1979

Chromosome numbers in Boykinia and allies (Saxifragaceae) RICHARD J. GORNALL Deparftnetzf of'Bofatzy, Leicesfer Urziversity, United Kitzgdorn, LEI 7RH

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BRUCE A. B O H M ' Departmetzt of Botany, Utziversity of Britislz Cohttnbin, Vancouver-, B.C., Catzncln V6T lW5 AND

ROYL. TAYLOR Botarzical Gnrdetz, Utziversioi of Brifish Colurnbin, Vcrncouver, B.C., Catzacln V6T I W5 Received August 29, 1982

GORNALL, R . J . , B . A. BOHM,and R. L . TAYLOR. 1983. Chromosome numbers in Boykitzin and allies (Saxifragaceae). Can. J . Bot. 61: 1979-1982. Chromosome numbers are reported for the first time for Boykirrin itzfertnedin (212 = 14), B. tnnjor (212= 28), Peltoboykirzin wntni1nbei (211 = 22), and Sirksdorfin (=Hetnieva) rnt~itt~citlifolin(211 = 14). Additional counts are provided for eight other species of Boykitzia (including Telesotzi.~),Pelfoboj~kitzin,Sitksdor-jn, and Bolarzdrn. Previous literature reports are reviewed and some errors pointed out. Chromosome numbers do little to resolve the problem of generic limits, with the exception of supporting the segregation of Peltoboykinin. They do, however, support the recognition of Boykinin intermedia as a species distinct from B. tnnjor.

GORNALL, R. J . , B. A. BOHMet R. L. TAYLOR. 1983. Chromosome numbers in Boykitzia and allies (Saxifragaceae). Can. J. Bot. 61: 1979-1982. Les auteurs prksentent les premitres determinations du nombre chromosomique de Boykitzin intertnedin (2tz = 14), B. ttznjor (211 = 28), Peltoboykitzin wntntznbei (212 = 22) et Suksdorfin (= Hetnievn) ranutzculifolin (212 = 14). Des dknombrements additionnels sont prCsentCs pour huit autres esptces de Boykinin (incluant Telesotzix), Peltoboykitzin, Suksclorfin et Bolnrzdrn. Les donnees publiees anterieurement sont cornmentees et quelques erreurs sont signalees. Les nombres chromosomiques n'aident pas beaucoup a resoudre le probltme de limites gkneriques, sauf qu'ils appuient la separation du genre Peltoboyki~zin. Cependant, ils contribuent 5 appuyer I'acceptation de Boykinin itztertnedin comme esptce distincte de B. major. [Traduit par le journal]

Introduction Boykinia and its relatives belong to the herbaceous subtribe Saxifraginae of the family Saxifragaceae. As treated by Engler (1928), Boykinia contains nine species, but generic limits are somewhat controversial and evidence from all sources is needed to resolve the problems (Spongberg 1972). The relatives included in this study are the segregate genera Peltoboykinia and Telesonix and the closely related genera Bolandra, Hemieva, and Suksdorja, the latter two often united under Suksdorja. Another closely related genus, Sullivantia, has already been studied by Soltis (1980). Apart from the two Peltoboykinia species, which are native to Japan, all other species studied are North American, with nine in the west and one, Boykinia aconitifolia, in the east. This study presents cytological data for 12 of the species; counts for four of them, Boykinia intermedia, B. major, Peltoboykinia watanabei, and Suksdorja ranunculifolia (= Hemieva), are reported for the first time. Method Both meiotic and mitotic divisions were observed. For 'Author to whom all correspondence should be addressed.

meiosis, buds were fixed in 3:l absolute alcohol : glacial acetic acid for at least 24 h. Pollen mother cells were squashed in 1% acetocarmine (Darlington and La Cour 1976). Mitosis was observed in root tips. Initially the technique used involved an 8-h pretreatment with a-monobromonaphthalene at 5OC followed by hydrolysis for 10 min in 1 N HC1 at 60°C, staining with Feulgen's solution for 3 h at 20°C, and finally squashing in 45% acetic acid. Later, however, far better preparations were obtained by pretreating with 8-hydroxyquinoline for 3 h, followed by hydrolysis as before, but then staining and squashing in aceto-orcein.

Results A summary of the chromosome numbers of the species examined in this study is given in Table 1; full details of the collection localities are also provided.

Discussion Previous counts reported in the literature There is some confusion surrounding the counts made by Hamel (1953) for Boykinia rotundifolia and B. occidentalis (= elata). His drawings and text assigned 2n = 14 to B. rotundifolia and 2n = 26 to B. occidentalis, while in his summary table these numbers were transposed. Subsequent works quoting counts for Boy-

CAN. J. BOT. VOL. 61. 1983

TABLE1. Chromosome numbers and collection data for the species examined in this study. Voucher specimens are deposited at DAO or UBC as indicated. Alternative generic placements for certain of the species are shown in parentheses

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Pelroboykinia tellitnoicies (Maxim.) Hara 2r1 = 22 (Boykrnin) Ex J. W. Goethe University Botanic Garden, Frankfurt, Gornall 343 (UBC), 2r1 = 22; ex Munich Botanic Garden, Gomall 400 (UBC), 2r1 = 22 Peltoboykinia wararlabei (Yatabe) Hara 2t1 = 22 (Boykirlia) Ex J. W. Goethe University Botanic Garden, Frankfurt, Gornall 328 (UBC), 2n = 22 Boykirlia aconitijolia Nutt. 2r1 = 14 North Carolina, Macon Co., Chatooga River, Gornall 342 (UBC), 2r1 = 14; ex J. W. Goethe University Botanic Garden, Frankfurt, Gomall 404 (UBC), 2r1= 14; ex F. Schiller University Botanic Garden, Jena, Gomall 401 (UBC), 2r1 = 14; ex Karlsruhe University Botanic Garden, Gomall 402 (UBC), 211 = 14; ex St. Andrews University Botanic Garden, Gornall403 (UBC), 2r1 = 14 Boykinia inrermedia (Piper) G. N. Jones 2r1 = 14 Washington, Grays Harbor Co., Humptulips, Gornall 23 (UBC), 211 = 14 Boykinia major A. Gray 2n = 28 Idaho, Idaho Co., Rackliff Creek by Selway River, Gornall 335, 337 (UBC), rl = 14; Montana, Missoula Co., Lolo Pass, Gornall 202 (UBC), 211 = 28; Oregon, Douglas Co., upper reaches of Rogue River, Gomall 38 (UBC), 2r1 = 28; Oregon, Jackson Co., Rogue River, Gomall 45 (UBC), 2t1 = 28; Califomia, Eldorado Co., Kyburz, Gornall 92 (UBC), 2t1 = 28 Boykinia occidenralis Torr. & Gray 2r1 = 14 British Columbia, North Vancouver, Capilano Canyon, Gornall 1 (UBC), t~ = 7; Washington, King Co., Snoqualmie Falls, Gomall 216 (UBC), rz = 7; Oregon, Clatsop Co., Saddle Mt., Bohm 1293 (UBC), t~ = 7; Oregon, Tillamook Co., Glenwood, Taylor & Gilbert 4658 (DAO), 2t1 = 14; Oregon, Douglas Co., Diamond Creek, Taylor & Gillett 4669 (DAO), 2t1 = 14; California, Del Norte Co., Gasquet, Straley 1753 (UBC), n = 7; Califomia, Siskiyou Co., Happy Camp, Gomall 65 (UBC), n = 7; Califomia, Siskiyou Co., 46 miles SW of Happy Camp, Taylor, Baker et al. 4449 (DAO), 2t1= 14; ex Rancho Santa Ana Botanical Garden, Taylor 64C-17 (DAO), 2rz = 14; ex Copenhagen Botanic Garden, Taylor 63C-176 (DAO), 2n = 14 Boykinia rorundijolia Parry 2t1 = 14 California, Los Angeles Co., Crystal Lake, Gornall98 (UBC), 2r1 = 14; California, Los Angeles Co., Hwy. 39 bridge over N Fork San Gabriel River, Gomall 101 (UBC), 2t1 = 14; Califomia, San Bernadino Co., Icehouse Canyon, Gornall 105 (UBC), 2n = 14; ex Uppsala University Botanic Garden, Taylor 63C-175 (DAO), 2n = 14 Boykinia heucherijormis (Rydb.) Rosendahl 2n = 14 (Te1esoni.r) Alberta, Jasper National Park, Miette Hot Springs, Calder & Spicer 3381 1 (DAO), 2n = 14; Montana, ParkCo., PineCreekLake, Gomall 193 (UBC), n = 7; Utah, Cache Co., Mt. Magog, Gomall 132 (UBC), n = 7; Wyoming, Fremont Co., 1.2 miles NE of Shoshone National Forest Boundary on Hwy. 28, 'Taylor & Gillett 4726 (DAO), 2n = 14; Nevada, Clark Co., Charleston Peak, Gomall 1 15 (UBC), n=7 Boykinia jamesii (Torr.) Engl. 2n = 14 Colorado, Pike's Peak, Gornall405 (UBC), 2t1 = 14

(Telesorlix)

Bolandra oregana S. Watson 2n = 14 Washington, Asotin Co., Buford Creek Canyon, Taylor & Gillett 4625 (DAO), 2n = 14; Washington, Klickitat Co., Klickitat, Taylor & Gillett 4640 (DAO), n =7

GORNALL ET AL

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Suksdotjia rnt~r~nculifolin (Hook.) Engl. 2n = 14

(Hettlievn. Boykit~in)

British Columbia, Lower Arrow Lake, 6 miles N of Fauquier, Gornall 332 (UBC). n = 7; British Columbia, 19 miles W of Sheep Lake, near Rossland, Gornall33 1 (UBC), t~ = 7; British Columbia, Fraser River Canyon. Saddle Rock near Yale, Bohm 1132 (UBC), tl = 7; British Columbia, Manning Provincial Park, Blackwall Mt., Bohm 1 144 (UBC), t~ = 7

(Hettlievn) = 14 British Columbia, Kootenay Lake, Crawford Bay, Bohm 1 1 12 (UBC), t~ = 7; British Columbia, Kootenay River, Coma Linn Dam, Taylor & Sherk 4878 (DAO), 2n = 14; British Columbia, 22 miles W of Sheep Lake, near Rossland, Gornall 329 (UBC), tl = 7; British Columbia, 19 miles W of Sheep Lake, near Rossland, Gomall 330 (UBC), t1 = 7; British Columbia, Yahk, Calder & Spicer 32855 (DAO). 2t1 = 14; Montana, Flathead Co., Flathead River, Bad Rock Canyon, Taylor & Gillett 4605 (DAO), 2n = 14, n = 7

Suksdorja violncen A. Gray 2n

kinia have apparently used the summary table as their source, e.g., Munz (1959) and Hitchcock et 01. (1961). It is fortunate, if ironic, that B . occicletltcl1i.s does in fact have 2n = 14 in all collections counted: indeed, this count appears in a later report by Hamel (1958). Boykitzia rotutldifolia, however, does not have 211 = 26 as in Hamel's (1953) table but rather 211 = 14 as in his text and illustration. A further error made by Hamel (1953) is the count of 211 = 12 for Boykitlia acotlitifolicz. Checks on all collections revealed that 2t1 = 14, confirming the count made by Soltis (1980). Finally, Spongberg's 1972 report of 211 = 28 for Bojkitzia jamesii is an erroneous citation of Hamel's (1953) count of 2n = 14 (S. A. Spongberg, personal communication). The latter count was confirmed in the present study. Regarding the other previously counted species, the results found here agree with the published data, viz. 2n = 14 for Boykinia heucheriformis (Packer 1968); 2n = 22 for Peltoboyki~liatellimoides (Skovsted 1934; Hamel 1948, 1953); 211 = 14 for Suksdor-a violacea (Packer 1968); and 2n = 14 for Bolandra oregana (Hamel 1958; Ornduff 1962). The remaining species for which counts are available is Boykinia richardsonii. It is a native of Alaska and the Yukon. Plants from the arctic slope of the Brooks Range have been counted as 2n = 84 (Johnson and Packer 1968), and as 211 = ca. 84 (Packer and McPherson 1974). However, plants from the Alaska Range have been counted as 2n = 36 (Knaben 1968). Until confirmed the deviant count of 2n = 36 (Knaben 1968) for B. richardsonii should be regarded as suspect. Species in the group under consideration for which counts have yet to be reported are Boykinia lycoctonifolia (from Japan), Hieronymusia alchemilloides (from Bolivia and Argentina), and Bolandra califortlica (from California).

Taxonomic implicatiotls The base number x = 7 is very common in the subtribe Saxifraginae and is probably ancestral (Schoennagel 193 1; Skovsted 1934; Hamel 1953). The fact that almost all the species studied here share this number sheds little light on the problem of generic limits. The closeness of the relationship between Boykitzia and Sullivantia even extends to an identical karyotype (Soltis 1980). However, the base number of x = 11 in both species of Peltoboykinia, together with a distinctive morphology, including peltate leaves, supports their separation from Boykitlia as a distinct genus (Hara 1937). The origin of this number is obscure, and relationships may possibly lie in the direction of Saxifraga where some sections, e.g., Miscopetalum, Triclactylites, Neplzrophyllum, Dacryloides, and Diptera, have x = 11 as a common base number (Hamel 1953). Like Peltoboykinia, many of these sections also have uniseriate, glandular trichomes. Other peltate-leaved genera, Peltiphyllum and Astilboides, have 2n = 34 (Hamel 1948, 1953). At the species level and below, it has previously been customary to treat Boykinia intermedia as a variety of B. major (e.g., Hitchcock et al. 1961). However, the difference in chromosome number (Table I ) , together with the differences in morphology (Hitchcock et al. 1961) and in flavonoid chemistry (Gornall and Bohm 1980), strongly support the specific distinction originally recognised by Jones (1936). DARLINGTON, C. D., and L. F. LACOUR.1976. The handling of chromosomes. 6th ed. Allen & Unwin, London.

ENGLER, A. 1928. Saxifragaceae. In Natiirlichen Pflanzenefamilien. 2nd ed. Vol. 180. Edirerl by A. Engler and K. Prantl. W. Engelmann, Leipzig. pp. 74-226.

GORNALL, R. J., and B. A. BOHM. 1980. The use of flavonoids

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CAN. J. DOT. VOL. 61, 1983

G. 1968. Chromosome numbers of flowering plants in the taxonomy of Boykinia and allies (Saxifragaceae). KNABEN, from Central Alaska. Nytt Mag. Bot. (Oslo), 15: 240-254. Can. J. Bot. 58: 1768-1779. HAMEL,J . L. 1948. Notes preliminaires a une etude caryo- MUNZ,P. A. 1959. A California flora. University of California Press, Berkeley. logigue des Saxifragacees. 1. Les chromosomes de Pelriphyllurn pelrarurn (Torr.) Engl. et de Boykinia relli~~loides ORNDUFF,R. 1962. Documented chromosome numbers of plants. Madrono, 16: 266-268. (Maxim.) Engl. Bull. Mus. Hist. Nat. (Paris), 20: 198-200. J. G. 1968. I.O.P.B. chromosome number reports 1953. Contribution a l'etude cyto-taxinomique des PACKER, XVII. Taxon, 17: 287. Saxifragacees. Rev. Cytol. Biol. Veg. 14: 1 13-3 13. J. G., and G. D. MCPHERSON. 1974. Chromosome 1958. Materiaux pour l'etude caryo-taxonomique des PACKER, numbers in some vascular plants from Northern Alaska. Saxifragacees. 5. Le noyau et les chromosomes samatiques Can. J. Bot. 52: 1095-1099. du Bolandra oregana S. Wats. Bull. Mus. Hist. Nat. SCHOENNAGEL, E. 193 1. Chromosomenzahl und Phylogenie (Paris), 30: 522-524. der Saxifragaceen. Bot. Jahrb. Syst. Pflanzengesch. PflanHARA,H. 1937. Two new genera of Saxifragaceae in Japan. Bot. Mag. 51: 250-253. zengeogr. 64: 266-308. A. 1934. Cytological studies in the tribe SaxiHITCHCOCK, C. L., A. CRONQUIST, M. OWNBEY, and J. W. SKOVSTED, THOMPSON. 1961. Vascular plants of the Pacific Northwest. frageae. Dan. Bot. Ark. 8: 1-52. 3. University of Washington Press, Seattle. SOLTIS,D. E. 1980. Karyotypic relationships among species JOHNSON, A. W., and J . G. PACKER.1968. Chromosome of Boykinia, Heuchera, Mirella, Sullivanria, Tiarella, and Tolmiea (Saxifragaceae). Syst. Bot. 5: 17-29. numbers in the flora of Ogotoruk Creek, N.W. Alaska. Bot. SPONGBERG, S. A. 1972. The genera of Saxifragaceae in the Not. 121: 403-456. southeastern United States. J . Arnold Arbor. Harv. Univ. JONES,G. N. 1936. A botanical survey of the Olympic Peninsula, Washington. Univ. Publ. Biol. 5: 1-286. 53: 409-498.

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