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A New Phylogeny-Based Generic Classification of Costaceae (Zingiberales)

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A New Phylogeny-Based Generic Classification of Costaceae (Zingiberales) UC Berkeley UC Berkeley Previously Published Works Title A new phylogeny-based generic classification of Costaceae (Zingiberales) Permalink https://escholarship.org/uc/item/1v06n4pb Journal Taxon, 55(1) ISSN 0040-0262 Authors Specht, Chelsea D. Stevenson, Dennis Wm. Publication Date 2006-02-01 Peer reviewed eScholarship.org Powered by the California Digital Library University of California 55 (1) • February 2006: 153–163 Specht & Stevenson • Generic classification of Costaceae A new phylogeny-based generic classification of Costaceae (Zingiberales) Chelsea D. Specht1,2 & Dennis Wm. Stevenson1 1 New York Botanical Garden, Bronx, New York 10458, U.S.A. 2 Current address: University of California, Department of Plant and Microbial Biology, Berkeley, California 94720, U.S.A. [email protected] (author for correspondence) Recent cladistic analysis of multiple molecular data from chloroplast and nuclear genomes as well as mor- phological data have indicated that a reclassification of the family Costaceae is necessary in order to appro- priately reflect phylogenetic relationships. The previously described genera Tapeinochilos, Monocostus, and Dimerocostus are all upheld in the new classification. Monocostus and Dimerocostus are found to be sister taxa. The large pantropical genus Costus is found to be polyphyletic and is thus divided into four genera, three of which are new (Cheilocostus, Chamaecostus, Paracostus). Costus has now a more concise generic concept including morphological synapomorphies previously absent due to the polymorphic nature of the prior non- monophyletic assemblage. Of the three new genera, one (Paracostus) was previously recognized as a subgenus of Costus. Cheilocostus comprises several Asian taxa and is sister to Tapeinochilos, whereas Chamaecostus comprises entirely neotropical taxa and is sister to a neotropical Monocostus + Dimerocostus clade. A reeval- uation of the traditional taxonomy indicates that floral characters and pollination syndromes commonly used to identify groups exhibit homoplasy when analyzed in a cladistic framework and are thus unreliable as taxo- nomic indicators. KEYWORDS: Costaceae, Costus, Zingiberales, classification, phylogenetic taxonomy. family (e.g., Thiselton-Dyer, 1898), but in later treat- INTRODUCTION ments was reduced to subgeneric status within Costus Costaceae is one of the most easily recognizable (Schumann, 1904). Floristic treatments that included groups within the Zingiberales, distinguished from other species of Cadalvena either did not distinguish subgen- families within the order by well-developed and some- era (Koechlin, 1964, 1965; Hepper, 1968) or treated times branched aerial shoots that have a characteristic Costus and Cadalvena as subgenera within Costus monistichous (one-sided) spiral phyllotaxy (e.g., Kirch- (Maas, 1972, 1977). Maas (1972) described several new off & Rutishauser, 1990). Its close relationship with species from South America which he included in C. Zingiberaceae is evidenced by its former placement as a subgenus Cadalvena. No new species from Africa have subfamily within the larger Zingiberaceae family. The been added to Cadalvena since Schumann (1904) and, in placement of Costaceae within Zingiberaceae was large- fact, many of the species included in Schumann’s treat- ly based on broad similarities of inflorescence and floral ment have been subsequently synonymized with the characters. Tomlinson (1962) suggested that, although generic type, Costus spectabilis. these types of characters may indicate common ancestry, In addition to Cadalvena, Schumann (1904) recog- they are not sufficient to overcome the morphological nized four other subgenera in Costus in his treatment of and anatomical differences that warrant independent global Zingiberaceae (including the Costoideae): familial rank of the two lineages which had been pro- Eucostus (= C. subg. Costus according to the current posed by Nakai (1941). International Code of Botanical Nomenclature, ICBN, Costaceae sensu Tomlinson (1962) consisted of four Greuter & al., 2000), Metacostus, Epicostus and Para- genera: Costus, Monocostus, Dimerocostus and Tapeino- costus. These same subgenera were maintained by chilos. Costus, which contains the majority of the species Loesener (1930). In Maas’ treatments of neotropical and maintains the greatest morphological diversity, is Costaceae (1972, 1977), a formal division between the pantropical with its greatest diversity centered in the two subgenera that are found in South America (Costus, neotropics (c. 40 spp.); 25 species occur in tropical Afri- Cadalvena) was maintained. In addition, subgenus ca and about five species in southeastern Asia. A separate Costus was divided into two separate sections: Costus genus, Cadalvena, was recognized when Costoideae was sect. Ornithophilus and Costus sect. Costus. These sec- part of Zingiberaceae (Scitamineae) (Fenzl, 1865) and tions were based upon characters of the labellum and was maintained in some subsequent treatments of the reflected two distinct floral forms associated with polli- 153 Specht & Stevenson • Generic classification of Costaceae 55 (1) • February 2006: 153–163 nation syndromes. Taxa placed in Costus sect. Ornitho- ed floral and geographic diversity. Tapeinochilos, philus all have a tubular labellum adapted to bird polli- Monocostus, and Dimerocostus are also maintained nation (Fig. 1.5), whereas those in Costus sect. Costus because the morphological distinction with which they have a broad labellum with a distinct exposed limb that were originally described corresponds to their monophy- appears to be adapted to pollination by bees. While all ly. A new generic classification of the family is present- neotropical Costus species can be placed rather easily ed and species lists are given for each of the newly into one of the two sections based on floral and inflores- described genera. Several lineages within Costus having cence characteristics, African and Asian Costus do not unique associations with pollinators are recognized. share this distinction of forms. Subsequent to Schumann (1904) and Loesener (1930), the division of African and Asian Costus into various subgenera has not been main- tained, with the exception of Costus spectabilis as sub- MATERIALS AND METHODS gen. Cadalvena (Lock, 1985). The most recent interfamilial classification for In contrast to the large pantropical Costus, the Costaceae is provided by a combined molecular and remaining genera of Costaceae (Monocostus, Dimero- morphological cladistic analysis that includes a represen- costus and Tapeinochilos) are restricted in distribution. tative sampling of Costaceae taxa (Specht & al., 2001; Dimerocostus and Monocostus are both restricted to the Specht, 2006). A summary of the current evolutionary neotropics, the former extending from Honduras in the hypothesis for Costaceae based on the recent phyloge- north to central Bolivia in the south and the latter known netic analysis is shown in Figure 1. This phylogenetic only from the Río Huallaga region of central Peru. hypothesis forms the basis for the revised taxonomic Monocostus is the only taxon to have a solitary flower in classification system. the axils of the leaves rather than a highly structured The formal generic classification is presented as inflorescence of spirally arranged bracts subtending sin- governed by the current ICBN. In addition, a list of syn- gle or paired flowers. Although differing in overall plant apomorphies is provided for each of the newly described morphology, Monocostus and Dimerocostus share a flo- genera and follows the discussion of the individual ral morphology that is at least superficially similar to that group. of Cadalvena. Tapeinochilos is restricted to the paleo- There was no major discrepancy between the molec- tropics where it is found primarily in New Guinea with a ular and morphological analyses, thus the topology few species extending into the surrounding Indonesian obtained from the combined analysis was selected for the islands and south to Queensland, Australia. Although new classification system. Morphological distinctiveness most closely resembling the Costus sect. Ornithophilus was also considered in the definition of new genera, as in floral form, the floral and inflorescence morphology of was adherence to previously recognized natural lineages. Tapeinochilos is distinct from that of other Costaceae. Major clades with strong support (as determined by Few species of Costus have distributions falling Bremer support, jackknife) and clear morphological syn- within the range of Tapeinochilos, and those that do have apomorphies are named. Within Costus, several clades a floral form of the type found in Cadalvena, Mono- are given informal recognition as having unique mor- costus and Dimerocostus. These Asian Costus species phological characteristics but are not designated formal- share with Tapeinochilos the tendency to undergo vege- ly due to the resulting basal paraphyly. tative branching, and both possess woody inflorescence bracts in contrast to the herbaceous and chartaceous bracts of the African and neotropical taxa. The bracts are often red, but can be dark brown or even black in color. TAXONOMIC TREATMENT Despite these similarities, no specific or formal affilia- Costaceae Nakai in J. Jap. Bot. 17: 203. 1941, tion between Asian species of Costus and Tapeinochilos basionym: trib. Costeae Meisn., Pl. Vasc. Gen.: Tab. had been proposed. Diagn.: 389, Comm.: 291. 17–20 Aug 1842. – Type: Recent phylogenetic analyses
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