Thursday, 20 December 2012

Strange violets.

Māhoe (Melicytus ramiflorus) is one of the plants I love to show to visiting botanists from temperate Europe or North America.  They invariably ask, "What family is it?" and the answer always surprises because they're used to Viola as the representative of Violaceae.  Here's a violet that's a tree, has radially symmetrical flowers, and fleshy fruit; the world truly is upside-down in New Zealand.

Māhoe flowers on twigs of a male tree, Karori, Wellington
Well, not so fast.  It's true Viola is the type genus of Violaceae, but who said types have to be typical?  The type simply determines the application of the name according to the rules of nomenclature.  It's the circumscription—the definition of the membership of the group—that outlines its overall collective characteristics.  Violaceae is a largely tropical family, and they're woody except for Viola.  Quite a few have radially symmetrical flowers and fleshy fruits (Mabberley 2008).

A "normal" violet (Viola sp.), France.
Māhoe might be New Zealand's commonest tree.  There's certainly lots of it around Wellington, where it's an important component of the still low second growth forests that, over the last 50 years, have replaced the gorse on hills around the city.

Māhoe flowers in pulses, all the trees in a location flowering in synchrony several times a summer (Powlesland et al. 1985).  Coffee does the same thing.  The flowers are small and borne on the twigs, a form of flower presentation called ramiflory.  They're scented, especially at night, but it's not a very pleasant smell, in my opinion.
Melicytus ramiflorus, male flower.  Note the vestigial ovary in the centre and the swollen connective at the back of each anther that functions as a nectary
Melicytus ramiflorus female flower.  Note the large stigma and the nectar produced from the connective at the back of each staminode.  Female flowers are about 2/3 the size of males.
The flowers are unisexual and produced on separate trees, so a whole tree is either male or female.  The connective of the stamens and (in female flowers) staminodes is also the nectary, and each produces a glistening drop as a reward for the pollinators.
Māhoe leaf skeletons
Their leaf veins are pretty tough, but the rest of the leaf decomposes freely, to leave exquisite lacy leaf skeletons on the forest floor.
Viola cunninghamii, Hooker Valley, Mt Cook National Park.
New Zealand has three native violets (like V. cunninghamii above), plus some introduced species.  We also have quite a few Melicytus; some are small trees and others are twiggy shrubs.  Some used to be classified in the genus Hymenanthera, but that was merged with Melicytus in the 1980s.  Recent research (Mitchell et al. 2009) shows there are two pretty clear-cut groups within the genus, although the authors didn't draw attention to it, and it might make sense to recognise Hymenanthera again.


Mabberley, D.J. 2008.  Mabberley's Plant-Book (3rd ed.).  Cambridge.

Mitchell, A.D.; Heenan, P.B.; Murray, B.G.; Molloy, B.P.J., de Lange, P.J. 2009.  Evolution of the south-western Pacific genus Melicytus (Violaceae): evidence from DNA sequence data, cytology and sex expression.  Australian Systematic Botany 22(3) 143–157.

Powlesland, M.H.; Philipp, M.; Lloyd, D.G. 1985.  Flowering and fruiting patterns of three species of Melicytus (Violaceae) in New Zealand. New Zealand Journal of Botany 23: 581–596

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