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Sunday, September 01st, 2013 | 

Dactylorhiza saccifera f. albiflora
Though Dactylorhiza fuchsii is developing albiflora forms more often than other European orchids, especially in certain regions (Ireland, some German regions), there are no mentions of white-flowered Dactylorhiza saccifera. Both species are diploid and related with each other. Exploring the marsh areas of the Smolikas mountain in mid-June in Northern Greece, I’ve seen this splendid Dactylorhiza saccifera f. albiflora on wet grounds, in a height of 1200 m, embedded in Marsh Horsetails (Equisetum palustre) and accompanied by Dactylorhiza baumanniana and Neottia ovata.
Dactylorhiza saccifera f. albiflora

In the same area there was also a white-flowered Gymnadenia conopsea with a slight purple hue.

Another highlight of the field trip to Northern Greece: three white-flowered Dactylorhiza cordigera on a clearing in the Vitsi mountain range near Kastoria. Those were surrounded by more than 1,000 cordigera plants with their characteristic deep purple colour.
Dactylorhiza saccifera f. albiflora

The picture of albiflora forms of late flowering orchids in Northern Greece was completed by a Dactylorhiza incarnata f. albiflora near the village of Chrisi:
Dactylorhiza incarnata

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Saturday, March 16th, 2013 | 

aho29_2 Two white flowered forms of Orchis are presented by Norbert Griebl in the latest edition of “Berichte aus den Arbeitskreisen Heimische Orchideen” (29/2012, 2, p.94-110). The contribution gives an overview of the seven Orchis species in Austria and shows their distribution maps. About Orchis spitzeli he notes: “In some years white or whitish plants appear at the location in Salzburg.” The paper has a photo of a second white-flowered plant, an Orchis mascula subsp. speciosa fo. albiflora. In the same edition of the “Berichte”, Adolf Riechelmann decribes his field trip to Ibiza and mentions an apochrome specimen of Ophrys dyris, found at the southern tip of the Mediterranean island. But the main article of the edition is contributed by Werner Hahn: In the footsteps of Christian von Steven. Searching orchids and pollinators in the Crimean mountains 2011 and 2012 – an exciting study of the orchid flora of the peninsula and as well as of a special chapter of the history of botany.

Tuesday, October 23rd, 2012 | 

On the occasion of a visit to Prague I looked up some specimens at the Charles University Herbarium (PRC). In order to help a friend, I searched for the holotype of a plant which was described by the Bohemian botanist Ignaz Friedrich Tausch as Ophrys purpurea (Flora; oder, (allgemeine) botanische Zeitung. Regensburg, Jena 1831) – now regarded as a synonym of Ophrys apifera or as Ophrys apifera var. tilaventina. The holotype was said to be in the herbarium in Prague, so I searched several packages of Ophrys specimen there, with the much appreciated help of PRC’s curator Jan Stepánek.

The holotype of Ophrys purpurea was not there, but I found an interesting specimen collected by the French botanist Jean Michel Gandoger (1850-1926):
Ophrys apifera
The description carries the information that Gandoger collected this plant in 1879 near Algier as Ophrys apifera f. elata, formerly described by Tausch as Ophrys purpurea:
Gandoger specimen

At the end of my visit I searched a further package of specimens with dried Orchis plants – hoping to find a albiflora specimen. Instead I detected a specimen collected by Tausch as Jan Stepánek confirmed by examining the hand-written label with the nomber “1470″ attached to the stipe of the plant:
Orchis mascula
A further label written by an unknown person has the information: “Orchis mascula L. vom berge Rhadisken bei Leitmeritz” – this information matches the catalogue of “Fundorte der Flora Boehmens nach weiland Professor Ignaz Friedrich Tausch’s Herbarium Florae Bohemicae alphabetisch geordnet von Johann Ott”, published 1859 in Prague:

So who was this Ignaz Friedrich Tausch? The Bohemian botanist was born on January 29th, 1793, in Udrči near Karlovy Vary. After his thesis about “De inflorescentia” (1835) he was director of the botanical garden of duke Canal de Malabaillas in Prague. He studied a broad spectre of plants and published “Bemerkungen über einige Arten der Gattung Paeonia” (1828) as well as his Flora Bohemiae (1831). Tausch was all his life rather poor, Stepánek told me. So he sold dried plants ot different herbariums. Tausch died on 8th September 1848 in Prague.

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Saturday, July 14th, 2012 | 

Most orchid species don’t like acid boglands – but there are two rare exceptions: One is geographically widely distributed from Belgium to Northwestern Germany and Scandinavia and is mostly addressed as Dactylorhiza sphagnicola. The other grows only in the Danish region of Thy: Just a few hundred meters behind the coastline of the North Sea there is a population of white-flowered orchids which have been described by Henrik Ærenlund Pedersen as Dactylorhiza majalis subsp. calcifugiens (in: Nordic Journal of Botany, 2004). In 2007, Sebastian Sczepanski and Karel Kreutz argued it would be more appropriate to regard these plants as a subspecies to Dactylorhiza sphagnicola – while Pedersen und Mikael Hedrén are viewing sphagnicola only as another subspecies of Dactylorhiza majalis. Apart from colour, the morphological differences of the single flowers of Dactylorhiza majalis subsp. majalis (left), calcifugiens (middle) and sphagnicola (right) are difficult to discern:
src="" alt="Dactylorhiza" />The spur of Dactylorhiza sphagnicola is a bit longer than that of D. majalis subsp. calcifugiens. And the leaves of the latter are spreaded in a broader angle than those of D. sphagnicola:
Dactylorhiza majalis subsp. calcifugiens
The single flowers don’t show any hue of purple, even the pollinaria lack Anthocyanin. There is rather some yellowish hue in the center of the flower, slightly reminding of Dactylorhiza incarnata subsp. ochroleuca. In contrast to other populations of albiflora forms, e.g. with Dactylorhiza fuchsii, there are no gradual differences in the loss of colour pigments – all the plants are consistent in the white colour of their flowers.
Dactylorhiza majalis subsp. calcifugiens
Visiting the region, I found calcifugiens at two places, one near the small fisher village of Lild Strand with only three plants, the other further to the south at a bog called Nissekaer with about 150 plants. Surrounded by dunes this place is a natural depression (danish: “kaer”) with a length of about 1500 and a width of about 250 meters:
Google Earth image of Nissekaer
In mid-June the orchids are just in the beginning of flowering. Most Dactylorhiza majalis subsp. calcifugiens are growing at the edges of the wet places, not in the midst of them as it is the case with Dactylorhiza sphagnicola in the Venn moors in Belgium. And the calcifugiens plants are quite smaller, reaching just a height of up to 31 cm. Neighbouring plants are Sphagnum palustre, Equisetum fluviatile; Eriophorum angustifolium, Menyanthes trifoliata, Vaccinium oxycoccus, Calluna vulgaris, Trientalis europaea and even Drosera rotundifolia – most of those plants are clear indicators of acid soil. Among the shrubs there is the dominant Myrica gale, which is used by the brewery of the near-by town Thisted.
Some calcifugiens plants show a broader labellum, indicating a possible hybrid influence of Dactylorhiza maculata – similar to the sphagnicola plants of the Venn region.
Dactylorhiza majalis subsp. calcifugiens
Among all the white-flowered orchids in the Nissekaer bogland I found two purple-flowered plants which might be a hybrid of Dactylorhiza majalis subsp. calcifugiens and Dactylorhiza maculata (left) and a Dactylorhiza maculata still in buds (right):
Dactylorhiza calcifugiens x maculata
As a visiting and possibly pollinating insect of Dactylorhiza majalis subsp. calcifugiens there was a species of the genus Syrphida – as I’ve seen also with Dactylorhiza sphagnicola in the Venn moor (left)

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Monday, June 18th, 2012 | 

Cephalanthera damasonium It is well studied that Cephalanthera damasonium belongs to those orchids which can live without chlorophyllum – together with other species of the tribe of Neottieae or the genus Epipactis. While exploring a mixed forest near Lahnstein (Rhineland-Palatinate) together with Ingo Beller of the Arbeitskreis Heimische Orchideen (AHO) Rheinland-Pfalz, we found a group of three albino plants in addition to three green-leafed Cephalanthera damasonium. Of the apochromic plants one has two flowers, one only one flower and one has no flower. Those albino plants receive their organic carbon with the help of fungi. A study of V. Tranchida-Lombardo, M. Roy, E. Bugot, G. Santoro, Ü.Püttsepp, M. Selosse and S. Cozzolino, published in 2010 in Plant Biology suggests that the albino Cephalanthera damasonium may be viewed as “an intermediate step in the evolutionary emergence of mycoheterotrophy”, or of the ability to be nourished both by fungi and photosynthesis. By means of genetic analyses the authors declare: “Albinos could be either permanent mutants, as suggested by phenotype stability over the years, or transitory phenotypic stages, in which genes involved in the photosynthetic pathway can switch off depending on micro-environmental conditions (e.g., the amount of C resources provided by the nearby fungal mycelia or tree roots) that prevent greening.”

Wednesday, May 30th, 2012 | 

Dactylorhiza fuchsii f. albiflora

In addition to the Irish region of The Burren and the Hesse location of Biebergemuend there is a further location where Dactylorhiza fuchsii tends to white-flowered forms in big numbers: In a birch grove near the village of Wolken in the upper Moselle valley there are several hundred plants with a clear trend to bright and white flowers. A count of a random sample resulted in 13 per cent of white flowers without any markings on the labellum. Further 38 per cent of the plants have a white base colour with pink marks. The differences in flower colours correspond with the results found in the other two regions and are even a little bit more accentuated. These results may give further evidence to the assumption that Dactylorhiza fuchsii is in the midst of an evolutionary process which also changes the phenotype of the species.

Percentages of flower colours in different locations
different locations of Dactylorhiza fuchsii

The following tableau shows the wide range of fuchsii flowers found on that location. The brightest forms also lack the Anthocyanine pigments in the pollinaria as the lowest row of examples and the following macro photo demonstrate.
Dactylorhiza fuchsii

Dactylorhiza fuchsii f. albiflora

The only other orchids in the forest are Platanthera chlorantha and an Epipactis spec. – while in the other two locations with abundant albiflora forms of Dactylorhiza fuchsii there are also the earlier flowering Orchis mascula (Burren) and Dactylorhiza majalis (Biebergemuend), both flowering in pink and both – as well as Dactylorhiza fuchsii – trying to attract pollinators with nectar deception. Among other plants in the birch forest, a former gravel-pit and now a nature reserve called “Kuhstiebel”, are Orthilia secunda, Fragaria vesca and Tussilago farfara. But the dominating plant there as well as in a nearby marsh area is Dactylorhiza fuchsii with mostly spotted leaves – even in the case of the white-flowered plants: Dactylorhiza fuchsii f. albiflora

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Tuesday, May 01st, 2012 | 

Two specialists of the orchid flora on Corsica, Wolfram Foelsche and Klaus Cord-Landwehr, have described a new Dactylorhiza species: Dactylorhiza cyrnea belongs to the group of Dactylorhiza maculata and shares its characteristics of a tetraploid set of chromosomes. In their article “Dactylorhiza cyrnea und die Taxa der Gattung Dactylorhiza auf Korsika”, published in the Journal Europäischer Orchideen (Vol. 44, Heft 1, April 2012), the authors review the findings of Dactylorhiza insularis, Dactylorhiza sambucina and Dactylorhiza saccifera on Corsica and note that there is no Dactylorhiza majalis confirmed for this Mediterranean island. The plants described as Dactylorhiza cyrnea is morphologically similar to the diploid Dactylorhiza fuchsii. They are rather elongated and grow in humid locations. The leafs are mostly vaguely spotted but may also be unspotted. The flowers have a markedly thin spur (contrasting the thick spur of D. saccifera) which is shorter than the ovary. Their colour is a bright pink, with a purple labellum pattern. The population described by the authors “offers a very consistent appearance” – but there are also white flowered plants occasionally, the authors note. The photo above on the right, generously sent to me by Wolfram Foelsche, is also included in the article.

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Sunday, October 23rd, 2011 | 

Anacamptis morio
Oops, that’s a quite special Anacamptis morio which Norbert Griebl has found near Sittendorf, southwest of Vienna. In addition to the absence of anthocyanins (which happens quite often with this species) the lip has the same form as the sepals, with green veins. Here, not only the colour of the flower has changed, but also its morphological pattern.

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Sunday, August 01st, 2010 | 

Cephalanthera rubraThis 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.

Cephalanthera rubraAt 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.

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Wednesday, June 16th, 2010 | 

Dactylorhiza fuchsii
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:

Percentage of Dactylorhiza fuchsii lacking Burren other regions
leaf markings 43 13
labellum markings 48 15
labellum anthocyanins 48 12
all floral anthocyanins 35 8
all anthocyanins in flowers, stem, leaves and bracts 25 6

(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:

Dactylorhiza fuchsii with
dark pink flowers 0
medium pink flowers 11
bright pink flowers 17
white flowers and line markings 3
white flowers and dot markings 17
white flowers without markings 2
total 50

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).
Colour varieties of Dactylorhiza fuchsii
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, Dactylorhiza fuchsii ssp. okellyi 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.

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