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THE IMPORTANCE of ECOLOGY for GENERIC and SPECIFIC DIFFERENTIATION in the ARACEAE-AROIDEAE HARALD RIEDL Natural History Museum, Vienna, Austria

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THE IMPORTANCE of ECOLOGY for GENERIC and SPECIFIC DIFFERENTIATION in the ARACEAE-AROIDEAE HARALD RIEDL Natural History Museum, Vienna, Austria 1980] THE IMPORTANCE OF ECOLOGY FOR GENERIC AND SPECIFIC DIFFERENTIATION IN THE ARACEAE-AROIDEAE HARALD RIEDL Natural History Museum, Vienna, Austria It is Meusel's (1951) merit to perature are slight and there is have pointed out the significance of no period of more or less extreme growth-habit for interpreting the drought even during the dry season; evolution of a particular group of Where subterranean parts alone are plants. In his paper he chose Ara­ responsible for the survival of an ceae and Lemnaceae as striking ex­ individual, periods of drought and/ amples to prove his point. While it or low temperatures can be bridged is rather difficult to translate the without any harm. German terminology he used for There is little doubt that extra­ those plants which produce per­ tropical Aroideae evolved from sistent parts above the ground, the tropical ancestors, and several tribes term "geophytes" fits well for all (for an updated system of the fam­ those which persist with their sub­ ily based on Engler, 1920, see Bog­ terranean parts alone. Among Ara­ ner, 1978) are confined to tropi­ ceae, rhizomatous and tuberous cal areas even today. There is, how­ geophytes are known. Subfamily ever, a marked preference for more Aroideae is composed almost en­ extreme habitats such as high alti­ tirely of members of the latter tudes, slopes, etc., even in these. group with the exception of plants Genera like Spathantheum are con­ growing in water or at least swampy fined to the upper levels of the An­ ground, like Lagenandra. While, ac­ des in Bolivia and Peru; for species cording to Meusel, intermediates like Asterostigma lividum (Lodd.) between rhizomatous and tuberous Engl. we find label data noted by geophytes are found in Colocasio­ the collector of the type cited by ideae, geophytes are rare or absent Engler, I.c., 'in den Waldungen des in the rest of the family. Corcovado an abschlissigen SteUen' Advantages of geophytic (in the forests of Corcovado on growth habit are obvious; it is pos­ steep slopes.) sible for the plant to survive during Being more experienced in ex­ unfavorable periods in a very re­ tratropical regions of the Old World duced state underground. The eco­ I may be excused if the ideas devel­ logical implications of this advan­ oped in this paper are centered on tage can easily be seen: not only tribe Areae with its mainly extra­ habitats different from those colo­ tropical distribution. The tropical nized by members of Araceae de­ species included here are further void of it can be conquered but the examples of ecological specializa­ whole area of distribution can be tion, often growing in mountainous extended considerably. Araceae are districts. In the whole group, Aris­ primarily a group of tropical and aema Mart. is not only the genus subtropical plants growing in areas with the greatest number of species in which seasonal changes of tem- by far within the group, but also 49 50 AROIDEANA [Vol. 3 with the widest ecological and geo­ from the coast and the northern de­ graphical range of distribution. Two clivities of the adjacent hills. The or three chracters can be mentioned number of female flowers is re­ as ~omparatively primitive: the duced to one here, the ovary is mul­ number of ovules in one ovary is tiovulate. The sterile apex of the greater than one as a rule, the fili­ spadix is reduced. Reduction of the form, sterile male flowers are devel­ whole plant seems to have gone oped upwards to the apex of the rather a long way towards Lemna­ spadix in some species (the osmo­ ceae already, though it is certainly phores on the club of Arum, for not among the ancestors of this instance, can be derived morpho­ family: small size, reduced number logically from the bases of extreme­ of leaves, which are broadly ellipti­ ly reduced staminodial flowers), cal resembling the first leaves after and some of the species are dioeci­ the cotyledon of other Araceae, the ous. Series Fimbriata Engl. with fairly small number of male flow­ numerous staminodial flowers up to ers, which are arranged in two ser­ the top of the spadix has a south­ ies on one side of the spadix all eastern Asiatic distribution in Ma­ indicate a long phylogenetic histo­ laysia and Indonesia in the zone of ry. It is growing in rocky or sandy the monsoon forests. places, in grassy vegetation, degrad­ Prime (1960) raised the ques­ ed forests, hedges and especially tion of whether genera with a very macchia-like scrub. This is a typical limited distribution within the habitat of several Mediterranean Areae are primitive relics or rather Araceae, members of the genus very young, advanced groups which Arum like A. pictum L.f. with a could not spread more widely dur­ very similar geographical distribu- ' ing the comparatively short time of tion, Helicodiceros muscivorus, or their existence. In the case of Heli­ the widespread Arum italicum Mill., codiceros Schott, the first alterna­ Dracunculus vulgaris Schott, Aris­ tive seems to be more likely for the arum vulgare Targ.-Tozz. or Biarum same reasons we mentioned in Aris­ tenuifulium (L.) Schott. aema. The whole distal part of the spadix is covered with subulate If we try to discover a general staminodial flowers. There are up pattern of ecological distribution to six ovules in each ovary, and fe­ in Aroideae, we find two principal male flowers are numerous. The on­ trends starting from intermediary ly species grows on coastal rocks on mesophilic plants, one leading from a few Mediterranean islands such as mesic to semiaquatic and aquatic Corsica, Sardinia and some of the habitats, another to dry habitats Baleares. which cannot be colonized by any Ambrosinia L., another mono­ other group of Araceae. In the first typical genus, on the other hand group, Cryptocoryne and Lagenan­ has two centers of distribution dra are the extremes which no which are separated by the whole longer have tuberous, but slender, width of the Mediterranean Sea and elongated rhizomes with secondary each of which is comparatively fibrous roots at the nodes. Their small: Corsica, Sardinia, Calabria distribution in the zone of mon­ and Sicily in Europe, Algeria and soon rainfall in southeastern Asia Tunisia in North Africa. In Africa, indicates that they can be derived it is confined to a short distance directly from tropical ancestors but 1980] RIEDL: ECOLOGY OF AROIDEAE 51 they show a number of unusual sist occasional drought for a short morphological specializations. Oth­ time at least. In Bulgaria, Velenov­ er Aroids don't go as far as that but sky found it most often near Pali­ prefer moist ground along river urus spina-christi, a shrub of hot, banks or in narrow ravines. The dri­ dry, stony places that gives very est localities colonized by Araceae little shadow itself. Arum nigrum are rocky or gravelly places in the Schott is similar ecologically, Mediterranean region with open growing on stony and bushy slopes vegetation and a resting period dur­ in Yugoslavia not too far from the ing summer. To illustrate my point coast, so that it always receives a I shall give a short account on eco­ certain amount of d~mp air. Other logical as a parallel to taxonomical species of the Mediterranean region differentation within the genus with similar ecological demands are Arum. A. dioscoridis Sibth. et Sm., A. Arum creticum Boiss. et Heldr. conophalloides Kotschy ex Schott and A. pictum L. f., mentioned al­ and A. hygrophilum Boiss. Accord­ ready before, stand apart morpho­ ing to label-data cited by Engler, logically from all the other spe­ they grow near rivulets, springs cies of the genus, but fit well into or waterfalls, A. hygrophilum also the pattern of ecology displayed by near mountain tops where water other Aroids growing in rocky from melting snow is always avail­ places in the Mediterranean region. able. Either humid soil or damp air, They both have a very limited dis­ commoniy both, are essential even tribution, the first one being con­ for these species, which are growing fined to Crete and Karpathos, and in normally dry areas. may well be regarded as ancient Arum is one of the few genera relics. The most widespread species, within the family thrt reaches the on the other hand, A. italicum, temperate region of Europe, with A. grows in similar places, but has a maculatum L., while A. orientale much wider ecological range includ­ M.B. subsp. alpinum (Schott et Ky.) ing moist places. It is a matter of H. Riedl is found in eastern and personal opinion whether it should east-Central Europe and A. italicum be split into a number of closely in Western Atlantic Europe. There related species with a higher degree seem to be no great differences in of ecological specialization, or whe­ ecology between A. maculatum and ther Engler's example should be fol­ A. italicum in England, where the lowed. Arum italicum var. interme­ latter species has its northern limit. dium Mutel was collected along riv­ It is interesting to note that it ulets on Corsica by Briquet, var. changes its ecological preferences concinnatum (Schott) Engl. subvar. considerably outside the Mediter­ wettsteinii (Hruby) Engl. and sub­ ranean region. Prime, l.c., writes var. marmoratum (Schott) Engl. in "The rare lords-and-Iadies grows in moist places on Crete by Reverchon. rather similar situations to the com­ According to Engler, the typical mon ' plant, and in some localities variety is especially common on the two grow together. It is a plant moist, loamy soil. Certainly , it is of light shade, liking a little shelter not a species of shady forests but and requiring a moist, well-drained, prefers open country, hedges and fairly calcareous soil." The time of shrubby vegetation.
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