SUMMARY GEOGRAPHIC OF A John Sempie c. CHROMOSOME NUMBER NORTH REPORTS FOR AMERICAN ASTERS AND GOLDENRODS (ASTERACEAE: ASTEREAE)^ Abstract A microcomputer system that facilitates record keeping of text-based nomenclatural and cytotaxonomic files is described. The system was used to maintain records on asters and goldenrods. More than 4,800 chromosome number reports for asters (Aster, Virgulastery Virgulus) and over 2,000 reports for goldenrods {Solidago excluding Euthamia) were analyzed. By far the greatest number of chromosome number reports (2,110) were for Ontario, Canada. The numbers of counts for each taxon (species, subspecies, or variety) for each province or territory in Canada and state in the United States and for of Mexico were determined. The total number of chromosome number reports and all estimated numbers of taxa (species, subspecies, and varieties) of asters and of goldenrods by province, territory, and state were compared and are presented graphically. Both asters and goldenrods achieve their greatest morphological diversity in the eastern United peaking North Carolina. The numbers of counts for asters and goldenrods States, in m = each area were 0.70 excluding Ontario; 0.97 including Ontario) as were the highly correlated fairly (r = numbers of taxa counted including Ontario 0.73), but the percentages of taxa counted per area were not strongly (r = 86% correlated (r 0.49). Asters show more diversity in western North America than goldenrods, and of the areas had more aster than goldenrod taxa. My work on members Astereae estimated various of the Walter to include about 90 species and more than 140 and Canada, Walter subspecies varieties in the United Chromosome and Mexico. numbers have inspiration are acknowledged, and this States, gratefully him been reported for most of these taxa (see Sempie 1984, and additional references cited below); et al., = taxa have a chromosomal base rmmber of x all common Asters and common many Polyploidy in both asters and gold- goldenrods are in 9. is More babitats throughout most North America north enrods. than 6,900 indivlJuaJ cljromusome of of Mexico. Aster {Aster of Jones, number reports exist for Aster sens. lat. and Sol- L. sens, lat, 1980a; Aster, Virgulus (synonym: Lasallea) idago in North America alone. In order to keep Raf. & and and have Virgulaster Sempie of Sempie Brouillet, track of these reports to conveni^-nt ac- system microcomputer 1980a, and Sempie 1989) are estimated to cess to the data, a of hard- et al., include about 145 and more than 225 ware an d cribed species Some geographic generalizations about subspecies and Canada, the United below. varieties in numfxT chromosome reports for asters and gold- States, and Numerous Mexico. America were North arrived using ported chromosome numbers many of these enrods in at the for & presented below taxa Sempie 1980b, and addi- (see Brouillet, below) & Methods AND MATERIALS and (Sempie 9, 8, 7 occur Aster sens. str. in = and 13 Brouillet, 1980a); base numbers of 21 jc HARDWARE occur numbe in Vir^ulaslen and base 9, W "m ' and includes 5, 4 occur Virgulus. Aster sens. lat. in MHz computer (8/25 25 personal clock speed) numbers and Fnginccring Sciences This research was supported by a Natural ^JJ^i^^,^";!"^^ Biology iiversity Carh. 95-109. Rot. 79: 1902. Missouri Ann. 96 Annals of the Garden Missouri Botanical V with a 387/25 math coprocessor, eight megabytes (TRIBCNTS-3 [= puWished by Semple et of memory, a 100 megabyte hard drive (15 msec. al., 1989], to sp. [= published to species VGA access time), and a Zenith color monitor. level only, subspecies identification not de- (Waitronics Ltd. (Waterloo, Ontario N2J 2Y9, termined at the time of publication]) Canada) is one of a number of local companies LA. glaucode^ 2n=9II US WY. 2. Albany NW (IBM assembling microcomputers clones) that are Co.: Snowy Range, 3 mi of Centennial, sold at reduced prices to the University of Waterloo LHartman 3045-1 (RM) (Hartman'77) for research purposes.) I La. 3. pilosus-J var. Lpilo.-I [epithets sometimes US 2n=32 abbreviated] AL. Cleburne-Cal- SOFTWARE & LS houn SE count of Anniston, Ch line; software used and & All to tabulate analyze the 6305-1 [= Semple Chmie- collectors, chromosome count data were widely com- available lewskQ (PILOGEO-2 [= published by Sem- prod & Chmielewsk 985] ple as var. i I 1 DESQview (Q Lpriceae-I: corrected terdeck Office Systems, Santa Monica, California [= Semple 1989 noted that the et col- al. 90405, memory U.S.A.). This management pro- lection on which the 1985 report was based gram allows the user run more than one program to was reidentified as var. pilosus~\) a time and at to transfer screen data from one 2n=18 US 4. LSol. rigida-l ssp. Lglabrata-I cf. program another to regardless of the compatibility TN. LJRB [= Coffee Co.: Manchester, //?. of the individual programs. For example, a personal & Beaudry] DeSebn 57-472-1 (Beaudry'63, runnm the background, while word processing and data as LSol. jacksonii-l var. Lhum.-' [= var. entries are being performed two synonym in additional pro- grams. ssp bratal) location Raw data were entered askSam un into using files Version 4.1b (Access Stored Knowledge Sym- geography map on because via species identifi- bolic Access Method; askSam Systems, 1989). Each cation was uncertain time record was add- at was report entered as ASCII ed text without to field file] designation in the following order: taxon (binomial 5. LSol. simplex-l Lrandii-J var. ssp and subspecies or variety if appropriate); chro- CDN Lmonticola-1 2n=36 PQ. Megantic Co.: mosome number reported in standardized format & LjRB Black Lake, [= Beaudry] Cinq- J,R. "2n=9ir' "2n=18" or (e.g., differentiating mei- & Mars 56-433-1 as (Beaudry Chabot'59, from otic mitotic reports); country "CDN, (e.g., LS. R&S'87, LS. ran.-" US, MEXICO''); randii-l; as glut, province, territory, or state (post- code var. Lran.-I [= Solidago gludnosa subsp. abbreviation, al e.g., *'0N"); county, regional synonym of municipality, parish, or district "Waterloo randii var. randii, which is a (e.g., Reg.Mun."); simplex details of location data "Hwy-7 S. var. monticola]) (e.g., W km "L" 4 of Kitchener- rdside Hitrh"V f-olUr-t/^^ o^^ The symbols and "-P' (keyboard characters number; collection location of voucher(s) ah- 192 and ah-2 were inserted to not 17, respectively) if WAT; abbreviated publication data including any replace non-ASCII printer codes indicating that the synonym used "Anderson enclosed These con- (e.g., et al.'74, as A. word(s) should be h., italicized. n=32 ?^ as in the original publication the report veniently allow specific letters or words to be con- was listed as n 32 under the name Aster verted to when between askSam hes- italics shifting text perius), and any and used miscellaneous comments WordPerfect, The two symbols can be (e.g., observations about vouchers or a summary(ies) as query characters searching data bases. of in published comments about Any can be the count or voucher, of the mformation a data in file such as corrected The askSam, identifications). retrieved using Query following are the subroutine in examples of report and any any entries (in quotes and exactly individual report can be edited at as entered) with explanatory comments in brackets time. The Query subroutine can be used to also [=1 added here but not included the datp obtain a designated in tally of the reports containing filf^- I 1 search {PRINT}'') '*LA. lanceolatu^ fields (e.g., L LA.alpigenusJssp.LaIpigenu^2n=36US '^A. Boulean searches can made be also (e.g., Mono GA. Co.: Tioga E Pass, 0,25 mi 48 of lanceolatu^ {AND} 2n Uanceolatu^ var. & LS Yosemite Natn Pk, Hd {NOT} CDN ON 8688-1 data Any {TAL}"). of the or all ? Volume Number 79, Semple 1 97 1992 Chromosome Number Reports for Asters and Goldenrods & can be transferred directly or indirectly to a word- 1977; Jones, 1978; Powell Powell, 1978; Dawe & & Wordperfe Murray, 1979; Morton, 1979; Kail Pinkava, Wordpe & version feet Corp., Orem, Utah 1979; Pringle, 1979; Dawe Murray, 1980; 84057, U.S.A.) using either a subroutine askSam Weed in & & by saving the results of a Query operation as a Semple, 1981; Dawe Murray, 1981; & Keil new ASCII & by text or using the screen capture Pinkava, 98 Harriman, 98 Sem- file Parfitt 1 1 1 ; 1 ; DESQ & pie, 1981; Semple 1981; Kapoor et Gervais, al., & Wordperfe 1982; Love L6ve, 1982a, Semple, 1982; b; & convenience Semple It is this in searching and ex- Brammall, 1982; 1983; files Brouillet, & & changing information between data base programs Chmielewski Semple, 1983; Dean Chambers, and word-processing makes files that the system 1983; Hill, 1983; Sherif et 1983; Semple al., et being discussed useful for systematists. 1983a, Strother, 1983; Ward, 1983; Wind- al., b; & number ham Totals for the of reports for each taxon Schaack, 1983; Allen, 1984; 1984; Hill, Q Jones, 1983; Morton, 1984; Mulligan, 1984; San- entered into a matrix (taxon by political region) derson et 1984; Semple, 1984; 1985; al., Allen, & & Houle Brouillet, 1985; Chmielewski Semple, Q^ & (Borland International, Scotts Valley, California 1985a, b; Semple, 1985; Semple Chmielewski, 95066, The number U.S.A.). total of counts for 1985; Chmielewski, 1986; Sundberg, 1986; Sund- number Ward & & region (country; province, territory, or state), the Brouillet Labrecque, 1987; Chinnappa number of taxa and the number of counted and Chmielewski, 1987; Chmie- uncounted & lewski 1987; Lamboy, 1987; et Ringius al., & grand total for all count reports were calculated Semple, 1987; Semple Chmielewski, 1987; Va- & as part of the matrix of 28,890 data The hidy 1987; Heard Semple, 1988; Lam- fields. et al., & Q boy, 1988; Semple, 1988; Campbell Medley, was & facilitated using the Video and Switch options 1989; Legault 1989; Nesom, 1989 Brouillet, DESQ & Semple 1989; Brammall Semple, 1990 et al., & Q Chmielewski Semple, 1990; Semple 990 et al., 1 A & taneously. compiler program could not be used Semple Chmielewski, 1991). These data were with askSam, name entered under the taxon used the source in Also frequently consulted were askSam on publication unless the name was known to be a files synonym name oximate for another included the data in 345 kilobytes (Kb) and 200 Kb, respectively) and The on asters had 4,884 records and was files. file a Kb literature file. These were necessary for chro- about 553 in size at the time of writing, and under the goldenrods had 2,099 records (including file synonyms. Each nomenclature includes ASCII non-North American reports) and was about 257 file text data on some basionyms, synonyms, types, phylo- Kb, In cases, corrections found in later pub- genetic were relationships, excluded taxa, and miscel- lications incorporated into the original data laneous comments. The base record. vouchers for the majority of the my reports not published by laboratory were not seen as part of this study. Therefore, the data base DATA BASE may have some due biases to misidentificatlons. t'ata on chromosome number were no vouchers are known some and reports ob- Also, for reports, tamed from the literature and nearly 600 unpub- the original identifications had to be accepted on lished reports from on com- or the reports rejected as unconfirmable. studies certain species faith piexes my being chromosome investigated by laboratory. Data For certain taxa, a corrected count ^ere taken from 121 publications (see references and a comment noting this adjustment were in- m & Semple 1980b and Semple eluded the data record. For example, prior Brouillet, et al., in file iyo4; & and members Morinaga Fukashima, 1931; Hig- 1978 reports for of Aster sect. to all & *^^tham, 1936; Smith, 1965; Johnson Packer, Dumosi subsect. Heterophylli were published re- & 1968; Packer, 1968; Mulligan Cody, 1971; an assumed chromosomal base number of fleeting = Kapoor, 1972; Mulligan 1972; Andreasen Jones (1978) demonstrated that the chro- et aL, 9, jc & Eshbaugh, number 1973; Witherspoon 1974; mosomal base for subsect. //f'/r/-o/>/zj//i et al., is = & & = Keil Stuessy, 1975; KeU Pinkava, 1976; x Therefore, older published counts of a 8. & Hartman, "2n=8II 1977; Kapoor, 1977; Keil Pinkava, 9 were entered as cf." in the data matrix. 98 Annals of the Garden Missouri Botanical n=9" No with "as appended to the publication data. count reports were included in the data files These corrections did not affect the summaries for goldenrods from Prince Edward Island. Only 30 presented in this paper, but could influence other reports for asters and six for goldenrods from manipulations of the matrix* Mexico were entered into the files. In other cases, an "unusual" chromosome num- Tentative estimates were made of the num- total down ber report for a taxa was entered as published, ber of taxa to varietal level for each province comment with a that further investigation re- and territory in Canada, each state in the United is By quired. "unusual" I mean, for example, the States, and for Mexico (Table These are pre- 1). one count out of 40 or more that was not in sented graphically for asters and goldenrods in agreement with the other counts for the taxon.' Figures 2 and 4, respectively. Overall, the number number of aster taxa and the of goldenrod taxa GRAPHICS AND STATISTICS for each area correlate rather highly 6.7), but ( the maiority of the reeions (86%) have more aster xMjiaps J .V were prepared usmg the graphi_i.cs program u n j t,han goldenrodi taxa (/tT-able i \ T-rhue diversi•t* y off 1). Coreldraw Version 1.02 (Corel Systems Corpo- lAnn r n t^^a for asters and goldenrods greatest in the ^^.- ^ r is 1600 a r\.. • tKalZr^ ration, Carling Ave., Ottawa, Ontario i ^ Mnt:c\ tt /^ eastern Umt- ed, . i Lronquis• t (ivoo) htates; previously, Windows run under The \'^^ goldenrods. great- "^^^ for ^°**'*^ '^^ *^ Version 2 higher "(Microsoft Inc^" Redmo'nd'; "or' ' ' numbers and est of species, subspecies, varieties Washing QMS for both asters and goldenrods occur in North Car- The ^^^ ^"^ estimated, respectively) few- ^ ""^^^ Pearson were ^ correlations calculated using SYS^ TAT est taxa occur in Alaska and the far northern areas (SYSTAT, Version 5.0 1800 Sherman Inc., across Canada. The western part of North America Ave., Evanston, 60201, Illinois U.S.A.). has far fewer goldenrod taxa than the eastern part. However, because the pattern for asters different Results and is Discussion several taxon-rich and subsections of asters sections A of 6,908 chromosome number total reports achieve their greatest diversity in the west, unlike and for asters goldenrods from Canada, the United any group of goldenrods. The number of taxa along and Mexico States, were included in the study the Pacific coast high primarily because of the is The (Table 1). results of the analyses of the data presence oi Aster Eucephalus and Aster secX. sect. matrix on chromosome number and num- Dumosi two reports Without these subsect. Foliacei. bers of taxa per political region are presented pic- groups, the general pattern of diversity in asters The torially in Figures 1-4. numbers total of re- would be quite similar to that in goldenrods. ports for each province and among territory in Canada, There was a great range (0% to 90%) each state in the United States, and Mexico all of geopolitical regions the percentage of taxa for in are given Table and in Figures and number 1 1 3 for Aster which one chromosome at least individual's sens. and Solidago, A The lat. respectively. total of had been determined (Table Figs. 2, 4). 1; 4,844 was reports entered askSam into the mean United data percentages of taxa from Canada, the on file asters. Ofthese, 1,520 were were for plants native and Mexico States, sampled cytologically The to Ontario. next^ largest numbers 49.1% The of reports for asters and 45.1% goldenrods. for Quebe were highest percentages were counted of taxa that and 144, 133 reports, respectively. There were for Manitoba and 90% and 75% in the Ontario: more than 100 reports each for CaUfornia, Colo- former, and 80% and 83% for aster in the latter Carolin were and goldenrod Percentages taxa, respectively. The number lowest lower Canada, in all other provinces and states. In States was from Delaware with only two reports, the numbers of taxa of counts and the percentages No count reports were included in the data counted were and Prince files lowest Newfoundland in from for asters Newfoundland and Labrador, and Edward 62 provinces, Island. Thirty-four out of was there only one report from Prince Edward more territories, states, and Mexico (54.8%) had Two Island. thousand sucty-four reports were on en- aster taxa counted but than goldenrod taxa, goldenrods irod 590 these, were for plants native The to Ontario. ounted per percentages area. Generally, the numbers of reports were from Quebec an clw Idenrod *^^'^ i^ arf*a oo,-^.-ii«.-j New York, California, and Colorado = with 144, 75, not example correlate strongly For 49) (r 73 and 58 No reports, respectively. other state five of the 23 Nevada estimated aster taxa for m the United States had more than 50 (22%) reports. taxa versus five of the seven goldenrod o CD ^ CD (D CD 3 {^ 1520 194 r\«^ > O CO 94 \ \^ - 0> CD ^ CD 132 '^V 3 26 ^ o CO co/> 1 Q. 3 Canada CD o /- 23 \ 33 U.S.A. 2822 CD 3 Mexi "^'^'^^'^ O [50 CD Q. 4844 cn Total DO \24L, 68 CD O Numbers Chromosome (0 of o Number Reports km 10CX} Figure 1. Numbers of chromosome number reports for Aster sens. lat. {Aster, Virgulaster, and Virgulus) in Canada, the United States, and Mexico. Only small portions of CD Alaska arid the Northwest Territories are indicated at the top of the map; Numbers CD i Quebec, and Mexico, respectively. and percentages of and numbers of taxa counted, Numbers chromosome number estimated numbers of taxa (species, subspecies, varieties), Table of reports, 1 . taxa counted for the provinces and territories of Canada, states of the United States, and all of Mexico. Goldenrods Asters Esti- — mated Goldenrods Number Asters Number of taxa of taxa %of %of Total total . % 1 Number Number taxa of number of number Number of Esti- taxa of Esti- counted mated Counted counted taxa mated Counted counted reports area reports of taxa reports Geopolitical CANADA 62 13 3 65 23 13 8 95 39 72 26 17 Alberta -5 60 19 55 27 10 6 95 39 68 29 16 Columbia British 40 20 15 75 15 87 40 47 20 18 90 Manitoba -4 74 12 23 8 35 39 New 47 42 35 19 14 Brunswick -8 -14 14 9 14 2 9 20 6 Newfoundland -8 14 Labrador* -10 50 40 8 4 2 14 9 6 5 2 1 Northwest Territories -9 52 23 78 33 53 12 9 75 25 21 11 Nova Scotia 49 39 80 590 41 34 83 8 3 2,110 90 1,520 Ontario 8 10 28 10 10 18 Edward 1 1 Prince Is. 1 63 144 32 24 75 8 12 288 56 144 24 15 Quebec > 38 6 48 31 33 18 8 44 15 13 5 5 Saskatchewan 29 4 20 C/) 9 38 7 2 18 13 11 Yukon 51 1 Territories O 05 CO 11.7 50.4 (931) 18 9.8 47.8 2.7 2.7 Means (subtotals) (2,923) 35.5 (1.992) 19.1 O CD U.S.A. O 63 75 49 42 21 50 15 33 12 36 9 14 CD Alabama 43 32 47 15 16 8 6 75 31 7 3 1 o' Alaska 44 24 19 14 8 57 25 10 7 70 4 13 Arizona 45 63 66 33 33 15 45 30 33 15 Arkansas 0) -6 a 70 23 64 73 10 205 43 132 33 21 7 California CD 67 22 164 47 106 29 13 45 58 18 12 11 Colorado 26 51 18 27 12 44 8 24 6 25 3 19 Connecticut -5 Delaware 8 45 2 23 3 13 6 22 4 18 1 56 132 33 25 76 35 23 15 65 10 11 167 Florida 79 89 63 45 24 53 16 44 13 30 23 Georgia 1 -5 93 39 79 29 13 45 14 10 5 50 19 Idaho 109 69 94 46 29 63 15 23 7 30 23 33 Uliuiois 20 57 14 27 10 37 3 67 57 53 30 17 Indiana 56 9 19 9 47 8 9 46 42 27 15 51 Iowa -. ^' o CO Tari K Continued 1 CD Goldenrods Asters Esti- <0 mated Number of taxa ~ Number of taxa Asters Goldenrods % , %of of Total total % Number Number number of number Number of Esti- taxa of Esti- taxa of 3 Geopolitical area reports of taxa reports mated Counted counted reports mated Counted counted taxa counted CD Kansas 45 40 26 23 10 43 19 17 12 71 6 28 Kentucky 74 73 62 40 18 45 12 33 11 33 7 12 30 69 24 37 13 35 6 32 5 16 5 19 Louisiana Maine 57 49 42 26 14 54 15 23 11 48 3 6 Maryland 24 60 14 30 8 27 10 30 8 27 -5 82 59 44 34 20 59 38 25 16 64 9 Massachusetts Michigan 174 59 133 31 22 71 41 28 17 61 3 10 Minnesota 114 52 88 30 21 70 26 22 10 45 8 25 55 64 50 35 15 43 5 29 6 21 6 22 Missi>,sij)pi Mi 117 61 93 31 21 68 24 30 12 40 28 a^ouri 1 Montana 79 41 63 28 16 57 16 13 7 54 15 3 Nehraska 37 34 21 21 11 52 16 13 6 46 8 6 Nevada 39 30 32 23 5 22 7 7 5 71 16 49 > O W -4 New Hampshire 56 52 31 28 14 50 25 24 13 54 4 CD C/> New Jersey 46 73 19 43 11 26 27 30 16 53 13 27 CD ^ 3 New -8 ® Mexico 62 40 23 22 13 59 39 18 12 67 4 o New York 181 80 106 51 22 43 75 31 20 65 20 22 a o 3 North Carolina 171 107 124 56 30 54 47 51 27 53 5 1 CD o -2 North Dakota 51 29 32 16 7 44 19 13 6 46 3 Q. Ohio 49 49 35 27 11 41 12 22 3 14 5 27 CD 3 Oklahoma 38 55 23 28 17 61 15 27 13 48 13 1 O CD Oregon 209 46 194 33 20 Q. 61 15 13 6 46 20 15 Pennsvh'ania 103 62 84 39 16 41 19 23 12 52 16 33 11 CD Rhode Island 11 46 6 25 5 20 5 21 5 24 4 -4 "a o South Carolina 99 84 74 44 24 55 25 40 15 38 4 17 South Dakota 35 26 18 14 7 50 17 17 12 8 67 2 o Te niie-- »<> 92 78 42 40 18 45 50 38 22 58 2 13 Texas 83 67 68 29 20 69 15 38 12 32 -9 37 69 !'tah 35 52 24 9 38 17 17 11 6 55 13 Vermont 80 51 47 29 17 59 33 22 12 55 7 4 Virginia 154 80 127 38 27 -4 71 27 42 14 33 38 H^ f ^ ^ ^ r \- .^.^^U PO Table Continued. 1. Goldenrods Asters Esti. — mated Number Goldenrods Number taxa of taxa Asters of %of %of Total total . % 1 Number number number Number of taxa Number of Esti- taxa of of Esti- mated Counted counted reports mated Counted counted taxa counted Geopolitical area reports of taxa reports -11 42 22 48 46 42 34 10 29 6 12 5 Washington 35 10 32 6 19 2 16 42 66 32 34 12 West Virginia 40 27 68 27 28 14 50 12 18 68 86 116 Wisconsin > -15 2 58 16 Wyoming 76 54 58 35 15 43 18 19 11 Means 55.3 (2,822) 31.4 15.7 50.0 (1,127) 23.9 10.4 46.1 7.4 3.9 to (3,949) (subtotals) & o Canada U.S.A. 22.8 10.3 46.4 6.2 3.7 03 29.8 15.9 50.1 (2,058) Means 51.1 (4,818) (6,872) O (subtotals) CD MEXICO o Mexico 36 41 30 14 5 36 6 27 3 11 13 25 O 2,064 Totals 6,908 4,844 03 Means 111.4 50.9 78.1 28.7 14.7 49.9 33.3 22.8 10.1 45.9 5.9 4.0 <1) * Labrador part of the Province of Newfoundland; data on the island and the mainland portions of the province are listed separately for clarity. is 3 > O CO zr CD (/) 3 3 CD 8|d rs lOSOLUi and O CD o ^ Q. CD 3 rod CD a> 3J CD "O o a. CO o Figure Estimated numbers of taxa (species, subspecies, and varieties; shading) in Aster sens. lat. {Aster, Virgulaster, and Virgulus) and percentages of taxa for which a 2. romosome number has been determined (numbers the regions indicated). Numbers indicated are for both subregions of Ontario, Quebec, and Mexico, respectively. in 2 > s o CD O CD 0) o 0) Q. CD Figure 3. Numbers of cliromosome number reports for Solidago (excluding Euthamia) in Canada, the United States^ and Mexico. Only small portions of Alaska and the NoTlhwesl Territories are indicated; Yukon Territory not included in the map. Numbers indicated are for both subreglons of Ontario. Quebec, and Mexico, respectively. is ^