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Cytogeography of the Solidago rugosa Mill. Complex (Asteraceae: Astereae) in Eastern North America

1
Department of Biology, University of Waterloo, Waterloo, ON N2L 3G1, Canada
2
C & C International Inc., Kitchener, ON N2P 0A8, Canada
3
Department of Natural Sciences, Fairmont State University, Fairmont, WV 26554, USA
4
Faculty of Biology, Universitas Gadjah Mada, Bulaksumur, Yogyakarta 55281, Indonesia
*
Author to whom correspondence should be addressed.
Academic Editor: Daniel Potter
Taxonomy 2021, 1(4), 290-301; https://doi.org/10.3390/taxonomy1040023
Received: 24 September 2021 / Revised: 8 October 2021 / Accepted: 11 October 2021 / Published: 14 October 2021

Abstract

Chromosome numbers are reported here for the first time from 117 individuals of Solidago rugosa and S. fistulosa. Including 178 previously published reports for the two species plus S. latissimifolia, chromosome numbers have been determined from 295 individuals from 269 locations. Only diploids (2n = 18) were found throughout the range of S. fistulosa on the coastal plain in the eastern U.S.A. (44 counts). Diploids (2n = 18) were found in the northern portion of the range of S. latissimifolia, and tetraploids (2n = 36) and hexaploids (2n = 54) were found in the central and southern portions of the range (nine counts in total). Diploids (2n = 18) were found throughout the range of S. rugosa in much of eastern North America in four of the five varieties (northern var. rugosa, var. sphagnophila; southern var. aspera and var. celtidifolia). Tetraploids (2n = 36) were found in all four of these varieties and exclusively in var. cronquistiana in the southern high Appalachian Mountains. Hexaploids (2n = 54) were found in var. sphagnophila at scattered locations. One possible hexaploid in var. rugosa was found in the Allegheny Mountains. The diversity in ploidy levels was independent of the size of the range and the diversity of growing conditions among the three species of S. subsect. Venosae.
Keywords: cytotaxonomy; biogeography; goldenrods cytotaxonomy; biogeography; goldenrods

1. Introduction

The Solidago rugosa Mill. complex in the broad sense occurs in eastern North America and is a taxonomically difficult group of goldenrods making up Solidago subg. Pleiactila Raf. sect. Venosae (D. Don in Loudon) Nesom subsect. Venosae (G. Don in Loudon) Nesom (Semple and Beck 2021 [1]). The subsection includes three closely related species, S. fistulosa Mill., S. latissimifolia Mill., and S. rugosa, with four varieties recognized here and multiple basionyms: Solidago aestivalis Bicknell, S. aspera Ait., S. celtidifolia Small, S. edisoniana Mack., S. elliottii Torr. & A. Gray, Solidago elliottii Torr. & A. Gray var. ascendens Fern., Solidago elliottii Torr. & A. Gray var. pedicellata Fern., Solidago elliptica Ait., S. elongata Pépin, S. fistulosa Mill., S. latissimifolia Mill., Solidago mirabilis Small, S. rugosa Mill., S. rugosa var. cronquistiana Semple, S. rugosa var. glabrata Farwell, S. rugosa var. sphagnophila Graves, and S. villosa Pursh. The Solidago rugosa complex includes three to six species depending upon the classification scheme followed (Table 1). Fernald (1950 [2]) presented a treatment of members of the complex except for the much more recently described var. cronquistiana. He recognized three varieties in S. elliottii, no varieties in S. fistulosa, and five varieties in S. rugosa: var. rugosa, var. villosa (Pursh) Fern., var. aspera (Ait.) Fern., var. celtidifolia (Small) Fern., and var. sphagnophila. Earlier, Fernald (1936 [3]) dealt with the misapplication of the name S. altissima L. to members of the S. rugosa complex and acknowledged that var. sphagnophila might deserve species status as S. aestivalis. Cronquist (1947 [4], 1968 [5]) recognized three species in the complex and grouped the varieties of S. rugosa into two subspecies: a southern subsp. aspera and a northern subsp. rugosa. Cronquist (1980 [6]) referred to subsp. villosa, but this was likely a typographical error, since the taxon is clearly his subsp. rugosa of both Cronquist (1968 [5]) and Gleason and Cronquist (1991 [7]). Beaudry (1960 [8]) recognized four species based on limited cytological data that indicated normally a single ploidy level for each: S. aestivalis, S. aspera, S. celtidifolia, and S. rugosa (which included f. villosa (Pursh) Beaudry); S. fistulosa was not included in his study. Uttal and Porter (1988 [9]) determined that the older name S. latissimifolia was the same species as S. elliottii and should be used instead. This paper follows the classification presented in Semple and Cook (2006 [10]) and accepts the name S. latissimifolia and is most similar to Cronquist (1968 [5]) but includes var. celtidifolia, var. cronquistiana and does not recognize subspecies within S. rugosa.
Before the first author’s laboratory began working on the cytology of Solidago (Semple et al. 1981 [11]), 52 chromosome counts from 27 locations had been reported for all members of the S. rugosa complex (Table A1). The first report was by Goodwin (1937 [12]) for S. rugosa var. rugosa. Most of the early counts were published by Jean Beaudry of the Université de Montréal (Beaudry and Chabot 1959 [13]; Beaudry 1960 [8], 1963 [14], 1969 [15]). Beaudry’s contributions to our understanding of the genus are significant, but he adopted a “splitters” approach to recognizing species, which he apparently thought at the time should include little or no ploidy level variation. His very small sample sizes allowed him to maintain this position.
An investigation of the cytogeography of the complex was undertaken by the Semple Astereae Lab in the 1980s and early 1990s in hopes of clarifying what role ploidy level might play in taxon delimitation. The field work to gather more data continued until 2012. Herbarium specimens, including nomenclatural types, were examined in order to more clearly understand how each of the above cited authors had applied names to their cytovouchers collected as part of the cytogeographic investigation. The overall objective was a better understanding of taxon limits within the complex, an assessment of the appropriate taxonomic ranks for these taxa, and a large cytological sampling of the entire ranges of all taxa in order to determine cytogeographic patterns in the taxa recognized.

2. Materials and Methods

Meiotic counts were made from pollen mother cells dissected from buds fixed in the field in Carnoy’s Fixative 3:1 (absolute EtOH: glacial acetic acid) and subsequently stored under refrigeration in 70% EtOH until examined. Anthers containing pollen mother cells were dissected out of florets and squashed in 1% acetic orcein stain. Counts were made from freshly prepared slides using either a Zeiss RA Standard Microscope or a Nikon Microscope with phase contrast optics. Mitotic counts were made from root tip cells taken from transplanted wild rootstocks or from seedlings grown from fruits collected in the field. Root tips were pretreated in 0.01% colchicine or saturated paradichlorobenzene for 2–3 h, fixed in Acetic Alcohol Fixative (3:1/EtOH: glacial acetic acid), and hydrolyzed in 1 N HCl for 30 min at 60 °C before squashing. Counts were made from freshly prepared slides as above. Vouchers for all counts are deposited in WAT in MT (Thiers continuously updated [16]) with duplicates distributed to various other herbaria. Identifications were made by J.C.S.

3. Results

Cytovouchers for nearly all 178 previously published counts of Solidago fistulosa, S. latissimifolia, and S. rugosa were examined and identifications confirmed or corrected; these are listed in Appendix A Table A1, which includes a brief indication of geography, publication data, and indication of the name under which the counts were first published. Chromosome numbers were determined from an additional 117 individuals of the Solidago rugosa complex and are reported here for the first time in Appendix A Table A2. The distribution of all 41 diploid chromosome counts for S. fistulosa is shown in Figure 1. The distribution of the 9 diploid, tetraploid, and hexaploid chromosome counts for S. latissimifolia is shown in Figure 2. The distribution of all 242 diploid, tetraploid, and possible hexaploid counts for S. rugosa is shown in Figure 3 with ranges of each variety indicated.
A range of inflorescence branching patterns in S. rugosa were observed in the field. In one growth form, the inflorescence has very long arching lower branches. Type specimens and the majority of field collections of S. rugosa var. aspera and var. rugosa had large inflorescences with many long diverging branches with mostly secund branchlets with heads. Both varieties include some specimens with inflorescences with very short branches barely to slightly exceeding the subtending leaves. This is the supposedly diagnostic feature of var. villosa, but garden and greenhouse observations on transplanted individuals indicate that this trait is too unstable to be used as a diagnostic feature. It is common to find shoots of the same clone with the two extreme forms of inflorescence branching patterns. Plants with short-branched inflorescences are scattered through the range of var. rugosa. Thus, we treat var. villosa as a synonym of var. rugosa.
The type specimens of S. rugosa var. celtidifolia have ovate upper stem leaves and inflorescences that have very short lateral branches. Throughout the range of the variety, plants often have large inflorescences with long lower branches like inflorescences in var. rugosa and var. aspera. Intermediate plants between obvious var. aspera and obvious var. celtidifolia plants were also encountered in the field.
During field work, a number of collections were found that had leaves typical of var. aspera, but an inflorescence form that was like the “var. villosa” inflorescence with short lateral branches. The indument was sparse for either of these morphs. Plants with these characteristics did not fit well into any of the five varieties recognized by Fernald and Cronquist (Table 1). All these populations were found in the Carolinas and northern Georgia at the southern end of the Appalachian Mountains. Additional herbarium sheet collections were seen subsequently. All collections counted were tetraploid 2n = 36. These were placed in var. cronquistiana, named in honor of Art Cronquist.

4. Discussion

Including previously published reports (Goodwin 1937 [12]; Beaudry and Chabot 1959 [15]; Beaudry 1960 [8]; Beaudry 1963 [14]; Jones 1968 [17]; Beaudry 1969 [14]; Kapoor 1970 [18]; Anderson et al. 1974 [19]; Kapoor 1977 [20]; Semple et al. 1981 [11]; Morton 1981 [21]; Löve and Löve 1982 [22]; Semple et al. 1984 [23]; Semple 1985 [24]; Semple and Chmielewski 1987 [25]; Semple et al. 1989 [26]; Semple et al. 1992 [27]; Semple et al. 1993 [28]; Semple and Cook 2004 [29]; Semple et al. 2015 [30]; Morton et al. 2018 [31]; and Semple et al. 2019 [32]), 295 individuals have had chromosome numbers determined from 270 locations in total for S. fistulosa, S. latissimifolia, and S. rugosa (Figure 1, Figure 2 and Figure 3). Morphologically, S. fistulosa is the most distinct of the three species and has the smallest range of variation in leaf shape, leaf venation, and inflorescence branching pattern. Solidago fistulosa is strictly diploid with 45 counts from 33 locations throughout its range on the inner and outer coastal plains from New Jersey south to southern Florida and west along the Gulf Coast to eastern Louisiana (Figure 1). Habitats include “mostly wetter sandy soils, seepage areas, boggy grounds, edges of marshes and thickets, open pine woodlands, roadside ditches” at 0–100 m elevation (Semple and Cook 2006 [10]). Solidago latissimifolia includes diploids, tetraploids, and hexaploids encountered in the small sample of nine counts from nine locations from southern Nova Scotia to South Carolina; no counts have been reported from disjunct populations in peninsular Florida (Figure 2). Solidago latissimifolia has the smallest range of the three species and occurs in ”fresh and brackish swamps, thickets on the outer coastal plain” at 0–80 m (Semple and Cook 2006 [10]). Morphologically, S. latissimifolia might be confused with some individuals of S. rugosa var. sphagnophila due to convergence in leaf size, but the former has leaves that are glabrous and usually larger and less or not rugose (Semple and Cook 2006 [10]). Solidago rugosa includes diploids, tetraploids, and possibly hexaploids with counts for 242 individuals from 228 locations from much of eastern North America from Newfoundland west to Wisconsin and south to northern Florida and eastern Texas (Figure 3).
The four varieties of S. rugosa recognized here occur in overlapping ranges and include many individuals that can be difficult to assign definitively to a single variety. Solidago rugosa var. sphagnophila is the most morphologically distinct with its absence of hairs on at least the lower stems and often glabrous or glabrate leaves; upper stems can be glabrous/glabrate to moderately hairy in the inflorescence. Only diploids were found and reported here, while only hexaploids were reported by Beaudry (1960 [8], 1969 [15]); tetraploids were not found in the sample of 13 individuals from eight locations in outer coastal plain cedar swamps and bogs from Massachusetts to New Jersey, and no chromosome counts from disjunct populations in Virginia and North Carolina have been reported (Figure 3).
Although Cronquist (1947 [4], 1968 [5], 1980 [6]) grouped the more northern varieties into subsp. rugosa and the southern varieties into subsp. aspera, var. rugosa and var. aspera are often difficult to distinguish, with many specimens being only somewhat to moderately rugose, that is, the intervening leaf surface is elevated slightly above the plain of the supporting venation which itself is prominent on the undersurface. The difference may correlate with growing conditions more than with geographic location, with more plants in southern populations growing in seasonally drier habitats during longer growing seasons than the more northern populations. In fact, only some of the plants assigned here to var. aspera were considered to have very pronouncedly rugose leaves and these came from the drier open woods and full sun habitats encountered in the southeastern states of the U.S.A. Plants assigned to var. aspera that grew in shaded thickets with wetter soils sometimes had less rugose leaves and were close in appearance to more northern plants of var. rugosa. Beck et al. (2021 [33]) included 15 samples of S. rugosa in their phylogenomic investigation of S. ulmifolia, but did not assign the S. rugosa samples to varieties. Seven of the collections were made by J.C. Semple and were cytovouchers for diploids cited in this paper; eight were for collections by other researchers and were not cytovouchers but varieties could be assigned to five based on online images. All specimens of S. rugosa were placed in a single branch of the tree with 74% support. The single collection of var. sphagnophila from Massachusetts along with a single diploid collection of var. rugosa from New York were placed in the clade sister of the remainder of S. rugosa. The most derived branch of the S. rugosa samples included three diploid collections of var. celtidifolia from Louisiana, although two of these were biological duplicates. The remainder of the samples included a sample of var. rugosa from Ontario, samples of var. aspera from Arkansas and Mississippi, a diploid sample of var. celtidifolia from Mississippi, and samples of either var. rugosa or var. aspera from Arkansas, Missouri, and Louisiana. Thus, there was phylogenetic limited distinction among samples of vars. aspera, rugosa, sphagnophila and celtidifolia nor complete separation of northern and southern samples. Clearly, a much larger sampling of specimens of all varieties of S. rugosa from across the entire range of each variety must be undertaken before strong conclusions can be drawn about the phylogenetic separation of varieties in S. rugosa. Certainly, the limited data available do not support the recognition of northern and southern subspecies but do hint at some separation of varieties and raise the question of whether or not var. aspera really is distinct from var. rugosa.
There were some patterns to the distribution of diploids and tetraploids in S. rugosa with differences in the patterns between varieties. All 55 samples of var. rugosa from Canada were diploid. Tetraploid samples of var. rugosa were scattered among diploid samples in New England and further south with the two ploidy levels occurring in approximately equal numbers of samples in the southern portion of the range of var. rugosa. For var. rugosa, 90% of the 106 samples were diploid. No unquestioned hexaploid samples were found in var. rugosa; one possible sample reported here (Table A2) is shown in Figure 3 with a question mark because of uncertainty that the voucher and the plant counted came from the same wild plant. A second hexaploid count by Löve and Löve (1982 [22]) was reported under the name S. altissima L., but the CAN voucher of the collection number is a specimen of S. rugosa var. rugosa. Therefore, the count cannot be assigned to either species with certainty as the voucher in CAN may not be the source plant of the count. In contrast, 91% of the 66 samples of var. aspera were tetraploid with diploids scattered in the central and southern portions of the range of var. aspera from eastern Kentucky to Mississippi. In var. celtidifolia, 51% of the 37 samples were tetraploids with diploids and tetraploids occurring throughout most of the range of var. celtidifolia. Only tetraploids were found in the southern high Appalachian Mountains in var. cronquistiana. In var. sphagnophila, diploids were sampled at five locations in the middle portion of the range and hexaploids were sampled in four locations over the range, but no tetraploids were sampled, perhaps due to limited sampling of populations.
The diversity in ploidy levels in the three species of S. subsect. Venosae was independent of the size of the range and the diversity of growing conditions. Solidago latissimifolia had the narrowest and smallest range, which included diploids, tetraploids, and hexaploids, even though the sample size was small and included only acidic soils mostly in Eastern White Cedar swamps along the Atlantic coast. Solidago fistuosa included only diploids throughout its less narrow range from the sandy Atlantic Coastal plain of southern New Jersey to southern Florida and west to eastern Louisiana on the coastal plain in a mixture of wetter soil habitats. Solidago rugosa included diploids, tetraploids, and possibly hexaploids in a diversity of habitats across much of eastern North America with the ranges of the varieties being smaller and overlapping with at least one other variety. While the strictly tetraploid S. rugosa var. cronquistiana came from only higher elevation habitats in the southern Appalachian Mountains, these were not very different from the higher elevation habitats of some tetraploid samples of var. aspera. While each taxon in subsect. Venosae has a different range of distribution, the ranges of all taxa overlap with one or more other taxa. Ploidy level is thus neither correlated with a specific habitat nor associated with any particular taxon in subsect. Venosae.

Author Contributions

Conceptualization, J.C.S.; analyses of collections, J.C.S.; writing—original draft preparation, J.C.S.; writing—review and editing, J.C.S. and R.E.C.; field collecting and chromosome counting, J.C.S., J.Z., R.E.C. and B.A.S.; herbarium specimen examination, J.C.S. and B.A.S. All authors have read and agreed to the published version of the manuscript.

Funding

This work was supported by Natural Sciences and Engineering Research Council of Canada Operating and Discovery Grants to J.C.S., an Ontario Graduate Scholarship in Science and Technology and a Natural Sciences and Engineering Research Council of Canada Graduate Scholarship to R.E.C., and an Indonesian Faculty Improvement Grant to B.A.S.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

Not applicable.

Acknowledgments

The following people are thanked for their assistance in the field: R.A. Brammall, L. Brouillet, J.G. Chmielewski, D. and R. Cook, S. Heard, R. Keir, G. Ringius, I. Seiden, B. Semple, and K. Tereszchuk. Joan Venn is thanked for assistance in the WAT Herbarium.

Conflicts of Interest

The authors declare no conflict of interest.

Appendix A

Table A1. Previously published chromosome number determinations in Solidago rugosa complex from Canada and the United States, arranged alphabetically by species and variety. B = Beaudry; C = R.E. Cook; Ch = Chmielewski; T = K. Tereszchuk; M & V = J.K. Morton and J. Venn; S = J. Semple; S & S = J. and B. Semple; Su = Bambang Agus Suripto. Herbarium abbreviations from Thiers continuously updated [13].
Table A1. Previously published chromosome number determinations in Solidago rugosa complex from Canada and the United States, arranged alphabetically by species and variety. B = Beaudry; C = R.E. Cook; Ch = Chmielewski; T = K. Tereszchuk; M & V = J.K. Morton and J. Venn; S = J. Semple; S & S = J. and B. Semple; Su = Bambang Agus Suripto. Herbarium abbreviations from Thiers continuously updated [13].
Solidago fistulosa Mill.—2n = 9II U.S.A. Florida: Hillsborough Co., S & Wunderlin 2527 (WAT; Semple et al. 1981 [11]); Osceola Co., S et al., 5356 (WAT; Semple 1985 [24]; count cannot be definitively assigned to species because voucher specimens include both S. fistulosa and S. leavenworthii); St. Johns Co., G. Morton 4553 NY; Semple et al. 1984 [23]. North Carolina: Brunswick Co., G. Morton GPNC21 (not seen; Semple et al. 1984 [23]); Martin Co., S & Su 9739 (WAT; Semple et al. 1993 [28]).—2n = 9II + 1-3 supernumeraries. U.S.A. Florida: St. Johns Co., G. Morton 4550 (not seen; Semple et al. 1984 [23]), G. Morton 4551 (NY; Semple et al. 1984 [23]), G. Morton 4552 (not seen; Semple et al. 1984 [23]).—2n = 18U.S.A. Alabama: Mobile Co., S & Su 10128 (WAT; Semple et al. 1993 [28]). Delaware: Sussex Co., S & Ringius 7649 (WAT; Semple 1985 [23]). Florida: Calhoun Co., M & V NA16484 (TRT; Morton et al. 2018 [30]); Lafayette Co., S et al., 7447 (WAT; Semple et al. 1984 [23]); Okeechobee Co., M & V NA16531 (TRT; Morton et al. 2018 [30]); St. Johns Co., S et al., 7542 (WAT; Semple et al. 1984 [23]); St. Lucie Co., S 11734 (WAT; Semple et al. 2019 [31]); Washington Co., S 10952 (WAT; Semple and Cook 2004 [28]). Georgia: Coffee Co., S et al., 7416 (WAT; Semple et al. 1984 [23]); Echols Co., S 10879 (WAT; Semple and Cook 2004 [28]); Wilcox Co., S & B. Semple 11211 (WAT; Semple et al. 2015 [29]). Mississippi: Harrison Co., S & Su 10125 (WAT; Semple et al. 1993 [28]). New Jersey: Atlantic Co., S & Su 9506 (WAT; Semple et al. 1993 [28]); Cape May Co., B 57-221, 57-222, 57-223 (MT; Beaudry 1963 [14]); Ocean Co., B 57-222 (MT; Beaudry 1963 [14]), S 11812 (WAT; Semple et al. 2019 [31]). North Carolina: Carteret Co., M & V NA16561 (TRT; Morton et al. 2018 [30]); Jones Co., S & Su 9751 (WAT; Semple et al. 1993); New Hanover Co., B and Beal 57-408, 57-409, 57-410 (MT; Beaudry 1963 [14]). South Carolina: Clarendon Co., S & Su 9785 (WAT; Semple et al. 1993 [28]); Florence Co., S & Ch 6107 (WAT; Semple et al. 1984 [23]). Virginia: Southampton Co., S 11624 (WAT; Semple et al. 2019 [31]).—2n = 18 + 1 supernumerary. U.S.A. Florida: Osceola Co., S 7477 (WAT; Semple et al. 1984 [23]). 2n = 18 + 4 supernumeraries. U.S.A. South Carolina: Beaufort Co., EMH 57-704 (MT; (Beaudry 1963 [14])—2n = 18 + 7, 8, 9, 10 supernumeraries. U.S.A. Georgia: Ware Co., B 57-739 (MT; Beaudry 1963 [14]).
Solidago latissimifolia Mill. —2n = 9II U.S.A. Connecticut: New London Co., S & Su 9538 (WAT; Semple et al. 2019 [31]).—2n = 18 CANADA. Nova Scotia: Halifax Reg. Mun., Kapoor 69-109-1 (SMUH; (Kapoor’72, as S. elliottii var. ascendens). U.S.A. North Carolina: Harnett Co., B & Beal 57-378 (MT; Beaudry 1963 [14] as S. elliottii var. elliottii). Rhode Island: Washington Co., B & Palmatier 57-171 (MT; Beaudry 1963 [14] as S. elliottii var. ascendens), B & Palmatier 57-171 (MT; (Beaudry 1963 [14] as S. elliottii var. ascendens). —2n = 18II U.S.A. South Carolina: Chesterfield Co., S & Su 9778 (WAT; Semple et al. 1993 [28]).—2n = 36 U.S.A. Massachusetts: Plymouth Co., S & Brouillet 3553 (WAT; Semple et al. 1993 [28]). New Jersey: Ocean Co., S 11815 (WAT; Semple et al. 2019 [31]). —2n = 27II U.S.A. Rhode Island: Washington Co., S & Su 9544 (WAT; Semple et al. 1993 [28]).—2n = 54 U.S.A. Massachusetts: Plymouth Co., S & Su 9568 (WAT; Semple et al. 1993 [28]).
Solidago rugosa Mill. var. aspera (Ait.) Fernald—2n = 9II U.S.A. Mississippi.: Jasper Co., S. Jones 15167 (MISS; S. Jones 1968 [17] as var. rugosa). —2n = 18 U.S.A West Virginia: Monongalia Co., B 57-265 (MT; Beaudry 1969 [15] as S. aspera); Monongalia Co., B 57-265 (MT; Beaudry 1969 [15] as S. aspera).—2n = 36 U.S.A. Arkansas: Lafayette Co., Demaree 59-270-1 (MT; Beaudry 1969 [15] as S. aspera); Yell Co., M & V NA16268 (TRT; Morton et al. 2018 [30] as var. aspera). Delaware: Sussex Co., S & Ringius 7641 (WAT; Semple et al. 1992; to sp.). New Jersey: Ocean Co., B 57-215, 57-216, 57-217 (MT; Beaudry 1960 [8], as S. aspera). North Carolina: Henderson Co., B & Fautrier 62-109 (MT; Beaudry 1969 [15] as S. aspera), Semple 10834 (WAT; Semple and Cook 2004 [28]; as var. aspera); Richmond Co., S, Brammall & Hart 3045 (WAT; Semple et al. 1981 [11]; as var. aspera); Transylvania Co., S 10843 (WAT; Semple and Cook 2004 [28], as var. aspera). Oklahoma: Pushmataha Co., Semple & Heard 8264 (WAT; Semple and Chmielewski 1987 [25] to sp.). South Carolina: Beaufort Co., Horlbeck 57-709 (MT; Beaudry 1960 [8], as S. aspera). Virginia: Allegheny Co., Cook & Tereszchuk 357 (WAT; Semple et al. 2015 [29] as var. aspera); Prince William Co., Semple, Brammall & Hart 3069 (WAT; Semple et al. 1981 [11] as var. aspera).
Solidago rugosa var. aff. aspera—2n = 36 U.S.A. North Carolina.: Avery Co., B & Fautrier (MT; Beaudry 1969 [15], as S. aspera).—2n = 36 + 1 supernumerary. U.S.A. Rhode Island: Washington Co., B & Palmatier 57-172 (MT!; Beaudry 1960 [8]).
Solidago rugosa Mill. var. celtidifolia Fernald—2n = 18 U.S.A. Louisiana: Vernon Parish, M & V NA16423 (TRT; Morton et al. 2018 [30]). North Carolina: Wake Co., B 57-367 (MT; Beaudry 1960 [8]). Virginia: Southampton Co., Semple 11617 (WAT; Semple et al. 2019 [31]), Semple 11621 (WAT; Semple et al. 2019 [31]).
Solidago rugosa Mill var. cronquistiana Semple—2n = 18II U.S.A. North Carolina: Avery Co., S 11131 (WAT; Semple et al. 2015 [29] as var. cronqusitiana).
Solidago rugosa Mill. var. rugosa (as var. rugosa unless otherwise indicated)—2n = 9II CANADA. Ontario: Algoma Dist.: S & K. Shea 1877 (WAT; Semple et al. 1981 [11]); Brant Co., S 2393 (WAT; Semple et al. 1981 [11]) Glengary Co., Semple 2410 (WAT; Semple et al. 1981 [11]); Sudbury Dist.: S & Brammall 2840 (WAT; Semple et al. 1981 [11]). U.S.A. Massachusetts: Middlesex Co., Goodwin 698 GH; Goodwin 1937; cultivated in Rochester, NY and from source population noted on label). New Hampshire: Cheshire Co., Gregory 493 (NY; Anderson et al. 1974). —2n = 9II + 1 supernumerary. CANADA. Ontario: Parry Sound Dist.: S & Brammall 2379 (WAT; Semple et al. 1981 [11]). —2n = 18 CA. New Brunswick: Queen’s Co., M & V NA12517 (TRT; Morton et al. 2018 [30]). Newfoundland: M & V NA12202 (TRT; Morton et al. 2018 [30]), M & V NA12130 (TRT; Morton et al. 2018 [30]). Nova Scotia: Halifax Reg. Mun., Kapoor 69-70-1 (SMUH not seen; (Kapoor’70; as f. villosa); Richmond Co., S & Keir 4778 (WAT, MT; Semple et al. 1984 [23] to sp.); Yarmouth Co., S & Keir 4852 (WAT; Semple et al. 1984 [23] to sp.). Ontario: Algoma Dist., S & B. Semple 11083 (WAT; Semple et al. 2015 [29]), S & B. Semple 11096 (WAT; Semple et al. 2015 [29]); Bruce Co., Cook & Faulkenham 10 (WAT; Semple and Cook 2004 [28]), Cook & Faulkenham 9 (WAT; Semple and Cook 2004 [28]), Ch 2267 (WAT; Semple and Chmielewski 1987 [25] to sp.); Cochrane Dist., S & Brammall 2826 (WAT; Semple et al. 1981 [11] as var. villosa); Frontenac Co., S 10664 (WAT; Semple and Cook 2004 [28]); Grey Co., Cook & Faulkenham 15 (WAT; Semple and Cook 2004 [28]), Cook & Faulkenham 16 (WAT; Semple and Cook 2004 [28]), Ch 2271 (WAT; Semple and Chmielewski 1987 [25] to sp. Ch 2273 (WAT; Semple and Chmielewski 1987 [25] to sp.), Ch 2277 (WAT; Semple and Chmielewski 1987 [25] to sp.) Ch 2275 (WAT; Semple and Chmielewski 1987 [25] to sp.); Halton Reg. Mun.: Ch 2233 (WAT; Semple et al. 1989 [26] to sp.); Chi 2237 (WAT; Semple, Chmielewski and Lane 1989, to sp.); Lanark Co., Ch 2318 (WAT; Semple and Chmielewski 1987 [25] to sp.); Muskoka Dist.: Co., Semple & K. Shea Semple 2931 (WAT; Semple et al. 1981 [11]), Ch 2293 (WAT; Semple and Chmielewski 1987 [25] to sp.), Ch 2295 (WAT; Semple and Chmielewski 1987 [25] to sp.); Nipissing Dist.: Ch 2299 (WAT; Semple and Chmielewski 1987 [25] to sp.); Parry Sound Dist.: Co., Semple & Brammall 2889 (WAT; Semple et al. 1981 [11]); Renfrew Co., Ch 2313 (WAT; Semple and Chmielewski 1987 [25] to sp.), Ch 2314 (WAT; Semple and Chmielewski 1987 [25] to sp.); Timiskaming Dist.: Slater 1 (WAT; Semple et al. 1984 [23] to sp.), Slater 2 (WAT; Semple et al. 1984 [23] to sp.); Wellington Co., Ch 2004 (WAT; Semple and Chmielewski 1987 [25] to sp.), S 7397 (WAT; Semple et al. 1992 [27] to sp.), S & Cook 10650 (WAT; Semple and Cook 2004 [28]); Wentworth Reg. Mun.: Ch 2235 (WAT; Semple and Chmielewski 1987 [25] to sp.). Prince Edward Island: Prince Co., M & V NA12047 (TRT; Morton et al. 2018 [30]). Québec: Argenteuil Co., B, Blume & Chabot 57-689 (MT!; Beaudry’60 JCS’2019), B, Blume & Chabot 57-690 (MT; Beaudry 1960 [8]); Gaspésie, S & B. Semple 11422 (WAT Semple et al. 2019 [31]); W of Havre-St-Pierre, M & V NA15300 (TRT; Morton et al. 2018 [30]); Montcalm Co., Fr. Marcellin-Sylvio 55-191 (MT; Beaudry and Chabot 1959 [13]); N of Laniel, Ch 2310 (WAT; Semple and Chmielewski 1987 [25] to sp.); Papineau Co., B 55-200 (MT; Beaudry and Chabot 1959); Parc La Vérendry, B & Doucet 59-88 (MT!; Beaudry 1969 [15] as f. rugosa). U.S.A. Maine: Penobscot Co., S & Keir 4647 (WAT; Semple et al. 1984 [23] to sp.). Maryland: Cecil Co., S & Ringius 7617 (WAT; Semple et al. 1989 [26] to sp.). Michigan: Arenac Co., M & V NA10893 (TRT; Morton 1981 [21] to sp.). Massachusetts: Bristol Co., Seymour 57-28-1, -2, -3 (MT; Beaudry and Chabot 1959); Franklin Co., S 6871 (WAT; Semple et al. 1984 [23] to sp.). New Hampshire: Cheshire Co., S & Keir 4957 (WAT; Semple et al. 1984 [23] to sp.); Coos Co., Ringius 1583 (WAT; Semple et al. 1984 [23] to sp.); Grafton Co., S & Brouillet 3474 (WAT; Semple et al. 1981 [11]); Hillsborough Co., S & Keir 4942 (WAT; Semple et al. 1984 [23] to sp.). New York: Delaware Co., S 6817 (WAT, NY; Semple et al. 1984 [23] to sp.); Essex Co., Ringius 1536 (WAT; Semple et al. 1984 [22] to sp.), Ringius 1544 (WAT; Semple et al. 1984 [23] to sp.); Sullivan Co., S 6828 (WAT; Semple et al. 1984 [23] to sp.). Pennsylvania: Butler Co., R. Cook & D. Cook 55 (WAT; Semple and Cook 2004 [28]). Vermont: Addison Co., S 6898 (WAT; Semple et al. 1984 [23] to sp.); Chittenden Co., Ringius 1576 (WAT; Semple et al. 1984 [23] to sp.). Virginia: Bland Co., S 10735 (WAT; Semple and Cook 2004 [28]); Floyd Co., B & Fautrier 62-26 (M; Beaudry 1969 [15] as. S. aspera), B & Fautrier 62-27 (MT; Beaudry 1969 [15] as S. aspera). West Virginia: Marion Co., S 10687 (WAT; Semple and Cook 2004 [28]); Wisconsin: Crawford Co., S 9052 (WAT; Semple et al. 1989 [26] to sp.). —2n = 18 + 0, 2, 3 supernumeraries. CA. PQ: B & Doucet 59-89 (MT; Beaudry 1969 [15] as f. rugosa). —2n = 36 U.S.A. Arkansas: Logan Co., Demaree 59-265-2 (MT; Beaudry’69, as S. aspera). Delawate: New Castle Co., S & Ringius 7621 (WAT; Semple et al. 1989 [26] to sp.). Indiana: Porter Co., S & Brouillet 4526 (WAT; Semple and Chmielewski 1987 [25] to sp.). Maine: Franklin Co., S & Keir 4631 (WAT; Semple et al. 1984 [23] to sp.). Maryland: Washington Co., S & Ringius 7655 (WAT; Semple et al. 1992 [27] to sp.). Ohio: Ashland Co., R. Cook & D. Cook 36 (WAT; Semple and Cook 2004 [28]). Pennsylvania: Centre Co., S & Ringius 7603 (WAT; Semple et al. 1989 [26] to sp.). Vermont: Lamoille Co., S & Brouillet 3440 (WAT; Semple et al. 1981 [11]); Nicholas Co., S 10703 (WAT; Semple and Cook 2004 [28]).
Solidago rugosa Mill. var. rugosa (counts are uncertain due to voucher identity)—2n = 54 CANADA. Québec: Löve, Löve & Rousseau 6976 (CAN; Löve and Löve 1982; voucher is not for 6x count). U.S.A. New York: Chattaraugus Co., S & Ringius 7571 (WAT; Semple et al. 1992 [27] to sp.; identity of greenhouse collection sample could not be confirmed to be same as voucher; may be a field numbering error).
Solidago rugosa aff. var. rugosa—2n = 18 CANADA. Ontario: Hastings Co., S & Brammall 2925 (WAT; Semple et al. 1981 [11] as var. aspera). U.S.A. Virginia: Floyd Co., B & Fautrier 62-25 (MT; Beaudry 1969 [15], as S. aspera; leaves weakly rugose); West Virginia: Preston Co., B 57-268 (MT; Beaudry 1969 [15]).
Solidago rugosa Mill. var. sphagnophila Graves—2n = 54 CANADA. Nova Scotia: Shelburne Co., Kapoor 69-130-1 (SMUH not seen; Kapoor 1970 as S. aestivalis), Kapoor 69-130-4 (SMUH not seen; Kapoor 1977). U.S.A. Massachusetts: Nantucket Co., McKeever 64-1, -2, -3, -4, -5, -6, -7, -8 (MT; Beaudry 1969 [15], as S. aestivalis). New Jersey: Ocean Co., B 57-219 (MT; Beaudry 1960 [8]), B 57-220 (MT; Beaudry 1960 [8]). Rhode Island: Washington Co., B & Palmatier 57-177 (MT; Beaudry 1960 [8]).
Solidago rugosa aff. var. sphagnophila—2n = 54 U.S.A. New Jerseu: Ocean Co., cult. Montréal, B 57-220 (MT; Beaudry 1960 [8] as S. aestivalis); stem glabrous proximally, becoming moderately strigose by mid stem, to strigose in inflorescence).
Table A2. Previously unpublished chromosome counts for Solidago fistulosa and S. rugosa. S = J.C. Semple; Su = B.A. Suripto. All cytovouchers in WAT in MT (Thiers continuously updated) [13].
Table A2. Previously unpublished chromosome counts for Solidago fistulosa and S. rugosa. S = J.C. Semple; Su = B.A. Suripto. All cytovouchers in WAT in MT (Thiers continuously updated) [13].
Solidago fistulosa Ait.—2n = 18 U.S.A. North Carolina: Bladen Co., NC-242 N of Elizabethtown, 0.8 km N of US-701, S 11792.
Solidago rugosa Mill. var. aspera (Ait.) Fernald—2n = 18 U.S.A. Alabama: Bibb Co., NW of Lawley, US-82 5.2 km N of county line, S & Su 10161. Kentucky: Leslie Co., S of Stinnett, US-421 just S of bridge across river, S & Su 9616. Oklahoma: McCurtain Co., US-259 20 km S of county line, 5.0 km E of OK-144, S & Su 9989. South Carolina: Kershaw Co., S of Camden, US-521 4.4 km NW of Swift Creek, S & Su 9781-2.—2n = 18II U.S.A. New Jersey: Ocean Co., S end of Parkertown, by US-9, S & Su 9523.—2n = 36 U.S.A. Arkansas: Logan Co., AR-23 5.5 km E of US-72, 1.1 km E of Scott Co. line, S & Su 9979, SE of Booneville, AR-23 2.2 km E of Carolon Cemetery, S & Su 9978; Polk Co., US-270, NW of Mena, Cook & Terezchuk 225, AR-88, unmarked unpaved rest area (1st on AR side), Cook & Terezchuk 234; Sebastian Co., AR-28 14 km W of US-71, 4.6 km E of Canthron, S & Su 9982. Kentucky: Christian Co., SE of Dawson Springs, KY-109 0.8 km from KY-398, S & Su 10334; Laurel Co., Daniel Boone St. For., For.Rd-056 ca. 2 km W of KY-192, S & Su 9610, Rockcastle R., gravel bar at Pine Is. Branch, S & Su 9857. Louisiana: Rapides Par., SE of Gardner, LA-121 5.1 km N of LA-1199, 1.8 km SE of LA-28, S & Su 10094; Red River Par.: NE of Edgefield, LA-7 11.2 km S of county line, S & Su 10037. Missouri: Wayne Co., Co.Rd.-A 10.9 km SW of US-67, N of Williamsville, S & Su 9898, Co.Rd.-F 0.1 km E of US-67 7.8 km S of Greenville, S & Su 9896. Mississippi: Franklin Co., NW of Roxie, US-84 8 km NE of county line, 1.5 km W of MS-33, S & Su 10099; Itawamba Co., MS-23 10.1 km N of Tremant, S & Su 10177, MS-23 7.4 km N of county line, S & Su 10171; Monroe Co., S of Becker, MS-25 0.9 km N of MS-8, S & Su 10168. North Carolina: Alamance Co., NC-62 1.5 km N of Alamance, by Gunn Creek, S & Su 9717; Buncombe Co., US-19 at S city limits of Weaverville, S & Su 9695; Yadkin Co., WSW of Yadkinville, US-421 ca. 1 km E of US-21, S & Su 9710; Yancey Co., Spruce Pine, US-19E by-pass, S & Su 9697. Oklahoma: Flore Co., US-59 just N of Ouchita Nat’l For., S & Su 9988; McCurtain Co., N of Broken Bow, US-259 20.9 km N of US-71, S & Su 9992, W of Millerton, US-70 3.6 km E of Valient, S & Su 9998. Texas: Anderson Co., ca. 20 km NW of Palestine, US-287 just W of Farm Rd.-321, S & Su 10015, E of Palestine, US-84 17.6 km E of Loop Rd-56, S & Su 10018; Cherokee Co., US-84 4.9 km E of Neches River, W of Maydelle, S & Su 10020; Harrison Co., NE of Marshall, TX-43 7.2 km NE of US-59, S & Su 10027; Newton Co., NE of Burkeville, TX-63 2.5 km W of Sabine R., S & Su 10051-B. Virginia: Augusta Co., George Washington N.F, US-250 ca 0.2 km E of Calf Pasture Creek, Cook & Tereszchuk 383; Bath Co., US-220 ca 1 km S of VA-39, Cook & Tereszchuk 370.
Solidago rugosa Mill. var. aspera aff var. celtidifolia—2n = 36 U.S.A. Florida: Leon Co., N of Bradfordville, S of Felps, LA-67 2.5 km S of LA-422, S & Su 10109. South Carolina: Clarendon Co., US-521 1.1 km SE of Wilson, S & Su 9787.
Solidago rugosa var. aspera aff. var. rugosa—2n = 18 U.S.A. Virginia: Lee Co., US-421 just outside Duffield, S & Su 9622.—2n = 36 U.S.A. Arkansas: Logan Co., Magazine Mt., For.Rd.-1606 1.5 km W of AR-309, S & Su 9974; Sebastian Co., AR-28 14 km W of US-71, 4.6 km E of Cauthron, S & Su 9983; Washington Co., N of Brentwood, US-71 at rest area, bank of creek, S & Su 9961. Kentucky: Menifee Co., Red R. Gorge, KY-715 1.2 km E of Tower Rock Trail No. 206, deeply shaded woods, S & Su 9861. Louisiana.: East Feliciana Par.: S of Felps, S of LA-422, S & Su 10110; Webster Par.: Co., LA-159 8.4 km S of Leton (LA-2), S & Su 10035. Mississippi: Itawamba Co., MS-23 10.1 km N of Tremont, S & Su 10176. Oklahoma.: Flore Co. US-59 just N of Ouchita Nat’l For. boundary, S & Su 9986. Texas.: Rusk Co., US-84 6.4 km E of Reklaw, E of TX-204, S & Su 10024.
Solidago rugosa var. aspera × var. cronquistiana—2n = 36 U.S.A. Georgia: Habersham Co., GA-15 2.0 km N of Tallulah Falls, S & Su 9828 (population a mixture of var. cronquistiana and var. aspera. X cronquistiana plants).
Solidago rugosa Mill. var. celtidifolia Fernald—2n = 9II U.S.A. Alabama: Mobile Co., N of Theodore, Co.Rd.-59 ca. 2 km S of I-10, S & Su 10127. Mississippi: Harrison Co., US-46 12 km N of Lyman, just N of Nat’l For. boundary, S & Su 10122. North Carolina: Richmond Co., NC-177 just S of Hamlet, S & Su 9775. South Carolina: Chesterfield Co., US-1 just SE of US-52, S & Su 9779; Kershaw Co., S of Camden, US-521 4.4 km NW of Swift Creek, S & Su 9781-1.—2n = 18 U.S.A. Alabama: Autauga Co., just W of Prattville, AL-206 at US-82, S & Su 10157; Dale Co., US- 84 near Choctawhatchee, near Choctawhatchee R., S & Su 10153. Florida: Walton Co., FL-20 4.8 km E of Freeport, S & Su 10135. Louisiana: Calcasieu Par.; E of DeQuincy, LA-12 0.5 km E of county line, S & Su 10086; Rapides Par., N of Forest Hill, LA-112 2.1 km W of US-165, S & Su 10093; Taugipahoa Par.; LA-22 8 km E of centre of Ponchatoula, S & Su 10117. North Carolina: Granville Co., NE of Butner, W of I-85 on county rd., S & Su 9723.—2n = 18II U.S.A. North Carolina: Brunswick Co., NC-133 W of Wilmington, S & Su 9764; Jones Co., US-17 ca. 5 km NE of Maysville, S & Su 9754; Pender Co., Hampstead, NC-210 ca 4.1 km W of US-17, S & Su 9759.—2n = 36 U.S.A. Florida: Jackson Co., Co.Rd-286 1.5 km N of Calhoun Co. (SE corner) line, S & Su 10143. Georgia: Thomas Co., W of Thomasville, US-84 just W of US-84 bus.-W, S & Su 10149. Louisiana: Beauregard Par.: LA-12 SE of DeQincy, S & Su 10084; Caddo Par.: Co., S of junction of LA-2 and LA-1, S & Su 10029. Mississippi.: Loundes Co., Columbus, N end of town, US-45 just S of Chapman Rd., S & Su 10166. North Carolina: Martin Co., US-64 just W of Parmele, S & Su 9738-A. Oklahoma: Flore Co., US-59 just N of Ouachita Nat’l For., S & Su 9987. South Carolina: Colleton Co., NW of Jacksonboro, NC-64 ca. 2.5 km from US- 17, S & Su 9797. Texas: Newton Co., TX- 63 2.5 km W of Sabine River, NE of Burkeville, S & Su 10051-A; Orange Co., NE of Vidor, TX-12 ca 3 km NE of I-10, S & Su 10081; Polk Co., E of Livingstone, US-190 1.8 km E of Soda, 16.4 km E of TX-146, S & Su 10060.
Solidago rugosa Mill. var. celtidifolia aff. var. aspera—2n = 9II U.S.A. North Carolina: Chowan Co., NC-37 just N of Albermarle Sound Causeway, ca. 10 km SE of Edenton, S & Su 9744.—2n = 18 U.S.A. Alabama: Coffee Co., NNW of Enterprise, AL-167 13.3 km S of county line, S & Su 10154. —2n = 36 U.S.A. Louisiana: East Feliciana Par.: Co., E of Woodland, LA-432 1.5 km E of LA-67, S & Su 10112. Mississippi.: Itawamba Co., MS-23 7.4 km N of county line, S & Su 10170. South Carolina: Berkeley Co., SC-45 ca 5 km E of St. Stephen, Francis Marion Nat’l For., S & Su 9791; Jasper Co., NC- 462 2.8 km W of Cousawhatchie, S & Su 9806. Texas: San Jacinto Co., TX-150 3.6 km E of Shepherd, S & Su 10063.
Solidago rugosa Mill. var. cronquistiana Semple—2n = 36 U.S.A. NC: Avery Co., between Linville and Roseborough, E of Blue Ridge Parkway, Semple 11120, US-221 S of Linville, S & Su 9698, E of Linville, Grandfather Mt., ca. 5000 ft el., S & Su 9675; Mitchell Co., top of Roan Mt., ca. 6260 ft el., Roan High Bluff Rd. (For.Rd.-130), S & Su 9667.
Solidago rugosa Mill. var. rugosa—2n = 9II U.S.A. Pennsylvania: Lycoming Co., SE of South Williamsport, vicinity of rest area by junction of US-15 and Skyline Drive, S & Su 9480; Tioga Co., Cowanesque Conservation Area Rd., W of US-15, S & Su 9476.—2n = 18 CANADA. Ontario: Parry Sound Co., Hwy-522 at junction of Stumpy Bay Rd, 27.4 km W of Loring, Semple & B. Semple 11072.—2n = 18 CANADA. Québec: Abitibi-Témiscamingue Co., 0.3 km from Hwy-117 on rd to la Mine Doyon, Cook & Seiden 105, Hwy-109 between Ames and Rivière-Héva, Cook & Seiden 104; Outaouais Co., Reserve la Verendrye, off Hwy-117, Park Rd-14, Cook & Seiden 122. U.S.A. Connecticut: New Haven Co., US-1 W of Branford, S & Su 9532; New London Co., Waterford, swamp by Cross Roads Mall, S & Su 9541-1, Waterford Conservation Area, Nevins Brook, S 9190. Kentucky: Menifee Co., Boone Nat’l For., KY-715, S & Su 9463. Maine: Oxford Co., Hwy-2 5 mi W of West Bethel, Ringius 1597. Massachusetts: Barnstable Co., Scusset Beach St. Pk., along rd to beach, S & B. Semple 11033, Cape Cod, W of Harwich, Great Western Rd. 2.0 km W of Clough Rd., just E of Lothrop Ave, S & Su 9563, Orleans, S & Su 9557; Norfolk Co., Dover, Springdale Ave. N of town centre, Chenning Park, S & Su 9572, NW of Medfield, Farm St. WW of Centre St, by RR, Semple 11027. New Jersey: Burlington Co., SE of Chatsworth on county rd., S & Su 9515; Ocean 10334Co., E of Cedar Run, cedar swamp just S of Cedar Run Dock Rd., S & Su 9521. New York: Livingston Co., Webster Crossing, NY-15, S & Su 9472; Suffolk Co., East Islip, Post Office Rd., S & Su 9529.—2n = 36 US. Connecticut: New London Co., NE of Voluntown, Pachaug State For., S 9188.
Solidago rugosa var. rugosa aff. var. aspera—2n = 36 U.S.A. Massachusetts: Plymouth Co., Marion, US-6, S & Su 9548. Pennsylvania: York Co., PA-372 woods below bridge over Susquehana R., S & Ringius 7607.
Solidago rugosa Mill. var. sphagnophila Graves—2n = 9II U.S.A. Connecticut: New London Co., Waterford, swampy ground by Cross Roads Mall, S 9192-A, S 9192-B. Massachusetts: Barnstable Co., Cape Cod, W of Harwich, Great Western Rd. just E of Lothrop Ave., S & Su 9564; New Jersey: Atlantic Co., alongside rd. E of NJ-50 just S of US-40 and just N of Co.Rd.-669, S & Su 9505.—2n = 18 U.S.A. Massachusetts: Barnstable Co., Cape Cod, Orleans, S & Su 9556-3, NE of Orleans, junction of Mallard Lane and Skaket Beach Rd., S & Su 9554-2.

References

  1. Semple, J.C.; Beck, J.B. A revised infrageneric classification of Solidago (Asteraceae: Astereae). Phytoneuron 2021, 10, 1–6. [Google Scholar]
  2. Fernald, M.L. Gray’s Manual of Botany, 8th ed.; Van Nostrand: New York, NY, USA, 1950. [Google Scholar]
  3. Fernald, M.L. Contributions from the Gray Herbarium of Harvard University. No. CXIII. VI. Studies in Solidago. Rhodora 1936, 38, 209–228. [Google Scholar]
  4. Cronquist, A. Notes on the Compositae of the northeastern United States IV. Solidago. Rhodora 1947, 49, 69–79. [Google Scholar]
  5. Cronquist, A. Solidago. In The New Britton and Brown Illustrated Flora of the Northeastern United States and Adjacent Canada; Gleason, H.A., Ed.; Hafner Pub. Co.: New York, NY, USA, 1968; pp. 413–438. [Google Scholar]
  6. Cronquist, A. Vascular Flora of the Southeastern United States; Vol.I. Asteraceae; University of North Carolina Press: Chapel Hill, NC, USA, 1980. [Google Scholar]
  7. Gleason, H.A.; Cronquist, A. Manual of Vascular Plants of Northeastern United States and Adjacent Canada, 2nd ed.; New York Botanical Garden: Bronx, NY, USA, 1991. [Google Scholar]
  8. Beaudry, J.R. Studies on Solidago L. V. The Solidago rugosa complex. Can. J. Genet. Cytol. 1960, 2, 389–396. [Google Scholar] [CrossRef]
  9. Uttal, L.J.; Porter, D.M. The correct name for Elliott’s goldenrod. Rhodora 1988, 90, 157–168. [Google Scholar]
  10. Semple, J.C.; Cook, R.E. Solidago linnaeus. In Flora of North America. Asteraceae, Part 2. Astereae and Senecioneae; Flora North America Editorial Committee, Ed.; Oxford University Press: New York, NY, USA, 2006; Volume 20, pp. 107–166. [Google Scholar]
  11. Semple, J.C.; Brammall, R.A.; Chmielewski, J. Chromosome numbers of goldenrods, Euthamia and Solidago (Compositae-Astereae). Can. J. Bot. 1981, 59, 1167–1173. [Google Scholar] [CrossRef]
  12. Goodwin, R.H. The cytogenetics of two species of Solidago and its bearing on their polymorphism in nature. Am. J. Bot. 1937, 24, 425–432. [Google Scholar] [CrossRef]
  13. Beaudry, J.R.; Chabot, D.L. Studies on Solidago IV. The chromosome numbers of certain taxa of the genus Solidago. Canad. J. Bot. 1959, 37, 209–288. [Google Scholar] [CrossRef]
  14. Beaudry, J.R. Studies on Solidago L. VI. Additional chromosome numbers of taxa of the genus Solidago. Can. J. Genet. Cytol. 1963, 5, 150–174. [Google Scholar] [CrossRef]
  15. Beaudry, J.R. Etudes sur les Solidago IX. Une Troisieme liste de nombres chromosomiques des taxons du genre Solidago et de certains genres voisins. Nat. Can. 1969, 97, 431–445. [Google Scholar]
  16. Thiers, B. Index Herbariorum: A Global Directory of Public Herbaria and Associated Staff. New York Botanical Garden’s Virtual Herbarium. Continuously Updated. Available online: http://sciweb.nybg.org/science2/IndexHerbariorum.asp (accessed on 6 September 2021).
  17. Jones, S.B. Chromosome numbers in southeastern United States Compositae. Bull. Torrey Bot. Club 1968, 95, 488–489. [Google Scholar] [CrossRef]
  18. Kapoor, B.M. In IOPB chromosome number reports XXVII. Taxon 1970, 19, 437–442. [Google Scholar]
  19. Anderson, L.C.; Kyhos, D.W.; Mosquin, T.; Powell, A.M.; Raven, P.H. Chromosome numbers in Compositae. IX. Haplopappus and other Astereae. Am. J. Bot. 1974, 61, 665–671. [Google Scholar] [CrossRef]
  20. Kapoor, B.M. Further observations on the morphology of some Solidago species. Cytologia 1977, 42, 241–253. [Google Scholar] [CrossRef]
  21. Morton, J.K. Chromosome numbers in Compositae from Canada and the U.S.A. Bot. J. Linn. Soc. 1981, 82, 357–368. [Google Scholar] [CrossRef]
  22. Löve, A.; Löve, D. IOPB chromosome numbers of reports. LXXVII. Taxon 1982, 31, 766–768. [Google Scholar] [CrossRef]
  23. Semple, J.C.; Ringius, G.S.; Leeder, C.; Morton, G. Chromosome numbers of goldenrods, Euthamia and Solidago (Compositae: Astereae). II. Additional counts with comments on cytogeography. Brittonia 1984, 36, 280–292. [Google Scholar] [CrossRef]
  24. Semple, J.C. Chromosome number determinations in Fam. Compositae Tribe Astereae. Rhodora 1985, 87, 517–527. [Google Scholar]
  25. Semple, J.C.; Chmielewski, J.G. Chromosome numbers in Fam. Compositae, Tribe Astereae. II. Additional Counts. Rhodora 1987, 89, 319–325. [Google Scholar]
  26. Semple, J.C.; Chmielewski, J.G.; Lane, M.A. Chromosome numbers in Fam. Compositae, Tribe Astereae. III. Additional counts and comments on generic limits and ancestral base numbers. Rhodora 1989, 91, 296–314. [Google Scholar]
  27. Semple, J.C.; Chmielewski, J.G.; Chun, X. Chromosome numbers in Fam. Compositae, Tribe Astereae. IV. Additional reports and comments on the cytogeography and status of some species of Aster and Solidago. Rhodora 1992, 94, 48–62. [Google Scholar]
  28. Semple, J.C.; Jie, Z.; Chun, X. Chromosome numbers in Fam. Compositae, Tribe Astereae. V. Eastern North American taxa. Rhodora 1993, 95, 234–253. [Google Scholar]
  29. Semple, J.C.; Cook, R.E. Chromosome number determinations in fam. Compositae, Tribe Astereae. VII. Mostly eastern North American and some Eurasian taxa. Rhodora 2004, 106, 253–272. [Google Scholar]
  30. Semple, J.C.; Cook, R.E.; Owen, E. Chromosome numbers in Fam. Compositae, Tribe Astereae. VIII. Eastern North American taxa. II. Rhodora 2015, 117, 80–91. [Google Scholar] [CrossRef]
  31. Morton, J.K.; Venn, J.; Semple, J.C. Chromosome number determinations in Solidago (Asteraceae: Astereae). Rhodora 2018, 119, 341–348. [Google Scholar] [CrossRef]
  32. Semple, J.C.; Cook, R.E.; Morton, G.H.; Beck, J.B.; Laphitz, R.L. Chromosome number determinations in Fam. Compositae, Tribe Astereae. IX. North American taxa. II. Rhodora 2019, 121, 37–53. [Google Scholar] [CrossRef]
  33. Beck, J.B.; Markley, M.L.; Zielke, M.G.; Thomas, J.R.; Hale, H.J.; Williams, L.D.; Johnson, M.G. Is Palmer’s elm leaf goldenrod real? The Angiosperms353 kit provides within-species signal in Solidago ulmifolia s.l. bioRxiv 2021. [Google Scholar] [CrossRef]
Figure 1. Cytogeography of Solidago fistulosa in eastern North America.
Figure 1. Cytogeography of Solidago fistulosa in eastern North America.
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Figure 2. Cytogeography of Solidago latissimifolia in eastern North America; the identification of the northernmost diploid sample in Nova Scotia (red dot) was not confirmed.
Figure 2. Cytogeography of Solidago latissimifolia in eastern North America; the identification of the northernmost diploid sample in Nova Scotia (red dot) was not confirmed.
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Figure 3. Cytogeography of Solidago rugosa and its varieties in eastern North America.
Figure 3. Cytogeography of Solidago rugosa and its varieties in eastern North America.
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Table 1. Comparison of some classification schemes of the Solidago rugosa complex and related species.
Table 1. Comparison of some classification schemes of the Solidago rugosa complex and related species.
Fernald (1950 [2])Cronquist (1947 [4], 1968 [5])Beaudry (1960 [8])Semple and Cook (2006 [9])
S. fistulosaS. fistulosa S. fistulosa
S. elliottiiS. elliottiiS. elliottiiS. latissimifolia
S. rugosaS. rugosaS. rugosaS. rugosa
 subsp. rugosa  subsp. rugosa
  var. rugosa  var. rugosa  f. rugosa  var. rugosa
   (including var. villosa)
  var. villosa  var. villosa  f. villosa
  var. sphagnophila  var. sphagnophilaS. aestivalis  var. sphagnophila
 subsp. aspera  subsp. aspera
  var. aspera  var. asperaS. aspera  var. aspera
   (including S. celtidifolia)
  var. celtidifolia S. celtidifolia  var. celtidifolia
  var. cronquistiana
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