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A Revision of Malvaviscus (Malvaceae) Author(S): Billie L. Turner and Meghan G. Mendenhall Source: Annals of the Missouri Botanical Garden, Vol

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A Revision of Malvaviscus (Malvaceae) Author(S): Billie L. Turner and Meghan G. Mendenhall Source: Annals of the Missouri Botanical Garden, Vol A Revision of Malvaviscus (Malvaceae) Author(s): Billie L. Turner and Meghan G. Mendenhall Source: Annals of the Missouri Botanical Garden, Vol. 80, No. 2 (1993), pp. 439-457 Published by: Missouri Botanical Garden Press Stable URL: http://www.jstor.org/stable/2399792 Accessed: 18-06-2015 17:01 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]. Missouri Botanical Garden Press is collaborating with JSTOR to digitize, preserve and extend access to Annals of the Missouri Botanical Garden. http://www.jstor.org This content downloaded from 128.83.205.78 on Thu, 18 Jun 2015 17:01:01 UTC All use subject to JSTOR Terms and Conditions A REVISION OF Billie L. Turner2and MALVAVISCUS (MALVACEAE)l MeghanG. Mendenhall3 ABSTRACT Comprehensivereevaluation of herbariummaterial of the genus Malvaviscus yieldstwo widespreadspecies, M. arboreusof NorthAmerica and M. concinnusof South America,two localizedspecies, M. achanioides of Mexico and M. williamsiiof Peru and Colombia,and a widespreadcultivar of unknownorigin, M. pendulifiorus. In spite of recent attemptsto delimitand classify de Candolle divided Malvaviscus into two sections: its many specific and subspecifictaxa, Malvaviscus Achania and Anotea. Anotea was raised to generic (Malvaceae) remains enigmatic. The genus is highly rank by Kunth in 1846, and Achania is now variable morphologically and given to populational recognized as synonymous with Malvaviscus. forms. Furthermore, these forms intergrade pro- Generic treatmentshave not been in agreement. ducing an array of character combinations. An Schery's monograph (1942) included a complex of extreme taxonomic treatment of the genus would 11 varieties of M. arboreus as well as two additional recognize a horde of intergrading forms. Indeed, species that have since been transferred to Pa- this extreme infrageneric variation has led to the vonia. Other treatments, which have recognized proposal of over 50 specific names within Mal- up to a dozen species, occur in floras throughout vaviscus, though Schery (1942) recognized only the range of Malvaviscus (Robyns, 1966; Stand- three in his monograph of the genus. ley, 1923; Standley & Steyermark, 1949). How- Because of the common intergradation and re- ever, the only significanttreatment of the genus combination of character states, few characters since Schery has been the exemplary workof Fryxell withinMalvaviscus have proven useful for specific (1988) in his monograph of the Malvaceae of Mex- recognition. Modern interpretations of the genus ico. In his discussion of the taxonomic problems have generally recognized about a dozen overlap- withinMalvaviscus, Fryxell acknowledged the dif- ping taxa. However, we believe that the over- ficulty of clearly delimiting species: "There is a whelming majority of the species proposed under certain sameness of morphology that runs through Malvaviscus represent only two biologically sig- the genus, and clear-cut differentiatingcharacters nificant taxa, M. arboreus in North America and are lacking." Fryxell recognized six species of Mal- M. concinnus in South America. vaviscus in Mexico, three of which are maintained here. HISTORY OF THE GENUS CHROMOSOME NUMBERS The genus Malvaviscus was established almost 200 years priorto its monograph by Schery (1942). The first report of a chromosome count for Among the first generic names to include Mal- Malvaviscus was presented by Skovsted in 1935 vaviscus were Alcea, Althaea, and Malva. Lin- (Table 1). He reported a count of 2n = ca. 84 for naeus did not recognize the genus and placed its an unidentifiedcultivated species in Kew Gardens, species in Hibiscus in 1753. In 1759, Fabricius presumably obtained originally "from gardens on distinguished Malvaviscus based on a single spe- the Gold Coast." We take the species to be M. cies, i.e., Hibiscus malvaviscus L. (A discussion penduliflorus of the present treatment since of the typificationof Malvaviscus can be found in Skovsted noted: Taxon 17: 87, 1968.) In 1788, Swartz proposed Morphologically,the type under observation is distinctly to rename the genus as Achania. In 1824, A. P. differentfrom the wildgrowing M. arboreus Cav. Its 1 We are gratefulfor the encouragement and helpfulcomments of our esteemedcolleague, Paul Fryxell,throughout the courseof this study. Our investigationis based uponthe examinationof approximately3,200 specimenson deposit at the followinginstitutions: BRG (3), F (701), herb. Fryxell(2), GH (431), HUA (7), LL (200), MG (6), MO (783), NY (492), RB (6), SP (9), TEX (170), US (381). Nancy Webber providedthe illustration. 2 Plant ResourcesCenter, Department of Botany,University of Texas, Austin,Texas 78713, U.S.A. 3 Departmentof Botany,University of Texas, Austin,Texas 78713, U.S.A. ANN. MISSOURI BOT. GARD. 80: 439-457. 1993. This content downloaded from 128.83.205.78 on Thu, 18 Jun 2015 17:01:01 UTC All use subject to JSTOR Terms and Conditions 440 Annals of the MissouriBotanical Garden TABLE 1. Chromosomenumbers in Malvaviscus. Taxon 2n number Voucher/reference M. arboreusvar. arboreusre- ca. 56 Mexico. Veracruz:Fryxell & Bates 940; Bates (1976) portedas var. mexicanus M. arboreusvar. arboreusre- ca. 56 Mexico. Veracruz:Fryxell et al. 1676; Bates (1976) portedas var. mexicanus M. arboreusvar. arboreusre- ca. 28 Mexico. Jalisco:Fryxell, Bates & Blanchard 1575; Bates portedas M. pendulifiorus (1976) M. arboreusvar. drummondii ca. 28 U.S.A. Texas: Travis Co., Mendenhall 485; reportedhere M. pendulifiorusreported as ca. 84 Kew Gardens,where cultivated; Skovsted (1935) M. sp. M. pendulifiorus 86 India. Karnataka: wherecultivated; Krishnappa & Munirajappa (1982) cytologicalbehaviour indicates that it is probablya hybridalthough its originappears unknown.It is com- SPECIES CONCEPT where it is vegetatively monlycultivated in Trinidad In this treatment,species concepts in Malvavis- reproducedas it is apparentlycompletely sterile. cus are largely those of the senior author, the junior Krishnappa & Munirajappa (1982), however, author being new at the herbarium bench. Because reported M. penduliflorus to have a count of 2n fieldworkon the genus was limited,species concepts = 86, presumably a miscount of 2n = 84, or are based on morphogeographical considerations possibly an aneuploid clonal derivative of what and our experience with species recognitionin oth- would seem to be its original number, 2n = 84, er groups (e.g., Asteraceae, Fabaceae), both in the since other taxa in Malvaviscus appear to have fieldand as hypothesizedby herbarium evaluations. an ancestral base number of x = 14, if not x = We believe our specific delimitations in Malvav- 7. Bates (1976) listed a count of 2n = ca. 28 for iscus are populational in nature and are compa- M. penduliflorus, but we believe the voucher con- rable to what most workers referto as good species. cerned is more likely a morphological variant of We do not claim this to be true for M. penduli- the widespread, highly variable M. arboreus var. florus, for we suspect that the plants referable to arboreus, as conceived of here. this species are relatively uniform, mostly sterile Our report of 2n = ca. 14 pairs for M. arboreus cultivars, developed very early on by accidental or var. drummondii is to some extent biased because synthetic means. We have recognized M. pendu- of the report of 2n = 28 by previous workers. In liflorusas a "synthetic species," not occurring truth, the count would have been any number naturally in the wild except as it has persisted or between 12 and 15 pairs, the chromosomes being escaped cultivation by vegetative means, although very small and not formingneat bivalents. it is possible that occasional hybrids between M. In summary, only two taxa of Malvaviscus have penduliforus and M. arboreus might occur, as been counted with reasonable certaintyto date: M. noted under the discussion that follows each. arboreus var. arboreus with 2n = ca. 28 and ca. Our species concepts are similar to those of 56, and the cultivated M. penduliflorus withcounts previous revisional workers on Malvaviscus,name- of 2n = ca. 84 and 86 (Table 1). The base number ly Schery (1942) and Fryxell (1988). In spite of of the genus appears to be x = 14, if not 7. The the plethora of pressed specimens available for latter base would agree withthat found in the large, study, extraordinaryvariation, and widespread dis- closely related genus Pavonia, where some 35 or tributionof Malvaviscus, Schery recognized a sin- more species have been counted, all on a base of gle species with eleven varieties while Fryxell rec- x = 7, ranging from n = 14 to n = 56 (although ognized six species in Mexico, which suggests that some authors have reported occasional counts of he would likely have recognized less than ten taxa both n = 13 for a species that otherwise has been for the remainder of its distribution.In short, our counted as n = 14 (e.g., Pavonia zeylanica (L.) recognition of five species for Malvaviscus (in- Cav. with 2n = 52 (Krishnappa & Munirajappa, cluding M. penduliflorus) is conservative, standing 1980) vs. 2n = 56 (Dasgupta & Bhatt, 1976, between the evaluations rendered by
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