This year I had the chance to visit the white flowering form of Cephalanthera rubra in the Hesse part of the Rhoen some days earlier than last year. But first I followed a hint and looked up a place further in the South, near Ahlersbach. Quite near a path through the forest a white Cephalanthera rubra with a slight hue of pink! The pink colour is well visible in the buds, where the remaining pigments are more concentrated than in the opened flower.

At the second place near Huenfeld, characterized by an old beech mentioned by Marco Klueber in his great book about “Orchids in the Rhoen” the albiflora plants of Cephalanthera rubra are splendidly flowering. The Swedish botanist L. Anders Nilsson showed (in an article in Nature, 1984) that Cephalanthera rubra mimics the floral coloration of Campanula in the visual system of bees in order to be pollinated by them, especially by male bees of the genus Chelostoma. Since Cephalanthera rubra is flowering before Campanula, they are quite attractive for the bees. It would be interesting to see how bees are reacting to the albiflora forms of Cephalanthera rubra.

With regard to pollinators my visit on June 24th had a special highlight when I saw a wasp (Argogorytes mystaceus) pollinating Ophrys insectifera. The insect pseudocopulated two flowers in a timeframe of more than seven minutes.

The evolution of certain orchid species is far from being finished. The Burren, a region in County Clare at the Western coast of Ireland, illustrates this fact by its manifold colour varieties of Dactylorhiza fuchsii. In 1988, R.M. Bateman and I. Denholm, came to the result that the Burren populations of Dactylorhiza fuchsii show more often a lack of the purple pigment anthocyanin than plants in other regions of the British Isles:

(Source: R.M. Bateman/I. Denholm: A reappraisal of the British and Irish dactylorchids, 3. The spotted-orchids. In: Watsonia 17 (1988), p.332)
When exploring the fascinating area around the Lough Gealain and the Mullaghmore mountain these results seem to be quite realistic. In other areas as well there are many plants, which have bright or white flowers but still retain anthocyanins visible in the markings of the labellum. In a limited area of 40 square meters in the region of Rockforest, northeast of Corrofin, I counted 50 flowering Dactylorhiza fuchsii (in addition to 7 with buds) with the following characteristics:

Among the other plants in this area in the middle of a vast limestone pavement there are Orchis mascula, Geranium sanguineum, Rosa pimpinellifolia, Calluna vulgaris, Lotus corniculatus and Pteridium aquilinum.

The following image illustrates the broad variety not only of colours but also of the labellum forms of Dactylorhiza fuchsii in The Burren (some of the examples obviously showing a certain introgression with Dactylorhiza maculata). It becomes clear that most plants have less floral anthocyanins than continental populations of the species – for example the large forest populations in the French region Causses with its deep pink flowers. The pigments are first reduced in the sepals. This reduction continues in the base colour of the labellum. Then the markings of the labellum are reduced, often only a small rest is retained at the mouth of the spur. Even the very white flowers still have coloured pollinia but their colour is less intense. There is also a wide variety of labellum forms. Especially the central lobes largely differ. And there is the extreme case of a white flowering plant whose lateral lobes are reduced to a minimum (lowest row in the middle).

Most Irish and British botanists stress that the Dactylorhiza fuchsii var. okellyi (some authors view this as subspecies or even as species) must not be mixed up with the albiflora forms of fuchsii. Anne and Simon Harrap (Orchids of Britain and Ireland, 2005) are writing: “A lot of controversy surrounds okellyi” and explain: “In The Burren and elsewhere these classic white-flowered okellyi are just part of a population of plants with a variable flower colour”. Brendan Sayers and Susan Sex (Ireland’s Wild Orchids, 2008) stress that Dactylorhiza fuchsii var. okellyi flowers late, beginning in July. The photo in their field guide shows a flower with a labellum, which is deeply divided into three lobes. Charles Nelson (Wild Plants of The Burren and the Aran Islands, 2008) indicates that Dactylorhiza fuchsii f. okellyi flowers from June to August and has pure white flowers “without any pink tints or marks” and a “lip flat with 3 almost equal, deeply-cut lobes”. According to Pierre Delforge (Guide des orchidées d’Europe, 2005), who mentions a flowering period from May to July, the labellum has a maximum width of 8 mm (in contrast to fuchsii with 8-16 mm). Pat O’Reilly and Sue Parker (Wild Orchids in The Burren, 2007) have noted that “groups of pure-white orchids … are more likely to be O’Kelly’s Spotted-orchids than single plants, which might be just very pale examples of the Common Spotted-orchid”. When exploring the limestone pavements between Poulsallagh and Rockforest you’ll find lots of white-flowered Dactylorhiza fuchsii, rather small and with a pyramidal spike in the beginning of flowering, which are very similar to pink-flowered plants of the species – they should be considered as albiflora forms. Two times I found a pair of taller plants, very slender and with a distinct appearance of spike or flowers, which could be addressed as Dactylorhiza fuchsii var. okellyii.

Dactylorhiza maculata also tends to develop very pale flowers in The Burren. But most of them retain at least a faint marking on the labellum. The variety of Dactylorhiza maculata (the plants in Ireland are in general addressed as Dactylorhiza maculata ssp. ericetorum) seems to be not so wide as the variety of Dactylorhiza fuchsii. The relative frequency of Orchis mascula f. albiflora is in the same range as in continental Europe. Among thousands of plants – Orchis mascula being the most frequent orchid of the region – I’ve seen just two white ones. And there wasn’t one single albiflora form of Dactylorhiza incarnata or Dactylorhiza majalis ssp. occidentalis (on the Aran Island of Inisheer).

Compared with the relative stability of the other plants, the broad variability of Dactylorhiza fuchsii in The Burren clearly shows that this species is still in a state of an ongoing evolutionary process. It can only be speculated why Dactylorhiza fuchsii in The Burren prefers brighter or even white flowering – in the midst of an abundance of pink and purple flower colours on the meadows of the region.

Most white flowering orchids have a last hint of colour in its petals, its spur or ovary. In these cases, the ability to produce pigments is clearly reduced, but obviously not totally suppressed. Studying albiflora species of Gymnadenia odoratissima in the Dolomite Alps, some plants show a rest of rose colour (left) while others still have a violet hue (right) – especially visible in the buds and in the ovary. At the slope of a meadows near the Schlern mountain there are about ten plants of Gymnadenia odoratissima with more or less white flowers.

It seems that there are different pigmets suppressed in both cases. Only plants with all colour pigments suppressed have totally white flowers. That is also clearly visible with Gymnadenia conopsea at the Puflatsch alm of the Dolomite alps.

The Puflatsch Alm above the alpine village Siusi is famous for its colour varietes of Nigritella nigra ssp. rhellicani. As part of a field trip in the Dolomite Alps I had the chance to explore this region with an altitude of 1990 to 2150 metres for four days. Especially rich is the flora in the surroundings of the Arnica Huette, where thousands of Nigritella and Gymnadenia conopsea are flowering in mid July. Additional orchid species are Pseudorchis albida and – in depressed areas – Dactylorhiza majalis.

Among the flowering Nigritella colour varietes are quite common, even though I couldn’t find a totally white one. All the pale-yellow or white flowering plants still had single flowers with a slight rose hue on the edges of the lip. The most common colour variety is an inflorescence with light red flowers in the upper and pale flowers in the lower part.

Not rare at all are hybrids of Nigritella nigra ssp. rhellicani with Gymnadenia conopsea. Their carmine flowers are glowing in the meadows. Interesting was a hybrid of a colour variety of Nigritella nigra ssp. rhellicani with Gymnadenia conopsea, resulting in an inflorescence (left) much lighter than with common hybrids of both species (right):

Following the hint of an orchid friend, today I’m travelling to the Hesse part of the mountain region Rhoen. Northeast of Fulda, near Huenfeld, a path is leading through a long stretch of forest. Along a slope there are flowering countless Cephalanthera rubra. And directly at the path there are three plants with white flowers. Die first still has a slight violett hue. The second is totally white, but already slightly withering, and the third has only one, relative closed flower.

Along the slope there are more than one hundred Cephalanthera rubra, Gymnadenia conopsea and Epipactis atrorubens. There is one Gymnadenia conopsea flowering almost white, with a last hue of violett in the flowers.

South of Fulda and west of Bad Brueckenau we follow another hint and find a hilltop with an impressive  grassland vegetation. In the lower range there are numerous Gymnadenia conopsea. Farther above, some meters from the forest, there are Epipactis atrorubens flowering -- and among them some plants, which are not flowering red-brown, but yellowish-green! While strolling this wonderful slope, I’m seeing a white flowering Gymnadenia conopsea var. albiflora! It is a strong plant with more than 30 flowers, the upper ones still as buds. On the hill there are many butterflies and wild bees. The white flowering Gymnadenia is offering them with its spur filled with nectar as an interesting alternative.

Limodorum abortivum: Albiflora form (left; Photo: N.Griebl) and common form (right)

Limodorum abortivum is one of the rarest albiflora forms of orchids. Sometimes, literature is mentioning the existence of white flowering plants, e.g. Horst Kretzschmar states in his new “Die Orchideen Deutschlands und angrenzender Laender” (Wiebelsheim 2008), p.163: “In Southern Europe, there is a broad variation of flower colours, from white to purple to red, these colours have not been observed in Germany up to now, though.”  Norbert Griebl in Austria has sent me a photo of an albiflora form he found in northern Greece. He observed that the plant has a green stem and green sheathing leaves, “which prooves that Limodorum is not totally living saprophytic” (=myco-heterotrophic).

Compared with the violet stem colour of the regular form the green colour of the stem is indead striking. The existent chlorophyll is obviously covered by dominant anthocyanins. When these purple pigments are absent – as it is the case with the albiflora form – the green chlorophyll colour becomes clearly visible. A study published in 2006 (M. Girlanda, M. A. Selosse, D. Cafasso, F. Brilli, S. Delfine, R. Fabbian, S. Ghignone, P. Pinelli, R. Segreto, F. Loreto, S. Cozzolino and S. Perotto: Inefficient photosynthesis in the Mediterranean orchid Limodorum abortivum is mirrored by specific association to ectomycorrhizal Russulaceae. In: Molecular Ecology 15, 2006, S. 491-504) recognizes the existence of chlorophyll but stated that Limodorum abortivum’s photosynthesis “was found to be insufficient to compensate for respiration in adult plants”. It would be interesting to know how the albiflora orchid is behaving in this regard and if it is also dependent on nutrition by fungi.

The magazine of the Arbeitskreise Heimische Orchideen (AHO) has an article of Norbert Griebl in its latest edition giving an overview about the Dactylorhiza species in Austria. His contribution presents two photos of white varieties – a Dactylorhiza traunsteineri, the photo taken at lake Kochel in Bavaria,and a bright flowering Dactylorhiza incarnata, which is defined als Dactylorhiza incarnata f. ochrantha- with a yellowish accent in the lower part of the inflorescence but not as yellow as Dactylorhiza incarnata ssp. ochroleuca – Griebl views this taxon not as a subspecies but as a species of its own, since he argues that there are almost none hybrids between incarnata and ochroleuca.

In an article about maintenance of biotops in Rhineland-Palatinate Juergen Passin mentions a habitat near Vallendar on the Rhine “Orchis militaris, also var. albiflora”.

Sebastian Sczepanski und Peter Rolf schreiben in einem Beitrag für die AHO-Berichte: “Wie bei allen Dactylorhiza-, Orchis- und Gymnadenia-Arten, so treten auch bei D. praetermissa gelegentlich völlig weißblütige Pflanzen auf.” (Das Übersehene Knabenkraut, Dactylorhiza praetermissa (Druce) Soó, in Deutschland – ein Beitrag zur Orchidee des Jahres 2008. In: Berichte aus den Arbeitskreisen Heimische Orchideen, Jahrgang 25, Heft 1, 2008. S.37). Günther Blaich dokumentiert das Foto einer solchen Pflanze in Großbritannien.

Noch eine Fundmeldung von Orchis quadripunctata “albiflora” vom Monte Gargano bei Monte San Angelo – aufgenommen im April dieses Jahres und zu sehen bei TrekNature – auch diese Pflanze ganz weiß, ohne Punktzeichnung auf der Lippe.

Der Naturfotograf David Greyo hat im südfranzösischen Departement Aveyron eine Anacamptis morio dokumentiert, die neben Blüten in der Standardfärbung eine einzelne Albiflora-Blüte aufweist. Das ist selbst mit Blick auf die besondere Variationsneigung von morio eine verblüffende Entdeckung und wirft die Frage auf, ob einzelne Blüten einer Pflanze ihren eigenen genetischen Bauplan haben.

Photo: David Greyo

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