74 Araceae/Bromeliaceae

Schott, H.W. 1860. Prodromus systematis Aroidearum. Vindobonae: typis congregationis mechitharisticae.

Schott, H.W. 1984. leones Aroideae et reliquiae. Microfiche and index. (Nicolson, D.H., ed.) Zug: IDC AG.

Seubert, E. 1993. Die Samenmerkmale der Araceen und ihre Bedeutung fiir die Gliederung der Familie. Koenigstein: Koeltz.

Shaw, D.E., Cantrell, B.K. 1983. A study of the pollination of Alocasia macrorrhiza (L.) G. Don (Araceae) in southeast Queensland. Proc. Linn. Soc. N. S. W. 106: 323-335.

Silva, C.J. da 1981. Observa<;6es sobre a biologia reprodutiva de Pistia stratiotes L. (Araceae). Acta Amazonica 11: 487-504.

Simmonds, N.W. 1950. Notes on the biology of the Araceae of Trinidad. J. Ecol. 38: 277-291.

Solereder, H., Meyer, F.J. 1928. Systematische Anatomie der Monokotyledonen, Vol. 3. Berlin: Borntraeger, pp. 100-169.

Stockey, R.A., Hoffman, G.L., Rothwell, G.W. 1997. The fossil monocot Limnobiophyllum scutatum: resolving the phy­logeny of Lemnaceae. Am. J. Bot. 84: 355-368.

Takhtajan, A.L. 1982. See general references. Thanikaimoni, G. 1969. Esquisse palynologique des aracees.

Trav. Sci. Sect. Tech. Inst. Fr. Pondichery 5: 1-31. Thompson, S. 1982. Cyrtosperma chamissonis (Araceae): ecol­

ogy, distribution and economic importance in the South Pacific. J. Agric. Tradit. Bot. Appl. 29: 185-203.

Tillich, H.-J. 1985. Keimlingsbau und verwandtschaftliche Beziehungen der Araceae. Gleditschia 13: 63-73.

Tillich, H.-J. 1995. Seedlings and systematics in mono­cotyledons. In: Rudall, P.J., Cribb, P.J., Cutler, D.F., Humphries, C.J. (eds.) Monocotyledons: systematics and evolution. Royal Botanic Gardens, Kew, pp. 303-352.

Uhlarz, H. 1985. 1st Pinellia tripartita (Blume) Schott (Araceae-Aroideae) habituell anemophil geitonogam? Beitr. Bioi. Pflanz. 60: 277-291.

Valerio, C.E. 1984. Insect visitors to the inflorescence of the aroid Dieffenbachia oerstedii (Araceae) in Costa Rica. Brenesia 22: 139-146.

Vogel, S. 1963. Duftdriisen im Dienste der Bestaubung: iiber Bau und Funktion der Osmophoren. Abh. Math.-Naturwiss. Kl. Akad. Wiss. Mainz 1962 (10): 639-677.

Vogel, S. 1978. Pilzmiickenblumen als Pilzmimeten. Flora 167: 367-380.

Wang, J.-K. (ed.) 1984. Taro: a review of Colocasia esculenta and its potentials. Honolulu: University of Hawaii Press.

Wettstein, R. 1935. Handbuch der systematischen Botanik, 4th edn. Leipzing: Deuticke.

Williams, C.A., Harborne, J.B., Mayo, S.J. 1981. Anthocyanin pigments and leaf flavonoids in the family Araceae. Phy­tochemistry 20: 217-234.

Williams, N.H., Dressler, R.L. 1976. Euglossine pollination of Spathiphyllum (Araceae). Selbyana 1: 349-356.

Young, H.J. 1986. Beetle pollination of Dieffenbachia lon­gispatha (Araceae). Am. J. Bot. 73: 931-944.

Bromeliaceae

L.B. SMITH1 and W. TILL

Bromeliaceae Juss., Gen. Pl. 49 (1789), "Bromelieae".

Perennial rosette herbs with a short axis, more rarely with an elongated stem, or rosette trees, from few centimeters to more than lOrn, terres­trial, saxicolous, or epiphytic. Roots absorbing in terrestrials, modified to holdfasts in epiphytes and epilithes, rarely lacking. Leaves spirally arranged, dilated-sheathing at the base, undivided, entire or spinose-serrate, usually bearing peltate trichomes at least when young. Stipules lacking. Inflores­cence terminal, often becoming pseudolateral by sympodial growth, scapose or sessile, compound or simple, usually bearing brightly colored con­spicuous flower bracts or "petaloid" scape bracts. Flowers perfect or sometimes functionally uni­sexual (all Hechtia, Catopsis p.p.), usually actinomorphic. Perianth heterochlamydeous, the 3 sepals and 3 petals free or connate, the petals often with scalelike basal appendages and/or lon­gitudinal callosities. Stamens 3 + 3; filaments free, or joined to the petals and/or to each other; an­thers introrse, bithecal, 4-loculate, latrorsely de­hiscent. Ovary superior to inferior, 3-carpellate, 3-loculate; placentae axile, extending the length of the ovary or variously reduced; ovules anatro­pous, naked or with a chalaza! appendage; style 1, stigma usually 3-lobed; septal nectaries always present. Fruit usually a septicidal capsule or baccate; seeds winged, plumose, or naked; embryo small, cylindrical, situated laterally at the base of the copious starchy endosperm.

A neotropical family of 56 genera and more than 2600 species distributed from N America (Virginia) to southern America (Patagonia); a single species in W Africa.

CHARACTERS OccuRRING IN FEw GENERA AND SPECIES. Rheophytic plants in Pitcairnia and Guzmania; climbing ones in Pitcairnia; distichous phyllotaxis in Tillandsia, mainly in subgenus Diaphoranthema; petiolate leaves in Pitcairnia, Ronnbergia, Bromelia, and Cryptanthus, cuticular

1 Deceased. K. Kubitzki (ed.), Flowering Plants · Monocotyledons© Springer-Verlag Berlin Heidelberg 1998

Bromeliaceae 75

water absorption without functional peltate tri­chomes in Acanthostachys; lateral inflorescence in Dyckia; reduction to 1 flower in Tillandsia; pres­ence of glandular hairs in Puya, Deuterocohnia, and Pitcairnia; zygomorphic flowers in Pitcairnia and Tillandsia; !epidote petals in Pitcairnia, Dyckia, and Hechtia; paired petal appendages in the pitcairnioid genera Brewcaria and Steyer­bromelia; more than 1 vascular branch in each filament in Aechmea, Billbergia, Bromelia, Dyckia, Encholirium, and Vriesea; connectives with an apical appendage in Androlepis or an apical glan­dular outgrowth in Neoregelia eleutheropetala (Ramirez 1991); semiintrorse anther dehiscence in Dyckia and Vriesea; undivided stigma in Wer­auhia and Aechmea; additional apical loculicidal dehiscence of the capsules in Deuterocohnia, Dyckia, Encholirium, Fosterella, Hechtia, Pit­cairnia, and Puya; seeds with a strongly elongated chalaza! appendage in Acanthostachys.

VEGETATIVE MoRPHOLOGY. The roots are often much modified due to functional adaptations. Intracauline roots are present in species with a distinct stem (Fig. 19).

The usually erect stem ranges from extremely short in species with rosulate leaves to the stout 8-9.5-m-high columns of Brocchinia micrantha and Puya raimondii, it is usually branched and ligni­fied to various degree. Tillandsia usneoides is distinctive by its pendent stems without roots. Rhizomes may travel several meters underground (Bromelia, Deinacanthon, Neoglaziovia, Pit­cairnia, and Pseudananas) or be completely aerial as in species of Aechmea, Cryptanthus, Neoregelia, Tillandsia, and Vriesea.

Leaves vary from lingulate to narrowly triangu­lar or even filiformous, from fiat and soft to rigid and distinctly succulent. The leaf sheaths are usu­ally dilated but often merge in the leaf blades, rarely both are separated by a distinct petiole. The green coloration is often masked or ornamented by anthocyanin or by a dense indumentum of tri­chomes. In Tillandsioideae the leaves are always entire, while in Pitcairnioideae and Bromelioideae the leaves are predominantly spinose-serrate.

In Pitcairnia leaf dimorphism or even polymor­phism is observed in which the outer leaves are narrow and spiny and the inner ones broad and even pseudopetiolate. In some long-caulescent species one type may grade into the other, some­times repeatedly every year. In rosulate species the leaves often form a tank (phytotelm, Janetzky and Vareschi 1993; Krugel1993) which holds water, a feature that is lacking in caulescent species with

Fig. 19. Bromeliaceae. Schematic longitudinal section of an axis showing intracauline adventitious roots growing down­wards in the cortex and branching in the substrate. (Weber 1954)

trichoma! moisture and nutrient absorption. Dis­tichous phyllotaxis is rare (e.g., Till a ndsia subge­nus Diaphoranthema).

In terrestrial Pitcairnioideae, tank rosettes are lacking with the exception of Brocchinia species which have well-developed "pitchers". In Tilland­sioideae and Bromelioideae tanks of various types are common.

VEGETATIVE ANATOMY. A comprehensive and critical treatment was given by Tomlinson (1969), and modern data are supplied by Benzing (1980, 1990), who should be consulted also for functional and ecological aspects.

From the periphery to the center, the roots con­sist of a piliferous layer with well-developed root hairs in soil roots, of a multiseriate exodermis of suberized cells, of the outermost cortical layers immediately within the exodermis which are always differentiated as a wide sclerotic cylinder forming a conspicuous mechanical layer, of the middle cortex of thin-walled parenchyma, of the lacunose inner cortex, of a uniseriate thick­walled endodermis, of a 1-(2)-layered thin-walled

76 Bromeliaceae

pericycle, and of a stele with the vascular tissues and pith cells. The pith is strongly lignified in spe­cialized taxa (Strehl and Winkler 1983). The xylem always contains at least a few vessels with scalari­form perforations only or tracheids (Wagner 1977); only for Pitcairnia have specialized vessels been recorded (Cheadle 1955). The outer tissues are separated from the inner ones by the clearly defined endodermis. Intracauline roots are located in the stem cortex and differ from external roots in the absence of root hairs, exodermis, and lacunae. Roots of epiphytic species are often modified from the above scheme.

The stem anatomy is quite uniform within the family. A thin cuticle covers an epidermis which contains spherical silicia bodies. The narrow cortex has narrow, elongated, thick-walled, ligni­fied cells in the outer hypodermal layers which eventually are suberized in old stems; the inner hypodermal layers are transitional to thinner­walled, more isodiametric cells of the inner cortex. Periderm is developed from secondary meristem­atic layers. The central cylinder is delimited from the cortex by a narrow, 2-4-layers-wide zone of elongated cells which become sclerotic in older stems. The peripheral sclerenchyma is separated from the vascular tissue by a narrow, thin-walled layer, termed pericycle by some authors. The vas­cular bundles lie diffuse in the center but are more congested in the periphery of the central cylinder, and the bundles have a more or less distinct bundlesheath. The xylem contains mainly trache­ids, vessels are rare and less often encountered than in roots (Cheadle 1955; Wagner 1977). The sieve elements of the phloem contain protein crys­talloids of the Pile type in their plastids (Behnke 1981). The anatomical character of the stem is somewhat confused by the strong suppression of internodial growth and by the junction of adventi­tious roots. The arrangement of the stem bundles conforms to the meristele type, but the collateral vascular bundles are sometimes reversed and may be sickle-, crescent-, or annular-shaped. The stem of the Bromeliaceae is capable of transition to pri­mary radial growth through a cambial cap at the stem apex. This growth tissue apparently can con­tinue to develop after flower formation.

The fundamental character of the leaf of the Bromeliaceae is the small-celled and small­lumened epidermis with variously thickened walls. Another unique feature of the epidermis is that the greater cell dimension is often at right angles to the long axis of the leaf. Some species are distinguished by lateral walls strengthened by sinuous folds so that adjacent cells interlock with

one another. The entire surface of the epidermis has a smooth coating of the cuticle (Harms 1930). Epicuticular wax crystalloids of the Strelitzia type (Barthlott and Frolich 1983) occur in several spe­cies. Adequate protection and strength for the leaves of most Tillandsioideae and Pitcairnioideae seem to be provided by the epidermis. In Puya, Ananas, and others the epidermis is further strengthened by a sclerenchymatous hypodermis whose cells are generally longer than the epider­mal cells. These hypodermal cells often have brownish contents and sometimes contain sili­ceous crystal fragments. Opaque yellowish drop­lets of tannins also occur in the hypodermal cells and seem to be characteristic of the family.

Chlorophyll is generally not contained in typical palisade cells, which are completely absent from a number of species. Chlorophyll-bearing cells are nearly always separated from the upper leaf surface by water-storage cells and their position otherwise includes a great variety of types. Corru­gated leaves with longitudinal furrows on their lower side provide mechanical strength which is caused by vascular bundles and bundle-sheath fibers that make up the convex portion of the cor­rugations. The stomata usually lie on the sides, and the stalks of the trichomes lie in the bottom of the concave portion. In cross-section, leaves are often crescent-shaped, which enhances surfacial water conduction. In addition, such leaves are 51 times more resistant to bending than a flat blade of the same surface area.

The stomata of the Bromeliaceae are located almost entirely on the underside of the leaf, usu­ally at the same level as the epidermis, although raised accessory cells sometimes produce pseudo­stomata. Stomata are characterized by especially narrow lumina and the presence of at least 1 lat­eral and 1 polar pair of accessory cells. Guard cells seem to be relatively uniform throughout the fam­ily but the substomatal pores are distinctive enough for Robinson (1969) to have used them as part of a classification system for Navia. Stomatal movement is usually lacking, with the exception of Ananas; guard cells are not separated.

The peltate trichomes have a basic function in the absorption and retention of moisture by the leaves (Benzing 1980; Benzing et al. 1985). They consist of a foot cell, a stalk of few living cells and a shield of dead cells. In Brocchinia reducta, how­ever, the cells of the shield are alive at maturity. Simple hairs without a shield (Fig. 20A,B) are rare (Lindmania, Navia, Pitcairnia, Strehl and Winkler 1982). The most primitive forms of trichomes are probably simple (Lindmania, Navia, Pitcairnia)

Bromeliaceae 77

and stellate hairs (Pitcairnia, Puya) in Pitcairnio­ideae, the latter type with long, slender, slightly fused cells radiating from 1 central cell. Trichomes of most Pitcairnioideae, however, have an undi­vided central cell or a central cell group which differs in size and/or content from the peripheral cells. Pitcairnia, Lindmania, and Fosterella have primitive trichomes from which radiate long filamentous cells only slightly fused to each other (Fig. 20D,E). In Dyckia the central cells are sur­rounded by concentric circles of rounded cells that are only slightly elongated radially. In most Bromelioideae the cells are polygonal so that the outer and inner ones differ little (Fig. 20K). In several species the trichomes are differentiated into a disk and a wing, especially in Neoregelia and some Aechmea. However, in Bromelia and Orthophytum the trichomes are similarily primi­tive as in Pitcairnioideae. The trichomes of the Tillandsioideae, especially the genus Tillandsia, form the highest stage of development (Fig. 20F-I) and are constructed according to a regulary pat­tern. The stalk consists of 1-3 thin-walled cells (but up to 8 in Glomeropitcairnia) and an apical dome cell. The shield consists of 4 right-angled

Fig. 20A-K. Bromeliaceae. Epidermal hairs (A, B) and scales (C-K). A-C Navia fontoides. A Simple and B branched hairs. C Surface view. D, E Fosterella penduliflora. D Transversal section. E Surface view. F, G Tillandsia flexuosa. F Transversal section. G Surface view. H, I T. recurvata. H Transversal section. I Surface view. J, K Nidularium innocentii. J Transver­sal section. K Surface view. (Tomlinson 1969)

cells in the center surrounded by a ring of 8, then a ring of 16, and in many species an outer ring of 64 cells. The trichome may be fused to the epider­mis as far as the 8-celled ring. Although the cell division formula of 4 + 8 + 16 + 64 is the general rule, many exceptions occur, noteworthy being the 4 + 8 + 64 in Catopsis, Glomeropitcairnia, and some Vriesea. The shield varies from perfectly orbicular to extremely excentrical.

INFLORESCENCE STRUCTURE. Inflorescences are paniculate and are made up of racemes or spikes. Reductions of the axes lead to corymbs and heads and even to single flowers. The primary axis usu­ally flowers only once with a flowering period from a few days to over 1 year in large many­flowered species of Aechmea. In Deuterocohnia

78 Bromeliaceae

(except Abromeitiella) the inflorescence becomes woody, has a cambiumlike meristematic cylinder, and flowers continuously for several years, with buds opening at the beginning of each rainy season. Usually, the renewing shoots are formed during the period of flowering, thus providing additional vegetative propagation. A minority of Bromeliaceae is hapaxanthous and survives only by seed.

The sequence of flowering is usually acro­petalous. Exceptions occur in Canistrum, where the middle region flowers first, and in species of Aechmea subgenus Ortgiesia, where acropetalous and basipetalous flowering, as well as such as in Canistrum, can be found among closely related species.

FLOWER STRUCTURE. In general, the sepals are contorted with the right margin covered, and the median sepal is in abaxial position. However, in Brocchinia, Dyckia, and others the sepals are imbricate with both margins of the median sepal covered, and in Cottendorfia florida cochlear, where the median sepal covers both margins of the adaxial ones. Sepals vary from symme­trical to strongly asymmetrical (Racinaea, most Bromelioideae ).

The petals are nearly always regular but in many species of Pitcairnia and Werauhia, one is curved, forming a hood over the stamens. Form varies from linear to ovate; they may be entire, finely crenate or even denticulate and the apex may be round, retuse, apiculate, or mucronate. Color may be white, red, purple, rose, yellow, green, blue, or violet of various shades and intensities. The most interesting feature of the petals is ligular append­ages on the inside base which have been used for taxonomic classification. The structure may be a simple membranaceous scale or flap (Pitcairnio­ideae) or it may consist of 2 separate scales (Brewcaria, Steyerbromelia, Bromelioideae, Til­landsioideae) which are usually serrate or !acini­ate (Brown and Terry 1992). Harms (1930) suggested that the evolution of the 2 more or less separate scales is explained by pressure of the fila­ment on a single simple ligular primordium. Arrais (1989) suggested a staminodial nature of the petal appendages. The function of the append­age is not known with certainty. The early inter­pretation as a mechanism to avoid loss of nectar has been queried.

The stamens are usually free and contain 1 vas­cular bundle, rarely 2, or up to 6 in Encholirium (Arrais 1989). The filaments are linear, fiat, or thickened throughout or apically, or distinctly

succulent in the lower parts as in Racinaea. They may be plicate in several species of Tillandsia. Anthers are dorsifixed or basifixed, usually linear, rarely with an elongated connective (Androlepis).

The ovary is superior to inferior in the Pitcairnioideae, superior in the Tillandsioideae (except of Glomeropitcairnia), and inferior in the Bromelioideae. However, Bohme (1988) con­cluded from the position of the septal nectaries that all gynoecia of the Bromeliaceae are more or less inferior. The carpels are only weakly fused at the edges in Pitcairnioideae but distinctly in the other two subfamilies. Septal nectaries are devel­oped at various degrees and very rarely reduced in Araeococcus (Bohme 1988).

The style is slenderly cylindric and straight or curved according to the direction of the corolla (Pitcairnia), or short and stout or even lacking (Dyckia, Hechtia, Racinaea, Tillandsia).

Stigma morphology (Fig. 21) has proved to be a valuable taxonomic character with far-reaching consistency in Tillandsia subgenus Tillandsia (Gardner 1982, 1986b) and in the thecophylloid species of Vriesea (now Werauhia; Utley 1978, 1983). Brown and Gilmartin (1984, 1989b), as well as Schill et al. (1988), defined the types of stigma structure: free erect untwisted lobes (simpleerect), joined erect untwisted lobes lacking papillae and forming a cup (cupulate), conduplicately folded lobes twisted together spirally (conduplicate­spiral), bladelike lobes folded in an irregular convolute fashion (convolute-blade), and a prolif­eration of irregular growth in the marginal stig­matic line of each lobe (coralliform).

Pollen Morphology. The pollen grains (Fig. 22) are generally single, only in Androlepis and Hohen­bergiopsis are they known to be united in tetrads. The exine is subtectate and usually reticulate, less often scrobiculate or pertectate, with the sexine thicker than the nexine. Grains are either sul­cate or 2-4-porate to pantotreme. Erdtman and Praglowski (1974) found the porate type restricted to subfam. Bromelioideae (12 genera studied), al­though, apart from this, sulcate pollen is present in 11 genera of this subfamily; Aechmea being het­erogeneous in this respect. Subfamily Pitcairnio­ideae and Tillandioideae have invariably sulcate grains, although the data basis is still small (12 genera of Pitcairnioideae and three of Tillandio­ideae studied). With respect to both exine sculp­ture and number and position of apertures, a certain similarity between Bromeliaceae and Amaryllidaceae, especially subfam. Hypoxido-

Bromeliaceae 79

A,B

D,E

ideae, was noted by Erdtman and Praglowski (1974).

Since the time ofErdtman and Praglowski, more palynological data have accumulated (Ehler and Schill 1973), but broader systematic or anthe­cological relationships have not yet been per­ceived (Benzing 1994). However, Halbritter (1988, 1992) successfully demonstrated the value of the aperture morphology for classification at generic and subfamiliallevel. A simple sulcus with smooth margins is typical of Pitcairnioideae and Catopsis while porate pollen is restricted to Bromelioideae. A "distal germination area" seems to occupy an intermediate position and is typical of Pitcair­nia, Tillandsioideae, Billbergia, Cryptanthus, and Orthophytum. Inaperturate pollen is rare (Guzmania, Aechmea, Canistrum).

EMBRYOLOGY. The anther tapetum is secretory, pollen mother cell division is successive, the pollen grains are 2-celled, ovules are bitegmic and crassinucellate, the embryo sac is of the Polygonum type, occasionally proliferation of the antipodals occurs, endosperm is Helobial, and

c

F

Fig. 21A-F. Bromeliaceae. Stigma morphology. A Pitcairnea andreana, conduplicate-spiral. B Tillandsia castellanii, simple-erect. C Vriesea psittacina, convolute-blade. D Werauhia gigantea, cupulate. E Tillandsia cyanea, coralli­form. F Canistrum aurantium, conduplicate-spiral. Bar SOO[!m. (Photo G. Gortan)

embryogeny follows the Asterad type (J ohri et al. 1992). Polyembryony has been found in several species of Tillandsia subgenus Diaphoranthema (Subils 1973).

KARYOLOGY. Chromosome numbers have been tabulated by McWilliams (1974). The chromo­somes are very small; Pitcairnioideae have uni­form complements of equal-sized chromosomes mostly based on x = 25. The same base number predominates in the Bromelioideae, in which varying degrees of bimodality are observed. In Aechmea tillandsoides n = 21 was explained as having arisen through a combination of n = 25 and n = 17 (unique for Cryptanthus) at an early date (Marchant 1967); other numbers reported in this subfamily include n = 24, 36, and 48.

A,B

D,E

80 Bromeliaceae

Fig. 22A-F. Bromeliaceae, pollen grains. Pitcairneoideae: A Puya spathacea, simple sulcus. B Pitcairnia grafii, insulae type. Tillandioideae: C Tillandsia jucunda var. jucunda, operculum type. D Guzmania acorifolia, diffuse sulcus. Bromelioideae: E Lymania corallina, biporate. F Hohenbergia stellata, tri- to tetraporate. Bar lOf!m. (Photo H. Halbritter)

Uniformly bimodal complements characterize the Tillandsioideae, in which a greater variety of base numbers is reported, which include, apart from n = 25, also n = 16, 17, 27, and 32. Polyploidy is quite rare within the family (Bromelia, Fosterella, Guzmania, Nidularium, Pseudananas, Tillandsia, McWilliams 1974; Brown and Gilmartin 1986, 1989a; Till 1992) and the base number x = 25 appears to be paleopolyploid itself. Recent studies (Brown and Gilmartin 1986, 1989a) suggest a hybridization of 2 paleodiploids with n = 7 and n = 8, respectively, and a subsequent secondary hybridization step with the n = 8 lineage yielding x = 25. Decreasing base numbers are explained with aneuploid reductions. Elevated chromosome numbers, however, do not provide reliable evi­dence for polyploidy in the history of a taxon (Greilhuber 1995) as the isozyme numbers in Bromeliaceae are typical of diploids (Soltis et al. 1987).

PoLLINATION. Hummingbirds (Trochilidae) are the major pollinators ofbromeliads (Harms 1930; Utley 1983; Gardner 1986a; Kraemer et al. 1993; Araujo et al. 1994; Martinelli 1994; Seres and

Ramirez 1995) with Trochilinae predominant in montane regions. Honeycreepers (Coerebidae) prevail in highland areas. Species of Puya subge­nus Puya have the outer part of each branch sterile and this forms a distinct perch that is used by icterids (oriole-blackbird family).

Hymenoptera (bees, bumble bees) and Lepi­doptera (moths, butterflies) are listed as visitors by several authors (Harms 1930; Gardner 1986a; Till1992; Martinelli 1994). Hawkmoth pollination may be supposed for Tillandsia xiphioides with long and fragrant flowers.

Bat pollination is reported for Encholirium (Sazima et al. 1989), Alcantarea, Vriesea, and Werauhia (Utley 1983; Martinelli 1994) with glossophagine bats as the pollinator.

The size and structure of the septal nectary (Bohme 1988) and the amount and quality of the nectar offered (Freeman et al. 1985; Bernardello et al. 1991, 1994) are crucial for the pollination type and the visitors. In entomophilous flowers the septal nectaries are smaller and less nectar is offered than in ornithophilous ones. On the other hand, entomophilous flowers often enhance their attraction function by enlargement of the petals (salver flowers) and/or flower fragrance.

REPRODUCTIVE SYSTEMS. The majority of species in the family are certainly outbreeders. Self­fertility has been found in Tillandsia subgenus Tillandsia, and facultative autogamy in Tillandsia subgenus Diaphoranthema (Till 1992). Vriesea

c

F

Bromeliaceae 81

and Werauhia species are capable of inbreeding and numerous Guzmania species are self­pollinated. In the Bromelioideae, Acanthostachys pitcairnioides, Aechmea bromeliifolia, and Ronnbergia nidularioides are examples of self­pollinated species, whereas Ae. fulgens and Ae. nudicaulis appear not to be. Most of the species of Billbergia subgenus Helicodea seem to be inbreeders, whereas subgenus Billbergia, the more primitive group, is primarily outcrossing. Fosterella penduliflora, Pitcairnia andreana, P. xanthocalyx, and Puya mirabilis are a few examples of in breeders in Pitcairnioideae.

Natural hybrids appear to be rather rare under field conditions (Harms 1930) but have been found between several species of Tillandsia subgenus Tillandsia (Gardner 1982). The great abundance of artificial hybrids produced by horticulturists attests to the fact that genetic in­compatibility systems are of minor importance in isolating species under natural conditions. Differ­ent phenologies and tight adaptation to pollina­tors seem to be the most effective barriers against hybridization in the field (Araujo et al. 1994; Martinelli 1994).

FRUIT AND SEED. Detailed modern studies in the seed morphology have been made by Nemirovich­Danchenko (1983) and GroB (1988a). In the Pitcairnioideae the fruit is generally a septicidal capsule with an additionalloculicidal apical dehis­cence in some genera. The pericarp is not differen­tiated into exocarp and endocarp. Seeds are usually small and appendaged (Fig. 24H), rarely naked (Navia, Pepinia, Varadarajan 1986), the ap­pendage being a completely surrounding wing or the seeds are bicaudate. Tillandsioideae have cap­sular fruits with the pericarp splitting into a stramineous exocarp and a dark lustrous endocarp and have plumose seeds (Figs. 25G, 26G). In Glomeropitcairnia the outer integument forms plumose appendages at both ends, which suggests an intermediate position between Pitcairnioideae and Tillandsioideae. In Catopsis single hairs grow from the chalaza to form a folded coma. In the remaining genera of Tillandsioideae, the outer integument splits up into rows of elon­gated cells forming the pseudopappus (Fig. 26G). The seed coat formed by the inner integument usually consists of 2 cell layers and seems to have a characteristic structure for individual genera (Smith and Downs 1974). In the Bromelioideae the fruits are berries with a variously fleshy pericarp, the seeds unappendaged and often with a sarcotesta; Acanthostachys is a remarkable excep-

tion with seeds that have strongly elongated cha­lazal appendages.

Modern information about germination is found in GroB (1988b). In Pitcairnioideae and Bromelioideae, the primary root first appears with the formation of a crown of root hairs, but never in Tillandsioideae, where the formation of a crown of root hairs is rare. In the epigeal Pitcairnioideae the cotyledon moves out of the seed and becomes green while it remains in the seed in the two other subgenera which are both hypogeal.

DISPERSAL. Wind-borne seeds prevail in Pit­cairnioideae and are obligatory for Tilland­sioideae, both with capsular fruits. Seeds of Pitcairnioideae are less adapted to anemochory and are bicaudate or winged at most. The trans­port of the naked seeds of members of this sub­family, e.g., Navia, is unknown; perhaps rain serves as an agent. In Tillandsioideae an elaborate and very effective flight apparatus (pseudo­pappus) allows dispersal over long distances. The berry fruits of Bromelioideae are dispersed mainly by birds and less often by bats (Muller in MOller 1921). Ants appear to disperse seeds of several bromelioid genera. Dispersal in the vegetative state is common in Tillandsia usneoides, as it is used by birds for nest construction.

PHYTOCHEMISTRY. Flavonols, flavones (the latter as highly methoxylated exudates), triterpenes, and steroids have been isolated from leaves. The flavonoid pattern is similar to that of the Commelinaceae, with which the Bromeliaceae agree also in the occurrence of silica bodies (Hegnauer 1963, 1986). This reflects the isolated position of the family within the monocotyledons (Williams 1978). Cyanidin is the prevaling antho­cyanin, proanthocyaninidins seem to be absent. Flower fragrance includes the triterpenes cycloart-23-en-3~, 25-diol, citronellol, geraniol, and nerol (Hegnauer 1963). Cell wall UV fluorescence is due to the combined occurrence of p-coumaric acid, ferulic acid, and diferulic acid (Harris and Hartley 1980). Exudated gums contain galactose, xylose, and arabinose. Ananas comosus contains brome­lain, a mixture of at least 5 proteolytic enzymes; the odor of its fruit is mainly caused by undecatrien and undecatraen (Carle 1992).

SUBDIVISION AND RELATIONSHIPS WITHIN THE FAMILY. Bentham and Hooker (1883) have been the first to propose a modern subdivision with three tribes which have been widely accepted as natural units. Pitcairnioideae were soon further

82 Bromeliaceae

divided. Harms (1930) first recognized subfami­lies, including the fourth and newly erected Navi­oideae, to which he attributed a primitive status. The peculiarities of Navia were confirmed and extended on the genus Cottendorfia by Robinson (1969). The three-partition of Bentham and Hooker ( 1883) was accepted at subfamiliallevel by most researchers up to the most recent mono­graphs (Smith and Downs 1974, 1977, 1979).

Generally, the terrestrial Pitcairnioideae are considered as the least specialized group with epigeal germination, most primitive trichomes, predominantly superior ovaries, the capsular fruits, and the weakly elaborated seeds. Tillandsioideae with hypogeal germination, most highly elaborated trichomes, prevailing superior ovaries, capsules, plumose seeds, and the atmo­spheric growth habit, as well as Bromelioideae with hypogeal germination, moderately elabo­rated trichomes, inferior ovaries, and baccate fruits, are interpreted as derived lineages. Both Pittendrigh (1948) and Tomlinson (1969) have emphasized that the three subfamilies are best re­garded as separate derivatives of a common ances­tral group, so that distinct evolutionary trends must be sought within each subfamily. A mole­cular analysis has indicated the monophyly of Tillandsioideae and Bromelioideae and the sister group relationship of Bromelioideae and Pitcairnioideae, the latter supposedly being paraphyletic (Ranker et al. 1990). A cladistic study of Varadarajan and Gilmartin (1988a) led to the recognition of three monophyletic tribes within Pitcairnioideae: Brocchinieae, Pitcairnieae, and Puyeae (Varadarajan and Gilmartin 1988b). The relationships among the genera of Bromelioideae are insufficiently understood and the arrangement presented under Genera of the Bromeliaceae (below) follows the supposed progressions from sulcate to porate pollen, from a lacking to a large epigynous tube, and from appendaged to naked petals, but remains somewhat arbitrary.

AFFINITIES. The family Bromeliaceae is widely recognized as monophyletic (Gilmartin and Brown 1987). Synapomorphies of the family include stellate and scalelike multicellular hairs; solitary silica bodies; a conspicuously sepaloid and petaloid perianth; conduplicate-spiral stig­mas; and possibly the chromosome base number of x = 25. Molecular analyses support a concept of a close relationship between the Bromeliifiorae, Commelinifiorae, and Zingiberifiorae, which form a monophylum to which the palms are basal (Clark et al. 1993). At the family level, rbcL data indicate

that Bromeliaceae are closely related to Rapatea­ceae and Mayacaceae (Chase et al. 1993; Clark et al. 1993, Givnish et al. 1995), or to Commelinaceae (Duvall et al. 1993). A relationship to Velloziaceae as proposed by Dahlgren ( 1983) and Dahlgren et al. (1985) cannot be confirmed. Takhtajan (1980) is alone in his suspicion of a relationship with the agavaceous stock ofLiliales.

DISTRIBUTION, HABITATS, AND THE EVOLUTION OF EPIPHYTISM. Bromeliaceae are native to the tropical and subtropical regions of the neotropics extending to Virginia in N America and to north­ern Patagonia in S America. They range from sea level to over 4500 m in the central Peruvian Andes. Only Pitcairnia feliciana is endemic to western tropical Africa; its occurrence there is interpreted by long-distance dispersal. This pattern of distri­bution is equalled by Rapateaceae, with Marsch­alocephalus as the only African member. A few Bromeliaceae live far off the South American sub­continent on islands: Racinaea insularis on the Galapagos Islands, Greigia berteroi and Ochagavia elegans on the Juan Fernandez Islands.

The Brocchinieae are restricted to the Guayana Highlands, Pitcairnieae have their center in north­ern S America extending into Mesoamerica, the Antilles, and southeastern Brazil, and Puyeae have one center along the Andean cordillera and another in eastern Brazil; only Brewcaria and a few Puya species are represented in the Guayana Highlands. Hechtia is remarkable for its restric­tion to Mexico and Guatemala. Tillandsioideae have their center in northwestern S America ex­tending into Mesoamerica, the Antilles, south­western S America, and southeastern Brazil, to which latter area Alcantarea is restricted. Bromelioideae are centered in eastern Brazil. Some genera exhibit long-distance disjunctions like Greigia, Billbergia, some subgenera of Aechmea, Hohenbergia, and Neoregelia, or are wide-ranging like some subgenera of Aechmea and Bromelia. For detailed distribution maps at generic and subgeneric levels see Smith and Downs (1974, 1977, 1979).

Bromeliads are found in a great variety of habi­tats. Mesophytes prevail in environments with high humidity like lowland rainforests, montane forests, and cloud forests, while CAM xerophytes are able to settle in dry areas like savannahs, campos rupestres, semideserts, fog deserts and even on bare rock. For more details see Benzing (1980, 1990) and Rauh (1990).

The evolution of epiphytism has been explained controversely. Schimper (1888) concluded that

Bromeliaceae 83

evolution of the epiphytic life form had occurred within the rainforest itself as proceeding from the forest floor to the lower limbs, and then ultimately to the relatively dry upper canopy of the forest as certain bromeliads became more and more inde­pendent of roots through increasing specialization of the adsorbing trichomes. Pittendrigh (1948) believed in a moderate xeric ancestor which gave rise to both fully xeric and mesic derivatives. Tomlinson (1969), however, pointed out that the anatomical evidence may be in conflict with Pittendrigh's evolutionary interpretation. Medina (1974) proposed an intermediary position of a heliophilic C3 ancestor between completely meso­phytic and xerophytic derivatives. However, the disucssion has not yet come to an end (Benzing et al. 1985).

AssociATED BIOTA. An extensive literature has developed on the animals which live in bromeliad tanks and leaf bases (Kriigel1993). The spectrum of phytotelm inhabitants comprises animals and prokaryota, algae, myxomycota, and higher plants (Laessle 1961), and ranges from occasionals to organisms restricted to these microhabitats like the crustacea Elpidium bromeliarum or the frog Syncope antenori. Many malaria mosquito species use the bromeliad ~anks during their life cycles.

FossiLS. Six different specimens of fossils have been assigned tentatively to the Bromeliaceae. According to the evaluation by Smith and Downs (1974), for none of them is the association with the family proven, or at least likely. The probably only true bromeliad fossil is Karatophyllum bromelioides from Costa Rica (Gomez P. 1972).

EcoNOMIC IMPORTANCE. Ananas comosus, the pineapple, is the species of primary economic im­portance and is cultivated for its fruits worldwide in tropical regions. It also provides the proteolytic enzyme bromelain which is pharmaceutically used (Carle 1992). Tillandsia usneoides, Spanish moss, was formerly important, especially in Loui­siana, as a squrce of pillow and cushion stuffing, but has been supplanted by plastic. Neoglaziovia variegata, the car6a or carua, and Ananas lucid us, the curagua, are a source of heavy fiber in north­ern S America. Some Bromelia species have edible fruits (Arenas and Arroyo 1988; Correa and Bernal 1990). Many bromeliads are cultivated as ornamentals.

CoNSERVATION. Seven species of Tillandsia have been protected by appendix II of CITES. Many

more may be endangered by commercial collect­ing on a large scale, especially in Guatemala and Brazil. A main source of endangering, however, is the destruction of the natural habitats by unscru­pulous land use.

CONSPECTUS OF CLASSIFICATION OF BROMELIACEAE

I. Subfamily Pitcairnioideae Harms (1930). 1. Tribe Brocchinieae G.S. Varad. & Gilmartin (1988).

Genus 1. 2. Tribe Pitcairnieae K. Koch (1874).

Genera 2-10. 3. Tribe Puyeae Wittm. (1888).

Genera 11-16. II. Subfamily Tillandsioideae Harms (1930).

Genera 17-25. Ill. Subfamily Bromelioideae Harms (1930).

Genera 26-56.

KEY TO THE GENERA

1. Mature seed appendaged or if not (Pitcairnia in part and Navia and Brewcaria), then the fruit capsular and dehis­cent; ovary superior or largely so in most genera to infe­rior; fruit capsular or if not (Pitcairnia in part), then the seed appendaged; germination epigeal or hypogeal 2

- Mature seed unappendaged; fruit baccate; ovary inferior; leaves mostly spinose-serrate; indument almost always of obvious scales; germination hypogeal; plants often terres­trial (subfam. Bromelioideae) 27

2. Seed appendages entire or slightly divided (Brocchinia paniculata) or lacking (Pepinia aphelandriflora) and Navia); fruit usually dehiscent; leaves mostly spinose­serrate; indument of finely to scarcely divided scales; germination epigeal; plants usually terrestrial (subfam. Pitcairnioideae) 3

- Seed appendages finely divided and forming a coma, always present; fruit dehiscent; leaves always entire; indument almost always of obviously radially symmetric scales; germination hypogeal; plants mostly epiphytic (subfam. Tillandsioideae) 19

3. Plants monoecious or, if rarely dioecious ( Cottendorfia) or polygamodioecious (Dyckia maritima, D. selloa, and D. hebdingii), then the petals yellow or orange and plants native to northeastern and southernmost Brazil 4

- Plants dioecious with functionally unisexual flowers; petals rose or white; plants of Texas, Mexico, and northern Central America 16. Hechtia

4. Bases of the filaments separate from each other, but some-times individually adnate to the petals and sepals 5

- Bases of the filaments forming a tube and adnate to the petals; petals yellow to orange; plants of Brazil, Uruguay, Paraguay, and Argentina 15. Dyckia

5. Seeds obviously and persistently appendaged 6 - Seeds naked at maturity 17 6. Petal blades tightly spiraled after anthesis, broad, distinct

from the claw; leaf blades narrowly triangular, never con­tracted at base; ovary superior or slightly inferior; Andean plants of open slopes and summits from Costa Rica and Guayana to Chile and Argentina 13. Puya

- Petal blades remaining separate after anthesis or, if sometimes slightly spiraled (Deuterocohnia), then not clawed 7

84 Bromeliaceae

7. Ovary wholly superior; petals regular 8 - Ovary partially to wholly inferior, or, if superior, then the

petals zygomorphic (Pepinia and Pitcairnia spp.) 14 8. Petals naked or with paired appendages (Steyerbromelia)

9 - Petals each bearing a single basal scale; xerophytic plants

of the southern Andes from Peru to Chile, Argentina, and W Brazil 12. Deuterocohnia

9. Seeds with a falciform appendage; petal blades narrow, indistinct; plants of NE Brazil 14. Encholirium

- Seeds bicaudate-appendaged 10 10. Anthers basifixed, linear, coiled at anthesis, inner fila­

ments adnate to the base of the petals; leaf blades thin, more or less contracted at base; mesophytic plants of Mexico to Argentina and W Brazil 8. Fosterella

- Anthers subbasifixed to equitant, stout, straight; filaments usually free; leaf blades firm, not contracted at base; plants of the Guayana Highlands 11

11. Flowers perfect; sepals convolute, each with its left side covering the right side of the next one 12

- Flowers unisexual, dioecious; sepals cochlear with the abaxial overlapping both the adaxial, plants of NE Brazil

6. Cottendorfia 12. Petals naked; stigmas straight, erect; flowers mostly

pedicellate 13 - Petals with paired appendages; stigmas broad, strongly

contorted; flowers sessile; inflorescence compound, lax 4. Steyerbromelia

13. Anthers subbasifixed; petals brightly colored, more or less massed together after anthesis but not twisted; sepals large and firm 7. Connellia

- Anthers equitant; petals white or rose, separate after anthesis; sepals not over lOmm long, thin, flat

5. Lindmania 14. Petals large, naked, or appendaged, usually zygomorphic

and forming a hood over the anthers; sepals convolute with the left side of each overlapping the right of the next one; plants of Mexico and the West Indies to Argentina and Brazil 15

- Petals minute, regular; sepals cochlear with both adaxial ones overlapping the abaxial; plants of the Guayana Highlands 16

15. Seeds 2-tailed; foliage usually dimorphic to polymorphic; inflorescence a raceme, panicle, spike, or frequently strobiliform; flowers and floral bracts occasionally secund and reflexed; leaf blades often broadly elliptic, ovate, lan­ceolate, or oblanceolate 10. Pitcairnia

- Seeds winged; foliage monomorphic to rarely dimorphic, never polymorphic; inflorescence a raceme or panicle, never strobiliform; flowers and floral bracts always erect to spreading; leaf blades narrowly elliptic to lanceolate

9. Pepinia 16. Epigynous tube lacking; inflorescence open and definitely

branched 1. Brocchinia - Epigynous tube well developed; inflorescence sessile,

capitate 3. Ayensua 17. Sepals convolute; petals zygomorphic (Pepinia

aphelandriflora) 15 - Sepals cochlear with both adaxial ones overlapping

the abaxial; petals regular; ovary superior (in most species) to nearly inferior; plants of the Guayana Highlands 18

18. Petals naked; inflorescence scapose, pinnate, and more or less open or sessile and capitate 2. Navia

- Petals with 2 appendages; inflorescence long-scapose, simple, densely cylindric 11. Brewcaria

19. Ovary nearly or quite superior; seeds plumose on base or apex or largely on the base and only slightly on the apex

20 - Ovary only half superior; seeds equally plumose­

appendaged at both ends; flowers polystichous. Lesser Antilles, Trinidad, adjacent Venezuela

18. Glomeropitcairnia 20. Appendage of the seed wholly or largely basal, straight at

maturity 21 - Appendage of the seed largely apical ( = chalaza!), folded

at maturity; sepals strongly asymmetric in most species; flowers in at least slightly more than 2 ranks; leaves often cretaceous-coated on the inside. Florida, Mexico, and the West Indies to Brazil and Peru 17. Catopsis

21. Petal claws free or with very short tube exceeded by the sepals; flowers distichous in most species 22

- Petal claws conglutinated in a tube, equaling the sepals or, rarely, the petals entirely included in the sepals 26

22. Petals naked; inflorescence of 1 or more distichous­flowered spikes or racemes or rarely reduced to 1 or more polystichous-flowered spikes or to a single flower; plants of southern United States to Argentina and Chile 25

- Petals bearing scales ("lateral folds" of Mez) on the inside of the claw; inflorescence as in Tillandsia. Mexico and the West Indies to Argentina and Uruguay 23

23. Seed with the apical appendage divided into a short coma; petals linear long, fusiform, usually 10-15 times longer than wide, soon flaccid and drooping 21. Alcantarea

- Seed with the apical appendage minute and undivided; petals elliptical, usually 5-10 times longer than wide, usu­ally firm and remaining more or less erect after an thesis

24 24. Flowers with brilliant coloration in most species, bright

yellow, orange, or red, rarely dull to white, light yellow, or light orange; the adaxial petal pair arranged apically in respect to the abaxial; petal appendages tongue-shaped; stigma with the convolute blade type morphology, that is, 3 obviously spreading lobes covered with papillae

22. Vriesea - Flowers generally dull in color, white, greenish white, light

green yellowish green, yellow, or light orange; the adaxial petal pair arranged basally in respect to the abaxial; petal appendages dactyloid with 1-5 fingers of varying length; stigma with the cupulate type morphology, that is, 3 apical, capitate, cup-shaped lobes, without papillae

23. Werauhia 25. Sepals symmetric or if slightly asymmetric, then ovate

or lanceolate and broadest below the middle, free or variously connate; seeds usually with a distinct apical appendage 24. Tillandsia

- Sepals asymmetric, free or nearly so, broadest near apex, not over 12mm long; seeds without apical appendage

25. Racinaea 26. Petal claws always naked; spikes always polystichous­

flowered. Florida, Mexico, and the West Indies to Brazil and Bolivia 20. Guzmania

- Petal claws bearing scales on the inside; flowers polystic­hous rarely secund or distichous. Colombia to Peru

19. Mezobromelia 27. Sepals symmetric or nearly so 28 - Sepals asymmetric 51

28. Filaments forming a tube to which the fleshy petals are joined along their centers but with their margins free; sepals mostly free or nearly so; leaves very laxly and coarsely spinose-serrate 29

- Filaments not connate but sometimes adnate 31

Bromeliaceae 85

29. Sepals with soft, usually broad apices; inflorescences compound. Mexico and the West Indies to Argentina and Uruguay 36. Bromelia

- Sepals spinose-mucronate 30 30. Inflorescence simple, subsessile. Argentina

37. Deinacanthon - Inflorescence branched with terminal strobils, scapose. S

Mexico, Guatemala 49. Hohenbergiopsis 31. Terminal axes of the inflorescence visible 32 - Terminal axes of the inflorescence covered by leaves or

bracts 38 32. Petals naked; sepals 0.5-7mm long 33 - Petals appendaged; sepals mostly much larger 34

33. Inflorescence compound; sepals broadly ovate or oblong, 0.5-2mm long. Costa Rica and Trinidad to Amazonian Brazil 28. Araeococcus

- Inflorescence simple; sepals narrowly elliptic, 7mm long; flowers subsessile or pedicellate. Mount Itatiaia area in E Brazil 31. Fernseea

34. Petals zygomorphic or tightly recoiled and flowers sessile. W Mexico and Central America to Argentina and Uruguay

35 - Petals regular (actinomorphic) 36

35. Epigynous tube usually well developed 34. Billbergia - Epigynous tube shallow. W Mexico 33. Ursulaea

36. Petals erect. E Brazil 3 7 - Petals apically recoiled 33. Ursulaea

37. Flowers sessile 44. Quesnelia - Flowers pedicellate 35. Neoglaziovia

38. Inflorescence simple, strobilate; flowers solitary in the axil of each bract 39

- Inflorescence compound 46 39. Scape short or none; strobils nidular or axillary 40 - Scape well developed, obvious 44

40. Floral bracts foliaceous. NE Brazil 26. Orthophytum - Floral bracts bracteiform 41

41. Scape distinct, its bracts shorter than the floral bracts; petals naked. Mexico and Venezuela to Chile

30. Greigia - Scape none or very short 42

42. Epigynous tube shallow, crateriform (A. pitcairnioides). Brazil: Bahia 41. Acanthostachys

- Epigynous tube cylindric, deep. Chile 43 43. Sepals obtuse, carinate 29. Fascicularia - Sepals acute to attenuate, pungent to mucronate, not

carinate 32. Ochagavia 44. Scape erect, without bracts (A. strobilacea). S Brazil,

Paraguay, Argentina 41. Acanthostachys - Scape covered with bracts 45

45. Scape bracts foliaceous, scape erect. NE Brazil 26. Orthophytum

- Scape bracts bracteiform; scape prostrate. French Guiana and adjacent Brazil 56. Disteganthus

46. Inflorescence obviously compound with several strobils on an elongate floral axis 47

- Inflorescence pseudosimple with hands or fiat fascicles in the axils of large bracts 48

47. Floral bracts foliaceous, serrulate; strobils sessile or subsessile. NE Brazil 26. Orthophytum

- Floral bracts bracteiform, entire; strobils on distinct scapes. Mexico and Venezuela to Chile

30. Greigia 48. Outer bracts of the inflorescence foliaceous; sepals high

connate; petals naked. NE Brazil 27. Cryptanthus - Outer bracts of the inflorescence bracteiform, large, and

covering most of the flowers. E Brazil 49

49. Petals connate or agglutinated in a tube the height of the sepals 52. Nidularium

- Petals free or nearly so 50 50. Petals completely free; inflorescence often scapose

45. Canistrum - Petals short-connate at or a little above the base

46. Wittrockia 51. Ovaries coalescing to form a compound fruit; inflores-

cence simple, strobilate 52 - Ovaries always remaining distinct 53

52. Inflorescence with a small, inconspicuous coma, never producing basal shoots; plant propagating by elongate rhi­zomes; petals bearing vertical folds. Paraguay and adjacent areas 39. Pseudananas

- Inflorescence usually with a large conspicuous coma (lack­ing in A. monstrosus), often with basal shoots; rhizomes lacking; petals usually bearing well-developed scales. Probably native from Paraguay to the Amazon Basin, now pantropical 40. Ananas

53. Flowers pedicellate 54 - Flowers sessile or subsessile 59

54. Inflorescence nidular, simple in most species; petals naked. Amazonia, E Brazil 53. Neoregelia

- Inflorescence scapose 55 55. Sepals more or less connate, long-mucronate; petals

appendaged. E Brazil 47. Portea - Sepals free or unarmed 56

56. Inflorescence simple; sepals unarmed 57 - Inflorescence compound 58

57. Petals naked. Colombia 48. Pseudaechmea - Petals appendaged. Colombia and Guyana to NE Brazil

43. Aechmea subg. 2. Lamprococcus 58. Sepals 1.5-3mm long; inflorescence glabrous; petals

naked. Colombia to Suriname and Amazonian Brazil 28. Araeococcus

- Sepals 3.5-22mm long; inflorescence !epidote; petals appendaged. Mexico to Peru

43. Aechmea subg. 1. Podaechmea 59. Petals appendaged with well-developed scales 60 - Petals naked or with lateral folds or rudimentary or

reduced scales 66 60. Epigynous tube shallow or lacking; flowers in terete

strobils; inflorescence mostly pinnate and lax, rarely digi­tate or simple (H. littoralis). Antilles to Venezuela and Brazil 42. Hohenbergia Epigynous tube well developed; inflorescence various 61

61. Sepals unarmed 62 - Sepals armed. N and S America 43. Aechmea subg.

3. Aechmea, 43. Aechmea subg. 4. Ortgiesia, and 43. Aechmea subg. 6. Pothuava

62. Floral bracts attached basally, not decurrent nor forming pouches; flowers polystichous 63

- Floral bracts decurrent and forming pouches around the flowers; flowers often distichous. N and S America

43. Aechmea subg. 5. Platyaechmea 63. Inflorescence branched 64 - Inflorescence simple 65

64. Leaves 2-ranked; blades marked with spots or bands; floral bracts minute; ovules obtuse (Qu. marmorata). Brazil: Espirito Santo to Sao Paulo 44. Quesnelia

- Leaves more than 2-ranked or the blades concolorous; floral bracts large to lacking; ovules long-caudate. Colom­bia, Venezuela, Amazonian Brazil 43. Aechmea subg.

2. Lamprococcus 65. Ovules obtuse (no further distinction possible without

keying by species). E Brazil 44. Quesnelia

86 Bromeliaceae

- Ovules apiculate to caudate. Central America to Brazil and Argentina 43. Aechmea subg. 7. Macrochordion

66. Ovary deeply sulcate; inflorescence simple or compound. NE Brazil 51. Lymania

- Ovary evenly rounded 67 67. Inflorescence lax; axes visible 68 - Inflorescence dense 71

68. Inflorescence simple. Costa Rica to Peru 55. Ronnbergia - Inflorescence pinnately compound 69

69. Flowers minute; sepals not over 3mm long; ovules few; epigynous tube none. Costa Rica, Venezuela, Trinidad, Tobago, Guyana to Amazonian Brazil 28. Araeococcus

- Flowers small to large; sepals more than 3 mm long; epigy-nous tube distinct 70

70. Branches elongate, many-flowered; flowers perfect; anthers unappendaged. E and Amazonian Brazil and adjacent areas 54. Streptocalyx

- Branches short, digitately few-flowered; flowers function­ally unisexual on different plants; anthers appendaged. Central America: Guatemala to Costa Rica 50. Androlepis

71. Flowers 2 or more in the axil of each bract 72 - Flower single in the axil of each bract 73

72. Inflorescence involucrate; sepals only slightly asymmetric, unarmed or mucronulate. E Brazil 52. Nidularium

- Inflorescence strobilate; sepals strongly asymmetric, mucronate. E and Amazonian Brazil and adjacent areas

54. Streptocalyx 73. Petals naked or with lateral folds; bracts chartaceous or

coriaceous; leaf blades often petiolate. Costa Rica to Peru 55. Ronnbergia

- Petals bearing rudimentary or reduced appendages; bracts mostly thick and ligneous; leaf blades never petiolate; pollen sulcate. Mexico to Peru and Amazonian Brazil, E Brazil 38. Chevaliera

GENERA OF THE BROMELIACEAE

I. Subfam. Pitcairnioideae Harms (1930).

subfam. Navioideae Harms (1930).

Nearly always terrestrial or saxicolous. Roots present and largely functional. Leaves nearly always spinose-serrate. Ovary superior or largely so to rarely inferior (in Ayensua, many Brocchinia spp., and a few species of Navia and Pitcairnia). Pollen sulcate, sulcus simple except of Pepinia and Pitcairnia with a distal germination area. Stigma of the conduplicate­spiral type morphology except in Brewcaria, Cottendorfia, Hechtia, Brocchinia (in part), and Fosterella (in part). Fruit capsular or rarely indehiscent.

1. Tribe Brocchinieae G.S. Varad. & Gilmartin (1988).

Seeds elongate, apical and basal ends dissimilar, basal appendage variously modified, surface cellu­lar pattern of seed appendage and seed body alike; foliar trichomes symmetric with stalk of six or

more cells; leaf photoassimilatory tissue with abundant stellate chlorenchyma; leaves almost always entire (except Brocchinia serrata L.B. Sm.); inflorescence a raceme or profusely branched panicle.

1. Brocchinia Schult. & Schult. f.

Brocchinia Schult. & Schult. f., in Roem. & Schult., Syst. veg. 7(2): lxx, 1250 (1830); L.B. Sm., Ann. Mo. Bot. Gard. 73:689-721 (1986).

Terrestrial or rarely epiphytic, stemless to mas­sively caulescent; leaves rosulate; blades usually entire, glabrous or subglabrous; scape conspic­uous; inflorescence compound, mostly ample; flowers perfect, minute, pedicellate; sepals free, imbricate with the 2 adaxial covering the abaxial; petals free, naked; stamens included at anthesis; filaments more or less adnate to the sepals and petals; stigma conduplicate-spiral or simple erect; ovary from 1/3 to wholly inferior; placentae linear; ovules few, long-appendaged; capsule wholly septicidal or rarely loculicidal as well (B. ser­rata L.B. Sm.). Twenty one spp., Guayana Highlands.

2. Tribe Pitcairnieae K. Koch (1874).

Seeds usually spherical, lenticular, cuneate, po­lygonal, reniform, or ellipsoidal; if elongate, then apical and basal ends similar, surface cellular pat­tern of seed appendage and body often different; foliar trichomes symmetric or asymmetric with a stalk of 2 or 3 cells; leaf photoassimilatory tissue without stellate chlorenchyma but with abundant palisade and spongy elements or undifferenciated chlorenchyma; leaves entire to mostly spinulose or serrulate, typically mem­branous, occasionally coriaceous; foliar trichome arrangement irregular and nonoverlapping; epi­dermis smooth (except for some Pepinia); water­storage tissue peripheral only; hypodermal sclerenchyma absent (except for Ayensua and some Pepinia); inflorescence a globose head, spike, raceme, or panicle.

2. Navia Mart. ex Schult. & Schult. f. Fig. 23

Navia Mart. ex Schult. & Schult. f., in Roem. & Schult., Syst. veg. 7(2): lxv, 1195 (1830); L.B. Smith, Ann. Mo. Bot. Gard. 73: 689-721 (1986); emend. Steyerm., J. Bromeliad Soc. 39: 54 (1989).

Stemless or caulescent; leaves rosulate or in a dense spiral along the stem; blades entire or more

Bromeliaceae 87

Fig. 23A-E. Bromeliaceae-Pitcairnioideae. Navia acaulis. A Habit. B Flower with subtending bract. C Pistil. D Dehiscing capsule. E Seed. (Takhtajan 1982)

or less serrulate; scape usually lacking, rarely well developed; inflorescence usually capitate but sometimes paniculate; floral bracts conspicuous; flowers sessile or subsessile, perfect; sepals imbri­cate with both adaxial ones overlapping the abaxial, free to high-connate; petals free to high­connate, naked; stamens of the second series highly adnate to the petals; anthers linear; ovary usually superior but rarely to wholly inferior; style slender; placentae linear; ovules naked or nearly so; capsule usually dehiscent; seeds reticulate, naked or soon losing the vestigial wing. Ninety eight heliophytic spp., Guayana Highlands from Colombia to Suriname.

3. Ayensua L.B. Sm.

Ayensua L.B. Sm., Mem. N. Y. Bot. Gard. 18 (2): 29 (1969); emend. L.B. Sm. & Read, Phytologia 38: 137 (1977).

Caulescent, terrestrial and saxicolous in dense colonies; leaves densely imbricate along the stem; sheaths distinct; blades attenuate, not contracted at base, deciduous; scape lacking; inflorescence capitate; flowers perfect, sessile or subsessile;

sepals free, imbricate with the abaxial wholly in­cluded by the 2 adaxial; petals free, naked; sta­mens included, the 2 series unequal; ovary wholly inferior, crowned with a large epigynous tube; ovules caudate; fruit indehiscent on the outside, dividing into 3 equal valves inside. Only one sp., A. uaipanensis (Maguire) L.B. Sm., Auyan-tepui in Guayana Highlands of southern Venezuela.

4. Steyerbromelia L.B. Sm.

Steyerbromelia L.B. Sm., Acta Bot. Venez. 14: 8 (1984), nom. illeg.; Ann. Mo. Bot. Gard. 73: 689-721 (1986).

Large heliophytic herbs; leaves rosulate; sheaths short; blades narrowly triangular, not constricted at base, more or less spinose-serrate; scape devel­oped, stout; scape bracts subfoliaceous; inflo­rescence amply pinnate-paniculate, lax; flowers perfect, sessile; sepals imbricate, free; petals appendaged with vertically attached scales, free; stamens equal; ovary superior; stigmas broad, strongly contorted; ovules caudate. Three spp., open summits, Guayana Highlands.

5. Lindmania Mez

Lindmania Mez, in A. & C. DC., Monogr. phan. 9: 535 (1896); L.B. Sm., Ann. Mo. Bot. Gard. 73: 689-721 (1986).

Cottendorfia sensu LB. Sm. (1960), (1974); in part but not as to type.

Small to large heliophytic herbs; leaves rosulate; blades mostly entire but sometimes more or less serrulate; scape usually but not always developed; inflorescence compound or rarely simple; flowers perfect, small, white, or greenish; sepals con­volute, free; petals free, naked; filaments mostly free; anthers versatile; stigma simple-erect; ovary wholly superior, glabrous; style slender; placentae short, basal; capsule short, ovoid, septicidal; seeds slenderly fusiform, bicaudate. Thirty eight spp., open habitats, Guayana Highlands.

6. Cottendorfia Schult. & Schult. f.

Cottendorfia Schult. & Schult. f., in Roem. & Schult., Syst. veg. 7(2): lxiv, 1193 (1830).

Heliophytic herb; leaves rosulate; blades entire, pungent; inflorescence tripinnately paniculate, lax; branches densely polystichous-flowered; flowers unisexual, dioecious, sessile, small; sepals cochlear, the abaxial overlapping both the adaxial, free; petals free, naked; stamens included; fila­ments free; anthers ovate, mediofixed; ovary

88 Bromeliaceae

superior; style cylindric, short; stigmas 3, linear, erect; placentae subscutate, basal; ovules few, apically short-caudate. Only one sp., C. florida Schult. & Schult. f., restricted to xeric habitats in NE Brazil.

7. Connellia N.E. Br.

Connellia N.E. Br., Trans. Linn. Soc. Lond. Bot. 6: 66 (1901); L.B. Sm., Ann. Mo. Bot. Gard. 73: 689-721 (1986).

Stemless or short-caulescent; leaves rosulate, co­riaceous; blades spinose-serrate to entire; scape developed; inflorescence simple or compound; flowers perfect; sepals convolute, free; petals free, naked; stamens included; anthers basifixed; ovary wholly superior; style elongate; capsule septicidal; seeds long-caudate. Five spp., open slopes and summits, Guayana Highlands.

8. Fosterella L.B. Sm.

Forsterella L.B. Sm., Phytologia 7: 171, pl. 1 (1960); L.B. Sm. & Read, Bradea 6: 134-140 (1992).

Lindmania sensu Mez (1896), (1935); sensu L.B. Sm. (1960); in part but not as to type.

Mesophytic terrestrial herbs, stemless or nearly so; leaves rosulate; blades entire or weakly serrulate; scape always well developed; inflores­cence usually laxly paniculate; flowers small to minute, perfect; sepals convolute, free; petals free, unappendaged; stamens barely included to ex­serted; filaments free or the inner adnate to the bases of the petals; anthers basifixed, linear; pollen dry, wind-borne; stigma conduplicate-spiral or simple-erect; ovary superior; placentae linear the length of the cell; capsule septicidal, globose to subpyramidal; seeds narrow, bicaudate. (2n = 50, 100). Sixteen spp., mesophytic habitats, Mexico to Argentina and W Brazil.

9. Pepinia Brongn. ex Andre

Pepinia Brongn. ex Andre, Ill. Hort. 17: 32, pl. (1870); G.S. Varad. & Gilmartin, Syst. Bot. 13: 294- 299 (1988).

Foliage monomorphic to dimorphic; leaves nar­rowly elliptic to lanceolate; inflorescence a raceme or panicle; flowers and floral bracts erect to spreading; seeds winged to rarely naked. (2n = 50). Sixty eight species, terrestrial and saxicolous in open savannas and scrub forests. Guayana Highlands and adjacent lowlands, N Andes, Central America.

10. Pitcairnia L'Her. Fig. 24A-H

Pitcairnia L'Her., Sert. angl.: 7 (1788); nom. cons.; G.S. Varad. & Gilmartin, Syst. Bot. 13: 294-299 (1988).

Mostly terrestrial herbs but sometimes saxicolous or epiphytic, mostly stemless to long-caulescent; leaves polystichous or rarely distichous; blades

Fig. 24A-M. Bromeliaceae-Pitcairnioideae. A-H Puya mirabilis. A Habit. B Leaf. C Stamen. D Pistil. E Flower. F Style with stigma. G Capsule. H Seed. 1-M Puya berteroiana. I Habit. J Flower. K Stamen. L Pistil. M Ovary, cross-section. (Takhtajan 1982)

Bromeliaceae 89

from linear to broadly lanceolate and petiolate, usually strongly dimorphic, entire or serrate, sometimes deciduous; scape usually developed; inflorescence simple or compound; flowers perfect; sepals convolute, free; petals free, usually zygomorphic, naked or with a single scale or its remnant at base; stamens elongate, from slightly shorter to slightly longer than the petals; anthers linear; ovary from almost wholly sup­erior to wholly inferior; ovules numerous, appendaged or rarely obtuse; fruit a capsule or rarely indehiscent on the outside; seeds bica­udate or broad and winged. (2n = 50). Two hun­dred and eighty five spp., highly varied habitats. Mexico and the West Indies to Argentina and Brazil.

3. Tribe Puyeae Wittm. (1888)

Seeds usually spherical, lenticular, cuneate, polygonal, reniform, or ellipsoidal, if elongate, then apical and basal ends similar, surface cell­ular pattern of seed appendage and body often different; foliar trichomes symmetric or asym­metric with a stalk of 2 or 3 cells; leaf photo­assimilatory tissue without stellate chlorenchyma but with abundant palisade and spongy ele­ments or undifferentiated chlorenchyma; leaves strongly spinose-serrate, succulent or semisuc­culent; foliar trichomes in epidermal grooves arranged in overlapping tiers; epidermis with ridges and furrows; water-storage tissue periph­eral and internal; hypodermal sclerenchyma present.

11. Brewcaria L.B. Sm., Steyerm. & H. Rob.

Brewcaria L.B. Sm., Steyerm. & H. Rob., Acta Bot. Venez. 14: 10 (1984); L.B. Sm., Ann. Mo. Bot. Gard. 73: 689-721 (1986).

Large heliophytic herbs; leaves rosulate; blades narrowly triangular, spinose-serrate; scape erect, stout; inflorescence spicate, dense; flowers perfect; sepals imbricate; petals bearing 2 horizontally at­tached scales, free; stamens equal, included; stigma simple-erect; ovary slightly inferior; ovules exalate or very narrowly alate. Two spp., Guayana Highlands.

12. Deuterocohnia Mez

Deuterocohnia Mez, in Mart., Fl. bras. 3(3): 506 (1894); M.A. Spencer & L.B. Sm., Bradea 6: 141-146 (1992).

Abromeitiella Mez (1927).

Plants growing in ring formations or forming cushions or polsters; leaves densely rosulate, spinose-serrate; scape developed and the inflores­cence perennial, or scape lacking and the inflores­cence sessile; flowers perfect; sepals convolute, free; petals free, each bearing a single basal ap­pendage; stamens free; ovary superior, ovules ob­tuse to very short caudate; seeds fusiform, bicaudate. (2n = 50). Fourteen spp., extreme xero­phytic habitats, southern Andes, Peru, Bolivia, Argentina, and Chile.

13. Puya Molina Fig. 241-N

Puya Molina, Sag. stor. nat. Chili: 160, 351 (1782).

Stemless or more generally long-caulescent; leaves coriaceous; blades never constricted at base, coarsely spinose-serrate; inflorescence simple or paniculate; flowers perfect; sepals convolute, free; petals free, unappendaged or with a pair of vertical folds, twisted together in a spiral after anthesis; stamens generally somewhat shorter than the pet­als; filaments free; ovary superior or slightly infe­rior; style long and slender; capsule loculicidal and generally tardily septicidal as well; seeds large, bearing a dorso-apical wing. (2n = 50). One hun­dred and ninety spp., open slopes and summits from Costa Rica and Guyana to Chile and Argen­tina. Divisible into two subgenera, Puyopsis and Puya, according to absence or presence of sterile branch ends serving as perches for pollinating birds.

14. Encholirium Mart. ex Schult. & Schult f.

Encholirium Mart. ex Schult. & Schult. f., in Roem. & Schult., Syst. veg. 7(2): lxviii, 1233 (1830); L.B. Sm. & Read, Bradea 5: 296-311 (1989).

Coarse xerophytes; leaves spinose-serrate; scape well developed; inflorescence usually simple or rarely few-branched, of many pedicellate flowers; flowers perfect; sepals convolute or imbricate, free; petals convolute, free, naked; stamens about equal­ling the petals to much exceeding them; filaments free; ovary wholly superior; capsule short, stout; seeds each with a single dorso-apical wing much extended backwards. Twenty nine spp., xerophytic habitats, northeastern Brazil to Mato Grosso.

15. Dyckia Schult. & Schult. f.

Dyckia Schult. & Schult. f., in Roem. & Schult., Syst. veg. 7(2): lxv, 1194 (1830).

Prionophyllum K. Koch (1874).

90 Bromeliaceae

Xerophytes, often repent; leaves usually spinose­serrate, rigid; scape usually lateral; inflorescence amply paniculate to simple; flowers perfect and uniform to rarely dimorphic with some function­ally unisexual; sepals imbricate, usually free; petals imbricate; stamens included to exserted; filaments basally connate among themselves and with the petals, free or connate above; ovary wholly superior; ovules many, each with a single wing; capsule short and broad, septicidal and sometimes partially loculicidal as well, seeds nu­merous, broad, alate. (2n =50, 100). One hundred and twenty four spp., xeric habitats, Central and E Brazil, Argentina, Bolivia, Paraguay, and Uruguay.

16. Hechtia Klotzsch

Hechtia Klotzsch, Allg. Gartenzeitung 3: 402 (1835).

Stem short; leaves densely rosulate; blades coarsely spinose-serrate in most species, rarely serrulate or even entire; scape usually lateral; in­florescence paniculate, bipinnate or more divided, branches from capitate to lax and very long; flow­ers functionally unisexual on different plants; se­pals free; petals naked, usually free but sometimes joined to a ring of filaments by their centers; sta­mens exserted to included; anthers mostly ovate; stigma simple-erect; ovary from wholly superior to 3/4 inferior; capsule septicidal and loculicidal; seeds many, oblong, narrowly winged or some­times almost naked, often sculptured. (2n = 50). Fifty one spp., Texas, Mexico, and northern Central America.

II. Subfam. Tillandsioideae Harms (1930)

Largely epiphytic, caulescent, or acaulescent. Roots often functioning only as holdfasts and sometimes completely lacking. Leaves rosulate or fasciculate or distributed along stem, entire; indumentum of radially symmetric scales. Pollen sulcate, usually with a distal germination area, rarely inaperturate (Guzmania) or sulcus simple ( Catopsis). Stigma morphology variable, all types present. Ovary superior or nearly so except in Glomeropitcairnia. Fruit capsular, septicidal; seeds with a plumose appendage at base or apex or both.

17. Catopsis Griseb. Fig. 25

Catopsis Griseb., Nachr. Konig!. Ges. Wiss. Georg Augusts­Univ. [1] : 10, 12 (Jan 1864).

Fig. 25A-G. Bromeliaceae-Tillandsioideae. Catopsis ber­teroniana. A Fruiting plant. B Flower with subtending bract. C Pistil in longitudinal section, showing one row of ovules. D Cross-section of ovary, showing two rows of ovules in each locule. E Three ovules showing elongated integument below and distal tuft of hairs developing into terminal appendage of seed. F Dehiscing capsule. G Seed with long terminal append­age of hairs. (Smith and Wood 1975)

Leaves densely utriculose-rosulate, often creta­ceous-coated; scape conspicuous; inflorescence usually bipinnate, rarely simple or tripinnate, the branches polystichous-flowered; flowers per­fect or dimorphic and functionally dioecious at least as to the staminate plants (Mexico and Central America); sepals free, usually strongly asymmetric with the right side produced; petals free, naked; stamens included, the 2 series usually very unequal; anthers ovate or elliptic; ovary superior; style short or none; stigma conduplicate-spiral or simple erect; ovules few to several, apically long-caudate, a small basal coma of agglutinated hairs appearing as a spur on the funicle; seeds with a folded apical coma. (2n =

Brameliaceae 91

50). Seventeen spp., xeric (canopy) epiphytes, Florida, Mexico and the West Indies to Brazil and Peru.

18. Glomeropitcairnia Mez

Glomeropitcairnia Mez, Bull. Herb. Boissier II, 5: 232 (1905); Gilmartin & al., Syst. Bot. 14:339-348 (1989); Till, Halbritter & Gortan, J. Brameliad Soc. 47: 65-72 (1997).

Stemless; leaves rosulate, the epidermal scales with a limb of regularly concentric cells; scape erect; inflorescence of sessile or short-stipitate heads or short spikes, bipinnate or tripinnate; flowers perfect; sepals free; petals free, bearing 2 scales at base; stamens included; stigma papillose, of the convolute-blade type; ovary about half infe­rior; capsule imperfectly dehiscent; seeds with a pappiform coma at both ends. Two spp., heliophytic habitats, Lesser Antilles, Trinidad and adjacent Venezuela.

19. Mezobromelia L.B. Sm.

Mezobromelia L.B. Sm., Proc. Am. Acad. Arts 70: 151, pl. 1, figs. 10, 11 (1935); H. Luther, Bromelia 2: 3-5 (1995).

Leaves densely rosulate; inflorescence bipinnate; scape well developed; flowers polystichous or dis­tichous and deflexed-secund, subsessile or pedi­cellate, perfect; sepals subfree or connate; petals tightly conglutinated, bearing 2 scales on the in­side; filaments adherent to the petals; stigma con­duplicate-spiral; ovary superior. Nine epiphytic spp., mesophytic habitats, Greater Antilles to Peru.

20 .. Guzmania Ruiz & Pav.

Guzmania Ruiz & Pav., Fl. peruv. 3: 37, pl. 261 (1802). Sodiroa Andre (1877).

Stemless or rarely caulescent; leaves densely polystichous; scape usually conspicuous; inflores­cence mostly bipinnate or simple, its branches polystichous-flowered; flowers perfect; sepals free to high-connate; petals naked, agglutinated in a tube; stamens equal, more or less agglutinated to the petals; ovary superior; style elongate; stigma simple-erect or of the convolute-blade type; ovules numerous; seeds with a basal, usually brown coma. (2n = 50). One hundred and sixty seven species, mainly mesophytic habitats, Florida, Mexico, and the West Indies to Brazil and Bolivia.

21. Alcantarea (E. Morren ex Mez) Harms

Alcantarea (E. Morren ex Mez) Harms, Notizbl. Bot. Gart. Berlin-Dahlem 10: 802 (1929); J.R. Grant, Trop. subtrop. Pflanzenwelt 91: 7-15 (1995).

Leaves forming large rosettes, inconspicuously !epidote; inflorescence usually paniculate; flowers distichous, secund, perfect; sepals free, convolute; petals free, soon becoming flaccid, with 2 entire appendages; stamens and style exserted, stigma with the convolute-blade type morphology; ovary cryptically half-inferior; seeds with both basal and apical comas. (2n = 50). Twelve spp., mainly heliophilic epilithes in lowland and montane for­ests, SE Brazil.

22. Vriesea Lindl.

Vriesea Lind!., Bot. Reg. 29: pl. 10 (1843), nom. cons.; J.R. Grant, Trap. subtrap. Pflanzenwelt 91: 5-57 (1995).

Acaulescent, usually epiphytic; leaves rosulate; scape usually conspicuous; inflorescence usually of distichous-flowered spikes; flowers perfect; se­pals convolute, free or nearly so; petals free or connate in a tube much shorter than the sepals, bearing 2 scales at base, firm and erect; stamens included or exserted; stigma con­duplicate-spiral, simple-erect, or of the convolute­blade type; ovary nearly or quite superior; ovules many, usually caudate; seeds fusiform with a long straight basal coma. (2n = 50). One hundred and ninety-three species in mainly mesophytic habitats. Mexico and theW est Indies to Argentina and Uruguay. Divisible into two sections, Vriesea and Xiphion, according to characters of the flowers.

23. Werauhia J.R. Grant

Werauhia J.R. Grant, Trap. subtrop. Pflanzenwelt 91: 16-57 (1995).

Usually stemless; leaves rosulate, inconspicuously !epidote; inflorescence paniculate with often re­duced branches; flowers usually with nocturnal anthesis, often secund, sessile to pedicellate, usu­ally zygomorphic; sepals free, ecarinate; petals free, with 2 badsal appendages; stamens and style not exserted from the corolla; stigma cupulate, its cup-shaped lobes erect and without papillae. (2n = 50). Sixty six species, mainly epiphytic in mon­tane and cloud forests, Central and N Andes, northern S America, Central America, and Greater Antilles.

92 Bromeliaceae

24. Tillandsia L. Fig. 26

Tillandsia L., Sp. pl.: 286 (1753); W. Till, Sippendifferen­zierung innerhalb Tillandsia subgenus Diaphoranthema. Doct. Diss. Univ. Wien (1984); C.S. Gardner, Selbyana 9: 130-146 (1986); Gilmartin & G.K. Br., Am. J. Bot. 73: 387-397 (1986); R.S. Beaman, Systematics of Tillandsia subge­nus Pseudalcantarea. M.Sc. Thesis Univ. Florida (1989).

Dendropogon Raf. (1825). Allardtia A. Dietr. (1852).

Caulescent or acaulescent; leaves rosulate or dis­tributed along a stem, polystichous or distichous; blades lingulate to narrowly triangular or linear; scape usually distinct; inflorescence usually of dis­tichous-flowered spikes, rarely reduced to a single polystichous-flowered spike or to a single flower; flowers perfect; sepals convolute, free or equally or adaxially joined; petals free, naked; stamens of various lengths relative to the petals and to the pistil; stigma conduplicate-spiral, simple-erect, coralliform, or of the convolute-blade type; ovary superior; ovules many, caudate or obtuse; seeds with a plumose, white, basal appendage. (2n = 38,

F

Fig. 26A-G. Bromeliaceae-Tillandsioideae. A Tillandsia bar­tramii, flower, showing tubular corolla and imbricate sepals. B-G T. utriculata. B Habit. C Style with stigmas. D Part of inflorescence with three nearly mature fruits. E Ovule, the apex caudate, the micropyle below (at right). F Nearly mature seed, oriented as in E. G Mature seed with basal appendage of hairs, oriented as in E and F. (Smith and Wood 1975)

44, 50, 100). Five hundred and forty spp., extreme xeric to extreme mesophytic habitats, terrestrial or frequently epiphytic. Southern United States to Argentina and Chile. Divisible into six subgenera, Allardtia, Anoplophytum, Phytarrhiza, Diaphor­anthema, Tillandsia and Pseudalcantarea accord­ing to characters of petals, stamens, and styles.

25. Racinaea M.A. Spencer & L.B. Sm.

Racinaea M.A. Spencer & L.B. Sm., Phytologia 74: 152 (1993).

Mainly stemmless with rosulate foliage; inflores­cence usually compound; flowers hermaphrodi­tic, small, actinomorphic, distichously arranged, sessile to short pedicellate; sepals convolute, asymmetric and broadest towards the apex, usu­ally free; petals naked, filaments succulent at the base; ovary superior, style subsessile to short, stigma papillose, of the simple-erect type; seeds with a badsal appendage only. (2n = 50). Fifty six mesophytic spp., mainly in montane and cloud forests, C and N Andes, Mesoamerica, Greater Antilles, Guayanas, and SE Brazil.

III. Subfam. Bromelioideae Harms ( 1930)

Herbs, largely epiphytic, usually acaulescent. Roots often functioning as holdfasts. Leaves rosulate or fasciculate, usually spinose-serrate or -serrulate; indumentum of peltate scales of irregu­larly arranged cells or very rarely of multicellular, cylindric trichomes. Pollen sulcate with a distal germination area, bi- to poly-porate, or inaper­turate (Canistrum, some Aechmea). Ovary com­pletely inferior or nearly so (Acanthostachys); stigma of the conduplicate-spiral type morphol­ogy except in Cryptanthus and Orthophytum; ovules obtuse to long-caudate. Fruit baccate but often dry, indehiscent; seeds naked.

26. Orthophytum Beer

Orthophytum Beer, Flora 37: 347 (1854).

Stemless or caulescent; leaves polystichous; blades serrate; scape erect or lacking; scape-bracts folia­ceous; inflorescence bipinnate from several very dense heads or rarely simple; primary bracts folia­ceous or subfoliaceous; floral bracts large, pun­gent; flowers perfect; sepals free; petals free, bearing 2 scales; stamens shorter than the petals and the second series adnate to them; pollen 4-pored or ellipsoid without distinct pores, reticu-

Bromeliaceae 93

late; epigynous tube usually lacking; placentae central; ovules obtuse. Twenty four spp., saxi­colous and terrestrial on rocky ground, NE Brazil.

27. Cryptanthus Otto & A. Dietr.

Cryptanthus Otto & A. Dietr., Allg. Gartenzeitung 4: 297 (1836), nom. cons.; non Cryptanthus Osbeck (1757).

Stemless or caulescent; leaves rosulate or evenly polystichous along the stem; blades narrowly tri­angular or slightly contracted to subpetiolate at base, spinose-serrate; inflorescence sessile in the center of the rosette or at the apex of the stem, compound; flowers polygamous in some species at least; sepals much connate; petals connate by their claws, naked or with two callosities; stamens exserted from the throat of the corolla, usually highly adnate to the petals; pollen sulcate; ovules usually few, obtuse; placentae apical or median; epigynous tube none or very shallow; fruit small, rather dry; seeds small, flat. (2n = 34, 54). Forty one spp., xerophytic, saxicolous, or terrestrial, NE Brazil.

28. Araeococcus Brongn.

Araeococcus Brongn., Ann. Sci. Nat. Bot. II. 15: 370 (1841).

Stemless; inflorescence laxly paniculate; flowers perfect; sepals free to connate for half of their length; petals free, naked; stamens shorter than the petals; filaments free; anthers longer than the fila­ments; pollen grains ellipsoid, sulcate; epigynous tube none; style longer than the petals; stigma lobes scarcely twisted; ovules few (up to 10), borne at the top of the locule, long-caudate; seeds nar­rowly ovoid-cylindric, acute at both ends. (2n = 50). Six spp., mesophytic epiphytes, Costa Rica, Colombia, Venezuela, Trinidad, Guyana, Brazil.

29. Fascicularia Mez

Fascicularia Mez in Mart., Fl. bras. 3(3): 627 (1891).

Terrestrial or rarely on three trunks, stemless; leaves very densely rosulate; blades spinose­serrate; inflorescence sunk in the center of the rosette, simple; flowers perfect; sepals free; petals free, bearing 2 small scales near base; epigynous tube conspicuous; stamens included; filaments free; anthers linear, elongate, dorsifixed near base; pollen globose, sulcate, smooth; style elongate; stigmas not contorted; placentae central to subapical; ovules rather few, globose, short-

appendaged; fruit dry; seeds globose, naked. Five spp., Chile.

30. Greigia Regel

Greigia Regel, Gartenflora 14: 137 (1865).

More or less caulescent; leaves imbricate along the stem, partially or completely serrate or serrulate; scape very short; inflorescence lateral or central, simple or compound, subglobose; sepals free or connate; petals connate, usually naked; filaments adnate to the petals; anthers narrow, acute; pollen large, spherical, sulcate, smooth; fruit fleshy. Twenty eight spp., saxicolous in open ground or terrestrial in forest or paramo, Chile to Mexico and Venezuela.

31. Fernseea Baker

Fernseea Baker, Handb. Bromel.: 20 (1889); emend. E. Pereira & Moutinho, Bradea 3: 343, fig. (1983).

Stemless; leaves forming a dense pseudobulb; blades spinose-serrate; inflorescence simple, race­mose; flowers perfect; sepals free; petals free, naked; stamens free, included at anthesis; pollen sulcate; epigynous tube large; ovules numerous, unappendaged; fruit succulent; seeds unappend­aged. Two spp., alpine habitats, Brazil; Mount Itatiaia region in NW Rio de Janeiro and adjacent Sao Paulo.

32. Ochagavia Phil.

Ochagavia Phil., Anales Univ. Chile 13: 168 (May 1856).

Suffruticose, amply proliferating; leaves tank­forming; blades spinose-serrate; inflorescence sunk in the center of the leaves, simple; flowers perfect; sepals free; petals free, naked; stamens usually exserted; anthers subelliptic, dorsifixed; pollen eporate with a single longitudinal fold; epi­gynous tube very conspicous; placentae extending the whole length of the locule; ovules rather nu­merous, unappendaged; fruit crowned by the per­sistent sepals; seeds large, globose. Three spp., coastal cliffs, Juan Fernandez Islands, Chile.

33. Ursulaea Read & H.U. Baensch

Ursulaea Read & H.U. Baensch, J. Bromeliad Soc. 44: 206 (1994).

Podaechmea (Mez) L.B. Sm. & W.J. Kress (1990) in part, not as to type.

94 Bromeliaceae

Stemless; leaves rosulate, serrate; scape elongated; flowers perfect, subactinomorphic, pedicellate; sepals free, nearly symmetric, mucronate; petals apically recoiled, with two fimbriate scales at the base; stamens and style exserted; epigynous tube shallow; pollen sulcate; ovary inferior, placentae subapical, ovules caudate. Two spp., epiphytic and terrestrial, Mexico: Jalisco.

34. Billbergia Thunb.

Billbergia Thunb., Pl. bras. 3: 30 (1821). Helicodea Lem. (1864).

Fig. 27A-F

Stemless; leaves forming broadly funnelform to long-tubular rosettes; blades serrate or serrulate or rarely entire; scapebracts large, mostly red; in­florescence compound or simple; flowers perfect; sepals free; petals free, actinomorphic or slightly zygomorphic, bearing 2 scales near base; epigy­nous tube usually developed; stamens exserted, both series or the second adnate to the petals only as high as the scales; pollen sulcate, sometimes subporate as well (B. elegans Mart. ex Schult. & Schult. f.); style exceeding the stamens; ovules many. (2n = 50). Sixty two spp., terrestrial, saxi­colous, and epiphytic, Mexico and the West Indies (Bahamas) to Uruguay and Argentina. Divisible into two subgenera, Billbergia and Helicodea, ac­cording to the opening of the petals at anthesis, and somewhat to the form of the inflorescence.

35. Neoglaziovia Mez

Neoglaziovia Mez, in Mart., Fl. bras. 3(3): 426 (1894).

Stemless; leaves few; inflorescence simple, race­mose; flowers perfect; sepals free, unarmed; petals free; stamens included; filaments free; pollen sul­cate; ovules few, short-caudate; fruit succulent. Three spp., terrestrial and saxicolous in dry open areas or low thorn forest (caatinga). NE Brazil.

36. Bromelia L.

Bromelia L., Sp. pl.: 285 (1753).

Coarse terrestrials or rarely epiphytes, spreading by rhizomes; leaves coarsely spinose along the margins; inflorescence sessile or scapose, com­pound; sepals free to high-connate, unarmed; pet­als united into a tube by the filaments but their margins free, without appendages; stamens in­cluded, their filaments forming a tube; epigynous tube conspicuous to nearly lacking; fruit succu-

Fig. 27A-K. Bromeliaceae-Bromelioideae. A-F Billbergia nutans. A Plant with pollinator. B Flower. C Same, longitudi­nal section. D Stamen. E Style with stigma. F Base of petal with scales. G-L Nidularium scheremetiewii. G Habit. H Flower bud. I Opened ovary and floral tube. J Stigma. K Stamen. (Takhtajan 1982)

lent, relatively large; seeds few to many, flattened, naked. (2n = 50, 100). Forty nine spp., mostly in dry open ground but sometimes submesophytic, Central and S America.

Bromeliaceae 95

37. Deinacanthon Mez

Deinacanthon Mez, in A. & C. DC., Monogr. phan. 9: 12 ( 1896).

Stemless, rhizomatose; leaves squarrose-rosulate; blades spinose-serrate; inflorescence simple, sessile; flowers perfect; sepals free, spinose-mucr­onate; filaments connate with the petals; pollen sulcate; ovules numerous. Only one sp., D. urbanianum (Mez) Mez, dry open woodland to rocky crests, Paraguay, NW Argentina.

38. Chevaliera Gaudich. ex Beer

Chevaliera Gaudich. ex Beer, Pam. Bromel. 22, 257 (1856); L.B. Sm. & W.J. Kress, Phytologia 66: 77-79 (1989).

Large xerophytes, leaves rosulate, serrate; inflo­rescence strobilate, usually simple; floral bracts coriaceous or ligneous; flowers in many ranks; petals with reduced or lacking appendages; pollen sulcate. Twenty two spp., terrestrial in lowland and montane forests, rarely epiphytic, Northwest­ern S America, Central America, SE Brazil.

39. Pseudananas Hassi. ex Harms

Pseudananas Hassi. ex Harms, in Engl. & Prantl, Nat. Pflanzenfam. ed. 2, 15a: 153 (1930).

Plant propagating by elongate basal stolons; leaves rosulate; blades with spinose margins; scape evident; inflorescence simple, strobilate, the apical coma inconspicuous or lacking; flowers connate by their ovaries and forming a syncarp; sepals free; petals bearing lateral folds which over­lap the filaments; fruit large, succulent. (2n =

100). Only one sp., P. sagenarius (Arruda) Camargo, terrestrial on open ground to dense forest, from near sea level to 1400 m alt, Ecuador, Bolivia, Brazil, Paraguay, Argentina.

40. Ananas Mill.

Ananas Mill., Gard. diet. abr. ed. 4 (1754).

Plant not producing stolons; leaves rosulate; scape evident; inflorescence densely strobiliform, usu­ally crowned with a tuft of sterile foliaceous bracts, often producing slips at the base; flowers sessile; sepals free; petals free, each typically bearing 2 slenderly funnelform scales; stamens included; pollen ellipsoid with 2 pores; ovaries coalescing with each other and with the bracts to form a fleshy syncarp; epigynous tube short; placentas apical; ovules caudate. (2n = 50, 3n = 75). Eight

spp., terrestrial in humid or dry open or forested areas, native to S America south to Uruguay.

41. Acanthostachys Klotzsch

Acanthostachys Klotzsch, in Link, Klotzsch & Otto, Icon. pl. rar. 1: 21, pl. 9 (1840); Rauh & Barthlott, Trop. subtrop. Pflanzenwelt 39: 5-35 (1982).

Rhizomatose; leaves fasciculate or rosulate; scape well developed or lacking; inflorescence simple, densely polystichous-flowered; flowers sessile, perfect; sepals free; petals free, bearing 2 scales; stamens included; pollen biporate; ovary com­pletely inferior, the thickened style base of A. strobilacea (Schult. & Schult. f.) Klotzsch partially superior; epigynous tube none or crateriform (A. pitcairnioides (Mez) Rauh & Barthlott); ovules long-caudate; seeds with a strongly elongated apical appendage. (2n = 50). Two spp., epiphytic and saxicolous, E Brazil, Paraguay, NE Argentina.

42. Hohenbergia Schult. & Schult. f.

Hohenbergia Schult. & Schult. f. in Roem. & Schult., Syst. V eg. 7(2): lxxi, 1251 (1830).

Stemless; leaves rosulate; blades spinose-serrate; scape well developed; inflorescence of strobilate spikes; flowers perfect; sepals mostly very short­connate; petals with 2 appendages; stamens in­cluded, series I free, series II partly adnate to the petals; pollen with 2 or 4 pores; placentas mostly apical; ovules obtuse to long-caudate. Forty seven spp., terrestrial, saxicolous or epiphytic, West Indies to E Brazil. Divisible into two subgenera, Hohenbergia and Wittmackiopsis, according to ovule shape, leaf anatomy, and inflorescence branching.

43. Aechmea Ruiz & Pav.

Aechmea Ruiz & Pav., Fl. peruv. prodr.: 47 (1794); nom. cons. Macrochordion de Vriese (1853). Hoplophytum Beer (1854), incl. Platyaechmea (Baker) L.B. Sm.

& W.J. Kress. Lamprococcus Beer (1856). Ortgiesia Regel (1867). Pothuava Gaudich. ex L.B. Sm. & W.J. Kress (1989). Podaechmea (Mez) L.B. Sm. & W.J. Kress (1990).

Acaulescent, frequently propagating by rhizomes; leaves usually forming a tank, usually spinose­serrate; scape well developed; inflorescence com­pound or simple; flowers distichous or polystic­hous, sessile or stipitate, perfect; sepals free or

96 Bromeliaceae

connate, usually strongly asymmetric; petals free, bearing 2 more or less adnate basal appendages; stamens shorter than the petals, free or the second series more or less adnate to the petals; anthers dorsifixed; pollen with 2, 3, 4 or numerous pores, rarely sulcate; ovules mostly caudate; seeds small. (2n = 42, 50). One hundred and eighty two species, largely terrestrial or epiphytic in forest. Mexico and the West Indies to Argentina. Divisible into seven subgenera, Podaechmea, Lamprococcus, Aechmea, Ortgiesia, Platyaechmea, Pothuava, and Macrochordion, according to inflo­rescence form, floral order, pedicels, and sepal connation and armament.

Aechmea includes some very discordant ele­ments and is very likely of polyphyletic origin. Further research is likely to divide it with some parts becoming independent genera and others merging with genera now considered distinct.

44. Quesnelia Gaudich.

Quesnelia Gaudich., Voy. Bonite, Bot., Atl. pl. 54 (1842).

Stemless or rarely long-stemmed; leaves serrulate; scape developed, inflorescence usually simple; flowers perfect; sepals free or short -connate; petals free, bearing 2 scales near base; stamens shorter than the petals and generally included; filaments of the second series highly adnate to the petals; pollen ellipsoid, biporate; epigynous tube distinct; placentas central to apical; ovules obtuse, numerous. (2n = 50?, 100). Fifteen spp., terres­trial, saxicolous, or epiphytic, from sea strand to upland rainforest, E Brazil. Divisible into tw-o sub­genera, Quesnelia and Billbergiopsis, according to form of floral bracts and inflorescence.

45. Canistrum E. Morren1

Canistrum E. Morren, Belgique Hort. 23: 257 (1873).

Mostly epiphytic; leaves rosulate, armed; scape evident; inflorescence compound, densely corym­bose, involucrate with the bracts covering all but the apices of the flowers; flowers perfect; sepals free or nearly so; petals free, appendaged; stamens included; pollen hi- or multiporate; ovules obtuse or short-caudate, numerous; seeds many, sub­fusiform. (2n = SO). Ten spp., rainforests, E Brazil. Divisible into two subgenera, Canistrum and Cucullatanthus, according to petal and ovule morphology (Leme 1997).

1 Two species have recently been segregated to be members of the new genus Edmundoa Leme (Leme 1997).

46. Wittrockia Lindm.

Wittrockia Lindm., Ofvers. Forh. K. Sven. Vetenskapsakad. III. 24(8): 15, 20 (1891).

Leaves rosulate, serrate or serrulate; scape usually evident; inflorescence compound, very densely capitiform in the center of an involucre of sub foliaceous bracts; flowers perfect; sepals free or more or less connate; petals more or less connate or agglutinated near base, appendaged; stamens included; style long but included; placentas cen­tral or nearly; ovules obtuse. Eleven spp., terres­trial, saxicolous, and epiphytic, E Brazil.

47. Portea Brongn. ex K. Koch

Portea Brongn. ex K. Koch, Index Sem. Horti Bot. Berol. "1856", App.: 7 (1857).

Leaves rosulate, serrulate; scape bracts brightly colored; inflorescence compound; flowers slen­derly pedicellate, perfect; sepals more or less con­nate; petals free, bearing 2 fimbriate scales at base; stamens of the second series highly adnate to the petals; pollen with more than 5 pores; placentas various; ovules numerous, long-caudate. (2n = 50). Nine spp., saxicolous, terrestrial, or epiphytic in forest, E Brazil.

48. Pseudaechmea L.B. Sm. & Read

Pseudaechmaea L.B. Sm. & Read, Phytologia 52: 53, pis. 3, 4 (1982).

Leaves rosulate, serrulate; scape elongate; inflo­rescence simple, laxly racemose; pedicels very long; flowers polystichous, perfect; sepals subfree; petals naked; stamens included; filaments free; pollen biporate; epigynous tube none; placentas subapical; ovules caudate. Only one sp., P. ambigua L.B. Sm. & Read, native to tropical forest in NW Colombia.

49. Hohenbergiopsis L.B. Sm. & Read

Hohenbergiopsis L.B. Sm. & Read, Phytologia 33: 440 (1976).

Coarse leaves rosulate; blades spinose-serrate; scape developed; inflorescence compound, the ul­timate branches strobilate; flowers sessile, perfect; petals naked, attached to the filament tube but not to each other; pollen in tetrads; epigynous tube lacking; ovules caudate. Only one sp., H. guatemalensis (L.B. Sm.) L.B. Sm. & Read, on trees in dense wet forest. S Mexico, Guatemala.

Bromeliaceae 97

50. Androlepis Brongn. ex Houllet

Androlepis Brongn. ex Houllet, Rev. Hort. 42: 12 (1870).

~eaves rosulate, spinose-serrate; scape erect; mflorescence compound with simple or divided branches; spikes polystichous-flowered; flowers functionally unisexual on different plants, sessile; sepals free; petals short -connate, naked; stamens included; filaments short, adnate; anthers each bearing 2 foliaceous appendages at apex; pollen biporate, in tetrads; epigynous tube short; ovules numerous; borne along the upper part of the locule, acute. Only one sp., A. skinneri Brongn. ex Houllet, epiphytic in forest, S Mexico to Costa Rica, and Peru.

51. Lymania Read

Lymania Read, J. Bromeliad Soc. 34: 201 (1984); Leme, J. Bromeliad Soc. 39: 262-264.

Slenderly stoloniferous or stout-rhizomatous; leaves few, forming a crateriform or utriculate rosette, minutely serrulate; scape erect; inflores­cence simple or bipinnate, lax or dense; flowers sessile, polystichous, perfect; ovary deeply sulcate or alate; epigynous tube very short but distinct; sepals about half connate; petals free, naked; sta­mens included; anthers sagittate, apiculate; pollen porate, smooth; filaments connate or free the . ' mner series adnate to the petals; placentae su~apical; ovules obtuse or variously caudate; frmt globose, ridged. Six mesophytic spp., Brazil: Bahia.

53. Neoregelia L.B. Sm.

Neoregelia L.B. Sm., Contr. Gray Herb. 104: 78 (1934); I. Ramirez, Systematic Revision of Neoregelia subgenus Hylaeaicum, St. Louis: M.Sc. Thesis (1991).

Leaves rosulate, usually spinose-serrate; inflores­cence sunk in the center of the rosette, simple in most species, very densely capitiform, umbellate or corymbose; flowers perfect, pedicellate; sepals asymmetric; petals free or connate, naked; sta­mens and pistil included; ovules obtuse. (2n = 50). Ninety seven spp., epiphytic in forest to saxicolous and heliophytic, E Brazil to Amazonian S America. Three subgenera, Hylaeaicum Neoregelia, accord­mg to petal connation, pedicel morphology, and polle~ morphology. Hylaeaicum probably merits genenc status.

54. Streptocalyx Beer

Streptocalyx Beer, Flora 37: 348 (1854).

Terrestrial or epiphytic; leaves densely rosulate, serrate; scape elongate to almost none; inflores­cence 2-4-pinnate, laxly paniculate to densely subglobose; flowers sessile; sepals free to distinctly connate; petals free, naked; stamens and pistil in­cluded; pollen 2- or 4-porate; epigynous tube dis­tinct; seeds ellipsoid or ovoid, naked. (2n = 50). Twenty spp., mostly in forest, Eastern and Amazo­nian Brazil and adjacent areas. Probably to be merged with Aechmea.

55. Ronnbergia E. Morren & Andre

Ronnbergia E. Morren & Andre, Ill. Hort. 21: 120, pl. 177

52. Nidularium Lem.2 Fig. 27G-L (1874).

Nidularium Lem., Jard. Fleur. 4 (Misc.): 60, pl. 411 (1854). Andrea Mez (1896).

Stemless; leaves forming funnelform tanks, serrate or serrulate; scape short; inflorescence compound with flowers in flat fascicles and covered by large bracts; flowers sessile or rarely pedicellate, perfect; sepals more or less connate; petals naked or rarely with vertical folds, tubular­connate; stamens included; pollen biporate; ovules numerous, obtuse. (2n = 50, 100). Fifty spp., epiphytic or terrestrial in rainforest, E Brazil.

A revision of the genus is forthcoming (Leme in press).

2 Pa~t of t~e .genus is being segregated as a separate genus Camstropsts (mel. Andrea; Leme in press).

Lea_ves fasciculate or rosulate, often petiolate, entire or serrulate; scape well developed; inflores­cence simple, elongate; flowers sessile, perfect; se­pals more or less connate; petals free, naked or with lateral folds; stamens included, the second series free or partially adnate to the petals; pollen biporate; epigynous tube large; placentas apical. Twelve spp., epiphytic in rainforest, Costa Rica to Peru. Probably to be merged with Aechmea.

56. Disteganthus Lem.

Disteganthus Lem., Fl. Serres Jard. Eur. 3: pl. 227 (1847).

Caulescent or acaulescent, propagating by stolons and rhizomes; leaves petiolate, armed; scapes several, borne beneath the rosette, bracteate; inflorescence strobilate, simple; flowers perfect;

98 Bromeliaceae

sepals high-connate; petals naked, free; stamens included; pollen porate; epigynous tube large; pla­centas apical; ovules obtuse. Three spp., swampy forest, Surinam, French Guiana, and adjacent Brazil.

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