UPSIDE UPSIDE DOWN

Leo Davis always has some interesting insights from his orchid observations.  In this article he examines the position of the tepals (petals and sepals) in particular the Moose Orchid which he saw for the first time this year.

Have a close look, next season (winter to early summer) at some of our native lilies.  Start with the jolly bulbine lily (Bulbine bulbosa), no longer a true lily incidentally, because it now resides in family Aspodelaceae, along with the grass trees. You will find three yellow petals at 12, 4 and 8 o’clock and closely behind them three almost identical sepals at 2, 6 and 10 o’clock, so at first sight you see six apparently identical tepals (sepals and petals).  Move on to the rush fringe-lily (Thysanotus juncifolius), as described in Ann Prescott’s ‘It’s Blue With Five Petals’.  Clive Chesson is more up to date and tells me it is now T. racemoides.  Again it is no longer a true lily, now sitting in family Asparagaceae.  Here the tepals are noticeably different.  Three wide densely fringe edged petals will be found, if you view the flower face on, at 12, 4 and 8 o’clock.  The narrow non fringed sepals sit close behind at 2, 6 and 10 o’clock.  These are just a generalisations because if the flower turns only about 60o a sepal will be at the top.

Most orchids, while close relatives of the true lilies and the one time lilies, do not show these arrangements.  Let’s start with some that do.

In the large duck orchid (Caleana major) the petal at 12 o’clock, the dorsal petal, is modified, as in most, but as usual, not all, orchids, to become a labellum.  In this charmer the labellum takes the form of a duck’s head.  Its function is to snap down trapping a pollinator insect in the cup shape column below it, forcing it into contact with the sticky off white stigma and/or the yellow pollinia below it.  Look closely and you will find the other two narrow petals drooping at around 4 and 8 o’clock.  Two folded, twisted sepals can be clearly seen at around 1 and 11 o’clock.  The third sepal, at 6 o’clock, is tucked in behind the cup shaped column.  Note that, as with lilies, the top tepal is a petal.

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Caleana major, Knott Hill NFR Photographer: Leo Davis

The leek orchids (genus Prasophyllum) follow this pattern and also have their labellum at around 12 o’clock.  These orchid groups, which are up the right way, are said to be ‘not upside down’, using the technical term ‘non resupinate’.

Most orchids are ‘upside down’ and are called resupinate.  The whole flower rotates 180o, clockwise or anti I don’t know, at the embryonic stage.  But let’s start with somewhat of an exception with the sun orchids (genus Thelymitra) which do not have a petal modified as a labellum.  But they are indeed upside down.

Have a close look at the Thelymitra benthamiana flower.  Note that the three petals, at roughly 2, 6 and 10 o’clock, are in front of the three slightly larger but very similar sepals, at 12, 4 and 8 o’clock.

Note that the top tepal is a sepal.  The flower is upside down, that is resupinate.  In most orchids the petal at 6 o’clock would be modified to be a labellum.

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Thelymitra benthamiana, Scott Creek CP; Photographer: Leo Davis

The Arachnorchis (possibly Caladenia to you) stricta, from Sherlock, out in the mallee, is more typical of terrestrial orchids in SA.  It is upside down, that is resupinate, and has a petal modified to be a labellum.

The bottom petal has become a wide labellum, with fine edge combs and parallel rows of rich plum coloured calli covering its centre.  Out at roughly 3 o’clock is a narrow petal, the other invisible on the other side.  At the top, pressed tightly against the column, a sepal arches forward.  Two larger sepals extend down at around 5 and 7 o’clock.

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Arachnorchis stricta, Sherlock; Photographer: Leo Davis

When I saw my first, my only, moose orchid, this season, I was in such a state of excitement that it looked to me to be up the right way, that is to say upside down.

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Cryptostylis subulata, Stipiturus CP; Photographer: Leo Davis

Have a look.  Two narrow short roughly vertical petals at about 1 and 11 o’clock.  There are two sepals at just past 3 and just before 9 o’clock. That’s OK but where is the other sepal?  Are there it is, where it should be, at midday.  But hang on, it’s behind the flower stem (peduncle) and where is the column?

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Cyrtostylis subulata with labellum lifted; Photographer: Leo Davis

Holding the labellum up with a stick I found the column, the stigma and the pollinia, underneath the labellum.  The third sepal now appears to be at 6 o’clock.  And it all became clear.  This flower was up the right way (non resupinate) but it has turned forward, on its peduncle, by about 180o, to become upside down, but not in the manner of resupinate flowers, because it is back to front.  It is an inverted non resupinate flower.  Still with me?

Introducing Orchid Pollinators of Victoria: Part Two of Two Parts

Continuing last week’s blog, here is the completion of Rudie Kuiter’s Introducion, Orchid Pollinators of Victoria. In this section he discusses some factors of hybridisation and the value of regular observation by local people interested in orchids.

Click here for Part One

Lissopimpla excelsa is the pollinator of all members of Cryptostylis, but hybrids are not known, even when sympatric, thus a molecular mechanism is in place that prevents cross-fertilisation.  Hybrids in other orchid genera do occur and these usually are amongst closely related species. Several congeneric orchids attract the same male pollinator species, thus would be emitting the same kairomones, the scent that is a mimic of the female’s sex-pheromones, but normally these orchids are allopatric or have different habitat preferences.  The land clearing, frequent fires, changes of watercourses, gold-diggings are amongst many unnatural human habitat interference of recent times.  Historically in undisturbed natural habitats sibling orchid species that attracted the same male insect were not sympatric, not flowering at the same time or were in close vicinity to each other.  In disturbed sites the situation has changed, as closely related species may have become sympatric and hybridisation take place.  Spider-orchids that attract thyniid wasps with kairomones normally target a certain local species, but many allopatric species are know to share the same pollinator and readily hybridise where they became sympatric.  In Pterostylis greenhoods the known hybrids are also caused when different species attract the same pollinators.

Cryptostylis subulata 008
Cryptostylis subulata (Moose Orchid)

We still have much to learn to fully understand how adaptable the orchids are, the role insects play and how to interpret what we see.  Orchids are finely tuned to their world and can change and adapt in ways that most people seem to underestimate.  I requires observations of the same plants over many seasons to get a good understanding of their variability and  adaptability.  Unusual forms often show after a drought or fire may look like a new species, but soon change back to typical or normal within a few good seasons or after regeneration.  It is usually the local people taking an interest that see the changes in the same plants over time that dispute what the ‘on-the-fly’ taxonomist come up with.

Creatures evolved as part of an endless combination of life-forms, ranging from microscopic to the tallest tree, that together form an ecosystem in which all organisms depend on each other.  The climate, weather and other factors changes the environment constantly that influence the members differently, dome doing better than others, but it collectively maintains a balance.  Natural events such as a major fire or flood may benefit environments in areas as part of seemingly long cycles, but they are very short in evolutionary terms.  Unnatural man-made fires are very destructive as these are conducted much too frequently, wrong time of the year, and in the wrong place.  Not obvious, but also very detrimental is the use of insecticides that seems to effect the Diptera members the most.  Many of the important pollinators such as the fungus-gnats have gone locally extinct and most of the Pterostylis depend on them.  To work with the pollinators it is essential to have a good understand of the life-cycles of the insects involved and watch the flowers in the wild.  After witnessing Pollinator behaviours of the fungus-gnat on Pterostylis nutans countless times, the principal pollinator is easily recognised with other species.  Unfortunately few good areas to find orchids and learn about their pollinators are left.  Many are now rare and measures taken to protect them usually focuses on just a species.  To be effective their habitat area and surrounding needs to be cared for, letting the natives grow and have the natural canopy reform.  At least, habitats should be protected from further disturbances, especially by badly informed governmental environment departments with their fires.

Pollinator and Corunastylis archeri
Pollinator with pollinia

Note This book is solely based on first-hand observations on the orchid-pollinators in the wild.  Descriptions and comments are from many hours of watching each species over multiple seasons.

Apart from Orchid Pollinators of Victoria, Rudie Kuiter has produced several Victorian orchid books.  If you are interested in purchasing any please contact us.

Introducing Orchid Pollinators of Victoria: Part One of Two Parts

This week’s blog, Part One of Two Parts, is quoted directly from the introductory chapter (Pages 2&3) Orchid Pollinators of Victoria 4th Edition, 2016, Rudie Kuiter.  Over the years of photographing orchids and their pollinators, Rudie and his team have been discovering much of the hidden world of orchid pollinators.  In this first section he highlights the fallacy of the “one orchid-one pollinator” as well as touching briefly on the vast difference between the insects and their role in the ecosystem.

Click here for Part Two

INTRODUCTION

Amongst flowering plants, orchids have evolved in their own special reproductive ways.  Their pollen is massed as waxy packages, pollinia, unlike like (sic) other flowers that produce masses of fine pollen grains that mostly go astray.  The pollinia are relatively heavy and the usually small creatures need to be strong fliers for cross pollination (see image below).  Orchids evolved with amazing strategies to attract specific carriers in order to transfer pollen between flowers of their species only, and in this way eliminating the need to produce great quantities.  Various insects, many moths, bees and even birds have been documented as pollinators of orchids around the world (v.d. Cingel, 2001).  A number of uniquely different examples of orchids attracting insects for their pollination evolved in Australia, especially in the more temperate southern zones originating from Gondwana times.  With very few exceptions the Victorian orchids are terrestrial, ground-dwellers, that rely on small insects such as fungus-gnats, native bees, wasps, ants and many attract only the males by sexual deception.  In the case no pollinators visits, many species may self-pollinate as a back up.

When taking an interest in orchids it seems difficult enough to identify some species.  Usually one looks and admires the amazing flowers that may resemble an insect and can be difficult to recognise as a flower at first.  An insect on a flower may be thought of something that spoils a picture – until taking an interest in the visitor!

160219 Anita Marquart Melangyna collatus on Caladenia rigida
Hoverfly on Caladenia rigida (syn Arachnorchis rigida)

I first learned about the orchid pollinators in Orchids of South Australia by Bates & Weber, 1990, an excellent book by today’s standard, but few were seen over the years by just being there at the right time when photographing orchids.  During preparation of the book on Caladenia spider-orchids, certain issues developed from questionable statements made in scientific papers about wasp-pollinators.  Of particular concern was about the one-to-one relationship – how only one wasp-species would be involved with only one spider-orchid species – and suggesting populations that were thought to represent the same species comprised different taxa if not pollinated by the same wasp-species.  A very different story emerged when monitoring the local spider-orchid populations to find the answers and it became clear that there was much more to it.  A site in Wonthaggi with a very large colony of Caladenia dilatata proved to be perfect for this study and also to photograph pollinators as it produces flowers for about four months.  It was found that a local wasp-pollinator species typically flies for a little over one month, thus this need to be investigated further.  Scientific publications on wasp-pollinators were generally based on short-term experiments, and usually employing baiting methods – moving flowers and often taking them to different sites.  Responses included unnatural behaviour or attracting sibling wasps at a non-local site.  It is certainly true that a particular flower may attract only one species of wasp a t a locality and a certain time in the season, but this reflects a very small part of the picture.  It can be different in the long-term, at certain localities or with a season.

The main study site was in coastal dunes, where in a very large population of Caladenia dilatata produced flowers for over four months from September, and under favourable Summer-conditions into January.  At least three congeneric (belonging to the same genus) thynniid wasps species were involved in this population.  The flying times were up to about 6 weeks for each wasp species, that were separated or slightly overlapping.  The flowering times in other populations of C. dilatata in Wonthaggi and Wilsons Promontory were usually about one month in each, and at the corresponding times to the study site the wasps visiting were the same species.  A close sibling C. parva in the Wonthaggi heathland habitats and early flowering C. dilatata were pollinated by the same species, but the later flowering C. tentaculata by a thynniid wasp of the different genus.

160219 Anita Marquart Melangyna collatus with pollinia
Hoverfly with pollinia

Whilst a flower may attract only one wasp species, the kairomones (chemical omitted by the orchid to attract a pollinator)  of a species may vary between flowers within in a population or when allopatric (growing in different geographical regions), just like colour or morphology.  Variations maybe in relation to locality, weather conditions, or ground chemistry and available pollinator.  Pollinators may evolve over time, but adaptions usually require many seasons and this would vary with location.  The observations made over several seasons suggest that more than one congeneric insect is involved in pollination depending on local or seasonal conditions, especially after a long drought.  Chiloglottis gunnii populations in Langwarrin were checked for pollinators since a decade-long drought and no action was seen for many seasons.  When wasps finally made an appearance they comprised different taxa of Neozeleboria the first season, but only one became the common and principal pollinator the following seasons.  Thynniid wasps are very localised as females lack wings and rely on the males to carry them around during copulation and to provide food.  It limits their travelling and their homing range may comprise just a few hundred metres.  Thynniid wasp are very vulnerable in small reserves isolated by land-clearing, and certain species have gone locally extinct due to conducting burns.  Insects form a crucial and fundamental part of an ecosystem, but their importance is never considered in the planned burning, showing a complete lack of understanding by people in charge.  So little is known of ecosystem’s foundations, but controlled burning continues – ruining precious habitats.  Orchid species failing to produce seed pods is an indication that pollinators were absent, probably gone locally extinct.  Orchid species that have a sexual association with thynniid wasp pollinators are localised and usually have geographically variable flowers.  The situation is different with Cryptostylis spp. And their pollinating male wasp Lissopimpla excelsa, as females are a strong flyers.  The flowers of Cryptostylis are geographically uniform in each species and the wasp is widespread.  Both sexes are very distinctive in colour that show no variation.  They are active over Spring and Summer and are great travellers.

To be continued …

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