The Musaceae
An annotated list of the species of Ensete, Musa and Musella.
Introduction
This annotated list started life as a simple attempt to resolve inconsistencies and ambiguities in nomenclature of the ornamental Musaceae encountered in various standard horticultural texts. The scope has widened somewhat to attempt to include all published taxa, and their synonyms (including pre-Linnean names), the majority of which are never likely to be encountered in horticulture whether for fruit production or ornamental use. The information presented is summarised here.
The Musaceae has been exploited by man for food for millennia and contributes staple crops to many regions of the world. It is probable that the banana was one of the first fruit-bearing plants to receive attention in the early development of agriculture more than 4,000 years ago (the first development of agriculture is generally reckoned to have occurred about 10,000 years ago). A number of distinct groups of edible bananas have been developed from species of Musa. By far the largest and now the most widely distributed group is derived from Musa acuminata (mainly) and Musa balbisiana either alone or in various hybrid combinations. Dividing the genus into sections, Musa acuminata is now placed in the section Eumusa (Musa) 1 and Musa balbisiana in section Eumusa (Musa) 2. The next and much smaller group is derived from members of the section Australimusa and is restricted in importance to Polynesia. Of even more restricted importance are small groups of hybrids in Papua New Guinea; a section Eumusa (Musa) 1 group to which Musa schizocarpa has also contributed and a group of section Eumusa (Musa) 1 x section Australimusa hybrids.
In addition to the direct consumption of the fruit of Musa as food, the fruit and other parts of the plant have various other uses. A selection of links to websites dealing with economic botany and ethnobotany is given on the genus Musa page. Specific uses for plants are mentioned in the species pages.
Apart from Musa other members of the family are also consumed or used for various purposes. The young flowering stems of Ensete ventricosum are a staple source of human food in parts of Ethiopia. Ensete glaucum is cultivated, or perhaps merely used sometimes as a vegetable in south-east Asia. The genus Ensete also yields a useful fibre, its commercial cultivation for this purpose has been investigated, certain types have medicinal uses or religious significance and the large seeds are used for ornament. Musella lasiocarpa is also said to be consumed in China (often for animal food but also sometimes as human food) and has medicinal uses too.
The Musaceae is also exploited in ornamental horticulture; as a source of garden and patio plants, indoor pot plants and cut flowers.[The section name Eumusa is very commonly used in the literature but following article 32 of the the ICBN it is illegal and the section should properly be named section Musa. Eumusa is widely used to prevent confusion between the genus Musa and the section Musa. By convention taxa above the genus level are not commonly italicised.]
Musaceae nomenclature in standard horticultural texts and nursery and seed catalogues is ambiguous, inconsistent, incomplete and sometimes wrong. There are various reasons for this.
First, taxonomy is a dynamic discipline and the Musaceae are no different to other families in being subject to periodic revision by and sometimes disagreement among taxonomists. Changes periodically affect all levels of classification. For example, in the first edition (1987) of The Plant-Book Mabberley uses Cronquist's System for the arrangement of Angiosperm families and provides the following classification:
Subclass Zingiberidae Order Zingiberales Family Strelitziaceae Heliconiaceae Musaceae Lowiaceae Zingiberaceae
incl. CostaceaeCannaceae Marantaceae In the second edition of The Plant-Book Mabberley (1997) uses Kubitzki's System which alters the classification as follows:
Subclass Zingiberidae Order Zingiberales Family Musaceae
incl. Strelitziaceae & HeliconiaceaeLowiaceae Zingiberaceae
incl. CostaceaeCannaceae Marantaceae However, most authorities have maintained Strelitziaceae, Heliconiaceae as families separate from the Musaceae, e.g. Watson & Dallwitz (1992 onwards) and Berry & Kress (1991).
Recent comparative studies of plastid and nuclear gene sequences coupled with the application of cladistics is providing a new, somewhat controversial, ordinal classification of flowering plants (Bremer et al 1998). The Zingiberales have been only slightly affected by such studies (see Kress & Hahn 1997 and Genera Zingiberarum).
Superorder Zingiberanae Order Zingiberales Family Musaceae Strelitziaceae Lowiaceae Heliconiaceae Costaceae Zingiberaceae Cannaceae Marantaceae Links to particularly useful web pages for other families within the Zingiberales are in the table above.
There have also been changes at the genus level. The 1931 edition of Willis, the pre-cursor of Mabberley's Plant-book, lists Musa as the only genus in the Musaceae. Although Horaninow described Ensete as early as 1862 the genus did not receive widespread recognition until revived by Cheesman in 1947. Mabberley, Cullen and recent RHS publications still include only two genera in the Musaceae, Ensete and Musa. The situation of Musella remains somewhat controversial. Musella lasiocarpa has been round the taxonomic block, being placed first in Musa and then in Ensete before it was placed in its own genus in 1978. We await the final word on Musella.
The largest genus, Musa, has historically been divided into six sections viz., Australimusa, Callimusa, Ingentimusa, Eumusa (Musa) 1 & 2 and Rhodochlamys. These sections do not have formal taxonomic significance but they are useful in discussing relationships between bananas. The five sections have recently been reduced to three. Previously the 2n = 20 chromosome species were separated into the sections Australimusa and Callimusa and the 2n = 22 chromosome species were separated into the sections Eumusa (Musa) and Rhodochlamys. Studies by Carol Wong and colleagues in Singapore have revealed that genetic differences between each section in the same chromosome group are smaller than those within each section. This means that the traditional separation of the sections can no longer be substantiated. The studies of Wong et al do, however, maintain the separation between the 20 and 22 chromosome species. The 2n = 14 Musa ingens remains in its own section. The morphological differences that once supported the separation of the sections are no longer considered important in determining sectional status. It remains to be seen whether Carol Wong's results are substantiated.
Changes have been equally great at the species level and are by no means complete. Cheesman is the father of modern Musaceae taxonomy and it may come as a surprise that the proper definition of Ensete and a number of very significant Musa species including Musa acuminata and Musa balbisiana is as recent as Cheesman's work in the 1940's and 50's. Willis (1931) noted that Musa contained 80 species but he did not list them. This number has now been reduced considerably with the separation out of Ensete by Cheesman and the recognition that a number of Musa "species" were actually hybrids. A strong case has also been made for abandoning Linnean binomials that were applied mistakenly to sterile Musa hybrids cultivated for fruit and which still cause confusion today (see below). As a counter to these downward pressures the number of species is increasing again as new taxa are discovered and described.
For the record, most authorities now give the number of Musa species as 35 to 42 (59 are listed here) and the number of Ensete species as 7 to 9 (6 are listed here). Musella is probably monotypic. No one knows for sure the precise number of species in the Musaceae which is anyway not an especially important fact in itself.A second reason for the inconsistencies in the literature is that the family is difficult to study. Hot spots of Musaceae biodiversity are in areas that are difficult, sometimes even dangerous to travel and work in. It is difficult to obtain and prepare adequate herbarium material especially of the massive inflorescences. As a result, and despite its importance as a source of valuable food plants, the family is still incompletely known after some 200 years study. Studies of plant genome structure are being applied to the taxonomy of the Musaceae with fascinating results and are likely to result in further changes within the family. Some of the species mentioned here are vulnerable to reduction as further synonymy or even hybridity is uncovered. But new species await discovery and/or formal description. For example, new species are emerging from Borneo where, sadly, destruction of the rainforest facilitates their discovery. IndoChina is still botanically very little known and likely to yield new species. These are exciting times for those interested in the Musaceae.
Thirdly, Musaceae taxonomy is made still more difficult by the group's domestication. Today's fruit-bearing cultivars are hundreds or perhaps thousands of years and many mutations away from the chance hybridisation or mutation that originally gave rise to them. The cultivated bananas and plantains are sufficiently removed from their original species that they cannot usefully be assigned Linnean binomials. Recognising this, Simmonds and Shepherd proposed in 1955 an alternate, genome-based system for the nomenclature of the section Musa (Eumusa) bananas. Despite the simplicity of the product of Simmonds and Shepherd's system, as distinct from its application, and its widespread use by banana fruit specialists, there is an extraordinary reluctance to adopt it in ornamental horticulture. The application of what are effectively spurious or mis-applied Linnean binomials is therefore still widespread even in recent and, supposedly, authoritative horticultural publications. This would not be so bad if publications were consistent one with another in terms of nomenclature but some, notably recent RHS publications, are not even internally consistent. Where their use has been devalued if not actually discredited by Simmonds and Shepherd, the tendency to cleave to Linnean binomials as the only mechanism with which to fix a plant in the horticultural firmament is, at the very least, unhelpful and sometimes downright misleading.
Fourthly, natural hybridisation is also likely to cause confusion. Simmonds 1956 discovered "two Eumusa-Rhodochlamys hybrids in a few weeks of field work [in north east India and Indo-China implying] that such hybrids must be very numerous. They could well give rise to considerable taxonomic confusion, especially in a group as poorly understood as Rhodochlamys. Collections based on segregates could easily be taken for new species unless they were extraordinarily well documented." As well as straight hybrids there also exists the phenomenon in Musa whereby a small number of genes of one species can through crossing and backcrossing introgress into another. The products of hybridisation and introgression especially involving the Rhodochlamys may well have ornamental potential and with the renewed commercial introductions of Musaceae in the form of seed and suckers such mongrels may well find their way into commerce. And to highlight Simmonds' concerns, most of the new material entering horticulture today is extraordinarily poorly documented. No doubt as the result of such imports due to increased interest in the Musaceae as ornamentals we can look forward to a period of further confusion of taxa and nomenclature in the horticultural trade. But we can also expect some pleasant surprises; the species Musa laterita and Musa gracilis, the latter vulnerable in the wild in Malaysia, have recently been recognised as being securely in cultivation mislabelled as cultivars of Musa ornata.Fifthly, Musaceae taxonomic literature is liberally peppered with mistakes! The published works of authorities from Linnaeus through, for example, Baker, Schumann, Cheesman, Simmonds, Champion and Lock (and minor scribblers like me) all include errors. To err, of course, is human but it is also human to tend to trust that the great authorities must always have been correct in their published works. Not so. Mistakes were made and were and are repeated and thus became part of the paradigm.
This list now attempts to include all published names of taxa in the Musaceae and by giving their proper or "accepted" names to try to resolve inconsistencies in nomenclature in the horticultural literature. Information on Musaceae is widely scattered and modern standard horticultural references have only partial and frequently mis-named species lists and poor synonymy. I have tried clearly to identify accepted names, those that are properly applied to true botanical species, subspecies and varieties. I have also tried to identify names that are synonyms, spurious binomials and plain mistakes such as typographical errors and transliterations that occur in the literature.
I do not think a similar list appears elsewhere on the web; the lists at the GRIN and Mobot VAST databases are the closest but are currently incomplete. Projects such as the The International Plant Names Index (IPNI) and the Species Plantarum Project (why are there two projects like this?) will presumably eventually generate a definitive list of species of the Musaceae - along with everything else. The Index Kewensis is effectively now available online via RBG Kew's electronic plant information centre (EPIC). A problem with most of these sites is that they do not make it clear which names relate to real species (accepted names) and which are synonyms. This problem is largely resolved at the World Checklist of Monocotyledons website but this seems to me currently (April 2008) to give sometimes spurious accepted names and to contain some mistakes. I do not mean to imply that this website is mistake free!
In January 2000 Gerda Rossel made me aware of Champion 1967 a rather obscure French publication which includes a wonderful (but not complete) list of Musa binomials together with their accepted names. If it had not been for his publication's obscurity, and being written in French, Champion's name would be as well known as Cheesman's or Simmonds' to those interested in the Musaceae. Had I known of this publication at the outset this website would probably never have been started, it provides all the information I was originally looking for. And Häkkinen & Väre have recently (2008) produced a more or less definitive checklist list of Musa names rendering this website largely irrelevant.Recently the work of Georg Forster has come to my attention. Until now, Forster's work has been ignored by students of the Musaceae. Together with his father, Johann, Georg Forster travelled with James Cook on his second expedition to the Pacific (1772 - 75). Forster's De Plantis Esculentis Insulam Oceani Australis Commentatio Botanica of 1786 contains a list of 16 varieties of banana. These varieties were largely seedless edible bananas and thus of little consequence taxonomically.
Because some were given spurious Latin names (e.g. by Georg Forster) I have mentioned a small number of fruit-bearing cultivars. I have not made and I do not intend to make a systematic, cross-referenced list of hybrid names. Michel Porcher has made a brave attempt to provide just such a cross-referenced list of Musa cultivar names (Sorting Musa names). The books by Stover & Simmonds (1987) and Shanmugavellu et al (1992) contain much information on this subject. Good sources of information about the cultivated bananas can be found at the website of the Banana and Plantain Section of Biodiversity International (formerly INIBAP).An explanation of the form of the entries in this list is given here.
A summary of the information presented including a table sorting the genus Musa into its several, recently revised, sections is given here.CAVEAT EMPTOR!
This is a work in progress and is occasionally revised and updated.
There are still some bad links and spelling mistakes for which I apologise. I am correcting these, slowly.
In some places the information represents merely my opinion but these instances will reduce with time. I have given references and explained my conclusions but the work has not been peer reviewed and I must have made mistakes I am unaware of. I have acknowledged unpublished sources throughout but would particularly like to thank Gerda Rossel and Markku Häkkinen for generously supplying information and for corrections. Any mistakes that remain are mine.Review, comment and opinion on any and all aspects of this web site is invited. I would be grateful to receive any further information and literature references on the plants mentioned and species or subspecies that I have not mentioned. And since this list has pretensions to being authoritative, I would be especially grateful to receive corrections to my mistakes. All contributors will be acknowledged as I update the list.
Please e-mail me at drc@globalnet.co.uk
David Constantine.
The Musaceae - an annotated list of the species of Ensete, Musa and Musella, The genus Musa - an annotated list of species, The genus Musella - an annotated list of species are Copyright © 1999 onwards by David Constantine
The genus Ensete - an annotated list of species is Copyright © 1999 onwards by Gerda Rossel & David Constantine
The right of David Constantine and Gerda Rossel to be identified as the Authors of the Work(s) has been asserted by them in accordance with the Copyright, Designs and Patents Act 1998
The copyright of other sources for and contributors to this website is acknowledged as appropriate.
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last updated 22/10/2008