Aliso: A Journal of Systematic and Floristic Botany Aliso: A Journal of Systematic and Floristic Botany A Fistful of Polemoniaceae: New Names and Combinations A Fistful of Polemoniaceae: New Names and Combinations

New taxa, names, and combinations are provided for five genera in Polemoniaceae: Dayia , Giliastrum , Leptosiphon , Linanthus , and Loeselia . Combinations include the transfer to Dayia of two species formerly included in Ipomopsis and one in Bryantiella ( Dayia glutinosa , D. havardii , and D. sonorae ), as well as the return of Giliastrum stewartii to species rank. A new name is offered for Linanthus aureus when transferred into Leptosiphon : Leptosiphon chrysanthus and its subspecies, L. chrysanthus subsp. decorus . New combinations for infraspecific taxa are made available for Linanthus californicus (subspp. glandulosus and tomentosus ) and L. pungens (subspp. hookeri , pulchriflorus and hallii ). New subspecies are described for Linanthus bigelovii (subsp. johnsonii ), L. dichotomus (subsp. pattersonii ), L. maculatus (subsp. emaculatus ), L. watsonii (subspp. laccolithicus and dolomiticus ), and Loeselia glandulosa (subsp. sonorae ). Linanthus maricopensis is described as new. In addition, 10 lectotypes are designated, associated with Dayia , Leptosiphon , and Linanthus . , Linanthus maricopensis , Linanthus watsonii subsp. dolomiticus , Linanthus watsonii subsp. laccolithicus , Loeselia glandulosa subsp. sonorae , Polemoniaceae.

different tribes of Polemoniaceae (Porter and Johnson 2000): Gilia in tribe Gilieae V.E. Grant and Giliastrum in tribe Loeselieae J. M. Porter & L.A. Johnson. Further, species of Giliastrum are characteristically perennials, with rotate corollas, the corolla lobes longer than the fused portion and have a unique, zonopororate-pertectate pollen type. Species of Gilia, in contrast, are annuals, with funnelform corollas, the lobes shorter than the fused portion and have a more common zonocolporate-striate pollen type.
Giliastrum's current circumscription includes nine species (Porter and Johnson 2000). One, Giliastrum purpusii (Brandegee) J.M. Porter, incorporates three subspecies. A reevaluation of these subspecies in light of their interrelationships and relationship with the Argentine species, G. castillanosii J.M. Porter, based on comparative DNA sequencing of the chloroplast trnL-F region and nuclear ribosomal ITS region (J. M. Porter and V. W. Steinmann, unpubl.) and comparative morphology has prompted a change in nomenclature.

LEPTOSIPHON AUREUS
Polemoniaceae has experienced more than its share of taxonomic and nomenclatural turmoil. Taxon circumscriptions at all ranks have fluctuated wildly over the past 200 years (compare Bentham 1833Bentham , 1845Gray 1870Gray , 1878Grant 1959;Porter and Johnson 2000). This has contributed to a complex and bewildering nomenclature. Because many of the original names predated the requirement for type specimens, the precise application of many names has been a matter of supposition, as lectotypification has not been accomplished for many species. Recently, a series of investigations into the phylogeny and generic boundaries in Polemoniaceae (Johnson et al. 1996(Johnson et al. , 2008Porter 1997;Bell et al. 1999;Bell and Patterson 2000) have made it very clear that Linanthus in the traditional sense (e. g., Grant 1959;Patterson 1993) is not monophyletic. In their revised classification of Polemoniaceae, Porter and Johnson (2000) divided Linanthus into two genera, resurrecting and expanding Leptosiphon Benth., in addition to Linanthus. As part of this change, they transferred Linanthus aureus (Nutt.) Greene into Leptosiphon. They were unaware that their combination was a later homonym of Leptosiphon aureus Benth. ex E. Vilm. Unfortunately, these two names do not refer to the same species.
Here we correct this error by providing a legitimate transfer of Gilia aurea Nutt. into Leptosiphon and clear up nomenclatural confusion involving the species known as Leptosiphon aureus.
Scattered across the arid lands of the southwestern USA and adjacent Mexico is a delicate spring annual with striking goldenyellow flowers, filamentous stems and opposite, palmate leaves. Although originally included in the genus Gilia by Thomas Nuttall (1848), for the last century it has been known as Linanthus aureus (Greene 1892;Fig. 6). While there seems no controversy concerning the application of Nuttall's name, there has been no clear identification of type material. Nuttall's protologue indicates that a collection by W. Gambel, made in (or around) Santa Barbara, California, was the basis for Gilia aurea. Three specimens are believed to be those of W. Gambel, GH 78819, K 000769037 and PH 1068920/12188. The specimen in Gray Herbarium states, ''Gilia *(1 Chrysantha *aurea, St. --Barb Angels,'' in Nuttall's hand and ''Nutt., Gambel'' in Gray's hand. This specimen was presented to GH by E. Durand in 1866 (fide annotation). The mount at K states, ''Gilia * aurea, Sta. Barbara, in Nuttall's hand and includes three plants. The specimen at Philadelphia Academy of Sciences states '' G. chrysantha, Nutt., Calif., Durand.'' The GH and PH specimens have the same source, the Durand Herbarium, where much of Nuttall's materials, including some of Gambel's collections ultimately resided. A large collection of Nuttall's materials is also found at K. While the GH specimen bears a label in Nuttall's hand, the location-St. Angels (Los Angeles)-is in conflict with the protologue. It is possible that the protologue was in error, as the change in location on the GH sheet was made by Nuttall, and subsequent comments by Gray (1870: 264) suggest that there was a question whether the specimen was collected in Santa Barbara or Los Angeles. Note that no other historic or modern collections have been made at Santa Barbara, but the species is historically known from Los Angeles. Regardless, the specimen at K is not in conflict with the protologue and most closely matches the illustration and description provided by Nuttall. The lectotype is therefore designated as K 000769037. The mounts at GH (78819) and PH (1068920/12188) are here interpreted as isolectotypes (material from the same W. Gambel collection).
Within the second edition of Les Fleurs de Pleine Terre (1866), Elisa de Vilmorin (director of Vilmorin-Andrieux and Co.) discusses five species under the generic name Leptosiphon. The third of these is '' Leptosiphon jaune d'or.-L. aureus, Benth.'' No indication of the source of the epithet aureus, other then the name Bentham is provided. However, it is also relevant to note that Vilmorin described the flowers in some detail: ''Les fleurs, très-nombreuses, disposées en corymbe, sont entourées chacune par un calice dont les 5 segments, de même forme que les feuilles, se confondent avec elles; le tube de la corolle est long d'un centimètre et demi; sa couleur est d'un blanc jaunâ tre analogue à celle de la partie inférieure du limbe; ce limbe est à 5 divisions ovales, étalées et d'un jaune doré; chacune de ces divisions est marquée à la base de deux petites taches purpurines qui forment par leur ensemble une jolie petite couronne; étamines et pistil saillants, de couleur orangée' ' (pp. 470-471). The description indicates that this species has a long, narrow corolla tube approximately 1.5 cm long, as do many species of Leptosiphon. This description clearly excludes Nuttall's Gilia aurea, but is consistent with a number of other species, including the yellow-flowered Leptosiphon acicularis (Greene) Jeps. Bentham's (1833) treatment of Polemoniaceae, in which he described the genus Leptosiphon, does not include the epithet aureus. However, in Plantae Hartwegianae (1849), Bentham provides the name Gilia (Leptosiphon) micrantha Steud. var. aurea, based upon Hartweg 234. Interestingly, of this taxon Bentham says only, ''nec aliter a species differre videtur'' (not otherwise different from the species it seems), providing no description. This apparently indicates that Bentham did not accept the taxon as different from G. micrantha. Given that Vilmorin cited Plantae Hartwegianae often, this is unquestionably the source of the epithet aureus. Subsequent editions of Les Fleurs de Pleine Terre, e.g., the 3 rd -5 th editions (1870,1894,1909), continued to recognize L. aureus, but also stated, ''syn. lat. Gilia lutea Steud. var. aurea Hort.'' (pp. 594,549,591,respectively). In addition to the citation, an illustration is also provided of Leptosiphon jaune d'or ( Fig. 7).
Brand's monograph of Polemoniaceae (1907) also includes many of Vilmorin's names. Like Gray's treatment in the  (1878) This rather lengthy explanation provides the justification for proposing that Gilia micrantha var. aurea Benth, nom. nud., and Leptosiphon aureus Benth. ex E. Vilm. are the same taxon. This conclusion is further supported by collections at K (000750977) and P (04591512). The former is a collection of Hartweg, identified as G. micrantha var. aurea, the latter identified as Leptosiphon aureus Benth. (Hortus Parisiensis, 1869). Both of these specimens represent the same taxon, Leptosiphon acicularis (Greene) Jeps. Although the name L. acicularis has been applied for over a century, it is now evident that the earliest legitimate name for this taxon is L. aureus. This also means that Linanthus (Gilia) aureus cannot be transferred to Leptosiphon using the original epithet. Here we provide a new name:
Navarretia aurea (Nutt.) Kuntze, Revis. Gen. Pl. 2: 433 (1891 Fam. 250: 131 (1907). Linanthus aureus (Nutt.) Greene var. decora (A. Gray) Jeps., Man. Fl. Pl. Calif. [Jepson] 803 (1925 The nomenclatural changes proposed above are unfortunate but necessary. Unfortunate in the sense that two names for well-known species formerly of Linanthus (Linanthus acicularis and L. aureus) cannot retain their specific epithets when transferred to Leptosiphon. This change also results in the confusing situation in which the name Leptosiphon aureus remains a legitimate name of priority; however, it refers not to the species formerly called Linanthus aureus, but to the species formerly called Linanthus acicularis. While this is unsatisfying, it is the only recourse aside from a proposal to conserve, which would have no guarantee of success.

A NEW SUBSPECIES OF LINANTHUS BIGELOVII
Field studies and examination of herbarium collections, with special attention to the type, has revealed considerable variation across the range (western Texas to California) of Linanthus bigelovii (A. Gray) Greene. This variation has resulted in some degree of confusion and mischaracterization concerning L. bigelovii and differences between it and L. jonesii (A. Gray) Greene. Specifically, populations of L. bigelovii in California have correctly been described as glabrous and glaucous (occasionally a few glandular trichomes may be present); however, those from the eastern portion of the range, including the type, are sparsely beset with glandular trichomes, particularly on the pedicels, but also sparsely so on the calyx (Fig. 8). Linanthus jonesii, with its coarsely glandular pedicels and calyx, is easily distinguished from L. bigelovii in California, but the distinction is less clear in Arizona, where L. bigelovii tends to also have glandular trichomes. The glandular trichomes of L. bigelovii are uniseriate with globular terminal glands and differ from the coarse multiseriate glandular trichomes with flattened, tack-like terminal glands found on L. jonesii (Fig. 8). In contrast with the eastern populations, the populations of L. bigelovii in California have glabrous pedicels and calyces. A somewhat more subtle difference is found in seed morphology. Seeds of L. bigelovii from the eastern portions of its range are small, elliptical to oblong in outline, usually notched at the hilum, and sometimes with narrow winged margins (Fig. 9A). Importantly, sometimes the seeds lack the winged margins and are nearly indistinguishable from those of L. jonesii (Fig. 9C). Linanthus bigelovii from California has seeds that are slightly larger, elliptical to oblong in outline, sometimes with very narrow wing-like edges, but generally lacking the notching at the hilum ( Erect, taprooted annual 4.5-47 cm tall, simple to more often freely, dichotomously branched; cotyledons 5.2-9.5 mm long, linear or narrowly linear-lanceolate, sheathing, sparsely ciliate villous proximally. Stems nearly glabrous throughout, or with a few eglandular and rarely glandular trichomes at the nodes; nodes generally opposite throughout. Leaves 1.4-6.5 cm long, simple and linear or linear-filiform at the lower nodes, 3-cleft essentially to the base into linear-filiform segments at the more distal nodes, the central lobe the longest, leaf base somewhat sheathing, very sparsely ciliate-villous, stipitate-glandular or glabrous at the leaf base. Inflorescence cymose, diffuse and branching, more or less regularly dichotomous, the terminal flower subsessile or pedicellate, the pedicel 0.6-10 mm long, commonly closely subtended by a pair of leafy bracts, similar to the leaves, 5-30 mm long, simple entire and linear or 3-lobed, the lobes linear. Flowers vespertine (nocturnal), closed during the day, opening in the late afternoon or evening and remaining open through the night. Calyx 8-18 mm long, the green, herbaceous costae terminating in unequal lobes; the lobes 2.4-6.9 mm long, apex acute acicular or mucronate; the tube 6-11 mm long, the conspicuous white-hyaline intervals of the tube as wide as or generally wider than the green and herbaceous costae, glabrous abaxially, sparsely to moderately villous, sometimes also glandular adaxially on the distal tube and proximal lobes. Corolla 8-16.3 mm long, the lobes (2.5-)4-7.5(-9) mm long, 1.7-5.2 mm wide, white, generally flecked with purple, maroon or brownish on the abaxial lobe margin exposed in bud, and a band of purple adaxially in the upper tube or throat, glabrous. Stamens equally or sub-equally inserted 3.5-5.6 mm above corolla insertion, generally 2 mm or more above the apex of the ovary; filaments slender, glabrous, 0.6-1.1(-1.6) mm long; anthers 0.5-0.7 mm long, included; pollen yellow. Ovary 1.9-3.5 mm long, 0.6-1.4 mm in diameter, glabrous; style 0.4-0.8 mm long; stigma lobes 1.2-2.1 mm long. Capsule cylindroid, 7-11 mm long, 1.5-2.8 mm in diameter, dehiscing from the apex down 1.8-2.5 mm, held for some time within the slightly accrescent calyx, the valves remaining attached toward the base at least for a time after dehiscence. Seeds 12-21 seeds per locule, 0.6-1.5 mm long, 0.4-0.7 mm in diameter, pale, more or less ellipsoid, angular, not reniform nor much indented at the hilum, often with very narrow marginal wings along the angular edges, the hilum generally along one of the angled edges, inconspicuous, swelling when wet, and becoming somewhat mucilaginous. 2n 5 18. Fig. 8. Glandular trichomes from the calyx (above) and pedicels (center) of Linanthus bigelovii (A. Gray) Greene subsp. bigelovii and from the calyx of L. jonesii (A. Gray) Greene (below). Note the uniseriate stalk with globular, multicellular distal gland of L. bigelovii subsp. bigelovii (above and center) and the multiseriate stalk and bi-tiered, flat-topped, multicellular, terminal gland of L. jonesii (below). Scale bar 5 0.5 mm. VOLUME 32 (2) Polemoniaceae: New Names and Combinations Key to the Subspecies of Linanthus bigelovii   In the course of reinvestigating the Linanthus species of Arizona, an unusual new taxon was discovered in the central portion of the state. In the past, these populations have consistently been identified as Linanthus dichotomus, owing to their large corollas. However, these populations lack three diagnostic traits of L. dichotomus: (1) inflated and densely pilose filament bases; (2) seeds with a pith-like, whitish and finely cellular-reticulate seed coat; and (3) a glabrous and somewhat glaucous calyx. Rather, the filaments are undilated and glabrous (Fig. 11), seeds are reniform, brown and rugose, and the calyx is coarsely glandular. In particular, the coarse glandular trichomes are similar to those of L. jonesii. The glandular trichomes of L. jonesii have stalk cells that are multiseriate, bearing large, tack-like terminal glands that are multicellular ( Fig. 8). They differ from the trichomes of L. bigelovii which have uniseriate stalks with terminal, 1-8-celled, globular glands (Fig. 8). What distinguishes these plants from L. jonesii are the exceptionally large corollas and the tendency for glandular trichomes of the calyx and pedicel to have uniseriate stalks. Because both L. jonesii and this largeflowered taxon are sympatric throughout the range of the new taxon, we believe that there must be isolation between the two. We therefore treat the new taxon at the rank of species:  . 12).
Erect, taprooted annual 6-25 cm tall, simple to more often freely, dichotomously branched; cotyledons 2.5-4.8 mm long, linear-filiform, sheathing, glabrous or very sparsely ciliate villous proximally. Stems sparsely but coarsely stipitateglandular throughout, more densely stipitate-glandular at and just below the nodes and in the inflorescence; nodes generally opposite, rarely subopposite or alternate. Leaves 8.9-33 mm long, simple and linear or linear-filiform (rarely 3-lobed and the central lobe longest), leaf base somewhat sheathing, coarsely stipitate-glandular on the adaxial proximal surface and margin (ciliate). Inflorescence cymose, diffuse and branching, more or less regularly dichotomous, the pedicel of the terminal flower 1.5-13.5 mm long, coarsely stipitateglandular, closely subtended by a pair of leafy bracts; bracts 9-32 mm long, simple, entire linear-filiform, very rarely 2(-3)cleft into linear-filiform segments, the central lobe the longest, petiole to 2 mm long, coarsely stipitate-glandular. Flowers opening in the late afternoon, or vespertine (nocturnal), closed during the day, opening in the evening and remaining open through the night. Calyx 8-13.2 mm long, the green, herbaceous costae terminating in unequal lobes, 2-7.9 mm long, the tube 5.4-7.5 mm long, the conspicuous white-hyaline intervals of the tube wider than the costae, glandular (the trichomes coarse, with multiseriate stalks and multicellular terminal glands) abaxially, sparsely to moderately villous and glandular adaxially on the distal tube and lobes. Corolla 14-30 mm long, the lobes 7.9-20.6 mm long, 5.4-10.2 mm wide, cream to white, generally flecked with purple abaxially and a band of purple adaxially in the upper tube, glabrous. Stamens equally or subequally inserted 4.4-6.2 mm above corolla insertion, filaments slender, glabrous, 1.1-1.6 mm long, anthers 1.1-1.5 mm long, included, pollen yellow. Ovary 2-2.5 mm long, 0.8-1.0 mm in diameter, narrowly pyriform to cylindric, glabrous, style 0.6-1.3 mm long, stigma lobes 1.4-2.5 mm long. Capsule cylindric, 4.5-9.1 mm long, 1.8-2.5 mm in diameter, held for some time within the slightly accrescent calyx, the valves remaining attached toward the base at least for a time after dehiscence.
Seeds 18-32 seeds per locule, ca. 1 mm long, 0.4 mm in diameter, pale, more or less angular reniform, the hilum indented and white margined, seed coat rugose, swelling when wet, and becoming somewhat mucilaginous.

A NEW SUBSPECIES OF LINANTHUS DICHOTOMUS
The distinction between Linanthus dichotomus Benth. and L. bigelovii (A. Gray) Greene has long been recognized and the traits used to diagnose them are straightforward, if somewhat cryptic. Linanthus dichotomus has large flowers (corolla lobes 10-16 mm long, Patterson 1993), the epipetalous stamens are inflated and densely pilose at their point of attachment (Fig. 13), and the outer seed-coat is pith-like, whitish and finely cellular-reticulate (Fig. 14), unchanged when wetted (not producing mucilage). By contrast, L. bigelovii has smaller flowers (corolla lobes 7-8 mm long, Patterson 1993), the epipetalous stamens are narrow and glabrous at their point of attachment (Fig. 10), and the outer seed-coat is smooth to rugose (at 103) brownish, either angled and narrowly winged or reniform, becoming slightly mucilaginous when wetted. Even so, it seems that there has been some confusion between the two species. For example in the Intermountain Flora, Cronquist (1984) describes the seeds of L. bigelovii as ''sometimes approaching those of L. dichotomus in structure.'' Examination of materials of both Linanthus dichotomus and L. bigelovii has revealed a series of populations in California, Nevada and Baja California, Mexico that possess small flowers like those of L. bigelovii (and have been identified as such) but display the seed (Fig. 14) and stamen features of L. dichotomus (Fig. 15). These populations represent a smallflowered phase of L. dichotomus and are here described as a new subspecies:  (2) Polemoniaceae: New Names and Combinations 0384700E, desert sage scrub/juniper woodland transition zone, 10 Apr 2008 (holotype RSA) (Fig. 16).
Plants 1-19(-22) cm tall; cotyledons linear, sometimes slightly broader distally, 3.5-12.5 mm long, 0.3-0.4 mm wide. Stems glabrous, the primary axis (0.3-)2-18 mm long, 2-7(-8) nodes to the terminal flower; lateral (secondary branches dimorphic, those of the lower nodes to 12 or more cm long, 2-4 nodes to the terminal flower; branches of the uppermost nodes 0.3-6.5 cm long, one node to terminal flower. Flowers vespertine, opening in evening and closing in daylight. Calyx 7-15 mm, membrane much wider than herbaceous costae, the membrane sinus V-shaped or somewhat truncate. Corolla narrowly funnelform, 13-17.5 mm long, tube 6.8-10 mm, purple more distally but orifice white or cream, lobes 4.6-11 mm long, white with light purple shading on abaxial margins. Stamens inserted 2-2.5 mm above the base (insertion) of the corolla, diverging adjacent to the apex of the ovary; filaments 2-2.4 mm long, dilated and villous at the insertion, anthers 1-1.6 mm long.    Porter. VOLUME 32(2) Polemoniaceae: New Names and Combinations  Fam. 250: 144 (1907). This taxon is based upon an H. Bolander collection of 1867. We have been unsuccessful at locating this collection; however, Brand's description suggests that the corolla is larger than the taxon we describe here and, as such, we consider var. uniflora to be synonymous with subsp. dichotomus. Even so, without a type the application of the name is uncertain. The third and final name is Gilia dichotoma var. integra M.E. Jones, Contr. West. Bot. 12: 53 (1908 A. Johnson (Patterson 1993;Patterson and Porter 2012) have paid little attention to infraspecific variation. This is unfortunate, as significant differences in a number of traits corresponding with geography and distribution of genetic variation have been well documented (Eastwood 1904;Gordon-Reedy 1990;Schultz and Soltis 2001). Gordon-Reedy (1990) recognized five subspecies based on a comparative study of trichome type, density and distribution. In 2001, Schultz and Soltis contrasted population genetic divergences with morphological variation in L. californicus. This study found great inconsistency between the patterns of genetic similarity and the subspecies delimitations of Gordon-Reedy (see also Schultz 1992), and concluded that there were three genetically-based morpho-geographic entities. We follow the subspecies delimitation of Schultz and Soltis, but differ in one aspect of nomenclature. Schultz and Soltis (2001) Bot. Gaz. 37: 447 (1904 Bot. Gard. 6: 438 (1910 Although Eastwood (1904) states that these collections were at CAS, none are there currently. We believe that all collections were destroyed in the fire that destroyed the California Academy of Sciences, following the great San Francisco earthquake of 1906. Interestingly, collections from all four of these sites were recollected by F. Grinnell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. californicus  Field studies in Imperial County, California, have led to the discovery of an unusual series of populations of Linanthus maculatus (Parish) Milliken. These populations are atypical in their geographic isolation from previously known populations, being 79.5 air mi (128 km) south of the nearest known population (Patterson 1989;Fig. 17). Moreover, they are uniform in their corolla coloration, lacking the characteristic maroon spotting at the base of each lobe (Fig. 18). This trait has been one of the key diagnostic features for the species and the basis of the specific epithet. Moreover, there are differences in corolla morphology of these southern populations, which include the presence of reverse herkogamy, a consequence of the much shorter style (Fig. 19). We believe that these fixed differences are a manifestation of the genetic isolation and divergence that the southern populations have experienced. Ephemeral-annual herb, taprooted, spreading or erect, 0.9-3.5 cm tall, to 5 cm in diameter; cotyledons sheathing, ovate to narrowly ovate, 0.9-1.5 mm long, 0.4-0.8 mm wide, apex cuspidate. Stems openly branching, beginning at the first or second node, coarsely spreading, villous with uniseriate, multicellular (2-6 cells), eglandular trichomes, the terminal cell acute. Leaves alternate, above the first two to three opposite, more or less distant, nodes fleshy and thickened distally and abaxially, (1.7-)2.5-5 mm long, 0.7-1.8 mm wide, sessile, simple, entire, narrowly oblanceolate or oblong, mucronate, the base sheathing, marginally ciliate villous, trichomes like those of the stems. Inflorescence open and 6diffuse, compoundcymose, composed of 1-3-flowered cymes, terminating the branches, sessile or subsessile, pedicels 0.2-0.7 mm long; floral bracts similar to upper cauline leaves, simple and entire, fleshy and thickened distally and abaxially, 6sheathing the stem, 2.5-4.5 (2) Polemoniaceae: New Names and Combinations  narrowly oblanceolate to spatulate, obtuse but mucronate, 0.8-1.5 mm long, 0.4-0.8 mm wide, the proximal portion of the lobes with herbaceous costae, thick and fleshy distally, sparsely villous abaxially, sparsely and minutely glandular-villous adaxially (particularly proximally), with a broad hyaline margin that is ciliate villous, the calyx-tube very short (0.3-0.8 mm long). Corolla broadly and shortly funnelform or salverform, 3.5-5 mm long, white to cream; the lobes 1-2 mm long, 1.2-2.2 mm wide, cordate to truncate-cuspidate, entire but wavy, recurved during late anthesis, glabrous; tube yellow to white, 2.3-3 mm long, glabrous. Stamens equally inserted 0.9-1.5 mm above the point of corolla insertion, free portions of filaments 0.9-2 mm long, somewhat dilated near the insertion, glabrous; anthers 0.4-1 mm long, 0.2-0.55 mm wide, slightly exserted; pollen yellow. Nectary deeply 5-lobed, the lobes obtusetriangular, 0.1-0.22 mm thick, and 0.2-0.33 mm broad. Ovary triangular-ovoid to globose, 0.5-1 mm long, 0.5-0.8 mm in diameter, glabrous; style (not including stigma lobes) 0.4-0.6 mm long; stigma lobes 0.6-0.75 mm long; 5 ovules per locule. Capsule ellipsoid, 1.5-2.7 mm long, 1-1.7 mm in diameter. Seeds 3-5 per locule, 0.7-1.1 mm long, 0.45-0.6 mm wide, ovoidal-reniform, sometimes angular, constricted at the hilum, seed coat dark red-brown, unchanged when wetted, not producing mucilage. Linanthus maculatus subsp. emaculatus occurs on dune margins, bases of sand ramps, sandy or coarse, opaque-white, decomposed granite soils of washes and on flats near wash margins; 900-1100 m, associated with Colorado Desert riparian vegetation in western Imperial and adjacent San Diego Counties, California. Flowering (March-)April(-May).
Etymology.-The epithet emaculatus (Greek 5 without spots) refers to the absence of the spots at the proximal end of the corolla lobes.

INTRASPECIFIC COMBINATIONS IN LINANTHUS PUNGENS
Similar to the situation observed in Linanthus californicus noted above, infraspecific variation within Linanthus pungens has received little attention in recent years (but see Schultz and Patterson 2012). This is remarkable given that 12 infraspecific taxa have been recognized historically (Hooker 1837;Gray 1870;Parish 1899;Brand 1907;Rydberg 1913;Peck 1936;Jepson 1943;. Variation across the wide range of L. pungens is profound but remains imperfectly characterized. Here we recognize four morpho-geographic taxa at the rank of subspecies. The type of L. pungens is from the plains at the eastern foot of the Rocky Mountains, in Colorado (see below), but the species ranges to the Pacific Northwest and to the mountains of northern Baja California, Mexico (Fig. 21). Two subspecies, as interpreted here, are more restricted in their range and two are quite widespread. Linanthus pungens subsp.  (1986) states that according to Wherry the location is actually the forks of the Platte River, in Lincoln County, Nebraska. However, both of these locations are well outside of the current range of the species. Goodman and Lawson (1995) indicate that James's collection was made in ''late June or early July along the South Platte River, anywhere from Ft Morgan, Morgan County, Colorado to Denver'' (p. 272), a more reasonable location; holotype (as here interpreted) NY. Key to the Subspecies of Linanthus pungens 1. Leaves mostly simple and entire, with only a few, distal, palmate leaves and those with highly reduced lateral lobes (,1/3 the length of the porrect, central lobe); floral bracts trifid; plants with only eglandular, villous trichomes; distributed along the Peninsular Ranges of southern California and Baja California, palmatifid (to pinnatifid) leaves, few if any proximal nodes bearing simple, linear leaves; floral bracts usually 5 or more, lobed; plant vestiture variable, often with both eglandular, villous trichomes and stipitate glands, but glandular trichomes always present; distribution mostly north and east of the Peninsular Ranges . . . . . . . . . . . . . . . . . . . . . . 2. 2. Calyx apparently glabrous abaxially or sparsely glandular-puberulent only at the very base; along the east slope of the Rocky Mountains and adjacent Great Plains, extending through Wyoming to Montana, eastern Idaho, and northeastern Utah .  abaxially, often also with eglandular-villous trichomes; occurring in the intermountain region, Colorado Plateau, extending to western Idaho, Oregon, Washington British Columbia, Canada to California . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. 3. Flowering stems erect and woody; leaves alternate distally, leaflets 1-1.8 mm wide, linear to linear-lanceolate or subulate; the intermountain region, Colorado Plateau, extending to western Idaho, Oregon, Washington, British Columbia, Canada to California, but not the Snake River Valley . . . . . . . . . . . . . . . . . . 4. 39 Flowering stems sprawling to mounded, herbaceous; leaves opposite distally, leaflets 0.2-0.6 mm wide, soft-filiform; NW Great Basin, Snake River Drainage . . . . . . . . . . L. pungens Wilken and Porter 2005). This morpho-geographic race appears to be restricted to limestone and dolomite. Eastern Arizona hosts another phase with small, 6-merous flowers and compact habit. This phase also appears to be edaphically restricted, in this case to basalt. Here we describe these two taxa as new subspecies.
Etymology.-The epithet dolomiticus is derived from the geologic substrate to which this subspecies appears to be restricted, dolomite and limestone. Loeselia glandulosa (Cav.) G. Don is the most broadly distributed species of Loeselia L. It ranges from Santa Cruz County in southern Arizona, through Mexico and Central America, to Colombia and Venezuela. This distribution spans a remarkable array of habitats ranging from temperate oak woodland, to tropical pine-oak forest, cloud forest, and tropical deciduous forest. While a number of infraspecific taxa have been recognized (e.g., Porter and Johnson 2000), one phase has gone unrecognized.
In the northwestern extreme of the species range, Loeselia glandulosa occurs on rocky outcrops in the shade of Quercus hypoleucoides A. Camus, Q. arizonica Sargent and Cercocarpus montanus Raf. of interior chaparral. To the south, it occurs in a diversity of habitats, including cloud forest, lowland tropical scrub, tropical deciduous forest and tropical pine-oak woodlands. While the interior chaparral habitat has much in common with tropical pine-oak woodlands, e.g., Madrean origins, the southern climates are significantly different (J. M. Porter and S. M. Namoff, in prep.). Associated with the different environments are a suite of morphological differences that distinguish this phase from the typical form of L. glandulosa (from the eastern trans-volcanic belt region, in the Querétaro-Hidalgo region of central Mexico). The new taxon (a new subspecies of L. glandulosa, see below) differs from subsp. glandulosa in having linear to linear-lanceolate leaves (subsp. glandulosa has lanceolate to lance-ovate leaves), nectar guides composed of magenta streaking on the lower corolla lobes (subsp. glandulosa has magenta streaking on all lobes and varying degrees of spotting on the lower two), and a short and ramified habit (subsp. glandulosa is relatively tall and less branched).
The difference in leaf morphology is subtle, but significant. Examination of a sample of 31 leaves taken from herbarium collections of Loeselia glandulosa subsp. sonorae (collections with an asterisk cited under Representative specimens below) and subsp. glandulosa, analyzed using elliptical Fourier analysis followed by principal component analysis of Fourier coefficients (momocs [Bonhomme et al. 2014] and ade4 [Thioulouse et al. 1997;Dray and Dufour 2007] libraries under the R environment [R Development Core Team 2008]), shows that the leaves of subspp. sonorae and glandulosa not only differ in shape, but that there are significant differences in the centroids of each (based on 95% confidence ellipses surrounding the centroids; Fig. 24 Suffrutescent perennial to 35 cm tall, usually with more than one primary axis. Stems light to dark brown or reddish, younger stems green; branches and leaves mostly alternate or occasionally with opposite nodes proximally and alternate distally; internodes 0.5-6.0 cm long, shorter in the inflorescence than below. Stems generally glabrate near base, becoming sparsely to densely puberulent to villous above with both antrorse-arcuate, eglandular trichomes, or straight, gland-tipped trichomes. Lower leaves sparsely puberulent or scabrous on upper sides and veins beneath; upper stems, peduncles, leaves and outer bracts glandular villous to glandular-puberulent and often with eglandular trichomes as well. Leaves bright green above, lighter beneath, with slightly raised, white, adaxial veins, lanceolate to linear-lanceolate, apex acute to acuminate, shortly petiolate, the blades attenuate to cuneate proximally, serrate-cuspidate except along the proximal margins where essentially entire; inflorescence leaves shorter and narrower than the primary leaves, grading into the cymule bracts. Inflorescence (synflorescences) diffuse, composed of solitary or paired, bracteate cymules borne terminally and on axillary shoots or in leaf axils of primary stems. Bracts polymorphic, mostly opposite, outermost herbaceous, flattened, linear, serrate with 3-5(-6) attenuate-aristate teeth per margin, occasionally entire, glandular-villous throughout; transitioning to innermost bracts that are entirely hyaline or green and herbaceous along the distal central vein, strongly keeled, coarsely serrate along the distal third but otherwise entire, bearing a few glandular or eglandular trichomes along the midrib. Calyx largely polysymmetric, entirely or largely hyaline, slightly herbaceous and green or anthocyanic on the distal lobes. Corolla monosymmetric (sometimes only slightly so), 16-22 mm long, corolla tube 8.5-10.5 mm long, exserted from the calyx and perceptibly bent, with the distal corolla angled geotropically, lobes 8-11 mm long, 2.3-4.2 mm wide, spatulate or narrowly oblanceolate and clawed proximally, claws somewhat conduplicate and channeled below, sinuses of lobes unequal, the three upper lobes erect and spreading, the two lower lobes spreading forward, lobe apex rounded to truncate and glandular-ciliate, midvein and lateral vascular strands branching or simple, but without anastomoses. Stamens equally inserted just below level of lowest sinus, filaments glabrous, long-exserted, arching upward, emerging together from lowest corolla sinus, becoming curled and reflexed to the corolla orifice following anthesis. Ovary oblongoid, with a small tuft of glandular trichomes at the apex; style straight, long-exserted, subequal corolla; stigmas 0.8-1.0 mm long. Capsule ellipsoid, 3.2-4.0 mm long, dehiscing between valves in upper half at maturity. Seeds 1.5-1.7 mm long, borne 2 (or 3) end to end in each locule, narrowly winged at the ends, flattish, rounded at one end, slanting-truncate at the other where seeds abut.
Occurring along the northern Sierra Madre Occidental, Loeselia glandulosa subsp. sonorae is associated with canyons, rocky slopes and crevices, of limestone, basalt, diorite and rhyolite, in oak woodlands (interior chaparral), and pine-oak forests, from 240 to 2100 m elev.
Although Loeselia glandulosa subsp. sonorae has been collected since the late 1800s, it has been treated as the typical VOLUME 32 (2) Polemoniaceae: New Names and Combinations VOLUME 32 (2) Polemoniaceae: New Names and Combinations 85