The Musaceae

An annotated list of the species of Ensete, Musa and Musella.

This website 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 widened to attempt to include all published taxa, and their synonyms, the majority of which are never likely to be encountered in horticulture whether for fruit production or ornamental use.  The information presented in this website, listing all known species and sorting the genus Musa into its several, recently revised, sections is summarised here.

The Musaceae has been exploited by humans 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.  Musa acuminata is part of a complex of subspecies and closely related species but Musa balbisiana appears genetically to be rather isolated within the genus.  The next and much smaller group is derived from members of the section Callimusa (species previously treated as section Australimusa) is restricted in importance to Polynesia.  Of even more restricted importance, but not lesser interest, are small groups of hybrids in Papua New Guinea; a section Musa group to which Musa schizocarpa has also contributed and a group of section Musa x section Callimusa (previously 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 bananas from the genus 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.

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 by Mabberley uses Cronquist's System for the arrangement of Angiosperm families and provides the following classification:

In the second edition of The Plant-Book Mabberley (1997) uses Kubitzki's System which alters the classification as follows:

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).

Links to particularly useful web pages for other families within the Zingiberales are in the table above.

There have also been substantial 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 lasiocarpa remains somewhat controversial.  The plant has been round the taxonomic block, being placed first in Musa, then in Ensete and back to Musa before it's monotypic status was recognised, at least by some, around 1978.  We await the final word on Musella lasiocarpa.

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.  The Linnean binomials that were applied mistakenly to these sterile Musa hybrids cultivated for fruit were rejected as valid names in the 1950's but are still in use and 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, the last as recently as 2002.

For the record, most authorities now give the number of Musa species as 35 to 42 (as of November 2002, 54 are now listed here) and the number of Ensete species as 7 to 9 (8 are listed here).  Musella is 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 have already begun to cause changes within the family.  Traditionally, the genus Musa was sorted into five sections, Ingentimusa, Australimusa, Callimusa, Musa and Rhodochlamys.  The sections were defined on the basis of chromosome number but strongly influenced and sometimes overridden by morphology.  Recently, studies by Carol Wong and colleagues in Singapore (Wong et al 2002) 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 now sorted into the sections Callimusa and Musa respectively.  The Ingentimusa remain as the third "group" comprised solely of Musa ingens.  The remarkable morphological differences that once supported the separation of the sections are no longer considered important in determining sectional status.  Once again in the taxonomic history of the genus, the stress given to morphological features has ultimately proved illusory.  We can expect further changes.  Some of the 54 species mentioned here are still very poorly known and some are vulnerable to reduction as further synonymy or even hybridity is uncovered.  But new species await discovery and/or formal description.  The Musaceae are large plants and one would have thought difficult to miss.  Yet, a new species was described from Indonesia in 1998, two, from Sabah and China, in 2000, two, from Papua New Guinea and Vietnam, in 2001 and thee more from China & Vietnam and the Philippines in 2002.  Undescribed species are also reported from Borneo, north-east India, Thailand and Vietnam, parts of which are still botanically very little known, and still more may be expected when Cambodia, Laos, Myanmar (Burma) and Indonesia are properly explored.  These are exciting times indeed 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, whether derived from the Musa (Eumusa) or Callimusa (previously Australimusa) sections, 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.  The "product" of Simmonds and Shepherd's system, as distinct from its application, is simple and is in widespread use by banana fruit specialists although there has been, and there still 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.  To complicate matters further there has been something of a breakaway from Simmonds and Shepherd's "classical" system by banana scientists in the Far East.  They have adopted a sort of hybrid approach, re-instituting Linnaean binomials but superimposing genome nomenclature.  I am not quite sure on what grounds this breakaway has been made.

Fourthly, natural and artificial hybridisation causes confusion.  Simmonds 1956 discovered "two Eumusa-Rhodochlamys hybrids in a few weeks of field work [in north east India and Indo-China and thought] 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 apparently a small number of genes of one species can introgress into another.  In the context of wild populations this introgression can be recognised for what it is.  Out of context, hybrids and introgressed plants may look like "new" species.  As part of an extensive research programme on Musa cytogentics, a large number of wide crosses, especially involving the Rhodochlamys, were made in Trinidad in the 1950's.   Several of these hybrids were recognised as having ornamental potential and released to botanic gardens and interested growers and thus found their way into commerce.   Unfortunately, the careful documentation that accompanied those releases seems to have been lost.  And to highlight Simmonds' concerns noted above, 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 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.  Examples are provided by Musa sikkimensis, Musa coccinea and Musa basjoo.

• Musa sikkimensis was introduced to cultivation in 1998 as Musa hookeri.   Although the nomenclature was quickly corrected the "wrong" name, Musa hookeri, is still in wide circulation for this material. 
  
• There has been confusion in the literature between Musa uranoscopos Loureiro and Musa coccinea Andrews for some 200 years.  In 2002, Ai-zhong Liu and colleagues and then Argent & Kiew formally and finally (?) proposed Musa coccinea Andrews as the correct name.
  
• Since its "discovery" by von Siebold in 1830 Musa basjoo has been said to be indigenous to the Ryukyu Islands of southern Japan and cultivated there for fibre.   In fact, Musa basjoo is a Chinese species and is useless for fibre.   The Japanese Fibre Banana, the ito-basho, is a form of Musa balbisiana.

A final and rich source of ambiguities is that, when it comes to nomenclature, wholesale and retail nurserymen and seedsmen, with honourable exceptions, are as erratic as they are inventive in naming their plants.

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 w3 Tropicos databases are the closest but are currently incomplete.  Projects such as the IPNI Plant Names Project and the IOPI Species Plantarum Project (why on earth are there two projects like this?) will presumably eventually generate a definitive list of species of the Musaceae - along with everything else.   At present (June 2002), the IPNI site has good coverage of the Musaceae but the IOPI site is very poor indeed.   The Index Kewensis is not directly available on the web (personally I think it should be) but is effectively accessible by virtue of its inclusion in IPNI.

In January 2000 Gerda Rossel made me aware of Champion 1967 a rather obscure French publication which includes a wonderful (but not quite 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 Champion's publication at the outset this website would probably never have been started, it provides all the information I was originally looking for.

Although 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.  Such a task would make the effort put into this current website seem facile.  Nonetheless, Michel Porcher has made a brave attempt to provide just such a cross-referenced, and multilingual, list of Musa names (Sorting Musa names) but this is currently incomplete.  The books by Stover & Simmonds (1987) and Shanmugavellu et al (1992) contain much information on this subject. 

In response to questions asked by those seeking more information on plants purchased from nurseries in the UK I have included a page on Musaceae offered for sale in Europe and USA.  In addition to websites already mentioned, good sources of information about the cultivated bananas can be found at the INIBAP and Promusa websites (the home pages have links to other Musa sites) and at the IPGRI website.

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 being regularly revised and updated. 

There are still some bad links and spelling mistakes for which I apologise.  I am correcting these, slowly.

The information presented is currently based on a review of only some of the literature, often secondary sources.  In some places the information represents merely my opinion but these instances have reduced and will continue to 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.  In drawing any conclusions I have not reviewed any herbarium or living material nor have I yet sought directly the opinion of professional taxonomists.  However, especially in the case of Ensete much of the information now being included here results from the work of just such a professional, Dr. Gerda Rossel from the University of Leiden, The Netherlands.  I should also like to acknowledge 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 now 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.

Acknowledgements.

It is a pleasure to acknowledge the following people who have contributed to the development of this website:

C. E. S.
Dr. George Argent for copies his papers on Musa and for his time and for much information.
Timothy Chapman for sowing the seed and for providing some!
Admasu Tsegaye for information on Ensete ventricosum.
Prof. Hu Zhihao for information and photographs of Musella and other Chinese Musaceae.
Dr. James Waddick for references to Chinese Musa and Ensete.
Paul Spracklin for photographs of Ensete superbum and Musa basjoo.
Dr. Ho Huu Nhi for information and photographs of Vietnamese Musaceae.
Claudine Picq at INIBAP for back issues of InfoMusa and Musarama.
Rob Wagner for information on the Chinese origin of Musa basjoo.
Markku Häkkinen for information and photographs.
Clarence Hester for seed of Musa balbisiana.
Hideo Shimizu for information about Musa basjoo.
Liu Aizhong for information on Chinese Musaceae and for clarifying the Musaceae section in Flora of China.
Ganesh Mani Pradhan for information and photographs.
Mike Bingham for photographs of Ensete homblei.
Jain Linton, Taman Pertanian Sabah, for photographs of Musa suratii
Dr Jeff Daniells, Queensland Department of Primary Industry, for photographs of Musa ingens.
Shih-Hui (Sylvia) Liu at the Taiwan Forestry Research Institute, for images of Musa formosana.
Eric Schmidt of the Harry P. Leu Gardens, Florida for information and photographs of Musa sp. "Yunnan"

last revision 16 July 2003