1990 - The first description

PHRAGMIPEDIUM XEROPHYTICUM,
UNA NUEVA ESPECIE DEL SURESTE DE MEXICO

a new species from southeastern Mexico

Orquidea (Mex.) 12:1-10. 1990

Phragmipedium xerophyticum,

Miguel Angel Soto, Gerardo A. Salazar & Eric Hágsater

Abstract
The new species, Phragmipedium xerophyticum, is described. The plant apparently has no close relatives and is characterised by xeromorphic [drought-adapted] plants, elongated rhizomes, fleshy, short, rigid leaves, inflorescences with compressed racemes, small, white and pink flowers, linear petals, a cup-shaped, roundish, inflated labellum and a single-chambered (unilocular) ovary with parietal placentation. The possible relationships are discussed.

The AMO herbarium is constantly receiving plant material for identification. This material often contains very interesting samples and is a very valuable source for our work. Among the material collected by the group of collaborators of Dr Thomas Wendt from the Colegio de Postgraduados de Chapingo, we were surprised by the presence of a very peculiar plant that we were initially unable to identify due to the absence of flowers. It consisted of small leaf fans connected by long rhizomes and produced a very hairy apical inflorescence. These features suggested that it might be a Cypripedioidaea and our curiosity grew. Photocopies of the specimen were sent to a number of specialists, including Dr Robert Dressler, in the hope that they might provide clues to the identity.

Shortly afterwards, Rolando Jiménez from the AMO discovered a specimen from the same collection, but with a flower that confirmed that it was indeed a Cypripedioideae, probably a Phragmipedium.

Faced with such a find, we decided to organise an expedition to the site as soon as possible in order to obtain living material of such an interesting plant. The biologist Patricia Vera Caletti explained to us how we could get in touch with Mr Heriberto Hernández, the collector of the plant. She also pointed out to us that it is difficult to get to the region during the rainy season [to protect this new species, the exact location where it is found is omitted] as there is no passage for vehicles due to the high water level of the rivers.

After locating Mr Hernández, he kindly took us to the place where he had collected the plant we were interested in years before. The region where it was collected has very rich and diverse vegetation. In addition to the evergreen tall forest that one would expect in a warm and humid region at an altitude of 300 metres, there are also extensive oak, pine and liquidambar forests; a very impressive mosaic for any botanist. In certain parts near the rivers there are rugged karst areas that form sparse scree slopes, with sparse and rather xeromorphic vegetation. Small trees such as Bursera simaruba, Plumeria rubra, Pseudo-bombax ellipticum grow in the crevices where the humus accumulates and, despite the otherwise humid environment, the unusual presence of Beaucarnea, Yucca, Agave and Acanthocereus could be observed. This special biotope is the habitat of the plant we are now talking about.

The plant is by no means numerous and it took us a while to find the first specimens, which fortunately were in flower. We immediately realised that this was a new species of Phragmipedium, different from all previously described species. These are very small plants with very distinct vegetative propagation, and the flowers are also very small, white with pink speckles. The labellum is inflated, globose, delicate, incurved at the margin, has an uneven surface structure and is reminiscent of the labellum of some Cypripedium species or that of Paphiopedilum micranthum T. Tang & Wang and its relatives. The only Phragmipedium species with such a labellum are P. schlimii (Linden & Rchb. f.) Rolfe and P. besseae Dodson & Kuhn, but these species have very distinct broad petals. Their description can be found below:

Phragmipedium xerophyticum Soto, Sala zar &Hágsater, sp. nov.

Plant: Relatively small, clonally growing, rupicolous [growing on stone] herb, consisting of distichous (two-rowed) shoots and elongated rhizomes, about 15-20 cm tall and probably up to 1.0 m2 in size.

Roots simple or sparsely branched, slender, light brown or whitish, glabrous or hairy on the surface in contact with the substrate, strongly adherent to the bare rock or penetrating into the humus, originating only from the base of the shoots and absent from the rhizome, up to 0.8 mm thick and 11 cm long.

Rhizome very compact, elongated, almost straight, hard and brittle, consists of 5-12 internodes covered by as many leaf sheaths. Diameter 1-2 mm, length between the individual shoots 3-8 cm.

Rhizome sheaths numerous, scarious, conspicuously veined, brownish, tubular or somewhat cone-shaped, blunt or pointed, notched, loose, spaced or overlapping, chain-like, 6-9 mm long.

Shoots of 5-8 distichous (two-rowed) leaves, which are 3-4, occasionally up to 12 cm high and 6.5- 13 cm wide.

Leaves strongly convex, ribbon-like, tip unequal, obtuse, pointed, leaf surface crenate, very rigid, light green, once dead they seem to remain on the shoot for a long time, turning brown, basal leaves small, intermediate form between rhizome sheath and stem leaf, upper leaves gradually increasing in size, 3.5-12 cm long and 1.2-1.8 cm wide, approx. 1 mm thick.

Inflorescence apical, inflorescence formed of 2 internodes, elliptic in cross-section; an inflorescence with 2 panicles (rarely only one panicle), the upper panicle developing first, and only later the lower one, probably when the apical one no longer produces flowers; Hairy, with multicellular hairs of different lengths, reddish-brown, more abundant near the flower sheath, gradually less abundant towards the apex and also more ingrown until the surface is practically glabrous or slightly warty; total length of inflorescence 6.5-13.5 cm, 1- 1.3 mm thick.

Inflorescence bracts 1, about halfway up the pedicel, concave, tubular at the base, broadly rounded or with an extension at the tip, not divided, yellowish, densely hairy at the base, gradually with a few adhering hairs, medially with short cilia, glabrous apically, 8-15 mm long.

Rhachis very short with about 3-7 consecutive flowers, only one flower open at a time, about 12-15 mm long;

Perianth imbricate [roof-tile-like], two-lined, strongly conduplicate, cylindrical, with caudal, recurved and thickened apex, dark brown, hairy and ciliate, the hairs multicellular, reddish, 4-5-veined; when spreading (which is not possible without some damage) broadly triangular, 4-5 mm long, 5 mm wide, gradually smaller towards the apex.

Pedicel short, almost completely covered by the bracts, very stiff, inclined, hairy, the hairs are multicellular, 2.5-3 mm long, 0.8 mm thick in the centre, widening towards the break-off zone of the ovary.

Ovary unilocular (in cross-section basal, medial and apical), placentation parietal; three-axis, elongated, attenuated towards the base and apex, densely hairy, the hairs multicellular, pink, approx. 2.7-2.8 cm long, 1.5 mm at the thickest point.

Flower small, showy, not fragrant, very similar to Cypripedium californicum, 1.3-2.5 cm high, 1.5-2.0 cm in diameter.

The perianth falls off and hangs down, although the flower is apparently fresh; it is white with a pink tinge and very weakly structured.

Sepals keeled, the lateral ones usually completely fused into a synsepalum, occasionally the fusion is not complete, the abaxial surface is smooth, the adaxial surface is conspicuously hairy, the hairs are multicellular, denser and longer towards the apex and near the separation zone with the ovary.

Dorsal sepal pointing forwards, elliptic, apex pointed-subacute, crenate and slightly thickened, 7-8-veined, veins branched and anastomosing towards the apex, concave, 9- 14 mm long and 5-6.5 mm wide.

Synsepalum pointing downwards, suborbicular, obtuse (or, if the fusion is not complete, with 2 adjacent tips), tapering and thickened at the apex, 12-nerved, veins branched and anastomosing towards the apex, concave, 8-9.5 mm long and 8.5-12 mm wide.

Petals linear-band-like, pointed, arched, sometimes slightly bent downwards, not or only slightly twisted, margin wavy, five-veined, glabrous or ciliate at the base, 11-15 mm long, 2.5-3 mm at the widest point.

Labellum cup-shaped, bulbous, inflated, slightly furrowed along the veins near the opening, very delicate in texture; outer surface smooth, the inner surface near the base conspicuously hairy, with glandular, multicellular, bright purple, very attractive hairs, towards the underside of the labellum the hairs are much reduced, continuing along the midline and are white and apparently thinner and sparser and clustered; Basal margins (around the exit hole) erect, somewhat recurved and thickened; apical margin curved; entrance hole small, ovoid, ca. 2 x 3 mm; margins of labellum not very long. 2 x 3 mm; lateral lobes small, inwardly curved, broadly triangular, subacute, without thickening at margins, nor forming outgrowths (horns) or hollow, bulging areas, adherent to each other, ca. 3 mm. Entry and exit holes well defined; labellum surface without ‘windows’ or transparent areas; total length of the labellum 10-14 mm, 6-8 mm high, 7-9 mm wide.

Columna [gymnostemium] short, gynoecium and androeceum only 1-2 mm fused, almost completely covering the exit opening.

Stigma curved, downward-directed, fleshy and massive, consisting of an almost trilobed body with a horizontal surface in front of the staminode, provided with a less conspicuous longitudinal margin and with multicellular trichomes, which are longer near the middle part; the tip of the body is directed downwards and longitudinally serrate;

Stigma lobes forming a more or less fan-like, apical structure projecting beyond the staminodes, thick, concave, ovate-triangular; densely hairy on the abaxial surface, with tiny papillae on the adaxial surface, the lateral lobes reduced to two inconspicuous margins on the lower surface; 4-5 mm long, 1. 0-1.3 mm at the widest part.

The stamens have 2 fertile anthers and a petaloid staminode. Anthers conspicuously stalked (c. 1 mm) convex, broadly 3-lobed, apex subacute and rounded, incised lengthwise and glabrous on the outer surface, purple or violet; with an axial rib on the inner surface, with two groups of multicellular, long, pale purple hairs along the rib, the rest of the inner surface smooth; lateral lobes elongated and directed downwards, nearly square, with slightly reddened margins; overall 3 mm long, 4-5 mm wide.

Anthers 2, each at the end of an elongated stalk, broad, short, slightly curved, fleshy, glabrous except for the slightly papillose apex; the actual anther is perpendicular to the columna and connected to it by a very small area; ovoid-triangular, somewhat keeled, pointed or blunt, fleshy, white, the apex directed downwards and outwards; with 2 ventral, brown, sucker-like zones containing the pollinia, separated by a groove that continues into a depression in the apical part of the anthers; ca. 1 mm long.

Pollinia 2 in each anther, adhering to each other, forming an oval, bevelled structure, granular, yellowish, 0.5 x 0.4 mm.

Mature seed pod not known, the young pods similar to the ovary, but somewhat thicker.

Alte Fotografie von Mexipedium xerophyticum
Alte Fotografie des Habitats von Mexipedium xerophyticum
Alte Fotografie von Mexipedium xerophyticum
Alte Fotografie von Mexipedium xerophyticum

Holotype: Mexico: Oaxaca: jungle on the slope of the Gulf of Mexico, 320 m a.s.l., xerophytic vegetation of Agave, Beaucarnea, Bursera simaruba, Plumeria and Pseudobombax ellipticum, in the karst zone, surrounded by a tall evergreen forest and tropical holm oak forests; plant growing on rocks, rare, 6 September 1988, G.A. Salazar 3740, M.A. Soto. E. Yañez und H. Hernández, AMO! ISOTYP: K!

Other specimens: Mexico: Oaxaca: Same locality and date, G.A. Salazar 3742, M.A. Soto, E. Yañez and H. Hernández, US! Type locality, 24 September 1985, Heriberto Hernández G. 1602, CHAPA!

Distribution: Endemic to Mexico. So far only known from one location in the warm and humid region of Oaxaca.

Ecology: This is the most xeromorphic species of the genus. Its clonal dispersal is very remarkable, and over time several parts of the plant become independent. This must be an advantage for the population, as the development of seedlings in this apparently seasonal environment seems to be a rare event. This type of growth has also been described for other species of the subfamily Cypripedioideae with distichous leaf arrangement, such as Paphiopedilum druryi Bedd., Phragmipedium pearcei (Rchb.f.) Rauh and P. besseae. The habitat of Phragmipedium xerophyticum is in some respects similar to that described for Paphiopedilum druryi. The plants were observed in the upper, treeless parts of the scree, but never in completely exposed places. Instead, they were found on vertical walls with north and east exposure, in small crevices with humus accumulation or on bare rock. None of the plants observed received direct midday sun. The most robust plants grew in humus with Selaginellas and Pitcairnias. The region receives about 2500 mm of rainfall throughout the year with an average annual temperature of about 25° Celsius. There is a pronounced dry season in spring.

Flowering time: The plants flowered in September at the site, and the flowering period probably extends over the months of the rainy season. During collection, some small wasps were observed around a flower; however, they were not observed to enter the flower. The flowers are apparently not self-pollinating, although numerous seed pods have been observed.
The pollinator should be small, measured by the size of the entrance hole. Cribb (1987) has suggested that the very similar flowers of Cypripedioideae with spherically inflated labellum (Cypripedium irapeanum Llave & Lex., Phragmipedium schlimii, Paphiopedilum micranthum etc.) are the result of adaptations to similar pollinators.

Identification: The combination of very small, 1.5-2 cm white and pink-coloured flowers and the elongated rhizomes of the plant make it unmistakable. The other Mexican species of Phragmipedium, P. exstaminodium, is so different that it is impossible to confuse it, as it is a yellow-flowered, epiphytic plant with earth and chestnut tones, whose petals are 25-45 cm long. The other species of the genus with white-pink flowers is P. schlimii from Colombia, but it is easy to distinguish from it because the latter has broadly ovate to rounded petals.

Notes: The genus Phragmipedium is characterised by (1) the valvate sepals, (2) the absence of curved epidermal cells in the perianth, (3) the lateral lobes of the labellum which are fused together, and (4) synsepals which are larger than the dorsal sepal. All these features are present in the new plant. With the exception of the taxa of the P. caudatum (Lindl.) Rolfe complex, all species have one or more sterile leaf sheaths on the inflorescence; in P. xerophyticum the leaf sheath on the inflorescence usually forms a new flower raceme. With the exception of P. schlimii, all Phragmipedium species have two hollow areas near the base of the labellum; these hollow areas are also absent in Phragmipedium xerophyticum.

A very unusual feature of Phragmipedium xerophyticum is the presence of a single-chambered (unilocular) ovary with parietal placentation (seen in transverse sections at the base, centre and apex of the ovary chamber). This type of ovary occurs in Paphiopedilum and Cypripedium, but is not reported for any species of Phragmipedium, where the ovary has always been described as three-chambered and with axillary placentation. Atwood (1984) mentions that in Paphiopedilum and Cypripedium intermediate forms between one- and three-chambered ovaries occur (one-chambered in the centre, but with two or more chambers at the ends), so that highlighting this feature is not justified. The same author notes incomplete fusion between neighbouring locules in a specimen of Phragmipedium schlimii, suggesting that this is a first step in the development of parietal placentation.

With the exception of the unilocular ovary, all other features of Phragmipedium xerophyticum fit well with Phragmipedium, but it is not easy to place it in one of the proposed sections. The genus is divided into three sections according to Atwood (1984), Micropetalum, Phragmipedium and Lorifolia, although Garay (1979) divides the species assigned by Atwood to section Lorifolia into three different sections. The Phragmipedium section has flowers with very elongated petals that continue to grow for several days after unfolding. The flowers open simultaneously and the inflorescences are unbranched. Apart from the generic characteristics, this section has little in common with P. xerophyticum. The section Lorifolia is somewhat more similar, but still distinguishable, as it includes plants with elongated leaves whose flowers are basically green in colour but with a reddish tinge. The labellum is slipper-shaped, not inflated, has hollow areas and in some species well-developed hairs. The petals are elongated, often with urine-scented tips. Similarities with this section are limited to the presence of elongated rhizomes (e.g. P. pearcei), the branched inflorescences, and the general shape of the petals. There are some similarities with the two species of section Micropetalum (Phragmipedium schlimii and P. besseae; see Atwood 1984; Dodson and Kuhn 1981; Hegedus and Stermitz 1986). White flowers with pink hues are only known from Phragmipedium schlimii and xerophyticum. The inflated, unstructured labellum also only occurs in these three species. However, there are some remarkable differences between P. schlimii-P. besseae and P. xerophyticum. The structure of the inflorescence is completely different: Micropetalum has successive single flowers with large leaf-like bracts, quite unlike the paniculate inflorescences of P. xerophyticum with shortened flower racemes and inconspicuous, very hairy small bracts. The section name Micropetalum, small petal, refers to the presence of petals that are much smaller than in the other species of the genus and that have a rounded-oval shape, i.e. are not elongated. The new plant also lacks the ‘windows’ (‘transparent areas’) on the sides and back of the labellum. In P. schlimii and P. besseae the surface of the entire flower is conspicuously hairy, whereas in P. xerophyticum the hairy surfaces are confined to the outside of the sepals, although there are some trichomes [plant hairs] on the basal margin of the petals and on the inside of the labellum. The staminodes of P. xerophyticum are glabrous and resemble those of P. boissierianum (Rchb. f.) Rolfe (Lorifolia) rather than those of P. schlimii or P. besseae (which are also conspicuously hairy). Although the leaves of plants of section Micropetalum are generally shorter and broader than those of the other Phragmipediums and the plants have the appearance of Paphiopedilum, they do not achieve the compact, xeromorphic growth of P. xerophyticum. The strongly articulated leaf epidermis cells described for P. schlimii are apparently also missing. The similarity between the flowers of P. schlimii and P. xerophyticum could be due to the same causes that Cribb (1987) mentions to explain the inflated labellum in all Cypripedioideae genera.

Botanische Illustration von Mexipedium xerophyticum

The new Phragmipedium is geographically quite distant from its other relatives. The species with the closest distribution is P. exstaminodium from the section Phragmipedium. None of the Central American species appear to be closely related either.

This species is very interesting from various points of view, and it should be mentioned that the presence of a plant that is apparently endemic to the warm and humid zone of Mexico is an unusual event, as species from these regions normally extend at least as far as Central America.

Ethymology: The specific epithet, xerophyticum, alludes to the relatively dry habitat of this species (compared to the habitats of its relatives) and to its vegetative morphology.

Conservation status: Threatened with extinction. If we apply the rarity criterion described by Rabinowitz et al. (1986), it can be stated that it is a very rare plant, as its geographical distribution (as far as we know) is very limited (one site), it is restricted to a specific habitat (it only grows in exposed karst areas) and its populations are very small (no more than 7 clones are known). Phragmipedium xerophyticum is so rare in the wild that all known plants can be collected in the field in a few hours; any collection of wild plants could have very harmful effects. It is recommended to avoid collecting wild plants as much as possible, even for scientific work. Some plants have already been distributed to recognised breeders and it is hoped to distribute seeds to specialised nurseries and botanical gardens to avoid collection pressure on natural populations. Recently (CITES Plants Committee, 7th meeting of the Conference of the Parties in Lausanne, Switzerland, 9-20 October 1989) all Paphiopedilum and Phragmipedium species were included in CITES Appendix I. Although this measure is probably very radical, it seems appropriate at least for such rare plants as the Mexican Phragmipediums.

Acknowledgements: We would like to thank Mr Heriberto Hernández for guiding us to the site and the local authorities for facilitating our stay in the region. Patricia Vera Caletti and Thomas Wendt provided us with information about the region. Elvira Yañez took part in the collection campaign with great commitment. Rolando Jiménez prepared the illustration. We would also like to thank Lucile McCook, Ed Greenwood and Fernando Chiang, who made important suggestions for the manuscript.

Botanische Illustration von Mexipedium xerophyticum
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