Gymnadenia conopsea x Nigritella rhellicani with white flowers

In the latest edition of the Berichte aus den Arbeitskreisen Heimische Orchideen (Jg 34, Heft 2, 2017, S.123-145), Norbert Griebl presents the diversity of alpine orchid hybrids on the meadows of the Golzentipp mountain (2317 m) in Eastern Tirol. Among them there is also a white-flowered hybrid of Gymnadenia conopsea and Nigritella rhellicani (Gymnigritella suaveolens). Griebl views two possible options:

  • a white-flowered Nigritella rhellicani, which can be rarely found at the Golzentipp, and a white-flowered Gymnadenia conopsea, which is not rare in this region, have hybridized
  • a spontaneous mutation of a hybrid affecting the production of color pigments.

Griebl sees a higher probability for the second case. This is supported by the fact that there have been no reports of such an Albiflora hybrid in other regions with color varieties of Nigritella rhellicani and white-flowered Gymnadenia conopsea, for example on the Seiser Alm in Southern Tirol – though these Suaveolens hybrids can be regularly seen there.

Orchids adapt to colour preferences of pollinators

This hypothesis has been formulated by Hannes Paulus in a contribution for the latest edition of the Journal Europäischer Orchideen (Hannes F. Paulus: Zur Bestäubungsbiologie der Gattung Ophrys in Nordspanien: Freilandstudien an Ophrys aveyronensis, O. subinsectifera, O. riojana, O. vasconica und O. forestieri. J. Eur. Orchideen. 49 (3-4): 427-471).

In this article the author studies both populations of Ophrys aveyronensis in Southern France and Northern Spain – the last one termed as Ophrys aveyronensis subsp. vitorica. According to Paulus it is just one species, because both are pollinated by the bee Andrena hattorfiana.

Paulus points to the fact that this bee is specialised on the widow-flower (Knautia). The pink inflorescence of this plant has the same colour as the perigone, i.e. the sepals and petals, of Ophrys aveyronensis. The expert of Ophrys is stating: It can be expected that this is not just a mere chance but an adaptation to the main nourishing plant of the pollinator. Knautia shows at the same time more deeply pink flowers as flowers tending to white.

This evidence is confirming the approach to also look for other plants when we search for reasons why albiflora forms of different orchid species are more often in certain places.

Trip reports and expert debates


After 21 years, the Orchid Conference of Sundheim, Germany, has found a new place: About 50 participants met in Kehl-Neumuehl in the beginning of October, in a Protestant community centre, only a few kilometres away from the traditional meeting place in the old “Stierstall”. The history of the conference has been described by Werner Hahn in the latest edition of Berichte aus den Arbeitskreisen Heimische Orchideen (Jg. 34, Heft 1, 2017, S.234-237).

This year, the program focused on travelling reports from Iran (Jean-Marc Haas), Croatia/Greece (Stefan Hertel) and Israel (Helmut Presser). Jean-Marc showed findings of Ophrys zagrica in an altitude of 2300 metres, Ophrys khuzestanica and Ophrys strausii as well as Tulipa stapfys in the midst of dry pebble, Fritillaria imperialis or Iris acutiloba subsp. longipetalis. Stefan followed the transition of Ophrys incubacea to Ophrys mammosa and presented Ophrys cephalonica from the island Kephalonica with its long narrow sepals. In the beginning of March, Helmut has explored Mount Karmel and Mount Meron. Among the species which grow there are Ophrys carmelii and Orchis galilaea.

Typical for the delight of orchid scientists in disputes was a discussion after a lecture of Wolfgang Wucherpfennig. He stressed his criticism on the taxon Ophrys lutea var. subfusca for an orchid flowering in Northern Africa. The description of Heinrich Gustav Reichenbach in 1851 – along with two not so significant sketches – had resulted in the fact, “that the name subfusca has been used for many different species in the following years”. The situation which has been developed is quite disturbing. Wucherpfennig pleaded anew: “The name subfusca should not be used anymore but should be viewed as a nomen dubium.” But Karel Kreutz pointed out that a herbar finding in the Reichenbach collection which is indicated as “Ophrys fusca” should be viewed as the holotype of Ophrys lutea var. subfusca. And he stated that he has found according plants in Algeria – documented in his paper C.A.J. Kreutz/L.Lewis: Typification of Ophrys lutea var. subfusca Rchb. f., invalidity of the name Ophrys murbeckii H.Fleischmann. In: Journal Europäischer Orchideen Vol. 46, 1/2014.

The participants persued their discussion for a while – without answering the question why this detail of science history should be so relevant. In the long term, questions of the preservation of orchid species are much more important than any taxonomic debates.

Albiflora couple in the valley of Kinzig

With Orchis mascula, the albiflora forms are not rare. But in many cases the pattern of violet points is preserved in the lips – even if the production of Anthocyanin is disturbed, the last reserves are obviously kept for this pattern. This year, a pair of Orchis mascula was flowering entirely white in the valley of Kinzig (in Hesse, Germany): Orchis mascula

The leaves of these plants are unspotted, while the violet-flowered Orchis mascula often have intense spots.

Nearby, Orchis purpurea was beginning to flower. The third orchid species on this meadow in the “Schdänerer Weiberch”, as it is called in the Hesse idiom at the beginning of the path, is Himantoglossum hircinum. In May, the lizard orchid is just beginning to develop its inflorescence.

A bit further on the path, at the edge of a small forest, there was another albiflora form of Orchis mascula, also without any violet color in the flower, but with a marked green stigmatic lobe. The green hue is also preserved in the spur.

Orchis_mascula f. albiflora

Orchis mascula f. albiflora

(with many thanks to Matthias Raschka for indicating me the location)

The dynamics of evolution on the Dactylorhiza meadow

Dactylorhiza fuchsii f. albiflora

Seven years after my first observations on a meadow in the valley of the small river Bieber in the northern region of the Spessart mountains I visit this special habitat again. The meadow with a wet trench in the middle is in full flowering. My estimation is that about 600 Dactylorhiza plants are flowering here: About 500 Dactylorhiza fuchsii, 50 already withered Dactylorhiza majalis and 50 hybrids Dactylorhiza fuchsii x majalis. Among the accompanying plants are Rhinanthus spec., Pedicularis palustris, Campanula patula and even a group of Arnica montana. Platanthera bifolia is just showing the first flowers.

Among the 500 Dactylorhiza fuchsii are about 400 plants with very bright flowers and a pattern of bright violet loops on the labellum. Furthermore, I count 22 albiflora forms with white flowers and non-spotted leafs. This is a rate of 4.4 per cent – much higher than you could expect if those were just spontaneous mutations. When I first visited this place on 20.6.2010 and 1.6.2012, there were less albiflora plants. Only about 80 of the 500 Dactylorhiza fuchsii have an intense violet colour. This meadow presumably has its own dynamics of evolution, developing increasingly bright forms of Dactylorhiza fuchsii.

Dactylorhiza fuchsii f. albiflora

Dactylorhiza fuchsii f. albiflora

There might be a relation with the learning experience of bees: With the earlier flowering Dactylorhiza majalis, many pollinators may have already made the experience that there is no nectar in the spur of violet orchid flowers. While developing brighter flowers, Dactylorhiza fuchsii might counteract this learning experience. Young honey bees still have to make this experience like this one on the flowers of the hybrid Dactylorhiza fuchsii x majalis:

Albiflora abundance on Sardinia

Anacamptis morio subsp. longicornu
Anacamptis morio subsp. longicornu

There are two forms of albiflora mutations with orchids:

  • the spontaneous growth of a white-flowered form as the consequence of a genetical defect in the process of creating anthocyanine pigments, traditionally called a freak of nature, Occurence: 1-5 among 1000 plants
  • populations of white-flowered mutations as the result of an evolutionary adaptation to environmental conditions, e.g. the concurrence of other violet-flowered food-deceptive orchids, occurrence: 10 to 500 among 1000 plants

I’ve found both forms on the Sarcidano plateau, a central region of Sardinia.

On lengthy hikes between Láconi, Ortuabis und Santa Sophia I’ve seen only one single albiflora form of Orchis mascula subsp. ichnusae, with the crimson marking of the flower labellum still preserved:

Orchis mascula subsp. ichnusae
Orchis mascula subsp. ichnusae

Before, I had already seen a single Orchis anthropophora without its typical flower colouring, in the forest of Domusnovas, on southern Sardinia:

Orchis anthropophora
Orchis anthrophora

Much more frequently are the white-flowered forms of Anacamptis morio subsp. longicornu in Sarcidano. Respectively one third of the overall several thousand plants in this region has the dark violet colouring, a bright violet (or rose) colouring or are white-flowered.

Anacamptis morio subsp. longicornu
Anacamptis morio subsp. longicornu
Anacamptis morio subsp. longicornu
Anacamptis morio subsp. longicornu

You can’t find this accumulation of albiflora forms in other regions on Sardinia which I’ve visited, neither at Domusnovas/Iglesias nor in the North or at Monte Albo. There, Anacamptis morio subsp. longicornu are consistently flowered in the regular violet. The albiflora forms of Sarcidano possibly have an evolutionary advantage. In this region there are also many Orchis mascula subsp. ichnusae giving pollinators as bees the learning experience that there is no nectar in the spur at flowers with this colour and form. In the other regions Orchis mascula subsp. ichnusae was less common or not present.

Anacamptis morio subsp. longicornu
Anacamptis morio subsp. longicornu

Albiflora orchid in Nepal

Photo: Bhakta Bahadur Raskoti

The Nepalese orchid expert Bhakta Bahadur Raskoti found an albiflora form of Neottianthe cucullata together with its regular form. This orchid is distributed in central and western Nepal at altitudes of 3700 to 5000 m. It’s flowering in August. The albiflora form has been described in 1995 by the Chinese botanist P.Y. Fu. The species has its western distribution limit in Poland.

Albiflora greeting from Lithuania

Before the end of the year I received special finding reports from Lithuania: Bernd Gliwa has found a white-flowered Dactylorhiza traunsteineri, only once in 2006. The location is in a lime marsh between Kaunas and Šiauliai. It’s quite remarkable that this Dactylorhiza species is quite rare to see with white-flowered forms.

Photo: Bernd Gliwa, 21/06/2011, near Dievogala/Lithuania

In this nature reserve he has also photographed this Dactylorhiza incarnata. In this area there are also Dactylorhiza incarnata subsp. ochroleuca (with W. Eccarius, Die Orchideengattung Dactylorhiza, 2016 it’s again a species of its own, D. ochroleuca), Dactylorhiza incarnata subsp. cruenta and Dactylorhiza majalis subsp. baltica.

Bernd Gliwa has also studied the dragonflies of Lithuania and just published a monograph about them: Lietuvos laumžirgiai.

Ophrys pigmentation demonstrates bilateral symmetry

While travelling in Attica, Greece, Marco Klüber photographed this special Ophrys helenae which demonstrates the bilateral symmetry of orchid flowers. The left half of the lip shows the regular red-brown coloring while the right half is hypochrome, with a partial loss of pigments. The lighter red hues are preserved, in addition to the chlorophyll in the lower edges.

Orchid flowers have a bilateral symmetry – as it’s the case with beetles or the human face elsewhere in nature. Other plants like the flowers of liliaceae have a radial symmetry with three or more mirror lines.

On his trip Marco also saw the albiflora form of Anacamptis papilionacea subsp. messenica (formerly subsp. heroica):

Wolfgang Eccarius presents his genus monograph about Dactylorhiza

Die Orchideengattung Dactylorhiza

Following the benchmark books about the genera Anacamptis, Neotinea and Orchis (together with Horst Kretzschmar und Helga Dietrich, 2007) as well as a monograph about the genus Cypripedium (2009), Wolfgang Eccarius now has finished his long lasting work and has published a compendium about the Dactylorhiza orchids.

The book closes a big gap and covers a genus of orchids which is especially rich in species and quite widespread in Europe. As he has established in former publications, the author at first gives a comprehensive introduction before portraying 37 species and 46 subspecies. Eccarius explains his methodological approach and offers a summary of the research history, beginning with the plant book of Otto Brunfels published in 1534. The book abstains from giving a system to identify the species  by morphological indicators. But a tree of the genus structure based on genetical research offers a good overview about the manifold Dactylorhiza orchids and their relations.

In his description, Eccarius has taken some main decisions. He abstains from describing varieties and forms, arguing that those terms are “highly problematic” with Dactylorhiza. “The main goal of the author was a genus structure wich matches logical principles as well as the needs of observations in the field.” He stresses that’s it’s more important to differentiate between the ten sections than defining species: “With Dactylorhiza, sections are much easier to define than species.” For example, the Fuchsiae form a section of their own, together with Dactylorhiza saccifera. The section of Majales comprise Dactylorhiza majalis, Dactylorhiza cordigera and Dactylorhiza elata.

Dactylorhiza majalis subsp. calcifugiens
Photo: Peter Zschunke, 16/06/2012, bei Glæde, Dänemark

It’s comprehensible that Eccarius views Coeloglossum viride as Dactylorhiza viridis. The Viridae are presented as a subgenus, in addition to the subgenus  Dactylorhiza. Other taxonomic decisions are more thought provoking. For example when the yellowish Early Marsh Orchid no longer is a subspecies of Dactylorhiza incarnata, but an own species Dactylorhiza ochroleuca – because there are only very few hybrids between both “which justifies the treatment as a single species in the view of the author”.

Difficult are the explanations about the white flowering Dactylorhiza fuchsii in Ireland, which are elevated by Eccarius to the status of a subspecies – while most experts view the okelly taxon of Dactylorhiza fuchsii as a variety. And the morphological description of the author is not quite helpful in the field: “The subspecies is different by its lower growth” – while the photos show rather high plants. And: “The white color of flowers is shown by whole populations and not only by single plants.”

But this is also valid for Dactylorhiza majalis subsp. calcifugiens, which is presented by Eccarius only as a synonym to Dactylorhiza sphagnicola. The book shows a photo of a plant from the German region of Celle which seems to be an albiflora form of Dactylorhiza sphagnicola, but which is quite different form the calcifugiens population in Northern Denmark.

Quite useful are the explanations about Dactylorhiza maculata, which is presented as a west and northern European species, distributed also in Northern Africa and Northern Asia. The color of flower is described as especially variable, from pure white to a soft and light purple.

Eccarius understands the tendency to a color dimorphism (or polymorphism) which is typical for the genus as functionally relevant. With Dactylorhiza romana, sambucina or incarnata this phenomenon serves as a factor, “to avoid quick learning experiences of polinating insects”. This matches with the regionally different tendency of Dactylorhiza fuchsii to develop albiflora forms.

The new book makes big progress in understanding the Dactylorhiza orchids. But for a full perception there is still a lot of research necessary.