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J. Mark Egger 

Herbarium, Burke Museum of Natural History and Culture 

University of Washington 

Seattle, Washington 98195-5325 


Lectotypification of the illegitimate name Castilleja brevifllora A. Gray 1 862 (non Benth. 
1846) is proposed in order to maintain Castilleja puberula Rydb. and to resolve the heterotypy of C. 
breviflora A. Gray. Castilleja breviflora A. Gray is then formally placed in synonymy under 
Castilleja flava S. Wats. 

KEY WORDS: Orobanchaceae, Castillejinae, Castilleja, Castilleja brachyantha Rydb., Castilleja 
breviflora A Gray, Castilleja puberula Rydb., Castilleja flava S. Wats., lectotypification, 
nomenclature, heterotype 

When A Gray first published Castilleja breviflora A. Gray (in 1862), he was unaware that G. 
Bentham (in 1846) applied this name in Castilleja to a species of Andean South America. Rydberg 
later realized this and in 1900 published the name Castilleja brachyantha Rydb. to replace Gray's 
name. Still later, in 1905, Rydberg described the morphologically similar Castilleja puberula Rydb., 
clearly implying that he regarded the two as different species. Eventually, C brachyantha was 
reduced to synonymy under Castilleja flava S, Wats, by Harrington (1954), while C puberula 
continues to be recognized as a bona fide species (e.g., Nelson & Harmon 1997; Seagrist & Taylor 
1998; Beidleman et al 2000; Hartman & Nelson 2001; Weber & Wittmarm 2012). Castilleja flava is 
a widespread species of moderately xeric habitats at middle elevations throughout much of the 
Intermountain West and is often associated with sagebrush communities. Castilleja puberula is an 
uncommon to rare alpine species endemic to four counties in the central Rocky Mountains of 
Colorado and likely evolved as a high elevation isolate of C. flava. 

When Gray established his Castilleja breviflora, he cited two elements, PI. Parry, No. 243 
(1861, headwaters of Clear Creek and the alpine ridges lying east of Middle Park, Colorado Territory, 
duplicates at DUKE!, GH!, IA-ISC!, NY!, P!) and an unpublished T. Nuttall name, "Euchroma 
breviflora, Nun. in herb. Philad." Based on the known range of the plant in question, Nuttall gathered 
his specimens in Idaho or Wyoming in 1834. Sheets bearing this name in Nuttall's hand (BM!, PH!) 
are clearly referable to C flava, while sheets of Parry 243 all bear plants identical to what is now r 
recognized as C puberula. This case of heterotypy complicates the modern usage of the name C. 
puberula, necessitating the lectotypification proposed below. 

Based on the fact that only Parry 243 is located at GH and is annotated by Gray as his 
Castilleja breviflora, it is likely the meager description in the protologue was based primarily on the 
Parry collection (Fig. 1). While most of the description could apply to either C flava or C puberula, 
Gray does mention that the plants are "spithamaea" or only one "hand-span" in height, the typical 
height of most flowering C. puberula plants (Fig. 2) but shorter than most C flava plants of similar 
development (Fig. 3), However, we know that Gray studied at least the BM specimen of Nuttall s.n, 
because it bears an annotation in his hand, 'Castilleja. Euchroma breviflora, Nutt, ined." (Fig. 4). 
That sheet contains one large, typical stem of C. flava but also several, much shorter, poorly 

Egger: Lectotypification of Castilleja breviflora 2 

developed or trimmed stems that may have influenced Gray's description as well. Also, according to 
Penneli (1936) it is likely Gray saw the PH specimen of Nuttatt s.n., though it lacks his annotation 
(Fig. 5). 

Until now Gray's Castilleja breviflora has not been lectotypified. Apparently, Harrington 
(1954) was the first to assign C. brachyantha to synonymy under C. flava, even though that species 
only partially corresponds to Gray's description of C. breviflora. Most authors working with the 
Rocky Mountain flora later adopted Harrington's synonymy. The isolectotype designated below was 
also annotated as C. flava by Castilleja authority F.W. Penneli in 1920. 

It should be noted that Rydberg's application of Castilleja brachyantha is somewhat different 
from that intended by Gray when he proposed his C. breviflora. Aside from linking his new name C. 
brachyantha with Gray's illegitimate C. breviflora. Rydberg (1900) cited only a single collection (P. 
Koch 10, Hoodoo Peak, Yellowstone National Park, Wyoming). This collection does not appear to be 
present at RM at this time, and its true identity has yet to be determined. However, I know of no 
verified specimens of C, puberula from either Montana or Wyoming, and the region in which Koch 
10 was collected is well within the known range of C flava (Holmgren 1984). In Rydberg's works 
published prior to the description of C. puberula, C. brachyantha was distinguished from C. flava by 
minor variations in the lower corolla lip. Following the publication of C puberula, Rydberg ( 1906, 
1917) distinguished C brachyantha from both C. puberula and C. flava based on stature and 
differences in the structure of the beak and lower lip of the corolla, but his application of the name C. 
brachyantha appears to most closely match plants now treated as C flava. 

Typification of Gray's Castillja breviflora with Nuttall's collection will assure that C. 
puberula, a name in current use, will remain available for this rare Rocky Mountain plant as C. 
puberula was described after both Gray's name and Rydberg's replacement name. This is not a 
concern with C flava, which was described in 1871, well before Rydberg's C brachyantha. 

CASTILLEJA FLAVA S. Wats, in King, Rep. Geol. Explor. 40th Parallel [Botany] 5: 230. 1871. 
TYPE: Utah. [Rich Co.:] Upper Bear River Valley, 7000 ft, Jul 1869, S. Watson 813 
(holotype: GH!, isotypes: NY!. US!, YU!). 

Castilleja breviflora A. Gray, Amer. J. Sci. 34: 338. 1862. Castilleja brachyantha Rydb., Mem. 
N.Y. Bot. Gard. 1: 360. 1900 (replacement name, not Castilleja breviflora Benth. in DC, 
Prodr. 10:534. 1846). LECTOTYPE (designated here): USA. [Idaho or Wyoming]. Rocky 
Mts., 1834, T. Nuttalls.n. (BM!, isolectotype: PH!). 

CASTILLEJA PUBERULA Rydb., Bull. Torrey Bot. Club 3 1:644. 1905. TYPE: Colorado. "Colorado 

Territory," 1872, C.C. Parry s.n. (holotype: NY!, isotypes: GH!, P!, PH!), 


I thank Kanchi Gandhi, John McNeill, James L. Reveal, and John H. Wiersema for 
consultation and advice concerning the disposition of these nomenclatural matters. I also thank 
Reveal in particular for assistance in earlier drafts of this paper, for his extensive comments in review 
of the present paper, and for his always timely and gracious assistance on matters botanical. Guy 
Nesom also provided very useful comments on the manuscript. I also thank the personnel of the 
herbaria cited in this paper for assistance with loans and BM, GH, and PH for permission to use 
images from their type collections. 

Egger: Lectotypification of Castilleja breviflora 3 

Figure 1. Mixed sheet at GH containing several collection of Castilleja puberula, including both an isotype of 
C. puberula Rydb. and a syntype of C. breviflora A. Gray. Note that both my annotation from 1996 and some 
of those by others are not fully accurate and that only the stems in the lower left obtained by Parry in 1861 
actually represent Parry 243. 

: Lectotypification of Castilleja breviflora \ 

Egger: Lectotypification of Castilleja breviflora 5 

r.flava, Angel Lake Road, East Humboldt Range, Elko Co., NV, 27 Jim 1995, M 

Egger: Lectotypification of Castilleja b 

Egger: Lectotypification of Castilleja breviflora J 

Egger: Lectotypification of Castilleja breviflora 8 


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