The Arum Species of Greece and Cyprus

Peter C. Boyce

Herbarium, Royal Botanic Gardens Kew, Richmond, Surrey, TW9 3AB, U.K

Abstract

A synopsis of the 14 species of Arum recorded for Greece and Cyprus with a key to all relevant infrageneric taxa.

The genus Arum consists of 26 species of tuberous herbs occurring from the Azores to western China and from Sweden to Morocco (Boyce, 1989, 1993). Since the publication of a monograph of Arum (Boyce, 1993) a further two, as yet undescribed, species have been discovered in Turkey. In addition this publication includes the recognition of A. orientale Bieb. subsp. sintenisii (Engl.) P.C. Boyce at specific rank as A. sintenisii.

So far 14 species of Arum have been recorded from Greece and Cyprus, representing almost all of the infrageneric divisions proposed by Boyce (1989). Over half of the species are recorded from the Greek islands and Cyprus; six species are known to occur on the mainland.

All Arum species in Greece and Cyprus belong to subgen. Arum (see Boyce, 1989, 1993). The other subgenus, Gymnomesium (Schott) Engl. (Engler, 1879), is restricted to the Mediterranean west of Italy.

Key to the Sections and Subsections

1. Stem consisting of a horizontal rhizomatous tuber with lateral adventitious shoots forming offsets which later become independent; established plants forming extensive spreading colonies. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . i. sect. Arum

Stem consisting of a vertically or horizontally orientated discoid tuber with peripheral adventitious shoots which sometimes form independent plants; established plants forming congested, compact colonies . . (ii. sect. Dioscoridea) 2

2. Spadix-appendix long-stipitate. . . . . . . . . . . . . . . 3

Spadix-appendix not so. . . . . . . . . . . . . . . . . . . 4

3. Peduncle longer than the petioles; inflorescence ± odourless; both staminodes and pistillodes with long, filiform, semirigid bristles . . . . . . . . . . . . . . a. subsect. Alpina

Peduncle shorter than the petioles; inflorescence with foetid odour; pistillodes and staminodes with subulate to long, slender-filiform, flexuous bristles

b. subsect. Dischroochiton

4. Staminodes and pistillodes well developed. . . . . . . . . . 5

Staminodes and pistillodes absent or poorly developed

f. subsect. Cretica

5. Spadix-appendix short-stipitate, more than 5 mm in diameter, spathe-tube interior white or stained purple in the upper portion, very rarely entirely purple. . . . . . . . . . . . 6

Spadix-appendix not stipitate, less than 4 mm in diameter, spathe-tube interior wholly purple d. subsect. Hygrophila

6. Peduncle ± equal to or exceeding the petioles; bristles of staminodes and pistillodes filiform, flexuous; inflorescence odourless. . . . . . . . . . . . . . . c. subsect. Tenuifila

Peduncle shorter than the petioles, occasionally ± absent; bristles of staminodes and pistillodes subulate, stiff; inflorescence with strong foetid odour

e. subsect. Poeciloporphyrochiton

 Key to Species

1. Plants occurring as dense extensive colonies. . . . . . . . 2

Plants occurring as scattered individuals or small, discrete colonies. . . . . . . . . . . . . . . . . . . . . . . . . . 3

2. Spadix-appendix ± slender-clavate, 1/4-1/2 as long as spathe-limb. . . . . . . . . . . . . . . . . . . . . 1. A. italicum

Spadix appendix massively to stoutly clavate-cylindric, subequal to more than 1/2 as

long as spathe-limb. . . . . . . . . . . . . . . . . . . . . . . . . . 2. A. concinnatum

3. Pistillodes and staminodes present. . . . . . . . . . . . . 4

Pistillodes and staminodes usually absent. . . . . . . . . 12

4. Spathe-limb blotched and spotted with purple on inner surface, inflorescence strongly foetid. . . . . . . 12. A. dioscoridis

Spathe-limb not marked on inner surface, inflorescence foetid or not. . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5. Staminodes and pistillodes subulate, spathe-limb interior deep purple. . . . . . . . . . . . . . . . . . . . . 6. A. nigrum

Staminodes and pistillodes filiform, spathe-limb interior as above or differently coloured. . . . . . . . . . . . . . . 6

6. Spathe-tube distinctly bicoloured on inner surface, purple above, pale green to white below. . . . . . . . . . . . . 7

Spathe-tube not so. . . . . . . . . . . . . . . . . . . . . 9

7. Spadix-appendix stout-cylindric, subequal to spathe-limb. . . . . . . . . . . . . 9. A. elongatum

Spadix-appendix slender-cylindric to cylindric-clavate, up to 3/4 as long as spathe-limb. . . . . . . . . . . . . . . . 8

8. Spathe-limb dark purple on inner surface, often paler towards the middle. . . . . . . . . . . . . . . . . 7. A. cyrenaicum

Spathe-limb evenly suffused with pale purple on inner surface. . . . . . . . . . . . . . . . . . . . . . . . . . 9

9. Inflorescence borne beneath leaves. . . . . . . . . . . . . 4. A. orientale subsp. orientale

Inflorescence borne at leaf level, sweet-smelling…5. A. sintenisii

9. Spathe-tube white or greenish white inside, sometimes flushed with pale purple above and along the margin. . . . . . . 10

Spathe-tube deep purple inside. . . . . . . . . . . . . . 11

10. Spadix-appendix generally slender long-stipitate; cylindric to clavate. . . . . . . . . . . . . . . . . . 3. A. alpinum

Spadix-appendix massive conic-cylindric, short-stipitate or stalkless. . . . . . . . . . . . . . . . . . 10. A. rupicola

11. Spathe-limb deep purple inside. . . . . 8. A. purpureospathum

Spathe-limb pale green with a 1-1.5 mm wide purple border. . . . . . . . . . . . . 11. A. hygrophilum

12. Spathe white, limb erect and cucullate at maturity; spadix-appendix deep purple. . . . . . . . . . . . . 13. A. idaeum

Spathe cream to yellow, limb reflexing at maturity; spadix-appendix mid-yellow to dark yellow. . . . . 14. A. creticum

Enumeration of Species

i. Sect. ARUM. Stem a horizontal rhizomatous tuber with lateral adventitious shoots forming offsets which later become independent; established plants forming extensive spreading colonies.

1. Arum italicum Miller, Gard. Dict. (ed.8), no.2 (1768). Schott, Syn. Aroid. 10 (1856) & Prodr. Syst. Aroid. 82 (1860); Engler in A. & C. DC. Monog. Phanerog. 2: 591 (1879); Hruby in Bull. Soc. Bot. Genève 4: 128 (1912); Engler in Engler, Das Pflanzenr., 73 (IV. 23F): 82 (1920); Mayo & Meikle in Meikle, Fl. Cyprus. 2: 1667 (1985); P.C. Boyce, The Genus Arum, 69 (1993).

Distribution: Greece: Mainland: Macedonia (Makedhonia) Peloponnese (Peloponnisos), Thessaly (Thessalia), Thasos, Thrace (Thraki). Islands: Corfu (Kerkira). Cyprus (Kypros): Troodos.

Of the four subspecies recognized for A. italicum in the recent revision (Boyce, 1993), only the typical subspecies is represented in Greece and Cyprus. The striking silver-grey venation usually associated with A. italicum subsp. italicum is rarely encountered in eastern Mediterranean populations.

Confusion between A. italicum and A. concinnatum Schott is possible, although the latter is readily distinguished by its more massive spadix-appendix.

2. Arum concinnatum Schott, Icones Aroid., t.39 & 40 (1859) & Prodr. Syst. Aroid., 84 (1860); Turland, Chilton & Press, Flora of the Cretan area, 158, map. 1366 (1993); P.C. Boyce, The Genus, Arum 79 (1993).

Distribution: Greece: Mainland: southern Peloponnese (Peloponnisos).

Islands: Chios (Khios), Crete (Kriti), Karpathos, Cos (Kos), Lesbos (Lesvos), Naxos, Rhodos (Rhodes), Samos, Simi. Cyprus (Kypros): Lapithos.

Note. Almost certainly on all of the eastern Aegean islands south of Lesvos.

Arum concinnatum has been much confused with A. byzantinum Blume (Blume, 1836), a species restricted to northwestern Turkey. Recent guide books (e.g. Polunin, 1980) have misnamed A. concinnatum as A. byzantinum. Arum byzantinum is a much smaller plant, with darker coloured spathes, a shorter, thinner spadix- appendix and narrower leaves lacking the cloudy, silver-grey markings typical of A. concinnatum.

Arum concinnatum is variable in spathe-limb coloration. On Crete forms ranging from very pale greenish white to plum-purple occur as mixed populations. Some of the forms have been recognized as distinct species (e.g. A. wettsteinii Hruby (1912)). In southwestern Crete A. concinnatum hybridizes with A. cyrenaicum producing plants intermediate between the parents.

ii. Sect. DIOSCORIDEA (Engler) P.C. Boyce. Stem a vertically or horizontally orientated discoid tuber with peripheral adventitious shoots which sometimes form independent plants; established plants forming congested, compact colonies.

a. Subsect. ALPINA P.C. Boyce. Spadix-appendix clavate, long- stipitate. Peduncle longer than petiole. Bristles of staminodes long, filiform, semi-rigid. Inflorescence ± odourless.

3. Arum alpinum Schott & Kotschy in Bot. Zeitung. 9: 285 (1851); Schott, Syn. Aroid., 12 (1856) & Prodr. Syst. Aroid., 91 (1860); Turland, Chilton & Press, Flora of the Cretan area, 158, map 1365 (1993); P.C. Boyce, The Genus Arum, 85 (1993).

Distribution: Greece: Mainland: Attica (Attiki), Ipiros, Macedonia (Makedhonia), Sterea Ellas. Islands: Crete (Kriti).

Long confused with A. maculatum L. (Linnaeus, 1753), A. alpinum is characterized by carrying its generally odourless inflorescences borne above the leaves. Arum maculatum has yet to be recorded from Greece or Cyprus, records from Crete are referable to A. alpinum or A. idaeum (see Greuter, 1984).

Arum alpinum is widespread, occurring from Portugal to southern Sweden and then east to northwestern Turkey. Given this extensive range the species is remarkably uniform in appearance. Isolated populations show some variation (e.g. spadix-appendix shape and inflorescence size) but the species is always easily recognizable.

b. Subsect. DISCHROOCHITON Schott. Spadix-appendix long- stipitate, usually stout. Peduncle shorter than the petiole. Pistillodes and staminodes very well developed, with long, filiform, flexuous bristles; Inflorescence smelling foetid.

4. Arum orientale Bieb., Fl. Taur.-Cauc. 2: 407 (1808); Schott, Syn. Aroid., 15 (1856) & Prodr. Syst. Aroid., 88 (1860); Engler in A. & C. DC., Monog. Phanerog., 2: 586 (1879); Hruby in Bull. Soc. Bot. Genève 4: 140 (1912); Engler in Engler, Das Pflanzenr. 73 (IV. 23F): 78 (1920); Mayo & Meikle in Meikle, Fl. Cyprus 2 1667 (1985); P.C. Boyce, The Genus Arum, 90 (1993).

Distribution: Greece: Mainland: Macedonia (Makedhonia).

Boyce (1993) recognized three subspecies for A. orientale. In Greece it is represented by the typical subspecies, which is widespread in eastern central Europe and the Balkans, extending through northern Turkey into the Caucasus and Crimea. Subsp. longispathum (Reichb.) Engl. (Engler, 1920) is restricted to Montenegro while subsp. sintenisii, endemic to Cyprus, is raised to species level elsewhere in this paper.

There are few collections of A. orientale known from Greece, all collected from the northern mainland.

5. Arum sintenisii (Engler) P.C. Boyce, stat. nov. A. orientale Bieb. subsp. elongatum (Steven) Engl. var. sintenisii Engl., Das Pflanzenr. 73(IV.23F):80 (1920). Type: Cyprus (Kypros), auf Triften bei Kythraea (Kythrea), 6 May 1880, Sintenis & Rigo 130 (holotype K!). A. orientale Bieb. subsp. sintenisii (Engl.) P.C. Boyce, The Genus Arum: 96 (1993).

Distribution: Cyprus (Kypros): Kythrea.

Boyce (1993) regarded this taxon as a subspecies of A. orientale. Since then, study of further material and access to living plants have convinced me that it is a distinct species. Arum sintenisii has a different mode of inflorescence presentation and a sweet, not foetid odour when in flower. Arum sintenisii is restricted to the vicinity of Kythrea in Cyprus where is occurs as extensive stands.

6. Arum nigrum Schott in Oesterr. Bot. Wochenbl. 27, 7: 213 (1857) & Icones Aroid., t.37, 38 (1859), & Prodr. Syst. Aroid., 81 (1860); Engler in A. & C. DC., Monog. Phanerog., 2: 586-7 (1879); Hruby in Bull. Soc. Bot. Genève 4: 137 (1912); Engler in Engler, Das Pflanzenr., 73 (IV. 23F): 74-5 (1920); P.C. Boyce, The Genus Arum 106, (1993).

Distribution: Greece: Mainland: Attica (Attiki) (doubtful), Evvoia (doubtful), south Macedonia (Makedhonia). Islands: Jura, Skiros.

Long thought to be restricted to Montenegro, A. nigrum is now known to be present but rare in Greece. The deep purple spathe- limb and black-purple to pewter-coloured spadix-appendix, combined with an exceptionally strong dung-like odour, make it easily recognizable. Unlocalized records of A. nigrum from southern mainland Greece reported by Halácsy (1904) have yet to be confirmed. Reports of A. nigrum on Samothraki (Akeroyd & Preston, 1987) are referable to A. elongatum subsp. elongatum.

7. Arum cyrenaicum Hruby in Bull. Soc. Bot. Genève 4: 159 (1912); Engler in Engler, Das Pflanzenr., 73 (IV. 23F): 81 (1920); El Gadi in Fl. Libya, 41: 236 (1977), Turland, Chilton & Press, Flora of the Cretan area, 159, map 1368 (1993); P.C. Boyce, The Genus Arum, 110 (1993).

Distribution. Greece: Islands: southwestern Crete (Kriti).

One of the most recent additions to the Greek flora, A. cyrenaicum was long thought to be endemic to Libya occurring between Tripoli and Benghazi. It has recently been discovered on Crete, but so far only three small populations have been found, each consisting of a few individuals. It is interesting to note that at one of the sites hybrids between A. cyrenaicum and A. concinnatum have been found.

8. Arum purpureospathum P.C. Boyce in Aroideana 10: 8 (1987); Turland in Bull. Alpine Gard. Soc. 57: 112-117 (1989), Turland, Chilton & Press, Flora of the Cretan area, 159, map 1370 (1993); P.C. Boyce, The Genus Arum, 113 (1993).

Distribution: Greece: Islands: southwestern Crete (Kriti).

Arum purpureospathum is one of the two Arum species endemic to Crete. It is restricted to a few sites in southwest of the island. The large, glistening, deep purple spathes are distinctive. Arum purpureospathum is protected by law in Greece.

9. Arum elongatum Steven in Bull. Bot. Soc. Mosc. 32,2: 67 (1857); Schott, Prodr. Syst. Aroid., 100 (1860); Hruby in Bull. Soc. Bot. Genève 4: 140 (1912); P.C. Boyce, The Genus Arum, 123 (1993).

Distribution. Greece: Mainland: Macedonia (Makedhonia).

Islands: Samothraki.

Arum elongatum occurs as the typical subspecies in northern Greece. Boyce (1993) maintained a second subspecies, subsp. alpinariae Mill & Alpinar (Mill & Alpinar, 1988), from Turkey. Arum elongatum is rare in Greece, represented by only a few collections. It is a striking species with relatively large, deep purple spathes, a long, rather stout spadix-appendix producing a powerful odour of horse dung. Records of A. nigrum on Samothraki (Akeroyd & Preston, 1987) are referable to A. elongatum subsp. elongatum.

c. Subsection TENUIFILA (Engler) P.C. Boyce (1989). Peduncle longer than petiole. Spadix-appendix shortly stipitate or sessile, slender to stout, occasionally massive. Staminodes and pistillodes moderately well-developed, bristles filiform, rather short. Inflorescence odourless.

10. Arum rupicola Boiss., Diagn. 1,13: 7 (1853); Schott, Syn. Aroid. 14 (1856) & Prodr. Syst. Aroid., 96 (1860); Engler in A. & C. DC., Monog. Phanerog., 2: 588 (1879); P.C. Boyce, The Genus Arum, 129 (1993).

Distribution: Greece: Islands: Lesbos (Lesvos).

Cyprus (Kypros): Troodos, Tripylos.

Arum rupicola is essentially an Asian species, occurring only on Cyprus and the island of Lesbos. Two varieties are recognized by Boyce (1993). The Greek and Cypriot plant, with a purple to dull brown spathe-limb and similarly coloured spadix- appendix, corresponds to the typical variety. Var. virescens (Stapf) P.C. Boyce (Boyce, 1993), has a greenish white spathe, pale lilac to grey spadix-appendix and is exclusively Asian.

d. Subsect. HYGROPHILA P.C. Boyce. Peduncle equal to or longer then the petiole. Spadix-appendix sessile, less than 4 mm diam. Staminodes and pistillodes with long, filiform, flexuous bristles. Spathe-tube interior wholly purple. Inflorescence odourless.

11. Arum hygrophilum Boiss., Diagn. 1,13: 8 (1853); Schott, Syn. Aroid., 13 (1856) & Prodr. Syst. Aroid., 98 (1860); Engler in A. & C. DC., Monog. Phanerog., 2: 589 (1879); Hruby in Bull. Soc. Bot. Genève 4: 157 (1912); Engler in Engler, Das Pflanzenr., 73 (IV. 23F): 77 (1920); Mayo & Meikle in Meikle, Fl. Cyprus, 2: 1665 (1985); P.C. Boyce, The Genus Arum, 147 (1993).

Distribution: Cyprus (Kypros): Kythrea, Kazaphani, Tala.

A distinctive plant with pale green spathes with a striking purple band along the spathe limb margin. Arum hygrophilum has a disjunct distribution. First described from the Syrian - Lebanese border, the plant also occurs on Cyprus and in northern Morocco.

e. Subsect. POECILOPORPHYROCHITON Schott. Peduncle much shorter than the petiole, sometimes apparently absent. Spadix- appendix shortly stipitate, cylindric to fusiform-cylindric or rather slender-cylindric, moderately stout. Staminodes and pistillodes well developed, bristles subulate, stiff, short to long. Inflorescence with foetid odour.

12. Arum dioscoridis Sm. in Sibthorp & Smith, Fl. Graec. Prodr. 2: 245 (1816); Schott, Syn. Aroid., 9 (1856) & Prodr. Syst. Aroid., 78 (1860); Engler in A. & C. DC., Monog. Phanerog., 2: 583 (1879); Hruby in Bull. Bot. Soc. Genève 4: 153 (1912); Engler in Engler, Das Pflanzenr., 73 (IV. 23F): 72 (1920), Mayo & Meikle in Meikle, Fl. Cyprus, 2: 1665 (1985); Boyce, The Genus Arum, 151 (1993).

Distribution: Greece: Islands: Chios (Khios), Rhodes (Rhodos). Cyprus (Kypros): Cap Greco, Kyrenia, Kythrea, Limassol, Nicosia, Sotia.

Four varieties of A. dioscoridis are recognized by Boyce (1993). Of these var. dioscoridis and var. cyprium (Schott) Engl. (Engler, 1920) are present on Rhodes, Chios and Cyprus. The varieties are distinguished by coloration of the interior of the spathe limb. Var. dioscoridis has spathes with large deep purple confluent blotches and spots overlaid with slightly paler purple staining to within a few centimetres of the spathe tip. Var. cyprium has pale green spathes with smaller, more-or-less discrete blotches and lacking any purple staining. When flowering both varieties usually produce a powerful smell of pig-dung.

f. Subsect. CRETICA (Engler) P.C. Boyce. Peduncle longer, or much shorter than petiole. Spadix-appendix sessile to obscurely stipitate, cylindric, laterally compressed, moderately stout. Staminodes and pistillodes absent, very rarely staminodes and pistillodes present in A. creticum. Inflorescence sweetly scented, more rarely odourless.

13. Arum idaeum Coust. & Gandoger in Bull. Soc. Bot. Fr. 63: 11 (1917); Turland, Chilton & Press, Flora of the Cretan area, 159, map 1369 (1993); P.C. Boyce, The Genus Arum, 162 (1993).

Distribution. Greece: Islands: Crete (Kriti).

Restricted to mountainous regions of Crete, A. idaeum has long been confused with A. creticum, usually being referred to as the high altitude form of that species. Arum idaeum typically has white spathes and a deep purple spadix appendix. However, the spadix appendix can vary, with forms displaying various degrees of yellow, ranging from a few patches to almost entirely yellow. Other interesting variation is found in the smell of the inflorescence. Most plants in cultivation in Europe have an odourless inflorescence but many plants investigated in the field produced a faint but readily discernible smell of violets and hyacinth.

14. Arum creticum Boiss. & Heldr. in Boiss., Diagn. 1,13: 9 (1853); Schott, Syn. Aroid., 11 (1856) & Prodr. Syst. Aroid., 94 (1860); Engler in A. & C. DC., Monog. Phanerog., 2: 590 (1879); Hruby in Bull. Soc. Bot. Genève 4: 151 (1912); Engler in Engler, Das Pflanzenr., 73 (IV. 23F): 69 (1920); Turland, Chilton & Press, Flora of the Cretan area, 159, map 1367 (1993); P.C. Boyce, The Genus Arum, 165 (1993).

Distribution. Greece: Islands: Karpathos, Kasos, Crete (Kriti).

A very striking species with large cream to lemon-yellow inflorescences borne well clear of the leaves. Widespread on Crete and occurring eastwards on Kasos, Karpathos and in southwestern Turkey, A. creticum is perhaps the most readily identifiable and certainly the best known of the Greek arums. The strong smell produced by the open inflorescence is usually described as sweet and freesia-like, but can vary from this to an highly unpleasent naptha-like odour.

ACKNOWLEDGEMENTS

I would like to thank Dr David Simpson and Dr John Dransfield for their helpful commments on earlier drafts of this manuscript. 

REFERENCES

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