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Chromosome Numbers in Some Cacti of Western North America-V Author(S): Donald J

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Chromosome Numbers in Some Cacti of Western North America-V Author(S): Donald J Chromosome Numbers in Some Cacti of Western North America-V Author(s): Donald J. Pinkava, Marc A. Baker, Bruce D. Parfitt, Mark W. Mohlenbrock and Richard D. Worthington Source: Systematic Botany, Vol. 10, No. 4 (Oct. - Dec., 1985), pp. 471-483 Published by: American Society of Plant Taxonomists Stable URL: http://www.jstor.org/stable/2419140 Accessed: 10-11-2015 17:03 UTC Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://www.jstor.org/page/ info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. American Society of Plant Taxonomists is collaborating with JSTOR to digitize, preserve and extend access to Systematic Botany. http://www.jstor.org This content downloaded from 156.40.216.1 on Tue, 10 Nov 2015 17:03:23 UTC All use subject to JSTOR Terms and Conditions SystematicBotany (1985), 10(4): pp. 471-483 K Copyright1985 by the American Society of Plant Taxonomists Chromosome Numbers in some Cacti of Western North America- V DONALD J. PINKAVA, MARC A. BAKER, BRUCE D. PARFITT, and MARK W. MOHLENBROCK Department of Botany and Microbiology,Arizona State University, Tempe, Arizona 85287 RICHARD D. WORTHINGTON Department of Biological Sciences, Universityof Texas, El Paso, Texas 79968 ABSTRACT. Documentedmeiotic and mitoticchromosome numbers and behaviorare reported for62 taxarepresenting 11 generaof Cactaceae from the southwestern United States and northern Mexico.Chromosome numbers for 12 speciesand six naturalinterspecific hybrids are firstcounts. New numberswere determinedfor four additional species. Chromosome counts of diploid and polyploidtaxa, including triploid and pentaploidhybrids, are all consistentwith the base number, x = 11. Translocationsare reportedfor the first time in theCactaceae. The Cactaceae comprise a large and complex stainability was based on 500-grain samples assemblage of New World succulent perenni- stained in aniline blue in lactophenol (Mane- als. Much of the complexity,particularly in the val 1936). The mitoticcount (Pereskiopsisporteri) Opuntioideae, arises from hybridization and was obtained fromroot tips fixed,stained, and polyploidy. The base number of the Cactaceae mounted according to the method of Parfitt is x = 11 and probably paleotetraploid (Lewis (1979). Nomenclature follows that of Benson 1980). Lewis (1980) reports only 14% of the (1982), Britton and Rose (1919-1923), Bravo- known species of cacti are secondary poly- Hollis (1978), Gibson and Horak (1978), Hunt ploids. Sato (1958) noted in his list of chro- (1967), and Pilbeam (1981). mosome counts for214 species and six varieties of cacti that 17.7% of the taxa were polyploid. RESULTS In our five-partseries on chromosomenumbers Chromosome numbers were determined for and behavior in westernNorth American cacti, 183 individual cacti representing62 taxa in 11 we have determinedthat polyploidy occurs in genera (table 1). First counts are reported for 37 of 68 taxa of Opuntioideae (52.4%) and in 14 12 species and six natural,interspecific hybrids; of 72 taxa of Cactoideae (19.4%), including hy- new counts are determinedfor fouradditional brids and taxa representedby both diploid and species. polyploid individuals. No polyploidy is known Not previously reported in our five-partse- for the most primitive subfamily, Peres- ries are counts for 33 taxa (figs. 1-36). At vari- kioideae. This reporton chromosomenumbers, ance with these counts are a hexaploid count hybridization,and polyploidy is an attemptto for Opuntiaschottii (Yuasa et al. 1973) and dip- clarify taxonomic and evolutionary relation- loid counts for 0. fulgidavar. fulgida(Pinkava ships among the Cactaceae. et al. 1973), 0. ramosissima(Pinkava et al. 1973), and 0. stanlyi(Yuasa et al. 1973). In this five- MATERIALSAND METHODS part series as a whole, chromosome numbers Flower buds were collected in develop- have been determined for 438 individuals of mental series from plants growing in native 149 taxa in 98 species in 21 genera (sensu Hunt habitatsor in cultivation.Buds were killed and 1967) of cacti. fixed for 24 hours in chloroform,95% ethanol, and glacial acetic acid (6:3:1), transferredto 70% DISCUSSION ethanol, and refrigerated. Anthers were One specimen (Ruffner27) of Opuntia lepto- squashed in acetocarmine and mounted in caulis was found to possess two translocations Hoyer's medium (Beeks 1955). Percent pollen (fig. 10) coupled with only 35.4% pollen stain- 471 This content downloaded from 156.40.216.1 on Tue, 10 Nov 2015 17:03:23 UTC All use subject to JSTOR Terms and Conditions 472 SYSTEMATIC BOTANY [Volume 10 ability.This is the firstreport of translocations elongate lowermosttubercles (as in 0. acantho- in the Cactaceae. Opuntia leptocaulisis a wide- carpa), but are somewhat fleshy (as in 0. spi- spread and variable taxon, thus far appearing nosior). as a tetraploid in the Chihuahuan Desert Grant and Grant (1971b) described the hy- (Fischer 1962; Conde 1975; Pinkava et al. 1977; brid, 0. spinosiorx 0. versicolor,and found per- Weedin and Powell 1978) and as a diploid in centages of morphologically good pollen for the Sonoran Desert (Pinkava et al. 1977; table four individuals froma population at Hough- 1). It is believed to hybridizewith otherspecies, ton Road (east of Tucson) to be 42%, 57%, 91%, formingtriploid hybrids with 0. kleiniaevar. and 97%. We found percentagesof pollen stain- kleiniae(Fischer 1962; Pinkava and Parfitt1982) ability for seven hybrids from a population and formingdiploid hybrids with 0. acantho- about 30 km southeast of Florence (north- carpa (table 1, fig. 2), 0. spinosior(table 1, fig. northeastof Tucson) ranged from21.0 to 90.9% 11), and 0. whippleivar. whipplei(Trushell pers. (x = 50.7%). comm.). Opuntia bigeloviivar. bigelovii,which repro- The hybrid,Opuntia acanthocarpa x 0. lepto- duces primarily from vegetative propagules caulis, is intermediate between the putative (easily detached stem segments),appears to be parents in stem diameter and branching pat- mostlya triploid(table 1; Pinkava and McLeod tern, perianth segment length and color (in 1971; Pinkava and Parfitt1982). Percentages of ours, yellow vs. yellow-orange or yellow- pollen stainabilityare low, ranging from1.0 to green), fruitsize and color (green vs. tan or 17.2% (- = 9.1% based on seven triploid indi- red), and seed size. The hybrid resembles 0. viduals) and 0.9 to 46.1% (x = 14.0% on 11 pre- acanthocarpain having slightlytuberculate and sumed triploid individuals from Maricopa wedge-shaped fruitsoften bearing 1-3 persis- County,Arizona). Surprisingly,two diploid in- tent spines and resembles 0. leptocaulisin hav- dividuals morphologically indistinguishable ing only 1-4 spines per stem areole and some- fromtriploid individuals were found growing what fleshyfruits that occasionally proliferate. togetherin Pinal County (table 1, fig. 6), each Percentage of pollen stainabilityis low (table with higher seed set (germinabilityunknown) 1). than the nearly sterile triploids but with low The hybrid,Opuntia leptocaulis x 0. spinosior, percent pollen stainability (12.4% for Baker is intermediatebetween its putative parents in 4586A). Because these diploids may be second- overall size, stem diameter,number of spines arily derived, furtherinvestigation is planned. (3-4) per stem areole, perianth segment length Opuntiafulgida var. fulgidais now known in and color (in ours, faded purple vs. yellow- apparentlyindistinguishable diploid and trip- green or brightpurple), and fruitsize and color loid forms(table 1, fig.8). We hypothesize that (yellow with red blush at base vs. red or yel- these are autotriploidsthat arose via the union low). It resembles 0. spinosiorin having an ir- of reduced and unreduced gametes. This is fur- regularlywhorled branchingpattern and mod- ther supported by the role of 0. fulgidain the eratelytuberculate fruits. origin of 0. kelvinensis.The common mode of Opuntia spinosioris another cholla that hy- polyploidy is throughthe formationand sexual bridizes with other diploid taxa: 0. acanthocar- functioning of cytologically unreduced ga- pa var. major(table 1, fig.3; Benson 1969, 1982; metes (deWet 1980). Pinkava and Parfitt1982), 0. leptocaulis(table Opuntia kelvinensis,as it is now named, has 1, fig.11), and 0. versicolor(table 1, fig.17; Grant long been considered a hybridbetween 0. ful- and Grant 1971b). gida and 0. spinosior(Peebles 1936; Benson 1940, The hybrid, Opuntia acanthocarpavar. ma- 1950, 1969, 1982; Grant and Grant 1971a). The jor x 0. spinosior,is intermediatebetween its predominantform (which includes the type of putative parents in branching patterns,width 0. kelvinensis),as we interpretit, is a triploid and length of stem tubercles,amount of uni- (table 1, fig. 9). Opuntia spinosioris a sexual formityof spine lengths in an areole, and peri- species characterizedby normal Polygonum-type anth color (brickred vs. yellow-orangeor bright gametogenesis and embryogenesis (Baker and purple). The fruits often bear 1-3 persistent Pinkava 1983), high percentpollen stainability spines and are somewhat wedge-shaped with and seed set, non-proliferatingfruits, and rig- This content downloaded from 156.40.216.1 on Tue, 10 Nov 2015 17:03:23 UTC All use subject to JSTOR Terms and Conditions 1985]
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