It is now ten years since I wrote the article "The Cryptocoryne albida group of Mainland Asia" in the "Liber Gratulatorius" on the occasion of the retirement of Prof. H. C. D. de Wit (Jacobsen, 1980). Very little new information about these species has in the meantime come to our knowledge; all the plants imported to Europe during the last 10 years have added only few new facts, most of the material confirming what we knew or suspected around 1980. Our information about the variation within the species has increased, but not more than what was to be expected.

The following account presents an abbreviated version of what was contained in the article 10 years ago, supplemented by the additional information obtained since then. The intervening time has also shown a need to be able to recognize some of the intraspecific variation found within the polymorphic species C. crispatula.

Introduction

The species of the Cryptocoryne albida group are rather homogenous. Also other species have narrow, strap-shaped leaves, but they do not have a similar spathe, i.e. with a relatively long tube, and a spirally twisted limb of the spathe which has circular to elongate markings on the surface, lacks a collar, and has a more or less speckled inner surface of the kettle wall, the wall also having circular, alveolar depressions in the upper part.

The main taxonomic characters for the delimitation of related species in Cryptocoryne are the shape and colour of the limb of the spathe. Each species also has its own characteristic leaf-form, but in most cases this alone is not enough for a definite determination of a species. In the species dealt with here, the differences between the species lie in the exact form of the spirally twisted limb of the spathe and the differences in the brown, red, and purple colours of the surface, features that seem more quantitative than qualitative. These characters are difficult to distinguish in herbarium specimens, and only after study of living material has it been possible to clear up the limits of variation between the different species and in most cases possible also to distinguish dried specimens.

The shape of female flowers, stigmas, and olfactory bodies, the number and shape of male flowers, the colouration of the inner side of the kettle wall, and the colour of the valve closing the opening of the tube vary continuously within the group and have been found to be of little use in the delimitation of the species. The three species, as interpreted here, have adjacent but not (or only slightly) overlapping distribution areas (Fig. 6).

The Cryptocoryne albida group has been the subject of several different opinions (Engler 1920, De Wit 1971, 1983, Rataj 1975). Earlier (Jacobsen, 1976, 1977, and 1985), I put forward the view that the species of Cryptocoryne were narrow endemics. However, for two of the three present species this opinion is not tenable. Viewing the present three species, and their variation, in connection with the rest of the species of Cryptocoryne, it might be asked if they could just as well be considered one species ? With e.g. three subspecies ? If someone insists this is the case, I have no convincing arguments in my favor. The point of view does, to some extent, depend on the taxonomical characters used and the weight they are given.

Engler (1920), obviously aware of the difficulties connected with a sectional division of the genus, forwarded a "dispositio gregum" and placed C. unilocularis in Unitubulosae (because of the apparently lacking tube of the spathe), while C. retrospiralis and C. crispatula were placed in Bitubulosae.

De Wit (1971) maintained C. albida and C. balansae as separate species but included under C. retrospiralis the varieties costata, crispatula, and tonkinensis, but later (De Wit, 1983) accepted both C. albida (incl. C. costata), C. balansae, C. crispatula (incl. C. tonkinensis), and C. retrospiralis.

Rataj (1975) distinguished 7 species (plus C. usteriana Engl., from the Philippines), placed in two sections, viz. Sect. Retrospiralae Rataj and Sect. Costatae Rataj, a division for which I have found no basis (Jacobsen, 1977) considering that C. retrospiralis ssp. albida, which is conspecific with C. costata, and C. korthausae were placed in two different sections; the species recognized by Rataj were C. retrospiralis ssp. retrospiralis and ssp. albida, C. crispatula, C. bertelihansenii, C. tonkinensis, C. balansae, C. costata, and C. korthausae.

Recently another narrow-leaved species from India, C. consobrina Schott, has come into cultivation, and it has proven to have a chromosome number of 2n = 36 like the here mentioned species (Sivadasan, 1985), but it is not included in the present context as it has a rough surface of the limb of the spathe and a collar. C. cruddasiana Prain from Burma, which bears some resemblance to the three mentioned species, also has a rough surface of the limb of the spathe, and is likewise not included here. The well known C. spiralis (Retz.) Wydl. from India, also has narrow leaves, but the spathe is very different in lacking a tube and having a limb of the spathe which has transverse, irregular ridges.

Morphology

The taxa in the C. albida group are aquatic or amphibious herbs with subterranean rhizomes that have a rosette of leaves. Vegetative propagation takes place by means of subterranean runners or by division or bud sprouting from the rhizome.

Leaves

The leaves are linear, strap shaped or lanceolate, green to brown, smooth to bullate (Figs. 7, 18). The length varies from 10 to 70 cm and the width from 0.2 to 4 cm. In the broader leaves there are conspicuous lateral veins parallel to the midrib. In narrower leaves the midrib is a main part of the leaf blade. The margin of the blade is entire to denticulate. In the monsoon season, C. retrospiralis has short, c. 15 cm long, terete leaves (Figs. 7B, 42), a feature otherwise known only from a few collections of C. crispatula, where they are 2-4 cm long.

Inflorescence

The spadix (Fig. 8) contains, as usual for Cryptocoryne, female flowers at the basis with vertical to almost horizontal stigmas. The olfactory bodies are shortly club-shaped and more or less lobed at the top. The male flowers at the top have a strongly mammillose surface. The kettle is constricted on the inside above the middle. The valve that closes the kettle is red spotted, as is the upper part of the inside kettle wall. The inside of the tube is also spotted, often showing a transition between the markings in the kettle and those on the limb of the spathe. The upper part of the inner side of the kettle wall has more or less circular depressions, alveoli, of unknown significance, but also found in other species, e.g. the C. beckettii group from Ceylon.

The limb of the spathe varies in length from 1 to 6 cm, and is more or less spirally twisted, sometimes becoming uncoiled (Figs. 8, 9, 10). The limb of the spathe is whitish to yellowish to greenish to grayish grounded. It is also so that the colour of the limb varies during flowering, being darker and more intense just after opening, and on the second and third day begins to fade. The markings on the limb of the spathe vary in form from small to large spots, from short, thin streaks to long, thick, irregular lines. The density of these markings varies from almost complete covering of the limb to nearly missing. Their colour varies from brown to red purple. A margin on the limb of the same colour as the markings may be present or absent. The shape and colour of the markings are characteristic for each species.

All three species flower from (October-) December to February, during the dry season. They are inhabitants of small or larger rivers and streams where they are submerged part of the year. At falling waters the long tube of the spathe ensures that the opening of the spathe reaches up above the water surface. Later on in the season, many of the plants near and on the river banks are completely emerse. After pollination, the fruit increases in size for the first few months and stays (mostly) hidden beneath the surface of the soil for about 3/4 of a year. At maturity the peduncle elongates 3 to 10 cm within about a week, the syncarpium rises above the surface of the soil, opens, and disperses the seeds (Fig. 33). The seeds are coated with a thin layer of a waxy appearance, which makes them float for a short time if liberated in water. This mechanism may serve to disperse the seeds only at falling waters, so that they do not travel too far, and may ensure that they are distributed below and near "normal" high water level.

Plants with fruit collected at Phu Khieo and Ban Nang Yon (Note: Misspelled: Ban Wangyon, 1991) (Thailand) in late February 1977 dispensed their seeds in November-December the same year under cultivation in Copenhagen.

Habitats

During a stay in Thailand in February 1977, I had the opportunity to study different Cryptocoryne populations. Three localities will be considered:

1. Mueak Lek (Fig. 11), c. 150 km NNE of Bangkok. Alt. c. 200 m. The 6-8 m wide river runs through the park near the main road. The water was rather calcareous and had a temperature of 24 C. Calcium carbonate incrustations covered almost everything that was under water, and in places with some water turbulence the result was formation of magnificent travertine dams across the river, creating small waterfalls. In the pools thus formed, a rich, submerged growth of Cryptocoryne crispatula was found (NJ 77-16). The largest plants were 50-70 cm long, with bullate, 30-50 cm long and 2-4 cm wide leaf blades (Figs. 12, 26). The leaves, except the youngest, were incrusted with a layer of calcium carbonate (these large-leaved plants represent what in the past has been named C. balansae).

Flowering specimens were found in places where the bottom of the river was raised to the surface of the water (leaves 20-40 cm, spathes 20-30 cm), and also on the banks at the top of the dams, where the leaves and the spathes were 10-15 cm long (Figs. 7 J, 31). The limb of the spathe of these plants was grayish grounded and had long purple markings.

2. Phu Khieo Wildlife Sanctuary, c. 350 km NNE of Bangkok. Alt. c. 700 m. The river, Huae Mae Chem, is 8-10 m broad, mostly filled with rather large, round stones and rocks (Fig. 13). The river is running quite rapidly and is less than 0.5 m deep during the dry season. The temperature in the water was 21 C. The Cryptocoryne were growing on the banks of the river in large quantities. The leaves were 10-20 cm long, 0.6-0.8 cm wide, green and smooth (Figs. 7 F - G, 16 B). In sheltered places they formed large carpets. Most of the plants were emerged and even though most of them had flowered a month or two earlier, a few could still be found in flower. Plants from this main course of the river formed the basis for the description of C. bertelihansenii, which is now recognized as a synonym of C. crispatula var. crispatula.

In a quiet, shaded tributary, just east of the main course, plants with a different foliage were found. The leaf-blades were almost copper-brown, bullate, 20-30 cm long, and 1-2 cm wide (Figs. 7 H, 16A). The limb of the spathe was yellowish with short, purple lines (Figs. 10 E, 39, 40). Some plants with foliage intermediate between the brown and green-leaved forms were found only a few meters from the brown-leaved plants. In these intermediate plants the limb of the spathe was grayish with longer, irregular purple lines, resembling those in the main course of the river.

On the west bank of the main course of the river was another small, shaded tributary, almost carved into the solid rock (Fig. 17). Here was a pool (4 × 15 m) with a depth of 0.2-0.5 m. The temperature of the water was 16° C. In cracks in the bottom of the pool plants with brown leaves were found, similar to those from the other tributary (both representing C. crispatula var. balansae). Just above the pool, in full sun, green-leaved forms grew abundantly between the rocks.

Material representing the different leaf-forms found at Phu Khieo (Fig.18) has been grown and flowered under identical growing conditions in Copenhagen. The leaf-forms are for the most part maintained constant under these cultivation conditions. All collections at Phu Khieo were made within 30 meters of each other and they are all regarded as representing C. crispatula.

3. In the Thai part of the Malay peninsula, south of Ranong, several localities with C. albida were visited, but the population at Ban Nang Yon (Note: Misspelled: Ban Wangyon, 1991), c. 100 km S of Ranong (Figs. 1, 22, 23, 24) is quite representative of the situation on the whole. Alt. c. 10 m. The river is about 4 m wide and runs rather slowly although there were some small rapids. The temperature of the water was 27º C. The river bed consisted of a mixture of sand and small stones. Emersed on banks in the river and on the river banks themselves were large stands of C. albida (NJ 77-81 a-k). The plants growing on open soil often had darker green or more reddish leaves while those growing close together and in the shade had lighter green leaves. Every transition between green leaves and marbled, red-brown were found. The leaves were more or less upright or flat on the ground, depending on the intensity of the light, those in full sun being flat on the ground.

Key to the species of the C. albida group:

1.	Limb rarely exceeding 2.5 cm in length, more or less recurved (to spirally twisted, with a short appendage). Opening of the limb usually distinctly dilatated. Markings on the limb irregular, line-like to spotted, brownish (to reddish). Leaves green to brown, 10-30 cm long, smooth, neither undulate nor denticulate at the margin.	C. albida
1.	Limb usually more than 3 cm long, more or less regularly spirally twisted. Opening of the limb of the spathe usually not broadly dilatated. Markings on the limb more or less regular, line-like and purple or larger red spots. Leaves 10-70 cm long, smooth to undulate to bullate or with denticulate margins	2
2.	Limb more or less open-spirally twisted, sometimes uncoiled. Markings on the limb irregular, line-like (to circular), purple. Leaves green to brown, smooth to undulate to bullate or with a denticulate margin, 10-70 cm long	C. crispatula
2.	Limb of the spathe usually more or less spirally twisted. Markings on the limb larger, more or less circular, reddish. Leaves green to brownish, smooth to somewhat undulate, 10-40 cm long	C. retrospiralis

Cryptocoryne albida Parker

Syn.: C. retrospiralis (Roxb.) Kunth ssp. albida (Parker) Rataj, C. costata Gagnepain, C. retrospiralis (Roxb.) Kunth var. costata (Gagnepain) De Wit, C. hansenii S.Y.Hu, C. korthausae Rataj.

Rhizome up to 0.8 cm thick. Stolons often with a metallic, blue-gray colour. Leaves lanceolate, green to brownish marbled, smooth (sometimes slightly undulate towards the margins), 10-30 cm long and 1-2 cm wide (Fig. 7 C, D); lamina with lateral veins; margin entire. Spathe 10-20 cm long; kettle 0.8-1.5 cm long; tube 5-15 cm long, slightly twisted; limb short, 1-2.5(-4) cm long, at anthesis sometimes more or less upright and spirally twisted, later becoming recurved (Fig. 9), colour grayish to cream, with reddish to brown, irregular to circular, elongate markings which sometimes may cover the surface completely; the markings may be missing. Female flowers 4-7 with more or less vertical, ovate stigmas. Olfactory bodies more or less rounded, often depressed in the centre, yellowish to greenish. Male flowers 80-120.

Chromosome number: 2n = 36.

Distribution: Southern Thailand and Burma (Fig. 6).

Habitat: At low elevations on sandy and stony banks of the rivers. Emersed when flowering (Fig. 8).

Nomenclatorial notes: The type specimens of C. albida, C. costata, C. hansenii, and C. korthausae are all so alike that it is not possible to find any differences between them: What has been called C. costata has a rather short, suddenly narrowed limb of the spathe, but can hardly be regarded even as a distinct form; cultivated type material of C. hansenii, grown in Copenhagen, proved inseparable from C. albida; type material of C. korthausae originated from the Botanical Garden in Munich, and this also proved indistinguishable from C. albida. Rataj (1975) distinguishes C. retrospiralis ssp. albida as different from both C. costata and C. korthausae, however, as all three are inseparable (and different from C. retrospiralis) there is no basis for maintaining them as separate.

Notes: The leaf-form in C. albida is rather constant; the colour varies from green to irregularly striped brown, marbled br own and almost evenly brown with some markings (Figs. 1, 3, 4, 5, 14, 15, 23, 24, 25). The species is distinguished by its short, more or less recurved limb of the spathe with the dilatated opening. The markings on the limb are brownish to reddish and their form varies from dots to irregular short line-like figures. The density of the markings as well as their size also varies and all transitions occur, from no dots at all to markings completely covering the limb. Also the length of the limb varies. It may be short, with only a very short appendage, or longer and twisted several times (Fig. 9). The total variation of the species does not exceed that found in the one locality at Ban Nang Yon, and it is not possible to distinguish between C. albida and C. costata as done by. e.g. De Wit (1971), Rataj (1975), and Jacobsen (1977).

Cryptocoryne crispatula Engler

Rhizome up to 1 cm thick, occasionally irregularly thickened. Stolons sometimes with a metallic blue-gray colour. Root system sometimes with a few upright roots. Leaves very variable, narrowly linear to lanceolate, green to brown, smooth to undulate to bullate, 10-70 cm long, and 0.2-4 cm wide (Figs. 7 E - K, 12, 16, 18, 29, 34); lamina in broader leaves mostly with conspicuous, lateral veins; margin entire to finely, irregularly denticulate (Fig. 28). In some forms, the leaves disappear during the monsoon period or become small, 2-4 cm long and terete (Fig. 27A, 35). Spathe 10-40 cm long; kettle 1-2.5 cm long; tube 6-30 cm long, slightly to somewhat twisted; limb (1.5-) 3-8 cm long, more or less spirally twisted (Fig. 10), colour yellowish to grayish to greenish, with short to long, usually purple (sometimes black purple) markings (before anthesis occasionally reddish), sometimes the markings can cover the surface almost completely. The markings may be missing, just like the limb may be almost white-grounded with only a few faint markings. Female flowers 4-6, with horizontal to vertical, round to ovate stigmas. Olfactory bodies more or less irregularly lobed. Male flowers 90-130.

Key to the varieties of C. crispatula:

C. crispatula Engl. var. crispatula

C. crispatula Engl., Pflanzenreich IV, 23 F, p. 247 (1920). - C. retrospiralis (Roxb.) Kunth var. crispatula (Engl.) De Wit, Aquarienpflanzen, p. 184 (1971). - Lectotype (selected by Rataj 1975): Harmand 65 (= 3355 = 3356), Se Lam Phao, 1876 (P, isotypes B, BM, K, P).

C. bertelihansenii Rataj, Rev. Gen. Cryptocoryne, Studie CSAV, c 3: 49 (1975). Type: Beusekom & al. 4055, Nam Phrom (= Phu Khieo), 10.12.1971 (L, isotypes BKF, C, K, MU, P).

Leaves: 10 - 30 cm long, 0.4 - 1.6 cm broad, smooth (always ?), upright, rather stiff, usually (?) emerged, margin entire, or serrulate, occasionally slightly undulate (Figs. 7 F - G, 18 D - E). Spathe with a short or long tube, 10 - 20 cm long, limb with a rather long, somewhat tight to open spiral (Figs. 10 A, C - D, F).

Notes: Within what is here called var. crispatula, there seem to be several forms that show a certain variation, but I have not been able to subdivide it further, without recognizing another 4 - 5 taxa. Apparently var. crispatula mostly grows emerged, and has comparatively stiff leaves, ending in a shorter or longer apex. Some forms have an entire margin, while others have a more or less irregularly serrulate margin. The length of the leaves varies from 10 - 30 cm, apparently always (?) green. Some plants have a relatively long tube of the spathe ((10-) 12 - 30 cm) while others have a rather short one (7 - 12 cm).

Var. crispatula seems to be found in the central and northern Thailand and northern Vietnam.

C. crispatula Engl. var. flaccidifolia Jacobsen, nov. var.

Folia 20 - 50 cm longa, 0.7 - 1.2 cm lata, plerumque submersa (?), flaccida, viridia vel rubro-fuscida, plana vel paulum undulata. Tubus spathae (15-) 20 - 30 cm longus; limbus helice aperta satis longa terminatus

Holotypus die 17 Februarii anni 1929 prope vicum thailandicum Kampong Takua Pa [Mistake: should be Kapong, Takua Pa] sub numero 17126 ab A. Kerr lectus, siccus in Herbario Silvario urbis Bangkok (BKF) sub numero 24260 depositus, isotypis ibidem ut in Herbario Kewensi (K) et Herbario Parisiensi (P).

Leaves 20 - 50 cm long, 0.4 - 1.2 cm broad, usually submerged (?), flaccid, green to red-brownish, smooth or somewhat undulate (Fig. 7 E, 30). Spathe with a long tube, (15-) 20 - 30 cm long, limb of the spathe with a rather long, open spiral.

Notes: Var. flaccidifolia has long, narrow, flaccid, and smooth or slightly undulate leaves and apparently grows submerged. It is more broad-leaved than var. tonkinensis but narrower than var. balansae. Var. flaccidifolia is found in India, northern Vietnam, and northern and southern Thailand.

C. crispatula Engl. var. balansae (Gagnep.) Jacobsen, stat. et comb. nov.

C. balansae Gagnep., Not. Syst. 9: 131 (1941). - Lectotype (selected by Rataj, 1975): Balansa 264 (= 415), Chan Moi, 21.1.1921 (P, isotype K). Paratype: Kerr 5020, Tan Chum, 7.3.1921 (BK, K, P).

C. longispatha Merr., Journ. Arn. Arb. 23: 156 (1042). - Holotype: Petelot 2681, Phan Me, 12.11.1939 (GH).

C. kwangsiensis H. Li, Acta Phytotax. Sinica 15: 109 (1977). - Type: R.C. Ching 8505, Nanning, 2.12.1928 (Chung-Shan Univ., isotypes NY, W). Paratype: Ching 8504 (Chung-Shan Univ.).

Leaves 20 - 70 cm long, 1 - 4 cm broad, usually submerged, rather flaccid, green to brownish, slightly to strongly bullate (Figs. 2, 7 H, J – K, 19, 31, 32). Spathe with a long tube, 10 - 30 (-40) cm long, limb of spathe with a rather long and rather open spiral (Fig. 10 B, E).

Notes: Var. balansae develops into the largest and most spectacular of all the varieties when growing under optimal conditions, i.e. with leaves up to 70 cm long in a submerged state, the same plant being, however, only 10 - 15 cm tall when growing emerged. Besides the differences in leaf-size caused by growing conditions, there are also some forms that do not become as large and have darker leaves, just as forms with narrower leaves exist.

Var. balansae is found in central Thailand, northern Vietnam, and southern China.

C. crispatula Engl. var. sinensis (Merr.) Jacobsen, stat. et comb. nov.

C. sinensis Merr. Sunyatsenia 3: 247 (1937). - Type: Morse 221, Lungvhow (NY, isotypes K).

C. yunnanensis H. Li, Acta Phytotax. Sinica 15: 108 (1977). - Type: Chin 60132, Meng-la (Bot. Inst. Yunnan). Paratype: Y.C. Li 2728 (Bot. Inst. Yunnan).

Leaves 10 - 20 cm long, 0.5 - 1.5 cm broad, usually emerged (?), green(ish), smooth to slightly undulate. Spathe with a rather short tube, 8 - 12 cm long, limb of spathe with a rather short spiral (Fig. 41).

Notes: Var. sinensis seems to be a form with rather short, broad, stiff leaves. It is not known to be in cultivation.

Var. sinensis occurs in southern China.

C. crispatula Engl. var. tonkinensis (Gagnep.) Jacobsen, stat. et comb. nov.

C. tonkinensis Gagnep., Not. Syst. 9:133 (1941). - C. retrospiralis (Roxb.) Kunth var. tonkinensis (Gagnep.) De Wit, Aquarienpflanzen, p. 189 (1971). - Lectotype (selected by Rataj, 1975): Balansa 2045, Baa Toi, Aug. 1887 (P). Paratype: Balansa 2044, Baa Toi, 8.2.1887 (L, P). - Another paratype: Balansa 2043, Riviere Noire, 29.11.1887 (L, P) represents var. crispatula and is drawn in De Wit (1983), Fig. 51.

Leaves 20 - 30 cm long, 0.2 - 0.4 cm broad, flaccid, green to brownish, smooth or slightly undulate, usually submerged (Fig. 7 I). Spathe with a long tube, 15 - 30 long, limb with an open spiral (?).

Notes: Var. tonkinensis is the variety with the most narrow leaves of all, viz. 0.2 - 0.4 cm broad, completely flaccid, unable to carry themselves, showing that it is almost constantly submerged. Var. tonkinensis is only rarely seen in cultivation and most of the cultivated plants referred to as "tonkinensis" represent var. crispatula or var. flaccidifolia.

Var. tonkinensis is found in northern Thailand and in northern Vietnam.

Distribution: Eastern India, Thailand, Laos, Vietnam, and southern China (Fig. 6).

Habitat: The various ecological races are adapted to the varying water-bearings in the rivers in which they grow. Some are submerged all (or almost all) the year round while others are completely emerged during flowering (Dec.-Feb.).

Notes:

The foliage of C. crispatula is extremely variable, but the species may be recognized by the mostly elongate, purple markings on the long, spirally twisted limb of the spathe.

Live material of C. crispatula collected at the Phu Khieo National Park showed a variation from the green, narrow-leaved form to the brown, bullate, broad-leaved form, the smallest leaves measuring 5 by 0.3 cm, and being smooth, the largest measuring 40 by 2 cm, being bullate (Fig. 18).

The spathes of the various forms have an overall morphological similarity, variation being found only in the form and density of the purple markings of the limb. The markings vary from more or less irregular lines to short, line like-dots (Fig. 10). Their density varies from almost completely covering the limb (Fig. 36) to nearly missing and the transition is continuous. In some forms the shape of the tube is slightly trumpet-like dilatated below the limb (Fig. 10 D - E), while in others it is more or less broadly to almost club-like dilatated from below the opening and into the opened spiral of the limb (Fig. 10 C).

It is important to note that the leaves can vary in size during the flowering season, and when studying herbarium material this should be kept in mind. This variability can be illustrated by the specimen NJ 77-38 from Phu Khieo, cultivated in Copenhagen. Fig. 27 A shows the specimen flowering in the beginning of November and Fig. 27 B shows the specimen in late December, a few days after the 14th spathe had withered. Thus, the actual development stage related polymorphism of a specimen under study, must be known before the limits of the variation can be outlined.

I would interpret C. crispatula as one species containing different ecological races, i.e. leaf-forms that are adapted to the water supply and the overall topography of the rivers in which they grow. In rivers with more constant water-bearings, and not too steep a fall, the more aquatic ones occur, viz. the long, narrow-leaved forms with undulate or bullate lamina. In rivers with a seasonal variation in the water-bearings, the more amphibious ones occur, characterized by having shorter, more or less smooth leaves. There are transitions between the various forms. In cultivation, all can grow emerged as well as submerged, but usually a given form grows better either submerged or emerged.

The ecological races of C. crispatula are widely used as aquarium plants. Their leaf-form is rather constant in cultivation and the races more or less suited for submerged growth. Ten years ago I was not very keen on a subspecific delimitation of C. crispatula. In later years I have been convinced of the need for recognition of some intraspecific taxa. As I then advocated, the variation within C. crispatula is very complex, with the plants known in cultivation at least half a dozen recognizable forms being at hand, some more readily distinguishable from the next than others. However, above I have tried at the varietal level to outline the most easily recognizable and most commonly cultivated ones. I know that some plants may only with some difficulty, unambiguously be referred to a definite variety. The number of varieties accepted is subjective, and in a number of years some alteration may have to be made, e.g. some more varieties recognized. This is not meant as an excuse for an insufficient treatment from my hand, but merely a recognition of the complexity of C. crispatula.

A number of plants with different leaf-shapes, especially forms with a more narrow leaf-blade, have appeared in the trade during the last 10 - 15 years, shipped via Bangkok. We do not know anything about their place of origin but judging from their shape it is likely that they come from central to northern Thailand.

Cryptocoryne retrospiralis (Roxburgh) Kunth

Syn.: Ambrosinia retrospirale Roxb., A. uniloculare Roxb., Cryptocoryne unilocularis (Roxb.) Kunth, C. roxburghii Schott, C. dalzellii Schott.

Rhizome stout, 0.5-1.5 cm thick, sometimes irregularly thickened. Stolons stout. The root system is strongly developed, with also upright roots that thicken on reaching above the soil, and the plant may form a carpet-like growth. Leaves linear to lanceolate, green, sometimes brownish, 15-40 cm long and 0.3-1.0(-1.5) cm wide (Fig. 7 A – B, 42); lamina smooth or somewhat undulate, main rib forming most of the blade especially in more narrow-leaved specimens, the lamina mostly without prominent lateral veins. Margin sometimes denticulate. Monsoon leaves terete, 10-15 cm long. Spathe 10-30 cm long, tube 5-20 cm long, slightly twisted; limb of the spathe (1-) 3-8 cm long, spirally twisted, occasionally short and more or less open, yellowish to greenish, with rather large reddish spots (Fig. 8, 21, Jacobsen 1982). Female flowers 4-7, with more or less horizontal, oval to roundish stigmas. Olfactory bodies irregularly lobed in the upper part. Male flowers 100-140.

Chromosome number: 2n = 36, 72.

Distribution: India (Fig. 6).

Habitat: Sandy and stony bottom in and along rivers, often in rivers that carry a lot of water during the rainy season (Fig. 20). More or less emerged during flowering (Oct.-Feb.).

Nomenclatural notes: During the years there has not been much controversy regarding the identity of C. retrospiralis in the strict sense. C. unilocularis has been described from India, its characteristics being a unilocular fruit (which must be a mistaken observation) and a limb of the spathe spirally twisted from just above the kettle, the limb being purplish with darker spots. This long spirally twisted limb is interpreted as a monstrous spathe of C. retrospiralis having non fused margins of the tube normally formed. C. dalzellii was described on the basis of some fruits and a leaf-fragment, the latter now being interpreted as belonging to another, non Araceous genus, and the fruits belonging to C. retrospiralis.

Notes: C. retrospiralis is the only species from the western part of India with a long tube of the spathe and a spirally twisted, smooth limb of the spathe; it is characterized by the large, more or less circular, red spots on the limb (Fig. 8). Only few live collections (7) have been available for study. However, herbarium material is well represented and many well-preserved specimens exist, giving a quite distinct picture of the species.

References

Engler, A. 1920. Das Pflanzenreich, IV. 23. F. Araceae - Aroideae. Leipzig.
Jacobsen, N. 1976. Notes of the Cryptocoryne of Sri Lanka (Ceylon). - Bot. Notiser 128: 179 - 190.
Jacobsen, N. 1977. Chromosome numbers and taxonomy in Cryptocoryne (Araceae) - Bot. Notiser 130: 71 - 87.
Jacobsen, N. 1980. The Cryptocoryne albida group of mainland Asia (Araceae). - Meded. Landbouwhogeschool Wageningen 19: 183 - 204.
Jacobsen, N. 1982. Cryptocorynen. - Kernen Verlag, Stuttgart.
Jacobsen, N. 1985. The Cryptocoryne (Araceae) of Borneo. - Nord. J. Bot. 5: 31 - 51.
Rataj, K. 1975. Revision of the Genus Cryptocoryne Fischer. - Studie ČSAV, č 3. Praha.
Wit, H. C. D. de 1971. Aquarienpflanzen. - Ulmer Verlag, Stuttgart.
Wit, H. C. D. de 1983. Aquariumplanten. – Hollandia, Baarn (4th. ed).
Sivadasan, M. 1985. Cryptocoryne consobrina, eine seltene Art aus Südindien. - Aqua-Planta 10: 3 - 7.

Legends to figures:

ap16_1_p.jpg med big

ap16_2_p.jpg small big

ap16_3_p.jpg small big

Fig. 1. Flowering plants of Cryptocoryne albida at the locality at Ban Nang Yon (Note: Misspelled: Ban Wangyon, 1991), Thailand

Fig. 2. C. crispatula var. balansae in an aquarium

Fig. 3. C. albida in an aquarium

ap16_4_p_bottom.jpg small big

ap16_4_p_top.jpg med big

Fig. 4 & 5. Emerged stands of C. albida at Klong Kam Phuam and Khlong Nang Yon, south of Ranong (South Thailand)

ap16_5_p.jpg small med big

Fig. 6. Distribution of the Cryptocoryne albida group

ap16_7_p.jpg small med big

Fig. 7. Leaves of C. retrospiralis. A emerged; B monsoon leaf. - C. albida: C - D. - C. crispatula: var. flaccidifolia: E submerged; var. crispatula: F - G emerged; var. balansae: H brown submerged, J green emerged, K green submerged; var. tonkinensis: I ( × c. 0.4)

ap16_8_p_a.jpg small big

ap16_8_p_b.jpg small big

Fig. 8. C. retrospiralis . A. Kettle cut open. B. Limb of the spathe. Cult. (× c. 2.2)

ap16_9_p_a.jpg small big

ap16_9_p_b.jpg small big

ap16_9_p_c.jpg small big

ap16_9_p_d.jpg small big

ap16_9_p_e.jpg small big

ap16_9_p_f.jpg small big

Fig. 9. C. albida. Variation in the limb of the spathe. Ban Nang Yon: A - E (NJ 77 81 a, b, g, h, i. Cult.: F (× c. 2.2)

ap16_10_p_a.jpg small big

ap16_10_p_b.jpg small big

ap16_10_p_c.jpg small big

ap16_10_p_d.jpg small big

ap16_10_p_e.jpg small big

ap16_10_p_f.jpg small big

Fig. 10. C. crispatula. Variation in the limb of the spathe. Var. crispatula: A Phu Khieo; D Nam Nao; C - F cult.; Var. balansae: B cult.; E Phu Khieo (× c. 2.2)

ap16_11_p.jpg small big

ap16_12_p_bottom.jpg small big

ap16_12_p_top.jpg small big

Fig. 11. Mueak Lek. Habitat of the broad-leaved, submerged form of C. crispatula var. balansae

Fig. 12. Mueak Lek. Submerged plants of the broad-leaved form of C. crispatula var. balansae

Fig. 13. Phu Khieo Wildlife Sanctuary. Downstream course of the main river with an abundant growth of C. crispatula var. crispatula emerged on the banks

ap16_13_p_top.jpg small big

ap16_13_p_bottom.jpg small big

Fig. 14 (upper right). Emersed stand of C. albida south of Ranong (South Thailand)

Fig. 15 (lower right). Submerse plants of C. albida at Klong Kham Phuam, South Thailand

ap16_14_p_l.jpg small big

ap16_14_p_r.jpg small big

Fig. 16. Phu Khieo Wildlife Sanctuary. C. crispatula: A. Submerged plants with copper-brown leaves of var. balansae, from the small tributary; B. Emerged plants of var. crispatula from along the main course

ap16_15_p.jpg small big

Fig. 17. Phu Khieo Wildlife Sanctuary. Small shaded tributary just west of the main course of the river with submerged growths of the brown-leaved C. crispatula var. balansae

ap16_16_p_top.jpg small big

ap16_16_p_bottom.jpg med big

Fig. 18 (upper left). C. crispatula from Phu Khieo Wildlife Sanctuary (left to right): 1. Two submerged plants of var. balansae; 2. Submerged plant intermediate between var. balansae var. crispatula; 3. Half submerged var. crispatula; 4. Emerged, shaded var. crispatula; 5. Emerged, sun exposed var. crispatula

Fig. 19 (lower left). C. crispatula var. balansae in an aquarium

ap16_17_p_top.jpg med big

ap16_17_p_bottom.jpg small big

Fig. 20 (upper right). C. retrospiralis and Lagenandra toxicaria in a tributary to the Parakkaduvu Riber in the Malappuram District (Kerala, India)

Fig. 21 (lower right). Limb of the spathe of a cultivated plant of C. retrospiralis from the Poona River, India

ap16_20_p_top.jpg small big

ap16_20_p_bottom.jpg small big

Fig. 22. Ban Nang Yon. Habitat of C. albida on the riverbank showing the very dense growths found in suitable places

Fig. 23 (upper left). Habitat at Ban Nang Yon. Stands of green and brown leaved C. albida in river bed

Fig. 24 (lower left). A brown leaved and two green leaved of C. albida at Ban Nang Yon

ap16_21_p_top.jpg small med big

ap16_21_p_bottom.jpg small big

Fig. 25 (upper right). C. albida with intensely red/brown coloured leaves in emersed cultivation

Fig. 26 (lower right). Uprooted plants of C. crispatula var. balansae from Mueak Lek: Left a submerse plant, right an emersed plant

ap16_22_p_l.jpg small big

ap16_22_p_r.jpg small big

ap16_23_p.jpg small big

Fig. 27. C. crispatula var. crispatula from Phu Khieo cultivated in Copenhagen: A in early November; B in late December

Fig. 28. C. crispatula var. crispatula with an irregularly denticulate leaf margin

ap16_24_p_top.jpg med big

ap16_24_p_bottom.jpg small big

Fig. 29. Emerged stand of C. crispatula var. crispatula from along the main course at Phu Khieo Wildlife Sanctuary

Fig. 30 (lower left). Emersed cultivated plant of C. crispatula var. flaccidifolia

ap16_25_p_top.jpg med big

ap16_25_p_bottom.jpg med big

Fig. 31 (upper right). Emersed plants of C. crispatula var. balansae from Mueak Lek on the river bank

Fig. 32. Submerse plants of C. crispatula var. balansae from Mueak Lek

ap16_26_p.jpg small med big

Fig. 33. Opened fruits of C. crispatula var. crispatula

ap16_27_p_tl.jpg small big

ap16_27_p_tr.jpg small big

ap16_27_p_br.jpg small big

Fig. 34. (upper left). Partly submerse growing plant of C. crispatula var. crispatula from Phu Khieo Wildlife Sanctuary

Fig. 35 (upper right). C. crispatula var. crispatula from Phu Khieo Wildlife Sanctuary, which has begun flowering and produce leaves after a resting period

Fig. 36 (lower). Spathe of C. crispatula var. crispatula

ap16_28_p_tl.jpg small big

ap16_28_p_tr.jpg small big

ap16_28_p_bl.jpg small big

ap16_28_p_br.jpg small big

Fig. 37/38 (upper). Emersed cultivated plants of C. crispatula var. balansae; limb of spathe

Fig. 39/40 (upper). Emersed cultivated plants of C. crispatula var. balansae from Phu Khieo; limb of spathe

ap16_30_p.gif small med big

Fig. 41: C. crispatula var. sinensis. 1. Plant; 2. Leaf; 3. Spathe; 4: Open kettle; 5. Fruit (From Flora Yunnanica, Vol. 2, 1979)

ap16_33_p.jpg small big

Fig. 42. C. retrospiralis with short, terrete leaves which are produced during our autumn and winter (short days)