Flora of North America species comparison

Varieties 3 (3 in the flora).

Cirsium remotifolium occurs from the Coast Ranges and valleys of the Pacific Northwest to the western slopes of the Cascade and Klamath ranges, south in the California North Coast Ranges to the San Francisco Bay region. It is closely related to the C. clavatum complex of the Rocky Mountains region. Both have a similar growth habit and some forms variably express the character of broadly scarious, lacerate-toothed phyllary margins. Gray, in naming Cnicus carlinoides var. americanus, included as syntypes both California and Colorado specimens. F. Petrak (1917) treated both the West Coast plants and those of the Rocky Mountains as Cirsium subsect. Americana, recognizing C. remotifolium with several infraspecific taxa plus two other species, C. callilepis and C. amblylepis from the West Coast, and four additional species from the Rocky Mountains. A. Cronquist (1955) rejected Petrak’s subspecies, treating C. remotifolium in a restricted sense, limited to plants of Washington and Oregon without dilated phyllary tips, and circumscribed C. centaureae broadly to include the Rocky Mountains and West Coast plants with dilated phyllary tips. Because of the frequent presence of dilated phyllary tips in C. remotifolium in the restricted sense, Cronquist acknowledged the likelihood of past introgression with C. centaureae in the broad sense.

J. T. Howell (1960b) recognized three species: Cirsium remotifolium, C. acanthodontum, and C. callilepis, the latter with four varieties collectively corresponding to the West Coast representatives of C. centaureae (in the sense of Cronquist). Because of the great similarity of the various West Coast plants and their intergradation, I see no value in recognizing two or more species.

The West Coast and Rocky Mountains plants are clearly related, but are separated by the Great Basin region and there is little chance of current genetic interchange. As is often the case with American Cirsium, genetic enrichment from past hybridization with other nearby species within their respective areas has likely been fertile ground for evolutionary diversification. Different species have contributed genes in the Pacific states and in the Rockies. I have chosen to recognize two geographically-based species complexes, each with intergrading races here treated as varieties. I treat the West Coast plants as C. remotifolium and the Rocky Mountains plants as C. clavatum.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Species ca. 200 (62 in the flora).

Only three genera in Cynareae are represented by native species in the New World, and of these Cirsium is by far the most widely distributed and diverse. Native species of Cirsium range from sea level to alpine and from boreal regions of Canada to the tropics of Central America. Members of the genus occur in a myriad of habitats including swamps, meadows, forests, prairies, sand dunes, and deserts.

Preliminary molecular phylogenetic studies by D. G. Kelch and B. G. Baldwin (2003) indicated that this diversity is the product of a rapid evolutionary diversification based upon a single initial introduction from Eurasia. Relationships among the North American species are apparently complex, and molecular studies have only begun to provide an outline of phylogeny for these plants. Although there has been a remarkable evolutionary and morphologic diversification in North American Cirsium, it has not been accompanied by very much divergence in the base sequences of genes commonly used to elucidate phylogenetic relationships. This suggests either that the diversification has been very rapid or that genetic markers in North American Cirsium mutate more slowly than in most other lineages.

Chromosomal diversification has accompanied the morphologic radiation of North American Cirsium. Many New World Cirsium species share the chromosomal base number of x = 17 that also predominates in most Eurasian species. Among the North American thistles, however, is a mostly descending dysploid series with chromosome numbers ranging from n = 18 to n = 10. Very few instances of polyploidy are known among New World Cirsium.

Cirsium species of remarkably different morphologies often are able to hybridize. Although in some hybrid combinations fertility is reduced, in others the formation of complex hybrid swarms indicates a lack of breeding barriers and the potential for emergence of novel character combinations. In the absence of adequate sampling and field observations, hybrids may go unrecognized—treated as distinct taxa or as variants of non-hybrid taxa, or left occupying the indeterminate folders of herbaria. In other cases hybridization has been invoked without much evidence as an explanation for Cirsium variants encountered in herbaria or in the field. Hybrid combinations are listed herein when evidence is convincing. Additional hybrids are likely to be found where the ranges of Cirsium species overlap. I have seen no documentation of hybridization between native American Cirsium species and introduced Eurasian taxa.

Much of the geographic range currently occupied by New World Cirsium species was greatly affected by the events of the Quaternary. Large areas were glaciated and other areas were vastly different during glacial episodes. The ancestors of thistles that currently occupy the high mountains of western North America were undoubtedly displaced elevationally and/or latitudinally during the recurrent glacial and interglacial episodes of the Pleistocene. Taxa that are currently isolated may have been in contact during glacial episodes with the opportunity for hybridization and genetic interchange. Episodes of prehistoric hybridization may have led to some of the character combinations found in modern American thistles, particularly in the western half of the continent. Current isolation and localized selection or genetic drift apparently have promoted differentiation of populations separated on mountaintop islands.

One of the most challenging aspects for a taxonomist studying New World Cirsium is the presence of species complexes that are apparently evolutionary works in progress. Some of the thistles, especially in the mountainous western part of North America, are frustratingly polymorphic with much overlapping variability and intergradation of characters. Early taxonomists, basing their work on a limited sampling of the morphologic diversity, named many of the forms as species, and the literature is rife with species names. The infilling that results from more collectors visiting more localities within the ranges of these complexes has blurred the boundaries between many of the proposed species and often added forms that do not “fit” the characteristics of named species. As I faced the challenges of preparing this treatment, I recognized that maintaining some of the named entities as species would, for consistency, require a further proliferation of species names. I have chosen to go the other way. Instead of proposing yet more ill-defined microspecies, I have chosen to recognize that the groups in question are rapidly evolving, only partially differentiated assemblages of races that have not reached the level of stability that is usually associated with the concept of species. Certainly much of such variation within the genus deserves a level of taxonomic recognition, or at least should be mentioned, but for those assemblages I think it much more prudent to recognize varieties—entities that may be expected to freely intergrade—rather than species.

Many problems remain to be worked out in North American Cirsium. Further investigation will undoubtedly reveal the need for refinement or major revision within some of the species groups. Studies that focus on variation within and among populations and on the biological basis for the variations are much needed. The field is open and the challenges are many.

Preparation of a workable key to Cirsium species has been frustratingly difficult. Extensive and overlapping ranges of variation in morphologic characteristics often require that a species be keyed two or more times. The resulting key is longer and more complex than I would prefer, and I have no doubt ignored, overlooked, or been completely unaware of variants that will not key out. Caveat clavitor!

The reputation of Cirsium has suffered greatly as a result of the introduction to North America of a few invasive weedy species from Eurasia. Cirsium vulgare (bull thistle) and C. arvense (Canada thistle—a misnomer) have long been despised as noxious weeds. In recent years C. palustre (European swamp thistle) has joined their ranks. Additionally, weedy Eurasian species of Carduus, Onopordum, Centaurea, etc., add to the public perception that all thistles are bad. Most North American native Cirsium are not at all weedy, and many are strikingly attractive plants. All are spiny plants that command respect, but they deserve a better reputation as one of North America’s evolutionary success stories.

Native Cirsium species have come under threat from biocontrol programs instituted to suppress populations of weedy introduced thistles. Beginning in 1968 the seedhead weevil Rhinocyllus conicus has been widely introduced in various areas of the United States and Canada, primarily to control weedy species of Carduus. S. M. Louda et al. (1997) reported that R. conicus has crossed over to several native species of Cirsium. They observed that the number of viable cypselae in infested heads was greatly reduced; e.g., heads of C. canescens infested by R. conicus produced 14.1 percent of the number of viable cypselae as in uninfested heads. Not all taxa are impacted as much as C. canescens, particularly those with later flowering phenology (Louda 1998). R. W. Pemberton (2000) reported that 22 Cirsium taxa in North America are known hosts of R. conicus. I suspect that the number is higher. During my field work I have observed that the heads of many Cirsium species are heavily parasitized, although I have not determined which of these are infested by R. conicus and which by native seedhead parasites. The long-term impacts of R. conicus and other biocontrol agents on native thistles, particularly rare taxa, remain to be determined.

Excluded species:

Cirsium canum (Linnaeus) Allioni was reported in 1924 from Massachusetts by C. H. Knowlton and W. Deane (Rhodora 26: 84) based on an 1899 collection from Weston. It has not been included in subsequent local or regional floras and apparently is not established in the flora area.

Cirsium scabrum (Poiret) Bonnet & Barratte was reported [as Cnicus giganteus (Desfontaines) Willdenow] in 1900 by A. Eastwood (Zoë 5: 59) based on a 1900 collection from Santa Cruz County, California. J. T. Howell (1959, 1960b) noted that Cirsium scabrum had not become naturalized. It apparently is not established in the flora area.

(Discussion copyrighted by Flora of North America; reprinted with permission.)