. Gray 1862 ; Watson 1871 ; and seed coat morphology, floral morphology, and experi- Eastwood 1909; Jepson 1925; Keck 1927; Chuang and Heckard mental hybridization studies, the five species comprising the 1991 , 1992 ). Prior to Keck’s (1927) treatment of Orthocarpus , 55 group were most recently treated as a distinct genus within species had been described and the lack of a distinct bound- Castillejinae (Chuang and Heckard 1991). Phylogenetic anal- ary between Orthocarpus and Castilleja had been made appar- ysis of cpDNA and nrDNA regions ( Tank and Olmstead ent; this was perhaps voiced best by Alice Eastwood (1909 p. 2008) support the monophyly of Triphysaria as a lineage 564) when she noted, “The line separating Orthocarpus and distinct from Orthocarpus , and indicate a sister group rela- Castilleja is not definitely fixed, and the species on the border tionship of Triphysaria and Castilleja ( Fig. 1 ; see discussion may suffer changes in name frequently.” Keck (1927) reduced of /Rhynchapetala below). Morphologically, the species of the genus to 25 annual species, moving all of the peren- Triphysaria are similar to the annual species that form the nial species to Castilleja . Chuang and Heckard (1991 , 1992 ) basal grade within Castilleja (also removed from Orthocarpus further reduced Orthocarpus to the nine annuals currently s.l. by Chuang and Heckard 1991). In addition, Triphysaria is comprising the genus, transferring Keck’s (1927) sects. primarily distributed in California west of the Sierra Nevada Castillejoides and Cordylanthoides to Castilleja and elevating his crest, where it is often sympatric with some annual Castilleja subg. Triphysaria to genus. Detailed morphological and cyto- species. Given the sister group relationship of Triphysaria logical analyses ( Chuang and Heckard 1991 , 1992 ) formed and Castilleja , we would be justified in expanding Castilleja the primary basis for the restriction of Orthocarpus to Keck’s to include this annual lineage. However, Triphysaria has a (1927) sect. Monosaccus , and molecular data support this number of synapomorphies that distinguish this lineage assertion, placing Orthocarpus as one of the basal lineages of from Castilleja . Therefore, rather than creating unnecessary subtribe Castillejinae (Fig. 1). The nine species of Orthocarpus confusion through another set of novel nomenclatural com- can be recognized by the morphology of the lower corolla lip, binations for the five Triphysaria species, we instead propose which is generally simple-saccate (or only minutely three- the unranked group /Rhynchapetala to express this relation- cleft), and the unequally cleft calyx in which the adaxial cleft ship (see below). Synapomorphies for Triphysaria include, 1) is cut more deeply than that of the abaxial calyx cleft. A revi- each of the four stamens are reduced to a single anther sac, 2) sion of Orthocarpus, including a detailed treatment of each of with the exception of the tiny (4–6 mm) corolla of T. pusilla , the nine species, has already been published ( Chuang and the throat of the corolla is abruptly indented, forming a dis- Heckard 1992 ). tinct fold below the lower corolla lip, and 3) the chromosome We provide the following node-based definition for the number is reduced to n = 11. A detailed nomenclatural treat- clade: Orthocarpus are the least inclusive clade that contains ment of the five species has already been published (Chuang O. luteus Nutt., O. tolmiei Hook. & Arn., and O. cuspidatus and Heckard 1991 ), including infraspecific taxa for two of the Greene. species. We provide the following node-based definition for the 6. Triphysaria Fisch. & C.A. Mey., Ind. Sem. Hort. Petrop. clade: Triphysaria are the least inclusive clade that contains 2: 52. 1836. Orthocarpus sect. Triphysaria (Fisch. & C.A. T. versicolor Fisch. & C.A. Mey. and T. eriantha (Benth.) T.I. Mey.) Benth. in DC. Prodr. 10: 535. 1846. Orthocarpus Chuang & Heckard. subg. Triphysaria (Fisch. & C.A. Mey.) Keck. Proc. Calif. Acad. Sci. Ser. 4, 16: 520. 1927.—TYPE: Triphysaria versi- color Fisch. & C.A. Mey. Additional Recognized Clades Annual herbs, 10–40 cm tall hemiparasitic; herbage green; /Rhynchapetala— We provide the following node-based roots forming haustoria along finer branches. Leaves linear to definition for this clade: /Rhynchapetela are the least inclu- linear-lanceolate, pinnately divided upwards on the stem into sive clade that contains Triphysaria versicolor Fisch. & C.A. many filiform divisions. Flowers arranged in a short or elon- Mey. (Triphysaria ) and Castilleja fissifolia L.f. (Castilleja ). gate spike. Floral bracts mostly pinnately lobed with 1–3 pairs The monophyly of Triphysaria and Castilleja is supported of filiform divisions. Calyx tubular-campanulate, subequally by molecular data ( Tank and Olmstead 2008 ), as well as a 2009] TANK ET AL.: CLASSIFICATION OF CASTILLEJINAE 193 number of morphological synapomorphies. The clade name, Bentham , G. 1836 . Adenostegia. Pp. 445 in A natural system of botany , ed. /Rhynchapetala , refers to the beaked upper petals that distin- J. Lindley . London : Longman, Rees, Orme, Brown, Green, and Longman . guish this clade from the remainder of Castillejinae, which Bentham, G. 1839 . Plantae Hartwegianae , London : W. Pamplin . are truly galeate, and is taken from the Greek word rhynch, Bentham, G. 1846 . Scrophulariaceae . Pp. 180–586 in Prodromus Systematis meaning beak or snout. In both Triphysaria and Castilleja the Naturalis Regni Vegetabilis ed. A. de Candolle . Paris : Victoris Masson . upper corolla lip is open at the tip, and there is no true galea. Bentham, G. 1876 . Scrophularineae . Pp. 913 – 980 in Genera Plantarum , eds. Throughout the literature, the use of the term galea to describe G. Bentham and J. D. Hooker . London : Reeve and Co. Breedlove, D. E. and L. R. Heckard . 1970 . Gentrya , a new genus of the condition of the upper petals in Castilleja and Triphysaria Scrophulariaceae from Mexico . Brittonia 22 : 20–24 . has been in error, and by naming this clade, we hope to pro- Cantino, P. D. and K. de Queiroz . 2007 . PhyloCode: a Phylogenetic Code mote the use of the term ‘beaked’ to describe this condition, of Biological Nomenclature . Version 4b. http://www.phylocode.org . as first suggested by Chuang and Heckard (1991) . Members Revised September 12, 2007. Chuang, T. I. and L. R. Heckard . 1972 . Seed coat morphology in Cordylanthus of / Rhynchapetala are also characterized by the presence of an (Scrophulariaceae) and its taxonomic significance. American Journal of expanded stigma that is either capitate or bilobed, and the ter- Botany 59 : 258 – 265 . minal attachment of the ovule to the placenta. Chuang, T. I. and L. R. Heckard . 1973 . Taxonomy of Cordylanthus subgenus /Pseudocordylanthus—— We provide the following node- Hemistegia (Scrophulariaceae) . Brittonia 25 : 135 – 158 . based definition for this clade: /Pseudocordylanthus are the Chuang, T. I. and L. R. Heckard . 1975 . Taxonomic status of Cordylanthus (subg . Dicranostegia) orcuttianus (Scrophulariaceae). Madroño 23 : 88 – 95 . least inclusive clade that contains Dicranostegia orcuttiana Chuang, T. I. and L. R. Heckard . 1976 . Morphology, evolution, and (A. Gray) Pennell and Chloropyron maritimum (Nutt. ex Benth.) taxonomic significance of the inflorescence in Cordylanthus A. Heller (Chloropyron ). This clade consists of one of the (Scrophulariaceae) . American Journal of Botany 63 : 272 – 282 . two lineages resulting from the disintegration of the traditional Chuang, T. I. and L. R. Heckard . 1986 . Systematics and evolution of Cordylanthus (Scrophulariaceae-Pedicularieae) . Systematic Botany Cordylanthus ( Fig. 1 ). The clade name, /Pseudocordylanthus , Monographs 10 : 1 – 105 . refers to the spathe-like calyx and laterally flattened, club- Chuang, T. I. and L. R. Heckard . 1991 . Generic realignment and synop- shaped corolla that this group shares with Cordylanthus sis of subtribe Castillejinae (Scrophulariaceae - tribe Pediculareae) . as defined here. The modified calyx was traditionally con- Systematic Botany 16 : 644 – 666 . sidered to be the primary diagnostic feature for the tradi- Chuang, T. I. and L. R. Heckard . 1992 . A taxonomic revision of Orthocarpus (Scrophulariaceae - tribe Pediculareae) . Systematic Botany 17 : tional Cordylanthus (Chuang and Heckard 1986, 1991 ), but has 560 – 582 . been shown to have evolved independently in Cordylanthus Cronquist, A. 1981 . An integrated system of classification of flowering plants , and /Pseudocordylanthus ( Tank and Olmstead 2008 ). New York : Columbia University Press . Members of the / Pseudocordylanthus clade are distinguished de Queiroz, K. 1988 . Systematics and the Darwinian Revolution . Philosophy of Science 55 : 238 – 259 . from Cordylanthus by an elongated spicate inflorescence, de Queiroz, K. 1992 . Phylogenetic definitions and taxonomic philosophy . the reduction of the androecium to two fertile stamens, and Biology and Philosophy 7 : 295 – 313 . the erect (as opposed to revolute) middle lobe of the lower de Queiroz, K. and J. Gauthier . 1990 . Phylogeny as a central principle corolla lip. However, both the elongated spicate inflorescence in taxonomy: phylogenetic definitions of taxon names. Systematic and the erect middle lobe of the lower corolla lip are ances- Zoology 39 : 307 – 322 . de Queiroz, K. and J. Gauthier . 1992 . Phylogenetic taxonomy . Annual tral characteristics in Castillejinae, having been modified in Review of Ecology and Systematics 23 : 449 – 480 . Cordylanthus. The reduction of the androecium to only two de Queiroz, K. and J. Gauthier . 1994 . Toward a phylogenetic system of bio- fertile stamens, a characteristic thought to be a modifica- logical nomenclature . Trends in Ecology & Evolution 9 : 27 – 31 . tion to increased specialization in pollination (Chuang and Eastwood, A. 1909 . Synopsis of the Mexican and Central American species of Castilleja . Proceedings of the American Academy of Arts 44 : 563 – 591 . Heckard 1986), is the only morphological synapomorphy for Farris, J. S. 1979 . The information content of the phylogenetic system . the clade. Systematic Zoology 28 : 483 – 519 . Fernald, M. L. 1898 . Notes upon some northwestern Castilleias of the Acknowledgments. The authors thank A. Whittemore, L. Struwe, parviflora group . Erythea 6 : 41 – 51 . F. Barrie and two anonymous reviewers for critical comments on the Ferris , R. S. 1918 . Taxonomy and distribution of Adenostegia . Bulletin of the manuscript. This research was supported by a Graduate Fellowship in Torrey Botanical Club 45 : 399 – 423 . Molecular Systematics from the University of Washington Department Gray, A. 1862 . Revision of the genus Castilleja . American Journal of Science of Botany, the Research Award for Graduate Students from the American II 34 : 335 – 339 . Society of Plant Taxonomists, the Karling Graduate Student Research Gray, A. 1867 . Characters of new plants of California and elsewhere, prin- Award from the Botanical Society of America, the Award for Graduate cipally of those collected by H. N. Bolander in the state geological Student Research from the Society of Systematic Biologists, Sigma-Xi survey . Proceedings of the American Academy of Arts 7 : 327 – 402 . Grants in Aid of Research from the University of Washington Chapter of Gray, A. 1883 . Contributions of North American Botany . Proceedings of the Sigma-Xi, and the Giles Award for Graduate Student Field Research from American Academy of Arts 19 : 1 – 94 . the University of Washington Department of Botany to D. C. Tank, and Gray, A. 1886 . Scrophulariaceae . Pp. 244 – 310 in Synoptical flora of North the NSF Doctoral Dissertation Improvement Grant DEB-0412653 to R. G. America: Volume II - Part I. Gamopetalae after Compositae . New York : Olmstead for D. C. Tank. Ivison, Blakeman, Taylor & Co. Greene, E. L. 1885 . Studies in the botany of California and parts adjacent . Literature Cited Bulletin of the California Academy of Science 1 : 66 – 155 . Greene, E. L. 1886 . Studies in the botany of California and parts adjacent . APG (Angiosperm Phylogeny Group) II . 2003 . An update of the Bulletin of the California Academy of Science 1 : 179 – 228 . Angiosperm Phylogeny Group classification for the orders and fami- Heckard, L. R. 1968 . Chromosome numbers and polyploidy in Castilleja lies of flowering plants: APG II . Botanical Journal of the Linnean Society (Scrophulariaceae) . Brittonia 20 : 212 – 226 . 141 : 399 – 436 . Heckard, L. R. and T. I. Chuang . 1977 . Chromosome numbers, polyploidy, Behr , H. H. 1855 . A new genus and species , Chloropyron palustre. Proceedings and hybridization in Castilleja (Scrophulariaceae) of the Great Basin of the California Academy of Sciences 1 : 62 . and Rocky Mountains . Brittonia 29 : 159 – 172 . Bennett, J. R. and S. Mathews . 2006 . 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Holmgren, N. H. 1971 . A taxonomic revision of the Castilleja viscidula Pennell, F. W. 1951 . Scrophulariaceae . Pp. 686 – 859 in Illustrated flora group . Memoirs of the New York Botanical Garden 21 : 1 – 63 . of the Pacific States ed. L. Abrams . Stanford : Stanford University Holmgren, N. H. 1973 . Five new species of Castilleja (Scrophulariaceae) Press . from the Intermountain Region. Bulletin of the Torrey Botanical Club Rydberg, P. A. 1917 . Flora of the Rocky Mts. and adjacent plains , New York : 100 : 83 – 93 . Published by the author . Holmgren, N. H. 1976 . Four new species of Mexican Castilleja (Subgenus Takhtajan, A. 1997 . Diversity and Classification of Flowering Plants , New Castilleja , Scrophulariaceae) and their relatives . Brittonia 28 : 195 – 208 . York : Columbia University Press . Holmgren , N. H. 1978 . Castilleja (Scrophulariaceae) of Costa Rica and Tank, D. C. and R. G. Olmstead . 2008 . From annuals to perennials: phy- Panama . Brittonia 30 : 182 – 194 . logeny of subtribe Castillejinae (Orobanchaceae) . American Journal of Holmgren, N. H. 1984 . Scrophulariaceae . Pp. 344 – 506 in Intermoun- Botany 95 : 608 – 625 . tain Flora , eds. A. Cronquist , A. H. Holmgren , N. H. Holmgren , J. L. Tank, D. C. , P. M. Beardsley , S. A. Kelchner , and R. G. Olmstead . 2006 . L.A.S. Reveal , and P. K. Holmgren . New York : New York Botanical Garden . Johnson review no. 7. Review of the systematics of Scrophulariaceae Jepson, J. L. 1925 . Manual of the flowering plants of California , Berkeley : s.l. and its current disposition. Australian Systematic Botany 19 : University of California Press . 289 – 307 . Judd , W. S., C. S. Campbell , E. A. Kellogg , P. F. Stevens , and M. J. Donoghue . Watson, S. 1871 . Botany . Vol. 5 of United States geological exploration of the 2007 . Plant Systematics: A Phylogenetic Approach. 3 , Sunderland : fortieth parallel , ed. C. King . Washington, D.C. : Government Printing Sinauer Associates Inc . Office . Keck, D. D. 1927 . A revision of the genus Orthocarpus. Proceedings of the Wettstein, R. 1891 . Scrophulariaceae . Pp. 39– 107 in Die Natürlichen California Academy of Science Series 4 16 : 517 – 571 . Pflanzenfamilien eds. A. Engler , and K. Prantl . Leipzig : Wilhelm Nesom, G. L. 1992a . Taxonomy of the Castilleja tenuiflora group (Scro- Engelmann . phulariaceae) in Mexico, with an overview of sect. Castilleja. Phytologia Wiggins, I. L. 1933 . New plants from Baja California . Contributions from the 73 : 389 – 415 . Dudley Herbarium 1 : 161 – 187 . Nesom , G. L. 1992b . Castilleja section Euchroma (Scrophulariaceae) in Mexico: Wiley, E. O. 1979 . Annotated Linnaean hierarchy, with comments on natu- new species and comments on other taxa . Phytologia 73 : 384 – 388 . ral taxa and competing systems . Systematic Zoology 28 : 308 – 337 . Nesom, G. L. 1992c . New species and taxonomic evaluations of Mexican Wolfe, A. D. , C. P. Randle , L. Liu , and K. E. Steiner . 2005 . Phylogeny and Castilleja (Scrophulariaceae) . Phytologia 72 : 231 – 252 . biogeography of Orobanchaceae . Folia Geobotanica 40 : 115 – 134 . Nesom, G. L. 1994 . A new species of Castilleja (Scrophulariaceae) from Young, N. D. , K. E. Steiner , and C. W. dePamphilis . 1999 . The evolution Chihuahua, Mexico . Phytologia 76 : 406 – 409 . of parasitism in Scrophulariaceae/Orobanchaceae: plastid gene Nuttall, T. 1818 . The genera of North American plants , Philadelphia : sequences refute an evolutionary transition series. Annals of the D. Heart . Missouri Botanical Garden 86 : 876 – 893 . Ownbey, M. 1959 . Castilleja . Pp. 295 – 326 in Vascular plants of the Pacific Northwest eds. C. L. Hitchcock , A. Cronquist , M. Ownbey , and J. W. Appendix 1. Synoptical classification of Castillejinae. A comparison of Thompson . Seattle : University of Washignton Press . the phylogenetic classification proposed here to the placement of species Pennell, F. W. 1935 . The Scrophulariaceae of eastern temperate North following the most recent traditional classification ( Chuang and Heckard America . Academy of Natural Sciences of Philadelphia Monographs 1 : 1986 , 1991 , 1992 ). Species that were included in the molecular phyloge- 1 – 650 . netic analyses of Tank and Olmstead (2008) are indicated with an asterisk Pennell , F. W. 1947 . Some hitherto undescribed Scrophulariaceae of the (*). The remaining species recognized by the authors are included in their Pacific states. Proceedings. Academy of Natural Sciences of Philadelphia respective genera based on the presence of synapomorphies as discussed 99 : 151 – 171 . in the text. Infraspecific taxa are not indicated.

Phylogenetic Classification Chuang and Heckard ( 1986 , 1991 , 1992 ) Castilleja Mutis ex L.f. (/Rhynchapetala ) Castilleja affinis* Hook. & Arn. Castilleja subg. Castilleja Castilleja alpicola T. I. Chuang & Heckard Castilleja subg. Colacus sect. Oncorhynchus Castilleja altorum Standl. & Steyerm. Castilleja subg. Castilleja Castilleja ambigua* Hook. & Arn. Castilleja subg. Colacus sect. Oncorhynchus Castilleja angustata (Rob. & Seaton) Eastw.Castilleja subg. Castilleja Castilleja angustifolia (Nutt.) G. Don. Castilleja subg. Castilleja Castilleja applegatei* Fernald Castilleja subg. Castilleja Castilleja aquariensis N. H. HolmgrenCastilleja subg. Castilleja Castilleja arachnoidea* Greenm. Castilleja subg. Colacus sect. Pilosae Castilleja arvensis* Schlecht. & Cham.Castilleja subg. Castilleja Castilleja aspera Eastw. Castilleja subg. Castilleja Castilleja attenuata* (A. Gray) T. I. Chuang & Heckard Castilleja subg. Colacus sect. Oncorhynchus Castilleja aurea Rob.& Greenm. Castilleja subg. Castilleja Castilleja auriculata* Eastw Castilleja subg. Castilleja Castilleja austromontana Standl. & BlumerCastilleja subg. Castilleja Castilleja beldingii* (Greene) Tank & J. M. Egger Clevelandia Castilleja bella Standl. Castilleja subg. Castilleja Castilleja brevilobata Piper Castilleja subg. Castilleja Castilleja brevistyla* (Hoover) T. I. Chuang & Heckard Castilleja subg. Colacus sect. Oncorhynchus Castilleja bryantii Brandegee Castilleja subg. Castilleja Castilleja campestris* (Benth.) T. I. Chuang & Heckard Castilleja subg. Colacus sect. Oncorhynchus Castilleja cerroana Edwin Castilleja subg. Colacus sect. Pallescentes Castilleja cervina Greenm. Castilleja subg. Colacus sect. Pallescentes Castilleja chambersii J. M. Egger & MeinkeCastilleja subg. Castilleja Castilleja chlorosceptron G. L. Nesom Castilleja subg. Castilleja Castilleja chlorotica Piper Castilleja subg. Castilleja Castilleja christii N. H. Holmgren Castilleja subg. Castilleja Castilleja chromosa* A. Nelson Castilleja subg. Castilleja Castilleja chrymactis Pennell Castilleja subg. Castilleja Castilleja chrysantha Greenm. Castilleja subg. Colacus sect. Pallescentes (continued) 2009] TANK ET AL.: CLASSIFICATION OF CASTILLEJINAE 195

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Phylogenetic Classification Chuang and Heckard ( 1986 , 1991 , 1992 ) Castilleja cinerea A. Gray Castilleja subg. Colacus sect. Pilosae Castilleja coccinea (L.) Spreng Castilleja subg. Castilleja Castilleja conzattii* Fernald Castilleja subg. Castilleja Castilleja covilleana Henderson Castilleja subg. Castilleja Castilleja crista-galli Rydb. Castilleja subg. Castilleja Castilleja cryptandra Eastw. Castilleja subg. Castilleja Castilleja cryptantha Pennell & G.N. JonesCastilleja subg. Colacus sect. Pallescentes Castilleja ctenodonta Eastw. Castilleja subg. Castilleja Castilleja cusickii* Greenm. Castilleja subg. Colacus sect. Pallescentes Castilleja dendridion G. L. Nesom Castilleja subg. Castilleja Castilleja densiflora* (Benth.) T. I. Chuang & Heckard Castilleja subg. Colacus sect. Oncorhynchus Castilleja dissitiflora N. H. Holmgren Castilleja subg. Castilleja Castilleja disticha Eastw. Castilleja subg. Castilleja Castilleja divaricata Benth. Castilleja subg. Castilleja Castilleja durangensis G.L. Nesom Castilleja subg. Castilleja Castilleja ecuadorensis N. H. HolmgrenCastilleja subg. Castilleja Castilleja elata* Piper Castilleja subg. Castilleja Castilleja elegans Ostenf. ex Malte Castilleja subg. Castilleja Castilleja elmeri* Fernald Castilleja subg. Castilleja Castilleja exserta* (A. Heller) T. I. Chuang & Heckard Castilleja subg. Colacus sect. Oncorhynchus Castilleja filiflora G. L. Nesom Castilleja subg. Castilleja Castilleja fissifolia L.f. Castilleja subg. Castilleja Castilleja flava S. Watson Castilleja subg. Colacus sect. Pallescentes Castilleja foliolosa Hook. & Arn. Castilleja subg. Castilleja Castilleja fraterna Greenm. Castilleja subg. Castilleja Castilleja fruticosa Moran Castilleja subg. Castilleja Castilleja galehintoniae G. L. Nesom Castilleja subg. Castilleja Castilleja genevievana G. L. Nesom Castilleja subg. Castilleja Castilleja glandulifera Pennell Castilleja subg. Castilleja Castilleja glandulosa Greenm. Castilleja subg. Castilleja Castilleja gleasonii Elmer Castilleja subg. Castilleja Castilleja gracilis Benth. Castilleja subg. Castilleja Castilleja gracillima Rydb. Castilleja subg. Castilleja Castilleja grisea Dunkle Castilleja subg. Castilleja Castilleja guadalupensis Brandegee Castilleja subg. Castilleja Castilleja haydenii (A. Gray) Cockerell Castilleja subg. Castilleja Castilleja hidalgensis J. M. Egger Castilleja subg. Castilleja Castilleja hirsuta M. Martens & Galeotti Castilleja subg. Castilleja Castilleja hispida* Benth. Castilleja subg. Castilleja Castilleja holmgrenii J. M. Egger Castilleja subg. Castilleja Castilleja hololeuca Greene Castilleja subg. Castilleja Castilleja hyperborea Pennell Castilleja subg. Castilleja Castilleja indivisa Engelm. Castilleja subg. Castilleja Castilleja integra* A. Gray Castilleja subg. Castilleja Castilleja integrifolia* L.f. Castilleja subg. Castilleja Castilleja irasuensis Oersted Castilleja subg. Castilleja Castilleja jiquilpana G. L. Nesom Castilleja subg. Castilleja Castilleja kaibabensis N. H. HolmgrenCastilleja subg. Castilleja Castilleja kraliana J. R. Allison Castilleja subg. Castilleja Castilleja lacera* (Benth.) T. I. Chuang & Heckard Castilleja subg. Colacus sect. Oncorhynchus Castilleja laciniata Hook. & Arn. Castilleja subg. Colacus sect. Oncorhynchus Castilleja lanata A. Gray Castilleja subg. Castilleja Castilleja lasiorhyncha* (A. Gray) T. I. Chuang & Heckard Castilleja subg. Colacus sect. Oncorhynchus Castilleja latifolia Hook. & Arn. Castilleja subg. Castilleja Castilleja lemmonii* A. Gray Castilleja subg. Colacus sect. Pallescentes Castilleja lentii N. H. Holmgren Castilleja subg. Castilleja Castilleja levisecta Greenm. Castilleja subg. Colacus sect. Pallescentes Castilleja linariifolia* Benth. Castilleja subg. Castilleja Castilleja lineariloba* (Benth.) T. I. Chuang & Heckard Castilleja subg. Colacus sect. Oncorhynchus Castilleja lineata Greene Castilleja subg. Colacus sect. Pilosae Castilleja linifolia N. H. Holmgren Castilleja subg. Castilleja Castilleja litoralis Pennell Castilleja subg. Castilleja Castilleja ludoviciana Pennell Castilleja subg. Castilleja Castilleja lutescens* (Greenm.) Rydb. Castilleja subg. Colacus sect. Pallescentes Castilleja macrostigma B. Robinson Castilleja subg. Castilleja Castilleja martinii Abrams Castilleja subg. Castilleja Castilleja mcvaughii N. H. Holmgren Castilleja subg. Castilleja Castilleja mendocinensis (Eastw.) PennellCastilleja subg. Castilleja Castilleja meridensis Pennell Castilleja subg. Castilleja (continued) 196 SYSTEMATIC BOTANY [Volume 34

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Phylogenetic Classification Chuang and Heckard ( 1986 , 1991 , 1992 ) Castilleja mexicana (Hemsl.) A. Gray Castilleja subg. Castilleja Castilleja miniata* Douglas ex Hook. Castilleja subg. Castilleja Castilleja minor* (A. Gray) A. Gray Castilleja subg. Castilleja Castilleja mogollonica Pennell Castilleja subg. Castilleja Castilleja mollis Pennell Castilleja subg. Castilleja Castilleja montigena Heckard Castilleja subg. Castilleja Castilleja moranensis Kunth. Castilleja subg. Castilleja Castilleja nana* Eastw. Castilleja subg. Colacus sect. Pilosae Castilleja nelsonii Eastw. Castilleja subg. Castilleja Castilleja nervata Eastw. Castilleja subg. Castilleja Castilleja nitricola Eastw. Castilleja subg. Castilleja Castilleja nivea Pennell & Ownbey Castilleja subg. Colacus sect. Pilosae Castilleja nivibractea* G. L. Nesom Castilleja subg. Castilleja Castilleja nubigena Kunth Castilleja subg. Castilleja Castilleja occidentalis* Torr. Castilleja subg. Castilleja Castilleja ophiocephala * Tank & J. M. EggerOphiocephalus Castilleja oresbia* Greenm. Castilleja subg. Colacus sect. Pallescentes Castilleja organorum Standl. Castilleja subg. Castilleja Castilleja ornata Eastw. Castilleja subg. Castilleja Castilleja ortegae Standl. Castilleja subg. Castilleja Castilleja pallescens (A. Gray) Greenm. Castilleja subg. Colacus sect. Pallescentes Castilleja pallida (L.) Spreng. Castilleja subg. Castilleja Castilleja papilionacea G. L. Nesom Castilleja subg. Castilleja Castilleja parviflora* Bong. Castilleja subg. Castilleja Castilleja parvula Rydb. Castilleja subg. Castilleja Castilleja patriotica Fernald Castilleja subg. Castilleja Castilleja peckiana* Pennell Castilleja subg. Castilleja Castilleja pectinata M. Martens & GaleottiCastilleja subg. Castilleja Castilleja peirsonii* Eastw. Castilleja subg. Castilleja Castilleja perelegans G. L. Nesom Castilleja subg. Castilleja Castilleja peruviana T. I. Chuang & Heckard Castilleja subg. Colacus sect. Oncorhynchus Castilleja pilosa* (S. Watson) Rydb. Castilleja subg. Colacus sect. Pilosae Castilleja plagiotoma* A. Gray Castilleja subg. Castilleja Castilleja porphyrosceptron G. L. Nesom Castilleja subg. Castilleja Castilleja praeterita* Heckard & Bacig. Castilleja subg. Colacus sect. Pallescentes Castilleja profunda T. I. Chuang & HeckardCastilleja subg. Colacus sect. Oncorhynchus Castilleja pruinosa* Fernald Castilleja subg. Castilleja Castilleja pseudohyperborea Rebr. Castilleja subg. Castilleja Castilleja pterocaulon N. H. Holmgren Castilleja subg. Castilleja Castilleja puberula Rydb. Castilleja subg. Castilleja Castilleja pulchella Rydb. Castilleja subg. Colacus sect. Pallescentes Castilleja pumila (Benth.) Wedd. ex HerreraCastilleja subg. Castilleja Castilleja purpurascens Greenm. Castilleja subg. Castilleja Castilleja purpurea (Nutt.) G. Don Castilleja subg. Castilleja Castilleja quirosii Standl. Castilleja subg. Castilleja Castilleja racemosa* (Breedlove & Heckard) T. I. Chuang & HeckardCastilleja subg. Gentrya Castilleja raupii Pennell Castilleja subg. Castilleja Castilleja revealii N. H. Holmgren Castilleja subg. Castilleja Castilleja rhexifolia Rybd. Castilleja subg. Castilleja Castilleja rhizomata N. H. Holmgren Castilleja subg. Castilleja Castilleja rigida Eastw. Castilleja subg. Castilleja Castilleja roei Crosswh. Castilleja subg. Castilleja Castilleja rubicundula* (Jeps.) T. I. Chuang & HeckardCastilleja subg. Colacus sect. Oncorhynchus Castilleja rubida Piper Castilleja subg. Castilleja Castilleja rubra (Drob.) Rebr. Castilleja subg. Castilleja Castilleja rupicola Piper ex Fernald Castilleja subg. Castilleja Castilleja salsuginosa N. H. Holmgren Castilleja subg. Colacus sect. Pilosae Castilleja saltensis Eastw. Castilleja subg. Castilleja Castilleja scabrida Eastw. Castilleja subg. Castilleja Castilleja schizotricha Greenm. Castilleja subg. Colacus sect. Pilosae Castilleja scorzonerifolia* Kunth Castilleja subg. Castilleja Castilleja septentrionalis* Lindl. Castilleja subg. Castilleja Castilleja sessiliflora Pursh Castilleja subg. Castilleja Castilleja sphaerostigma Eastw. Castilleja subg. Castilleja Castilleja spiranthoides Standl. Castilleja subg. Castilleja Castilleja stenophylla M. E. Jones Castilleja subg. Castilleja Castilleja steyermarkii Pennell Castilleja subg. Castilleja Castilleja stipifolia G. L. Nesom Castilleja subg. Castilleja Castilleja subalpina Eastw. Castilleja subg. Castilleja (continued) 2009] TANK ET AL.: CLASSIFICATION OF CASTILLEJINAE 197

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Phylogenetic Classification Chuang and Heckard ( 1986 , 1991 , 1992 ) Castilleja subinclusa Greene Castilleja subg. Castilleja Castilleja suksdorfii A. Gray Castilleja subg. Castilleja Castilleja talamancensis N. H. Holmgren Castilleja subg. Castilleja Castilleja tapeinoclada Loesner Castilleja subg. Castilleja Castilleja tayloriorum N. H. HolmgrenCastilleja subg. Castilleja Castilleja tenuiflora* Benth. Castilleja subg. Castilleja Castilleja tenuifolia Martens & GaleottiCastilleja subg. Castilleja Castilleja tenuis* (A. Heller) T. I. Chuang & HeckardCastilleja subg. Colacus sect. Oncorhynchus Castilleja thompsonii Pennell Castilleja subg. Colacus sect. Pallescentes Castilleja tolucensis Kunth Castilleja subg. Castilleja Castilleja trujillensis Pennell Castilleja subg. Castilleja Castilleja uliginosa Eastw. Castilleja subg. Castilleja Castilleja unalaschensis (Cham. & Schl.) MalteCastilleja subg. Castilleja Castilleja vadosa T. I. Chuang & HeckardCastilleja subg. Colacus sect. Oncorhynchus Castilleja variocolorata A. P. Khokhr.Castilleja subg. Castilleja Castilleja venusta Rzed. Castilleja subg. Castilleja Castilleja victoriae Fairbarns & J. M. EggerCastilleja subg. Colacus sect. Oncorhynchus Castilleja virgata (Wedd.) EdwinCastilleja subg. Castilleja Castilleja viscidula A. Gray Castilleja subg. Castilleja Castilleja wightii Elmer Castilleja subg. Castilleja Castilleja wootonii Standl. Castilleja subg. Castilleja Castilleja xanthotricha* Pennell Castilleja subg. Colacus sect. Pallescentes Castilleja zempoaltepetlensis* G. L. Nesom Castilleja subg. Castilleja Triphysaria Fisch. & C. A. Mey. (/Rhynchapetala ) Triphysaria eriantha* (Benth.) T. I. Chuang & HeckardTriphysaria Triphysaria floribunda* (Benth.) T. I. Chuang & HeckardTriphysaria Triphysaria micrantha* (Greene ex A. Heller) T. I. Chuang & HeckardTriphysaria Triphysaria pusilla* (Benth.) T. I. Chuang & HeckardTriphysaria Triphysaria versicolor* Fisch. & C. A. Mey. Triphysaria Chloropyron Behr (/ Pseudocordylanthus ) Chloropyron maritimum* (Nutt. ex Benth.) A. HellerCordylanthus subg. Hemistegia Chloropyron molle* (A. Gray) A. HellerCordylanthus subg. Hemistegia Chloropyron palmatum* (Ferris) Tank & J. M. EggerCordylanthus subg. Hemistegia Chloropyron tecopense* (Munz & J. C. Roos) Tank & J. M. EggerCordylanthus subg. Hemistegia Dicranostegia (A. Gray) Pennell (/ Pseudocordylanthus ) Dicranostegia orcuttiana* (A. Gray) PennellCordylanthus subg. Dicranostegia Cordylanthus Nutt. ex. Benth Cordylanthus capitatus * Nutt. ex Benth.Cordylanthus subg. Cordylanthus sect. Anisocheila Cordylanthus eremicus* (Coville & C. V. Morton) MunzCordylanthus subg. Cordylanthus sect. Anisocheila Cordylanthus kingii* S. Watson Cordylanthus subg. Cordylanthus sect. Anisocheila Cordylanthus laxiflorus A. GrayCordylanthus subg. Cordylanthus sect. Anisocheila Cordylanthus nevinii A. Gray Cordylanthus subg. Cordylanthus sect. Cordylanthus Cordylanthus nidularius J. T. HowellCordylanthus subg. Cordylanthus sect. Cordylanthus Cordylanthus parviflorus (Ferris) WigginsCordylanthus subg. Cordylanthus sect. Anisocheila Cordylanthus pilosus* A. Gray Cordylanthus subg. Cordylanthus sect. Cordylanthus Cordylanthus pringlei* A. Gray Cordylanthus subg. Cordylanthus sect. Cordylanthus Cordylanthus rigidus* (Benth.) Jeps.Cordylanthus subg. Cordylanthus sect. Cordylanthus Cordylanthus tenuis* A. Gray Cordylanthus subg. Cordylanthus sect. Cordylanthus Cordylanthus ramosus* Nutt. ex Benth.Cordylanthus subg. Cordylanthus sect. Ramosi Cordylanthus wrightii* A. GrayCordylanthus subg. Cordylanthus sect. Anisocheila Orthocarpus Nutt. Orthocarpus barbatus* J. S. Cotton Orthocarpus Orthocarpus bracteosus* Benth. Orthocarpus Orthocarpus cuspidatus* GreeneOrthocarpus Orthocarpus imbricatus* Torr. ex S. Watson Orthocarpus Orthocarpus luteus* Nutt. Orthocarpus Orthocarpus pachystachyus* A. Gray Orthocarpus Orthocarpus purpureo-albus* A. Gray ex S. Watson Orthocarpus Orthocarpus tenuifolius* (Pursh) Benth.Orthocarpus Orthocarpus tolmiei* Hook. & Arn. Orthocarpus