In 1991 the Cyclamen Society embarked upon a study of the subgenus Psilanthum Schwarz, more commonly referred to as the Repandum group of species. The aim was to establish the distribution of the various species and subspecies, study the environmental conditions in which they grow, and record statistical information about variations in leaf and petal dimensions, shape, patterns and colour. Representative herbarium collections were to be made. In addition, a number of plants were to be collected (under licence) for further study and to introduce into cultivation genetic material of interesting or desirable forms. A secondary activity was to be the collection of herbarium material and desirable forms of other Cyclamen species found in the areas visited.
In particular, the intention was to answer a number of outstanding
questions, such as those posed by Grey-Wilson (1988):
1. Does ssp.
repandum occur in the Greek Peloponnese, or do the dark-flowered plants
merely represent an extreme form of subsp. peloponnesiacum?
2. Does C. repandum occur anywhere in Crete and do pink-flowered
forms of so-called C. creticum actually represent plants that have
arisen initially by crosses between C. creticum and C. repandum?
3. Do all Cretan forms have a chromosome count of 2n=22.
In 1991 the Society sent an expedition to study C. repandum ssp. rhodense Grey-Wilson on Rhodes (Bailey, 1991) and to study C. creticum Hildebrand on Karpathos (Bailey, 1992). It also sent expeditions to the Peloponnese to study C. repandum ssp. peloponnesiacum f. peloponnesiacum Grey-Wilson, and C. repandum ssp. peloponnesiacum f. vividum Grey-Wilson in 1992 (Johnstone, 1992) and 1993 (Moore, 1993).
The Peloponnese and the island of Rhodes represent respectively the western and eastern ends of a chain of islands which are the remnants of a Tertiary mountain range, part of the Aegean land mass which became submerged in the Pliocene period some 5.2 million years ago. From west to east, these islands or near-islands are the Peloponnese, Kithira, Andikithira, Crete, Kassos, Karpathos, Saria, and Rhodes. In 1994, 1995 and 1996 the Society sent expeditions to Crete the largest of the islands to study Cyclamen creticum, and in 1997 an expedition will go to the island of Kithira to establish whether either C. repandum or C. creticum grow there.
Although Saunders (1979) described C. creticum as 'very abundant in Crete', Turland, Chiltern & Press (1993) gave a relatively restricted distribution with the majority of populations concentrated at the western end, limited populations in the east, and two isolated populations in the centre of the island. The Society's 1994 expedition (Aspland, 1994) visited the western end of Crete recording 20 sites, finding many more populations, and collecting 40 plants (4 graecum, 4 hederifolium & 32 creticum). In 1995 the drier eastern end of Crete was visited (Wiltshire & Dale, 1995), and C. creticum was found to be far more extensively distributed than had previously been recorded. 29 sites were visited and 28 plants collected (1 graecum, 3 persicum & 24 creticum).
The 1996 expedition was to cover the remaining central part of Crete. This area extended from Lefka Ori (The White Mountains) eastwards past Psiloritis (Mount Ida) to a point almost due south of the city of Iraklion, and represented some 50% of the land mass of the island. It was clear from the early planning stage that the expedition would only be able to 'sample' the area, not cover it thoroughly and in detail. Research whilst in the UK had produced a list of known sites and these provided a safety net. A number of other potential sites had been identified, from maps and walking books. However, the work of mapping the distribution of C. creticum required that time be spent investigating areas where there had been no previous sightings as it was as important to establish where it did not grow, as where it did grow. Thus there were days when no Cyclamen were seen. The team drove about 750 miles in two weeks, recording a total of 33 sites for creticum and graecum, carrying out surveys, and collecting 61 plants (46 creticum, 15 graecum).
Melvyn Jope, John Fielding and Martyn Denney flew from Gatwick to Iraklion in the evening of Friday 12th April arriving at their self-catering apartment in Rethymnon in the early hours of Saturday morning. As has been customary for a number of years, a normal package holiday had been booked on grounds of economy, and the fact that it provided the team with a base at which collected plants could be left in a cooler shaded environment. In fact 6 nights were spent in Rethymnon, the remainder being spent in remoter parts of the island in 'rooms'. The apartment had the bare essentials - a bedroom, living room with third bed, 'kitchen' and shower-room (cold water only most of the time).
Saturday morning was spent recovering from the flight and arranging a hire car. In the afternoon the team explored the Kouloukinas mountains, a low range on the north coast of the island between Rethymnon and Iraklion. Two sites for both creticum and graecum were recorded at between 300 and 600 metres. At the higher of the two sites large numbers of creticum were found amongst dense Pistachio scrub adjacent to a large stand of Arum concinnatum.
Sunday, the first full day was spent driving into Psiloritis from the north along the 22km road which leaves Anogia and crosses the northern flanks of the mountain to the Nidha Plateau and the Idaien Cave. Cyclamen creticum was found at 850m growing with the dwarf yellow and white, Orchis pauciflora. Other interesting plants on the Nidha Plateau included Corydalis rutifolia ssp. uniflora and Chionodoxa nana, however, the C. graecum reported by Lionel Bacon was not seen. This being the Greek Orthodox Easter Sunday, the team lunched on the traditional spit-roast lamb complete with all the bits (liver, kidneys and testicles!).
Cyclamen have never been reported from the Asteroussia Mountains on the coast of Crete directly south of Iraklion. These mountains had been clearly identified as an area which should be explored. The trip south across the island was uneventful other than the discovery of a site for C. creticum high above the road where some excellent pink flowered forms were found. The three-bedded 'rooms' in Pirgos were clean and comfortable but brought the second variation on the theme of Greek plumbing (a shower with scalding hot water but no cold). The hills around the town were green and lush with many species of Ophrys and Serapias but no Cyclamen. The Asteroussia Mountains stretching west towards Matala had a number of potential sites on both northern and southern flanks but these felt wrong, were too dry and trampled by goats, and nothing was found except some good colour forms of Gynandriris sisyrinchium and extensive stands of pale yellow Arum creticum in both its green and maroon stemmed forms. Whether C. creticum existed here before the mountains were so extensively grazed is pure speculation.
After a night in Agia Galini the team returned to Rethymnon along the southern flank of Psiloritis. Both creticum and graecum were found more or less as expected at various locations.
The sheer size of the area to be covered during the expedition meant that there was no possibility of returning to any site and on more than one occasion this lead to us arriving at our accommodation in the dark having found a particularly good Cyclamen site late in the afternoon. This happened this same afternoon. We came across a place where creticum grew in dense drifts under trees at the edge of cultivated land. The scent of the flowers was very apparent, carried on the late afternoon wind. By the time we had completed a 50 plant survey (Have you ever tried measuring Cyclamen petals in a howling gale?) it was coming on towards evening and we arrived back in Rethymnon very late.
There followed the most successful day of the entire trip. We decided to visit the Gipari Gorge. This uniquely runs roughly east-west, it is in the centre of the island and ends at the mountain village of Asigonia. We found creticum growing in very dense populations in the wooded river valley, under Quercus coccifera and Platanus orientalis with robust plants of Dracunculus vulgaris just coming into flower. Amongst the creticum there were some quite exceptional leaf forms and flowers with very long petals, some of which were collected: 96252 - petals 38mm long with 360° twist; 96256 - acuminate leaves 98mm x 62mm; 96258 - very bright silver hastate pattern; 96259 - silver leaves with a few flecks of green; 96260 - silver leaves. There was also an interesting plant (96261) which had foliate calyx and corolla. We returned to Rethymnon for the night and spent what was left of the evening packing the day's collections in moss, checking the earlier collections and preparing the whole lot to be left in the apartment for nearly a week.
Early the following morning, we headed south across the island towards the mountain village of Spili, finding further sites for both Cyclamen creticum and C. graecum on the way. After a brief pause to see the large colony of the scarlet endemic Tulipa doerfleri to the east of Spili on the Gerakari road, we made our way back to the south coast at Agia Galini. Over the next few days we explored the gorges and gullies which dissect the coast between there and where the Lefka Ori meet the sea in Sfakia to the west of Chora Sfakion. The south coast of Crete is noticeably warmer and drier than the north of the island, and C. creticum hides deep in damp shady places or at the bottom of the deep gorges, though even here it remains on the north side of rocks and cliffs. Some of the sites are well documented elsewhere (Turland, 1988; Rowland, 1991) and the team confirmed the existence of these populations in all bar one case - the Imbros Gorge.
Two sites were of particular interest: Site 96/23 between Plakias and Kato Rodakino had plants with leaves which were apparently thicker with a more leathery texture than usual. The plants collected here will require further investigation to establish whether they exhibit any morphological or chromosome variation. Site 95/25, the Aradena Gorge, had populations of creticum in very dark crevices, growing as a chasmophyte in limestone rock, and epiphytically in moss with a significant proportion of flowers in varying shades of pale pink (69D, 76B by RHS colour chart, or paler).
We had particularly wanted to find populations of C. creticum at high altitude. Grey-Wilson (1988) gives an altitude range of 0-1250m (up to 4,000 ft) for the plant. The 1994 expedition had found it at 1250m between Agia Irini and the Omalos Plain in Lefka Ori; the 1995 expedition had found it at 1050m above the Lassithi Plain in Dikti; in 1996 we had only found it at 850m in Psiloritis. We were now fortunate enough to find a site at 1000m on Mount Akones which is an outlier to the east of Lefka Ori (site 96/27). The site was very exposed and comprised a north-northwest facing gully with a few stunted trees of Acer sempervirens growing with mountain grasses which supported a flora of (amongst others) Anenome heldreichii, Phlomis fruticosa, Asphodeline lutea, Asphodelus aestivus, Arum idaeum, Daphne sericea, Crocus laevigatus, Hermodactylus tuberosus, Orchis quadripunctata and Ophrys fuciflora. The Cyclamen were only found growing in the gully where they sheltered under rocks which would probably be submerged in the stream in early spring, and around the stunted trees where they could make the most of the scant shade. Few plants were in flower.
After surveying a site for both creticum and graecum on the Askifou Plain, we headed north-west towards Chania - the plan being to spend a night in Platanias in order to further the arrangements for the Society's conference to be held there in September. On the way we found and surveyed an excellent population of C. creticum near the village of Agiou Pantes. The plants ranged around the edge of ancient Cupressus sempervirens woodland and included some excellent silvered leaf forms. Three plants were selected for collection; two with very distinct silver hastate patterns (96494 & 96496) and one with very heavy silver spotting (96497).
Returning along the north coast of the island to Rethymnon, we explored a series of limestone gullies to the west of the town but although these carried a fairly rich flora there was no sign of any Cyclamen and this was another occasion when the team agreed that although it was a likely habitat it simply felt wrong.
Our last evening and much of the following morning was spent back at the apartment checking the condition of the collected plants and herbarium specimens and packing both ready for the journey back to England. In the afternoon of Friday 26th April, we had time for one last trip to explore the north-west foothills of Psiloritis near the Arkadi Monastery at about 450m (1,480ft). Here we found C. creticum growing in a limestone ravine (which still had pools of water left from the torrent which flowed in winter) and in drifts in the wooded hillsides, accompanied by large quantities of Arum concinnatum.
None of the expeditions have seen anything to indicate the presence of Cyclamen repandum in any of its forms on Crete. There is thus no evidence that the 'pink' forms of Cyclamen creticum have arisen as a result of a cross between C. creticum and C. repandum. However, the species are clearly allied and in the Pliocene shared the same land mass. The Society's 1997 expedition to Kithira may contribute to this debate. Cyclamen repandum and C. balearicum have a chromosome count of 2N=20, whilst C. creticum is generally 2N=22. Collections made by the 1994 expedition near Agia Irini (site 94/17) west of the Omalos Plain and at Agios Stefanos (site 94/06) just south of the Rodopou Peninsula, have been subjected to scientific analysis and have been confirmed as having a chromosome count of 2N=20. The collection from Agia Irini confirms the findings of Greilhuber (1989), but as far as we are aware, 2N=20 chromosome counts have not previously been recorded from Agios Stephanos and it is considered that both sites warrant further investigation. The Society obtained permission from the Greek government to collect further plants from both sites in autumn 1996 in order to further this work, and we are greatful to George Handrinos for facilitating this.
The analysis which follows is based on field data collected during the three expeditions to Crete and at the one site on Karpathos in 1991. It must be borne in mind that it brings together statistics collected over four years and therefore differing seasons which may have effected the data collected and thus the conclusions which have been drawn. The data itself has been recorded by different personnel each year and therefore the subjective evaluation of such factors of leaf shape and leaf pattern may effect what is recorded. Still, it is doubtful if such infinitesimal inaccuracies invalidate the results.
At each site which is suitable for analysis, depending upon the size of the population, either 25 or 50 pairs of a leaf and a flower are collected from random plants. The following data is recorded: Leaf length, leaf width, leaf shape, leaf pattern, flower colour, petal length, petal width, degree of petal twist, and colour of leaf reverse. Leaf shape and pattern are determined by matching collected leaves against photographs of designated leaf standards.
The analysis of the field data is based on simple averaging - the author is not a statistician. For simplicity, leaf reverse has been treated as 'red' or 'green' rather than separating out 'pale red' etc. Flower colour has been treated as 'white' or 'pink'. In this context, pink includes all flowers which are not entirely white. This varies from petal colour which can be clearly identified as either 69D or 76B according to the RHS colour chart to such a pale pink as not to register. It also includes flowers with a pink nose or a pink flush.
The expeditions found C. creticum growing at almost every altitude
from 60m to 1250m (200 - 4,100ft).
There were 4 basic habitat types:
Phrygana
(mixed dwarf shrubs, often spiny and aromatic), generally on the northern side
of the island;
Mid to high altitude grassland, with an open situation and
the shelter of rocks;
Deciduous or Cupressus sempervirens woodland
at mid to low altitude in east-west inland valleys;
Southern gorges &
gullies.
The majority of populations were growing with an aspect in the northern half
of the compass. Where the aspect was basically southerly the plants were
growing in deep shade, usually within a gully, often with summer running water.
The degree of shade varied from 0%-90%, to some extent with altitude - the
higher populations being typically more open, possibly simply in the shelter of
rocks. There was no correlation between the percentage of shade and any other
factor, although a subjective view is that the larger leaves occurred in the
deepest shade.
Soil was generally a clay, usually Terra Rossa but sometimes
black/brown, sometimes overlaying limestone and usually with a humus layer of
decaying organic matter. The soil at the Agia Irini, 2N=20 site, differed in
being based on a schist. At most sites there was an identifiable amount of
moisture in the soil, and in some of the gully locations at certain times of the
year the plants would have been submerged in water.
Leaves varied dramatically in size, up to 162mm long by 148mm wide. There was a direct correlation between leaf length and leaf width. 9 sites had a ratio of leaf length to width of 1.1:1, 30 sites had a ratio of 1.2:1,and 3 sites had a ratio of 1.3:1. There was a very slight increase in leaf size as longitude increased i.e. from west to east, however, this was not carried through to Karpathos, where the average leaf size was amongst the 5 smallest found.
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| Leaf Shape 1 | Leaf Shape 2 | Leaf Shape 3 | Leaf Shape 4 | Leaf Shape 5 |
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| Leaf Shape 6 | Leaf Shape 7 | Leaf Shape 8 | Leaf Shape 9 | Leaf Shape 10 |
There was little correlation between longitude and leaf shape, except that west of Dikti there was a significant increase in the percentage of plants of Leaf Shape 1. Generally, there was no correlation between aspect and leaf shape, however at sites with a southerly aspect Leaf Shape 3 became predominant, at the expense of Leaf Shape 1. Except for a slight increase in the predominance of Leaf shape 1 the further north, there was no significant correlation between latitude and leaf shape.
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| Leaf Pattern A | Leaf Pattern B | Leaf Pattern C | Leaf Pattern D | Leaf Pattern E | Leaf Pattern F (All over silver wash) |
Leaf Pattern A predominated at the western end of the island, Leaf Pattern C
predominated in the centre of the island, and Leaf Pattern B predominated in the
eastern end of the island.
The photographs used as Leaf Patterns A, B and C
came from:
A - repandum album, B - repandum ssp. rhodense,
C - repandum ssp. peloponnesiacum
The predominance of a C.
repandum rhodense leaf pattern in the east supports a gradation along the
ancient Aegean land mass from rhodense in the east to peloponnesiacum
in the west. It is the only factor which supports this. However, contrary to
this, at the site in Karpathos, 87% of the population recorded had Leaf Pattern
C.
A percentage of plants had plain green leaves with no pattern, however
all these occurred to the east of longitude 24° 21'E (approximately,
Episkopi, half way between Lefka Ori and Rethymnon). At two sites at the
extreme eastern end of the island the percentage of plain leaves represented
over 40% of the population.
Leaves with a red reverse were found at all sites and formed the vast majority of plants recorded. Green backed leaves were found at only 12 sites, but at one site (94/02) formed 93% of the population. Sites where plants with green backed leaves were found generally had an aspect in the northern half of the compass, that is from East, through Northeast, North and Northwest. No green backed leaves were found at altitudes exceeding 680m (2,230ft). No green backed leaves were recorded from the site in Karpathos.
Across all sites, based on a total of 1628 plants recorded:
| Leaf Shape | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 |
| 28.5% | 7.9% | 23.8% | 1.9% | 1.0% | 1.4% | 8.5% | 3.4% | 4.2% | 19.2% |
| Leaf Pattern | A | B | C | D | E | F | No Pattern |
| 28.6% | 28.4% | 28.6% | 3.1% | 3.2% | 0.3% | 7.8% |
| Leaf Reverse | Red | Green |
| 90.4% | 9.6% |
The longest petals found were 38mm (1½ in) long; the widest were 8mm (1/3 in) in width. The average length varied from 16.8mm (2/3 in) at site 94/20 to 28.4mm (1 1/8 in) at site 96/02. Average width varied from 3.8mm (just over 1/8 in) at site 94/12 to 6.8mm (¼ in) at site 96/02. The degree of twist varied from 0° to 360°. There was no correlation between petal length and the width of petal, degree of twist or flower colour. The largest flowers recorded were all plain white, but this is probably not significant as white is in any case the predominant flower colour. The ratio of average petal length to average petal width varied between 3.5:1 and 5:1, except at the Karpathos site which had a ratio of 3:1 (17.6mm long by 5.9mm wide). At over 50% of sites the average ratio of petal length to width was 4:1.
The degree of petal twist correlated with a number of other factors:
With
the exception of one site (95/16) at 625m, all petals with a twist greater than
180° occurred at less than 400m altitude.
There is a significant
correlation between latitude and petal twist. No plants with a petal twist
greater than 180° were found north of 35° 16' 975'' N, and it is
significant that the further south the expeditions went, the greater the
proportion of highly twisted petals.
There is a very distinct correlation
between longitude and the degree of petal twist. With the exception of a small
number of plants (2%) at site 96/12 (Gipari Gorge), and 15% of plants at site
95/03 (north-west of Dikti), all plants with a twist greater than 180° were
at the eastern end of the island, that is east of longitude 25° 39' E (or,
east of Dikti). Here, most sites had a significant proportion of plants with a
petal twist greater than 180° and most had some plants with a twist of 360°.
The site on Karpathos had 6% of plants with a petal twist of 270°.
All
sites where plants were found with a petal twist greater than 180°, had an
aspect between due East, through North to Northwest. All sites with a southerly
or westerly aspect had plants with a petal twist of 180° or less
White flowers were found at all sites, whilst flowers with varying shades of pink were found at 85% of sites in proportions varying from 2% to 74% (site 94/08). There was no correlation between the proportion of pink flowers and any other factor. The Agia Irini, 2N=20 site (94/17) had 60% pink flowers. At the site on Karpathos, all flowers were white.
Across all sites, based on a total of 1628 plants:
| Petal Twist | 0° | 45° | 90° | 135° | 180° | 225° | 270° | 315° | 360° |
| 25.7% | 17.9% | 40.1% | 4.1% | 8.9% | 0% | 1.1% | 0.3% | 1.7% |
| Petal Colour | White | Pink |
| 79.6% | 20.4% |
Grey-Wilson, C. (1988) The Genus Cyclamen Royal Botanic
Gardens, Kew in association with Christopher Helm and Timber Press
Bailey,
R.H. (1991) The Society's expedition to Rhodes and Karpathos Cyc. Soc.
Journal. 15:46-50
Bailey, R.H. (1992) Cyclamen creticum on Karpathos Cyc.
Soc. Journal. 16:41-43
Johnstone, R. (1992) The Society's Peloponnese
expedition, April 1992 Cyc. Soc. Journal. 16:45-47
Moore, P. (1993)
Peloponnese expedition, April 1993 Cyc. Soc. Journal. 17:50-53
Saunders,
D.E. (1979) Cyclamen - A Gardener's Guide to the Genus Alpine Garden Society
Turland,
N.J.; Chiltern, L.; and Press, J.R. (1993) Flora of the Cretan Area HMSO The
Natural History Museum
Aspland, V. (1994) Expedition to Crete 1994 Cyc.
Soc. 18:42-44
Wiltshire, T. and Dale, H. (1995) Cyclamen Society Expedition
to Crete 1995 Cyc. Soc. 19:54-55
Bacon, L. (1979) Mountain Flower
Holidays in Europe. Alpine Garden Society
Turland, N. (1988) Mountains and
Gorges of Western Crete AGS Bulletin 56:46-57
Rowland, N. (1991) Orchids
of Southern Crete AGS Bulletin 59:297-304
Greilhuber, Prof. J. (1989)
Taxonomic relevance of karotype structure in Cyclamen L. subgen. Psilanthum
Schwarz Cyc. Soc. 13:57-62