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Morphology and anatomy of palm seedlings.

Abstract
Introduction
Historical Survey
Materials and Methods
Results
 I. Coryphoideae
 II. Calamoideae
 III. Nypoideae
 IV. Ceroxyloideae
 V. Arecoideae
 VI. Phytelephantoideae
Discussion
Acknowledgments
Literature Cited
Appendix


Introduction

The palm family (Palmae or Arecaceae) is one of the largest families of monocotyledons. The most recent estimate is that it contains 190 genera and 2364 species (Govaerts & Dransfield, 2005). These are widespread in tropical areas throughout the world, with the greatest concentration of species in America and Southeast Asia. Palms are usually very abundant in tropical ecosystems, especially in lowland and montane moist forests.

The Palmae are one of the most economically important families of plants to man, ranking after grasses and, in the tropics, equal with legumes. Apart from the well-known crops coconut (Cocos nucifera), oil palm (Elaeis oleifera), and date palm (Phoenix dactylifera), other species of palm provide numerous useful products such as food stuffs, fibers, and medicines (Balick & Beck, 1990). Because of this economic importance, and because of their abundance in tropical ecosystems, palms have received much attention from botanists. Numerous aspects of the family have been extensively studied, such as systematics, reproductive biology, economic uses, and biogeography. Nevertheless, some important aspects of palm biology remain to be investigated, including seedling biology.

The present study takes as a starting point that there has been no recent survey of germination and seedlings in the palm family, and the subject remains poorly understood. In this study, the seedling morphology and eophyll anatomy of 62 genera representing the six subfamilies of palms are described. Since almost all palms are propagated by seed, the subject of germination is clearly an important one.

Historical Survey

The scientific study of palm germination began in the early nineteenth century with the work of Martius (1823-1850). Martius recognized distinct types of germination in palms: "germinatio admotiva," in which the seedling develops adjacent to the seed, and "germinatio remotiva," in which the seedlings develop at some distance from the seed. This latter type was again divided into "germinatio remotiva tubulosa," in which the tubular cotyledonary sheath is open (e.g., Arenga, Phoenix) and "germinatio remotiva ocreata," in which the cotyledonary sheath extends, forming an ocrea (e.g., Brahea, Chamaerops). These three types of germination described by Martius have served as the basis for all future discussions of palm seedlings (Fig. 1) Mohl (in Martius, 1823-1850) described and illustrated in great detail the anatomy of embryos and germination of palm seeds.

[FIGURE 1 OMITTED]

Micheels (1889) proposed three germination types based on the attachment of the "embracing region" (cotyledonary sheath). His results, based on 33 taxa, are basically similar to the types of Martius, although he assigned new names: "Phoenix type" palms, with the sheath at the base of the cotyledonary stalk; "Sabal type" palms, with a tubular sheath; and "Dictyosperma type" palms, with a short sheath. Micheels' anatomical studies include a detailed description of the primary root (radicle), hyperphyll, and first and second cataphylls.

The most comprehensive historical review of germination studies in palms is that of Gatin (1906a), which includes information from the early works of Pliny and Theophrastus and the first illustration of a palm seedling, produced over 400 years ago (Camerarius, 1588). Gatin carried out detailed morphological and anatomical studies of embryos and the germination of Phoenix and Archontophoenix. He then surveyed 33 other genera and 58 species of palms. He observed a relation between the internal structure of the embryo and the germination pattern. The embryo can take one of three forms: the plumular-radicular axis can be straight or curved; the plane or orientation of the straight plumular-radicular axis can be parallel or oblique with respect to the axis of the embryo; and a straight plumular-radicular axis gives rise to nonligulate germination, whereas a curved plumular-radicular axis gives rise to ligulate (ocreate) germination. Gatin surveyed many aspects of palm seedlings (Gatin, 1904a): primary roots of palm seedlings (Gatin, 1904b), polyembryony (Gatin, 1905), chemistry of Borassus germination (Gatin, 1906b), anatomy of the hyperphyll (Gatin 1907a), roots of seedling palms (Gatin, 1907b), and morphology of germination (Gatin, 1908). He summarized all of his work in a comprehensive book on palms (Gatin, 1912).

Several other studies followed the germination types of Martius and Gatin without discussion. Significant contributions to the field were made by several researchers, including Zurawska (1912), who gave extensive morphological descriptions of germination and seedlings of 24 species of palms. Unfortunately, she seemed unaware of Gatin's (1906a) work, and her study does not include details of embryo structure. Yampolsky (1922), in a study of the oil palm Elaeis guineensis, gave a detailed historical review of studies on the leaf and haustorium in palms. Boyd (1932) carried out a survey of seedlings of all monocotyledons and included a few species of palms not treated by Gatin (1906a). She provided information on morphology and vascular tissues of the cotyledon, plumule, and roots, and described briefly the lamina of Cocos capitata (= Butia capitata).

In 1950, the French botanist Ginieis began a series of studies of palm germination and seedlings. These are referred to below under the relevant genera. Saakov (1954) studied germination of economically important palms. He concluded that palms with remote tubular (nonligular) germination are primitive, that admotive ligular palms are phylogenetically younger groups, and that remote ligular palms are intermediate between old remote tubular and early admotive ligular. He also suggested that morphological features of germination blur or disappear after the emergence of the second or third leaves.

Tomlinson (1960a, 1990) reviewed germination and seedling morphology, including illustrations and examples. He followed Martius's three germination types, but suggested that both Nypa and Phytelephas may have different patterns of germination. He discussed specialized germination types in Lodoicea, Cocos, Nypa, and Pigafetta.

Moore and Uhl (1973) attempted to use germination characteristics to understand phylogeny. They considered remote-tubular and remote-ligular germination types to be less-specialized groups, and the genus Livistona, which has remote-ligular germination, to be the most advanced of the Corypheae. The adjacent-ligular type corresponded to advanced groups, among them the Areceae.

Low (1976), in an extensive but overlooked work on palm phylogeny, reviewed seedling morphology of 84 species in 82 genera of palms. He made several original observations on morphological characteristics of germination. He listed several seedling characteristics and emphasized the number of cataphylls as a diagnostic characteristic. Uhl and Dransfield (1987) provided information on germination type and eophyll shape for most genera of palms. They followed Martius's basic germination types with a variation in the terminology: remote tubular (= germination remotiva tubulosa), remote ligular (= germinatio remotiva ocreata), and adjacent (= germination admotiva).

Tillich (1995) reviewed seedling morphology in all monocotyledons and demonstrated the usefulness of seedling characteristics in monocot systematics. In Tillich's review, the palm family formed an isolated group with basal characters. He also standardized germination terminology throughout the monocotyledons, and his terminology is used here (Fig. 2, Appendix). Tillich (2000) stressed the importance of the cotyledon and the nature of the first cataphyll versus the eophyll to define seedling types and evolutionary levels. He concluded that the ancestral seedling type in monocots is characterized by a compact cotyledon, one to several cataphylls, a short hypocotyl with inconspicuous collar, and a vigorously growing, branched primary root. He considered the seedling structure to be a key characteristic for detection of phylogenetic relationships.

[FIGURE 2 OMITTED]

Materials and Methods

Plant material (seeds and seedlings) representing all of the subfamilies were gathered from various sources, such as botanical gardens, private gardens, personal contacts, and fieldwork in Bolivia and Peru. Two taxa, Podococcus barteri and Iriartella setigera, were examined from herbarium specimens. The seeds were germinated and grown under standard conditions of light, temperature, and humidity. Identification of the source material was made in situ. Each sample consisted of several seedlings at different stages of development. Sixty-two genera representing all subfamilies were evaluated. Voucher specimens were made and are deposited at The New York Botanical Garden (Table I). Seedlings with the eophyll fully expanded were collected and fixed in FPA (formalin:proprionic acid:acetic acid, 5:5:90) and stored in 70% ethanol.

Seedling morphology was studied by direct observation using a hand lens or a Wild Heerbrugg MTr3 dissecting microscope. Anatomy of the lamina and petiole was studied by observation of transverse sections, epidermal peels, and leaf clearings using an Olympus Differential Interface Contrast Attachment model BH2-NIC microscope. Anatomical procedures follow Martens and Uhl (1984). Segments of 2 cm2 from near the base and apex of the lamina and petiole were processed through a dehydration series of absolute ethanol and toluene. Blocks were sectioned on an AO Spence 820 rotary microtome at 12-17 mm and stained with safranin and astra blue. Segments for epidermal peels were obtained from the upper side of the lamina. Images of anatomical features were taken with a Nikon FX-35 camera attached to the microscope. For morphological features, a Nikon Coolpix 990 digital camera was used. Images were processed in Adobe PhotoShop.

Results

Each sample consists of a complete seedling, eophyll transverse sections of proximal and distal sections, an eophyll epidermal peel, and eophyll clearing. The taxa descriptions within each subfamily and tribe (Uhl & Dransfield, 1987) are arranged in alphabetical order.

I. CORYPHOIDEAE

1. Corypheae

Acoelorraphe wrightii (Griseb. & H. Wendl.) H. Wendl. ex Becc.

Seed remaining above plumular-radicular node. Plumular-radicular axis straight in the same plane. Primary root stout and persistent, smooth; secondary roots simple; shoot-borne roots present; root hairs present. Hyperphyll elongate and smooth, connected halfway between cotyledonary sheath and coleoptile. Cotyledonary sheath distinct. Coleoptile apically open, short split opposite to seed. Cataphyll single; apex acute. Eophyll entire, linear-lanceolate; apex acute. Venation pattern costapalmate; leaf axis reduced; midvein distinct along blade; veins convergent at apex; transverse commissures widely separated from each other, connecting some longitudinal veins. Plication proximal and distal marginal folds induplicate. Epidermal cells rectangular; adaxial and abaxial anticlinal walls linear. Hairs absent. Stomata superficial, arranged in regular lines at intercostal regions; short terminal cells not overarching guard cells. Hypodermis single-layered, rounded regular cells at adaxial and abaxial sides; fibrous bundles at irregular intervals; fiber lumen wide. Chlorenchyma undifferentiated; spongy mesophyll with more than five layers; fibers appear to be restricted to hypodermal layer. Expansion cells single-layered with scattered fibrous bundles. Major veins associated with ridges, attached to adaxial hypodermis and abaxial expansion cells; distinct outer sheath (OS). Median veins free, equidistant; inner sheath (IS) single- or double-layered. Minor veins equidistant; IS single-layered; OS surrounding vascular bundle. Midrib adaxially prominent; single large vascular bundle. Marginal rib occupied by fibrous layers. Petiole transverse section crescent-shaped. Phloem strands two. Metaxylem vessel single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles; tannins abundant in full sacs.

Chamaerops humilis L.

Seed remaining above plumular-radicular node. Plumular-radicular axis on the same plane. Primary root straight and persistent; secondary roots simple; shoot-borne roots thick; root hairs present. Hyperphyll elongate, grooved lengthwise. Cotyledonary sheath apical opening, eventually splitting lengthwise, opposite to hyperphyll. Coleoptile undeveloped. Cataphyll single. Eophyll entire, broadly lanceolate; apex acute; fifth successional leaf splitting along adaxial ridge (Fig. 3A). Venation pattern palmate, convergent at apex; leaf axis reduced; midvein not distinct from other longitudinal veins; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal and distal marginal folds induplicate. Epidermal cells rectangular; adaxial and abaxial anticlinal walls linear. Hairs absent. Stomata slightly sunken, scattered on adaxial surface, abundant on abaxial surface; terminal cells short, occasionally elongate and overarching guard cells. Hypodermis single-layered, present at adaxial and abaxial sides; fibrous bundles at irregular intervals; solid bundles at ridges and grooves; lumen wide (Fig. 4A). Chlorenchyma undifferentiated; spongy mesophyll with more than five layers. Expansion cells single-layered; large rounded cells. Major veins associated with ridges, attached to adaxial and abaxial hypodermal layers; IS sclerotic, multilayered; OS not distinct. Median veins free and equidistant. Minor veins equidistant; OS surrounding vascular bundle. Midrib abaxially prominent; single large vascular bundle. Marginal rib with compact fibrous bundle. Petiole transverse section crescent-shaped. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles; tannins abundant, except in hypodermal and expansion cells.

[FIGURES 3-4 OMITTED]

Gatin (1906a) and Ginieis (1950, 1952a) have described germination in Chamaerops.

Chuniophoenix hainanensis Burret

Seed remaining above plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root persistent; secondary roots simple; shoot-borne roots present; collar disk and collar roots present. Hyperphyll medium length, smooth surface, slightly constricted at insertion to seed. Cotyledonary sheath distinct; texture rugulose. Coleoptile leathery, splitting opposite to seed. Cataphyll single, grooved. Eophyll entire, lanceolate; apex acute. Venation pattern palmate; leaf axis reduced; midvein not distinct from other longitudinal vascular bundles; veins convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal and distal marginal folds induplicate. Epidermal cells rectangular; adaxial and abaxial anticlinal walls slightly sinuous. Hairs present; few basal cells, associated with ribs. Stomata slightly sunken, scattered; short terminal cells not overarching guard cells. Hypodermis single-layered; large rounded or ellipsoid cells; parallel orientation; present at adaxial and abaxial sides. Chlorenchyma undifferentiated; spongy mesophyll with less than five layers. Expansion cells double-layered. Major veins associated with ridges, attached to adaxial hypodermis and abaxial expansion cells; OS distinct. Median veins buttressed to adaxial side. Minor veins equidistant; OS surrounding vascular bundle. Midrib abaxially prominent; single vascular bundle. Marginal rib lacking a vascular bundle or fibrous bundle. Petiole transverse section crescent-shaped. Phloem strands three. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles; raphids scattered, equidistant; tannins scarce.

Colpothrinax cookii Read

Seed remaining above plumular-radicular node. Plumular-radicular axis asymmetric, forming an angle. Primary root persistent; disk collar distinct; secondary roots branched; pneumatophores scattered; shoot-borne roots present; root hairs present. Hyperphyll short and smooth, without grooves or furrows; attachment area flat. Cotyledonary sheath extremely short. Coleoptile leathery; irregular splitting opposite to seed. Cataphyll single; basal part naked and smooth; distal part rough and grooved. Eophyll entire, broadly lanceolate; apex acute; splitting along adaxial fold. Venation pattern palmate; leaf axis reduced; midvein distinct; veins convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal and distal marginal folds induplicate. Epidermal cells rectangular or rhombohedral; adaxial and abaxial anticlinal walls linear; sclerotic walls; cuticle thick. Hairs present; multicellular base, associated or not associated with fibrous bundles. Stomata slightly sunken; short terminal cells overarching guard cells. Hypodermis single-layered, large ellipsoid cells; orientation parallel; present at adaxial and abaxial sides. Chlorenchyma undifferentiated; spongy mesophyll with more than five layers; mesophyll fibers forming compact bundles of about 20 strands, arranged at adaxial and abaxial sides; lumen small. Expansion cells double-layered cells, large, rectangular, and perpendicular; adjacent epidermal cells papillose. Major veins not associated with ridges; OS distinct. Median veins buttressed to adaxial side. Minor veins oriented toward abaxial side; OS surrounding vascular bundle. Midrib abaxially prominent, vascularized by grouped simple vascular bundles. Marginal rib not vascularized. Petiole transverse section crescent-shaped. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular and fibrous bundles; raphids equidistant; tannins in scattered full sacs.

Copernicia baileyana Leon

Seed remaining above plumular-radicular node. Plumular-radicular axis symmetric, straight. Primary root persistent; swollen collar absent; secondary roots simple; shoot-borne roots absent; collar roots present; root hairs present. Hyperphyll elongate with a single groove in adaxial side; connection to seed swollen; attached to distal part of cotyledonary sheath. Cotyledonary sheath elongate, splitting lengthwise; opposite to hyperphyll. Coleoptile absent. Cataphyll single; elongate. Eophyll entire, linear-lanceolate; apex acute, forming a short needle-like projection; margins spiny. Venation pattern palmate; leaf axis reduced; midvein not distinct from other longitudinal veins; veins convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal and distal marginal folds induplicate. Epidermal cells rectangular, elongate; adaxial and abaxial anticlinal walls linear. Hairs present; multicellular base associated with ribs. Stomata superficial, arranged into regular lines at intercostal regions; short terminal cells overarching guard cells. Hypodermis single-layered, present at adaxial and abaxial sides; fibrous bundles at irregular intervals. Chlorenchyma undifferentiated; spongy mesophyll with more than five layers; fibrous bundles at ridges. Expansion cells double-layered; fibrous bundles scattered; lumen small. Major veins associated with ridges; buttressed to adaxial hypodermis; IS sclerotic, multilayered; OS distinct. Median veins free and equidistant. Minor veins equidistant; OS surrounding vascular bundle. Midrib not prominent, vascular bundle single. Marginal rib with fibrous layers. Petiole transverse section crescent-shaped. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles; raphids equidistant; tannins abundant, in full sacs.

Corypha sp.

Seed remaining above plumular-radicular node. Plumular-radicular axis straight. Primary root straight; stout and persistent; secondary roots branched; shoot-borne roots absent; root hairs present; pneumatophores present. Hyperphyll elongate, grooved all around, swollen as it leaves seed. Cotyledonary sheath splitting lengthwise at side opposite to seed; conspicuous lenticels on cotyledonary sheath and hyperphyll. Coleoptile absent. Cataphyll single; opening apical. Eophyll entire, broadly oblanceolate; praemorse apex (Fig. 3B). Venation pattern costapalmate; leaf axis distinct; midvein distinct; veins diffuse at intercostal areas; transverse commissures abundant, closely arranged, some connect to longitudinal veins. Plication with proximal and distal marginal folds induplicate. Epidermal cells rectangular; adaxial anticlinal walls sinuous; abaxial walls linear. Hairs present; few basal cells associated with ribs. Stomata slightly sunken; short terminal cells overarching guard cells; lateral subsidiary cells ellipsoid. Hypodermal fibrous layer continuous. Chlorenchyma undifferentiated; spongy mesophyll with more than five layers. Expansion cells double-layered; fibrous bundles scattered; lumen small. Major veins not associated with ridges, attached to adaxial and abaxial hypodermal layers; IS sclerotic, multilayered; OS distinct. Median veins free; oriented toward adaxial side, occasionally connected to transverse commissures. Minor veins buttressed to adaxial side; OS restricted to abaxial end, u-shaped. Midrib abaxially prominent; single vascular bundle. Marginal rib with minor vascular bundle. Petiole transverse section crescent-shaped. Phloem strands one. Metaxylem vessels single; large. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular and fibrous bundles; raphids, abundant, equidistant; tannins scarce.

Gatin (1906a) has described germination of Corypha.

Cryosophila grayumii R. Evans

Seed remaining above plumular-radicular node. Plumular-radicular axis symmetric, straight. Primary root persistent; secondary roots branched; shoot-borne roots absent; root hairs absent. Hyperphyll moderate; single groove along adaxial side. Cotyledonary sheath single, longitudinally grooved; opening lengthwise, opposite to seed. Coleoptile absent. Cataphyll single; opening apical. Eophyll entire, broad, lanceolate; apex acute with a needle-like extension; about six or eight similar leaves borne before the segmented leaf; splitting side adaxial. Venation pattern palmate; leaf axis reduced; midvein not distinct from other longitudinal veins; veins convergent at apex; transverse commissures widely separated from each other, connecting some longitudinal veins and ending at intercostal areas. Plication with proximal and distal marginal folds induplicate. Epidermal cells rectangular, covered by thick cuticle; adaxial and abaxial anticlinal walls linear. Hairs free; few basal cells. Stomata slightly sunken, scattered; short and elongate terminal cells overarching guard cells. Hypodermal colorless layer absent, replaced by a layer of scattered subepidermal fibers. Chlorenchyma undifferentiated; spongy mesophyll with less than five layers; fibrous bundles restricted to ridges and grooves; lumen wide. Expansion cells single-layered, ellipsoid, perpendicular; adjacent epidermal cells papillose. Major veins not associated with ridges; OS distinct. Median veins buttressed to adaxial side. Minor veins free; oriented toward abaxial side; OS cap-shaped. Midrib abaxially prominent; vascular bundle single, simple. Marginal rib with fibrous layers. Petiole transverse section crescent-shaped. Phloem strands single. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular and fibrous bundles; raphids abundant, equidistant; tannins abundant.

Itaya amicorum H. E. Moore

Seed remaining above plumular-radicular node. Plumular-radicular axis symmetric, straight. Primary root straight and persistent; secondary roots simple; pneumatophores present; shoot-borne roots absent; collar roots present. Hyperphyll extremely elongate; single groove on adaxial side; connection to seed flat. Cotyledonary sheath leathery; opens longitudinally, opposite to hyperphyll. Coleoptile absent. Cataphyll single; open lengthwise. Eophyll entire, broad; apex acute with a needle-like extension; later apex splitting into irregular segments. Venation pattern palmate; leaf axis reduced; midvein inconspicuous; veins convergent at apex; transverse commissures abundant, closely arranged, connecting longitudinal or intercostal areas. Plication with proximal and distal marginal folds induplicate. Epidermal cells rectangular; adaxial and abaxial anticlinal walls sinuous; cuticle thick. Hairs present; basal cells few, not associated with ribs. Stomata slightly sunken, scattered at intercostal regions; short terminal cells overarching guard cells. Hypodermal colorless cells absent, replaced by a continuous fibrous layer; lumen small. Chlorenchyma undifferentiated; spongy mesophyll with less than five layers. Expansion cells single-layered; adjacent epidermal cells papillose. Major veins not associated with ridges, attached to abaxial epidermal layers; IS multilayered, sclerotic; OS distinct. Median veins free, toward abaxial side. Minor veins oriented toward abaxial side; OS surrounding vascular bundle. Midrib abaxially prominent; single multivascular bundle. Marginal rib with fibrous layers. Petiole not distinct. Phloem strands two. Metaxylem vessels two. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose distributed around vascular bundles.

Livistona chinensis R. Br.

Seed remaining above plumular-radicular node. Plumular-radicular axis straight and symmetric. Primary root straight and persistent; disk collar not distinct; secondary roots simple; shoot-borne roots present; collar roots present. Hyperphyll elongate, longitudinally grooved; connection to seed swollen. Cotyledonary sheath grooved. Coleoptile very short; apical opening. Cataphyll single; apical opening with a needlelike projection. Eophyll entire, broadly lanceolate; apex acute; three leaves similar to eophyll before first split leaf appears; splitting side adaxial. Venation pattern costapalmate; leaf axis distinct; midvein distinct; veins convergent at apex; transverse commissures widely separated from each other, connecting some longitudinal veins and intercostal areas. Plication with proximal and distal marginal folds induplicate. Epidermal cells rectangular; adaxial and abaxial anticlinal walls sinuous. Hairs free; basal cells few. Stomata superficial; short terminal cells overarching guard cells; two sets of lateral subsidiary cells. Hypodermis single-layered, present at adaxial and abaxial sides; fibrous bundles at irregular intervals; lumen small. Chlorenchyma undifferentiated; spongy mesophyll with less than five layers; mesophyll fibers in scattered bundles, equidistant from surface layer or toward abaxial side. Expansion cells double-layered; rounded cells; fibrous bundles scattered; adjacent epidermal cells papillose. Major veins associated with ridges, buttressed to adaxial hypodermis; IS multilayered, sclerotic; OS distinct. Median veins free and equidistant. Minor veins equidistant; OS surrounding vascular bundle. Midrib abaxially prominent; single vascular bundle. Marginal rib with fibrous layers. Petiole transverse section crescent-shaped. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose; distributed around vascular bundles and fibrous bundles; raphids equidistant.

Gatin (1906a), Zurawaska (1912), Mahabale and Kulkarni (1972), and Lothian (1959) have described germination of Livistona.

Nannorrhops ritchiana (Griff.) Aitchson

Seed remaining above plumular-radicular node. Plumular-radicular axis symmetric. Primary root stout, persistent, extremely long; secondary roots simple; shoot-borne roots present. Hyperphyll very short (ca. 1 cm. long), smooth; connection to seed flat. Cotyledonary sheath not distinct; hyperphyll at base of coleoptile. Coleoptile elongate; apical opening. Cataphyll single; apex acute; apical opening; some cataphylls with coleoptile-like extensions. Eophyll entire, linear-lanceolate; apex acute with a needle-like extension (Fig. 3C). Venation pattern palmate; leaf axis reduced; midvein not distinct from other longitudinal veins; veins convergent at apex; transverse commissures abundant, closely arranged, connected to longitudinal veins. Plication with proximal and distal marginal folds induplicate. Epidermal cells rectangular, elongate; adaxial and abaxial anticlinal walls linear. Hairs present; basal cells few. Stomata slightly sunken; scattered at intercostal regions; short terminal cells overarching guard cells. Hypodermis single-layered, rounded or ellipsoid cells; orientation parallel; present at adaxial and abaxial sides. Chlorenchyma well differentiated; palisade with two layers; spongy mesophyll with more than five layers. Expansion cells single-layered; slightly larger than epidermal cells. Major veins not associated with ridges, attached to adaxial and abaxial hypodermis; IS multilayered, sclerotic; OS distinct. Median veins buttressed to adaxial hypodermis. Minor veins equidistant; OS surrounding vascular bundle; radial OS present. Midrib not prominent, vascularized by grouped simple vascular bundles. Marginal rib with major vascular bundle. Petiole transverse section crescent-shaped. Phloem strands three. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles; raphids abundant, equidistant; tannins abundant in all cells, more concentrated in palisade cells.

Pritchardia remota (Kuntze) Becc.

Seed remaining above plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root persistent; secondary roots simple; shoot-borne roots present, protruding through cotyledonary sheath. Hyperphyll very short, smooth; connection to seed constricted. Cotyledonary sheath not distinct. Coleoptile leathery; opening apical; short slits opposite to hyperphyll. Cataphylls three, opening lengthwise; needlelike projection at apex; second and third cataphylls bifid. Eophyll entire, broadly lanceolate; apex hairy, acute. Venation pattern costapalmate; leaf axis distinct; midvein not distinct from other longitudinal veins; veins convergent at apex; transverse commissures widely separated from each other, connecting some longitudinal veins and intercostal areas. Plication with proximal and distal marginal folds induplicate. Epidermal cells rectangular; adaxial and abaxial anticlinal walls sinuous. Hairs present; few basal cells, associated with ribs. Stomata superficial, scattered at intercostal regions; short terminal cells overarching guard cells. Hypodermis single-layered, present at adaxial and abaxial sides; fibers at regular intervals; lumen wide. Chlorenchyma well differentiated; palisade single-layered; spongy mesophyll with more than five layers. Expansion cells double-layered; scattered fibrous bundles. Major veins associated with ridges, attached to adaxial hypodermis and abaxial expansion cells; IS multilayered, sclerotic; OS distinct. Median veins free and equidistant. Minor veins equidistant; IS multilayered, sclerotic; OS surrounding vascular bundle; radial OS present. Midrib abaxially prominent; single vascular bundle. Marginal rib with fibrous layers. Petiole transverse section crescent-shaped. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular and fibrous bundles; raphids abundant, equidistant; tannins abundant.

Rhapidophyllum hystrix (Pursh) H. Wendl. & Drude

Seed remaining above plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root persistent; collar disk distinct; secondary roots simple; shoot-borne roots protruding through cotyledonary sheath. Hyperphyll moderate, smooth; connection to seed constricted, attaching at midpoint between cotyledonary sheath and coleoptile. Cotyledonary sheath leathery. Coleoptile elongate, slightly grooved; apical opening, a short slit eventually appearing, opposite to hyperphyll. Cataphyll single, slightly furrowed; opening lengthwise; apex pointed. Eophyll entire, broadly lanceolate; apex acute (Fig. 3D). Venation pattern palmate; leaf axis reduced; midvein not distinct from other longitudinal veins; veins convergent at apex; transverse commissures widely separated from each other, connecting some longitudinal veins and intercostal regions. Plication with proximal and distal marginal folds induplicate. Epidermal cells rectangular; adaxial and abaxial anticlinal walls linear. Hairs present; few basal cells, associated with ribs. Stomata slightly sunken, scattered at intercostal regions; short terminal cells overarching guard cells. Hypodermis single-layered, at adaxial and abaxial sides; fibrous bundles at irregular intervals; lumen wide. Chlorenchyma undifferentiated; spongy mesophyll with more than five layers. Expansion cells double-layered; scattered fibrous bundles; adjacent epidermis papillose. Major veins associated with ridges, attached to adaxial hypodermis and abaxial expansion cells; IS multilayered, sclerotic; OS not well differentiated. Median veins free and equidistant. Minor veins equidistant; OS surrounding vascular bundle. Midrib abaxially prominent; single simple vascular bundle (Fig. 5A). Marginal rib with fibrous bundles. Petiole transverse section crescent-shaped. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, large, margins spinulose distributed around vascular bundles; raphids present; tannins abundant.

[FIGURE 5 OMITTED]

Clancy and Sullivan (1988) and Carpenter et al. (1993) have described germination in Rhapidophyllum.

Sabal minor (Jacq.) Persoon

Seed remaining above plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root persistent; secondary roots branched; shoot-borne roots absent. Hyperphyll elongate, connecting to lower part of coleoptile, forming saxophone-shaped projection below the primary root and plumule connection. Cotyledonary sheath short or absent. Coleoptile with basal part curved; leathery texture; opening apical. Cataphyll single; apex leathery. Eophyll entire, lanceolate; apex acute; nine leaves similar to eophyll appearing before first segmented leaf; splitting occurs at grooves on adaxial side. Venation pattern palmate; leaf axis reduced; midvein not distinct from other longitudinal veins; veins convergent at apex; transverse commissures widely separated from each other, connecting some longitudinal veins and intercostal areas. Plication with proximal and distal marginal folds induplicate. Epidermal cells rectangular; adaxial and abaxial anticlinal walls linear. Hairs present; few basal cells. Stomata slightly sunken, scattered; short terminal cells overarching guard cells. Hypodermis single-layered, present at adaxial and abaxial side. Chlorenchyma undifferentiated; spongy mesophyll with more than five layers; fibers in bundles at ridges and grooves; lumen wide. Expansion cells double-layered. Major veins not associated with ridges, attached to adaxial and abaxial epidermal layers; IS multilayered, sclerotic IS; OS distinct. Median veins buttressed to abaxial and adaxial sides. Minor veins buttressed to adaxial side; OS u-shaped. Midrib not prominent, squared; vascular bundle single (Fig. 5B). Marginal rib with minor vascular bundles and fibrous bundles. Petiole transverse section crescent-shaped. Phloem strands three. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular and fibrous bundles; raphids scarce, equidistant; tannins scarce.

Germination of Sabal has been described by Holm (1891) and Gatin (1906a).

Serenoa repens (Bartram) Small

Seed remaining above plumular-radicular node. Plumular-radicular axis symmetric, straight. Primary root stout and persistent; secondary roots simple, extremely elongate; shoot-borne roots present. Hyperphyll moderate, smooth; as in Sabal connecting almost directly to primary root; attachment flat. Cotyledonary sheath short; hyperphyll attached to lower part. Coleoptile apical opening with two opposite slits. Cataphyll single; opening lengthwise; apex with asymmetric split. Eophyll entire, lanceolate; seven leaves similar to eophyll appearing before first segmented leaf; splitting side adaxial. Venation pattern palmate; leaf axis reduced; midvein not distinct from other longitudinal veins; veins convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal and distal marginal folds induplicate. Epidermal cells rectangular; adaxial and abaxial anticlinal walls linear; cuticle thick and forming a continuous layer. Hairs present; few basal cells, associated with ribs. Stomata superficial, scattered at intercostal regions; short terminal cells overarching guard cells. Hypodermis single-layered, at adaxial and abaxial sides; fibers at irregular intervals. Palisade layer distinct; spongy mesophyll with more than five layers; fibrous bundles restricted to grooves; lumen wide. Expansion cells double-layered, ellipsoid, transversally arranged. Major veins associated with ridges, attached to adaxial hypodermis and abaxial expansion cells; IS multilayered, sclerotic; OS distinct. Median veins free, oriented toward abaxial side. Minor veins equidistant; OS surrounding vascular bundle. Midrib not prominent; single bundle. Marginal rib with fibrous bundle. Petiole transverse section crescent-shaped. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles; tannins abundant, scattered, mostly concentrated on palisade parenchyma.

Fisher and Tomlinson (1973) and Hilmon (1968) have described germination of Serenoa.

Thrinax excelsa Lodd. ex Griseb.

Seed remaining above plumular-radicular node. Plumular-radicular axis symmetric, straight. Primary root persistent; secondary roots simple. Hyperphyll moderate; single adaxial groove; connection to seed constricted. Cotyledonary sheath opening with a small slit at distal end. Coleoptile absent. Cataphyll single; opening apical; apex acute. Eophyll entire, linear-lanceolate; apex acute with needle-like projection; five leaves similar to eophyll before first split leaf appears; splitting side adaxial (Fig. 6A). Venation pattern palmate; leaf axis reduced; midvein present along blade; veins convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal and distal marginal folds induplicate. Epidermal cells rectangular to slightly rhombohedral; adaxial and abaxial anticlinal walls linear. Hairs present; few basal cells. Stomata superficial, arranged in single files at intercostal regions; elongate terminal cells overarching guard cells. Hypodermis single-layered, present at adaxial and abaxial sides; fibrous bundles at regular intervals; lumen wide. Chlorenchyma palisade layer distinct; spongy mesophyll with less than five layers. Expansion cells double-layered; short perpendicular rectangular cells; scattered fibrous bundles. Major veins not associated with ridges; IS multilayered, sclerotic; OS distinct. Median veins not distinct from major veins. Minor veins equidistant; OS surrounding vascular bundle. Midrib not prominent; simple vascular bundle. Marginal rib with fibrous layers. Petiole transverse section crescent-shaped. Phloem strands single. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles; raphids abundant, equidistant.

[FIGURE 6 OMITTED]

Trachycarpus sp.

Seed remaining above plumular-radicular node. Plumular-radicular axis symmetric, straight. Primary root stout and persistent; secondary roots simple. Hyperphyll moderate, covered by tegument; connection to seed flat. Cotyledonary sheath grooved, leathery; opening lengthwise down to node. Coleoptile absent. Cataphyll single; apex acute. Eophyll entire, lanceolate; apex acute with needle-like projection. Venation pattern costapalmate; leaf axis distinct; midvein not distinct from other longitudinal veins; veins convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal and distal marginal folds induplicate. Epidermal cells rectangular or slightly rhombohedral; adaxial and abaxial anticlinal walls linear. Hairs multicellular; few basal cells. Stomata slightly sunken, scattered at intercostal regions; elongate terminal cells overarching guard cells. Hypodermis single-layered, at adaxial and abaxial sides, occasionally interrupted by fibrous bundles. Chlorenchyma undifferentiated; spongy mesophyll with less than five layers; fibers in bundles restricted to ridges and grooves; lumen wide (Fig. 4B). Expansion cells double-layered. Major veins not associated with ridges, attached to adaxial epidermis and abaxial hypodermis; IS multilayered, sclerotic; OS distinct. Median veins buttressed to both abaxial and adaxial sides. Minor veins equidistant; OS surrounding vascular bundle. Midrib abaxially prominent, rounded; single multivascular bundle. Marginal rib with fibrous layers. Petiole transverse section crescent-shaped. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles; raphids present; tannins in full sacs scattered.

Gatin (1906a) and Depoux (1968, 1969) have described germination of Trachycarpus.

Trithrinax brasiliensis Mart.

Seed remaining above plumular-radicular node. Plumular-radicular axis symmetric, straight. Primary root stout and persistent; collar disk swollen; secondary roots branched; shoot-borne roots present; collar roots present. Hyperphyll elongate; single groove on adaxial side; connection to seed flat. Cotyledonary sheath opening lengthwise to node; splitting opposite to hyperphyll. Coleoptile undeveloped. Cataphyll single, plicate; apex acute. Eophyll entire, lanceolate; apex concave with sharp acuminate projection. Venation pattern palmate; leaf axis reduced; veins convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal and distal marginal folds induplicate. Epidermal cells rectangular; adaxial and abaxial anticlinal walls linear. Hairs absent. Stomata slightly sunken, scattered at intercostal regions; short and elongate terminal cells overarching guard cells. Hypodermis not distinct. Chlorenchyma undifferentiated; spongy mesophyll with more than five layers; fibers in bundles restricted to ridges and grooves; lumen wide. Expansion cells single-layered, slightly larger than epidermal cells, rounded, more conspicuous on distal portion of lamina. Major veins not associated with ridges, attached to adaxial and abaxial epidermal layers; IS multilayered, sclerotic; OS distinct. Median veins buttressed to both abaxial and adaxial sides. Minor veins abaxially buttressed; OS cap-shaped. Midrib not prominent; vascular bundle single, simple (Fig. 5C). Marginal rib with fibrous layers. Petiole transverse section crescent-shaped. Phloem strands single. Metaxylem vessels two. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles; raphids equidistant; tannins scarce.

Washingtonia filifera (Linden) H. Wendl.

Seed remaining above plumular-radicular node. Plumular-radicular axis symmetric, straight. Primary root stout and persistent; distinct swollen collar; secondary roots simple; shoot-borne roots; collar roots present; root hairs present. Hyperphyll short, smooth, attached to lower part of coleoptile; connection to seed flat. Cotyledonary sheath not distinct. Coleoptile opening apical. Cataphyll single; apical opening; apex acute. Eophyll entire, linear-lanceolate; apex acute with needle-like projection; nine or ten eophyll-like leaves borne before first split leaf; splitting along adaxial side. Venation pattern palmate; leaf axis reduced; midvein not distinct from other longitudinal veins; veins convergent at apex; transverse commissures abundant, closely arranged, connected to longitudinal veins. Plication with proximal and distal marginal folds induplicate. Epidermal cells rectangular; adaxial and abaxial anticlinal walls linear. Hairs absent. Stomata slightly sunken, scattered al intercostal regions; short terminal cells overarching guard cells. Hypodermis single-layered, present at adaxial and abaxial sides. Chlorenchyma undifferentiated; spongy mesophyll with more than five layers. Nonvascular fibers in layers, restricted to grooves; lumen wide. xpansion cells double-layered, large, ellipsoid, perpendicular. Major veins associated with ridges, attached to adaxial hypodermis and abaxial expansion cells; IS multilayered, sclerotic; OS distinct. Median veins free, oriented toward abaxial side. Minor veins equidistant; OS surrounding vascular bundle. Midrib not prominent, vascularized by grouped simple vascular bundles (Fig. 5D). Marginal rib with small fibrous bundle. Petiole transverse section crescent-shaped. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles; raphids equidistant; tannins abundant.

Gatin (1906a), Ginieis (1952b), and De Mason (1988) have described germination of Washingtonia.

SUMMARY FOR CORYPHEAE

Plumular-radicular axis straight; primary root straight and persistent; collar roots developed or not developed; hyperphyll elongate or contracted; cotyledonary sheath opening laterally or apically; coleoptile present or absent; cataphyll single in most taxa (three in Pritchardia); eophyll entire, lanceolate or linear-lanceolate, acute or lobed at apex; induplicate folding; venation palmate or costapalmate without obvious midvein. In Chamaerops, Livistona, and Trachycarpus, the plumular-radicular axis is straight and oblique to the axis of the embryo; the cotyledonary sheath opens apically; and a coleoptile is absent. Some genera have horizontally oriented eophylls with elongate petioles, for example, Cryosophila and Livistona. Other genera have erect eophylls with short petioles, as in Trithrinax, Chamaerops, Thrinax, Coccothrinax, Sabal, Acoelorraphe, and Trachycarpus.

2. Phoeniceae

Phoenix roebelinii O'Brien

Seed remaining above plumular-radicular node. Plumular-radicular axis symmetric, straight. Primary root persistent; secondary roots simple; collar roots present. Hyperphyll short; single groove at adaxial side; connection to seed flat. Cotyledonary sheath elongate, smooth; apical opening by a short slit; apex concave. Coleoptile absent. Cataphyll single. Eophyll entire, linear-lanceolate; apex with needle-like projection; seven eophyll-like leaves borne before split leaf appears; splitting at adaxial side, starting at proximal end. Venation pattern costapalmate; leaf axis distinct; midvein not distinct from other longitudinal veins; veins convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal and distal marginal folds induplicate. Epidermal cells rectangular; adaxial and abaxial anticlinal walls linear; wall thickened; cuticle thick. Hairs absent. Stomata slightly sunken; short terminal cells not overarching guard cells. Hypodermis single-layered, at adaxial and abaxial sides; cells rounded; large solid fibrous bundles at irregular intervals. Chlorenehyma undifferentiated; spongy mesophyll with more than five layers; fibers in bundles at adaxial and abaxial sides; lumen small. Expansion cells double-layered; epidermal cells papillose; scattered fibrous bundles. Major veins not associated with ridges, attached to adaxial and abaxial hypodermis; IS multilayered and sclerotic; OS distinct. Median veins free and equidistant. Minor veins equidistant; OS surrounding vascular bundle. Midrib not prominent; simple vascular bundle not centered and adjacent to fold. Marginal rib with fibrous bundle. Petiole transverse section crescent-shaped-shaped. Phloem strands one. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles; raphids present; tannins abundant, present in all mesophyll.

Germination of Phoenix has been described many times, beginning with the illustration of Camerarius (1588). Other descriptions are those of Sachs (1862), Gatin (1906a), and Ginieis (1951, 1957).

SUMMARY FOR PHOENICEAE

Plumular-radicular axis symmetric, straight; primary root persistent; hyperphyll short, single groove at adaxial side; cotyledonary sheath present, elongate, split lengthwise; coleoptile not developed; cataphyll single; eophyll entire, linear-lanceolate, apex with needle-like projection; costapalmate, induplicate.

3. Borasseae

Borassus sp.

Seed remaining above plumular-radicular node. Plumular-radicular axis symmetric, straight. Primary root persistent; secondary roots simple; pneumatophores at the base; collar roots borne on surface of flat collar disk. Hyperphyll smooth and covered by lenticels. Cotyledonary sheath covered by lenticels. Coleoptile absent. Cataphyll single, thick, robust; apical opening; apex curved inwards. Eophyll entire, broadly lanceolate; apex crenulated. Venation pattern costapalmate; leaf axis distinct; midvein not distinct from other longitudinal veins; veins divergent at apex; transverse commissures abundant, closely arranged, connected to longitudinal veins. Plication with proximal and distal marginal folds induplicate. Epidermal cells rectangular or rhombohedral; adaxial and abaxial anticlinal walls linear; cells adjacent to ridges with thickened walls (Fig. 7A). Hairs absent. Stomata sunken; short terminal cells overarching guard cells; two lateral subsidiary cells at each side of guard cells; both sets attached to the same terminal cells. Hypodermis single-layered; cells large, rounded; at adaxial and abaxial sides; fibrous layers or bundles at irregular intervals. Chlorenehyma well differentiated; palisade with two layers; spongy mesophyll with more than five layers; fibers in bundles at adaxial and abaxial sides; lumen wide. Expansion cells double-layered, rectangular or ellipsoid, perpendicular; adjacent epidermal cells papillose, with scattered fibrous bundles. Major veins not associated with ridges, attached to both hypodermal layers; IS multilayered; OS distinct. Median veins buttressed to adaxial side. Minor veins buttressed to adaxial side; OS u-shaped; some vascular bundles present at grooves. Midrib prominent, squared; vascular bundle single (Fig. 5E). Marginal rib with minor veins and fibrous layers. Petiole transverse section crescent-shaped. Phloem strands three, one large central flanked by two smaller strands. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles; raphids equidistant; tannins abundant, some in full sacs.

[FIGURE 7 OMITTED]

Gatin (1906a) and Dassanayake and Sivakadachchan (1973) have described germination of Borassus.

Hyphaene coriacea Gaertn.

Seed remaining above plumular-radicular node. Plumular-radicular axis symmetric, straight. Primary root persistent; secondary roots with distinct pneumatophores at base; shoot-borne roots absent. Hyperphyll elongate, grooved all around; connection point to seed slightly swollen. Cotyledonary sheath thick; apical opening; opposite short split; abundant lenticels covering hyperphyll and cotyledonary sheath. Coleoptile absent. Cataphyll single, plicate, curved inwards. Eophyll entire, linear-lanceolate; apex nearly closed and pointed. Venation pattern costapalmate; leaf axis distinct; midvein distinct; veins convergent at apex; transverse commissures widely separated from each other, connecting some longitudinal veins and intercostal areas. Plication with proximal and distal marginal folds induplicate (Fig. 8A). Epidermal cells rectangular or rhombohedral; adaxial and abaxial anticlinal walls linear; cuticle layer thick and uniform. Hairs multicellular, with multicellular base. Stomata sunken, scattered at intercostal regions; guard cells with distinct ledges; large substomatal chamber; in surface view short terminal cells overarching guard cells; two lateral subsidiary cells on each side of guard cells. Hypodermis double-layered, at adaxial and abaxial sides. Chlorenchyma well differentiated; palisade layer distinct; spongy mesophyll with more than five layers; fibers in solid rounded and ellipsoid bundles of approximately 14 strands, distributed at adaxial and abaxial sides; lumen wide. Expansion cells double-layered; adjacent epidermis papillose; scattered fibrous bundles. Major veins not associated with ridges; OS distinct. Median veins buttressed to both abaxial and adaxial sides. Minor veins buttressed at adaxial side and abaxial sides; OS u-shaped or cap-shaped; some vascular bundles present at grooves. Midrib prominent at adaxial and abaxial sides, vascularized by grouped simple vascular bundles. Marginal rib with minor veins and fibrous layers. Petiole transverse section crescent-shaped. Phloem strands three; large central flanked by two smaller strands. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles; raphids abundant, equidistant.

[FIGURE 8 OMITTED]

Latania loddegesii Mart.

Seed remaining above plumular-radicular node. Plumular-radicular axis symmetric, straight. Primary root persistent; secondary roots simple; pneumatophores at the base; root hairs present. Hyperphyll elongate, grooved all around; attachment to seed swollen. Cotyledonary sheath opening lengthwise; lenticels abundant, covering hyperphyll, cotyledonary sheath, and primary root. Coleoptile absent. Cataphyll single, plicate; apex acute. Eophyll palmate; linear segments; spiny margins; splitting side adaxial. Venation pattern palmate; leaf axis reduced; midvein present in each segment; veins convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal and distal marginal folds induplicate. Epidermal cells rectangular or rhombohedral; adaxial and abaxial anticlinal walls linear. Hairs present; multicellular base. Stomata sunken; guard cells with conspicuous ledges; short terminal cells overarching guard cells; two lateral subsidiary cells on each side of guard cells. Hypodermis single-layered, ellipsoid cells; orientation parallel; present at adaxial and abaxial sides. Chlorenchyma well differentiated; palisade with two to three layers; spongy mesophyll with more than five layers, about 12 layers on wider portion; fibers restricted to mesophyll, arranged in solid rounded bundles, at adaxial and abaxial sides; lumen wide. Expansion cells double-layered; adjacent epidermis papillose, with scattered fibrous bundles. Major veins not associated with ridges, buttressed to adaxial and abaxial hypodermal layers; IS multilayered, sclerotic, and fibrous; OS distinct. Median veins buttressed to adaxial side; IS multilayered, sclerotic, and fibrous. Minor veins buttressed to adaxial side, some associated with grooves; OS ushaped. Midrib prominent at adaxial and abaxial sides, vascularized by grouped simple vascular bundles. Marginal rib with fibrous bundle. Petiole transverse section crescent-shaped. Phloem strands three; large central flanked by two smaller. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles; raphids abundant, equidistant; tannins scarce.

Gatin (1906a) has described germination of Latania.

SUMMARY FOR BORASSEAE

Plumular-radicular axis straight; primary root straight and persistent; collar roots developed or not developed; hyperphyll elongate; cotyledonary sheath opening laterally; coleoptile absent; cataphyll single; eophyll entire or palmate; apex acute; venation palmate; marginal plication induplicate. Some Corypheae (Livistona, Trachycarpus, Chamaerops) and all Phoeniceae, all Borasseae, and all Caryoteae have similar germination patterns.

II. CALAMOIDEAE

1. Calameae

Calamus flagellum Griff.

Seed attached above plumular-radicular node. Plumular-radicular axis asymmetric, forming an angle. Primary root straight, vertical, persistent; collar disk distinct and swollen; secondary roots branched abundant; collar roots and root hairs absent. Hyperphyll inconspicuous; attachment surface flat; attached to base of coleoptile. Cotyledonary sheath not well developed. Coleoptile present, splitting longitudinally at opposite side of seed; texture rugulose. Cataphylls two; opening lengthwise; apex acute. Eophyll bifid; splitting along grooved side; segments sigmoid; petiole and leaf margins spiny (Fig. 6B, 6C). Venation pattern pinnate; leaf axis distinct; midvein not distinct from other longitudinal veins; veins convergent at apex; transverse commissures simple and bifurcate, connected mainly to secondary veins, abundant, and closely arranged. Plication with proximal marginal folds reduplicate; distal outer marginal fold reduplicate; inner fold induplicate; plication slightly sinuous (Fig. 8B). Epidermal cells rectangular; anticlinal walls dentate; intercostal cells slightly narrower and more elongate than costal cells. Hairs present; basal cells few. Stomata superficial, arranged in regular files at intercostal regions on both surfaces, more abundant in abaxial surface; terminal cells short and overarching guard cells. Hypodermal layer occupied by fibrous bundles arranged at irregular intervals, forming an almost continuous fibrous layer at adaxial side. Chlorenchyma not differentiated; spongy mesophyll with large cells, arranged in less than five layers; fibers arranged in large solid bundles of more than ten strands, these oriented toward abaxial side; fiber lumen small. Expansion cells arranged in single or double layers; cells elongate, ellipsoid, and rectangular, restricted to lateral flanks of major veins; fibrous bundles at outermost sides. Major veins associated with ridges, rounded and prominent; fibrous buttresses at adaxial side, attached to hypodermal layer; OS distinct at lateral sides of vascular bundle. Median veins free, oriented toward abaxial side; IS multilayered and sclerotic; OS lateral; adaxial fibrous buttress attached to hypodermal layer. Minor veins free, oriented toward abaxial side, not associated with grooves on abaxial folds; IS single-layered, surrounding vascular bundle; OS restricted to the adaxial end, forming a cap-shaped layer. Midrib abaxially prominent, distinct only at proximal section; adaxial side irregular; abaxial side rounded; highly vascularized by simple vascular bundles. Marginal rib occupied by subepidermal fibrous layers. Petiole rounded abaxially. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular and fibrous bundles; tannins scattered.

Germination of Calamus has been described by Gatin (1906a), Ginieis (1965), and Ilangovan and Padmanabhan (1993).

Pigafetta filaris (Gis.) Becc.

Seed remaining above plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root oblique, persistent; disk collar distinct and swollen, not as prominent as in Calamus; secondary roots branched; first internode elongate with shoot-borne roots; collar roots and root hairs absent. Hyperphyll inconspicuous. Cotyledonary sheath absent. Coleoptile distinct, splitting ventrally or laterally; tongue-like projection at apical end. Cataphylls two; apex acute. Eophyll bifid; segments linear-lanceolate; apex acuminate; spiny margins; splitting along grooved side. Venation pattern pinnate, convergent at apex; leaf axis short; midvein not distinct from other longitudinal veins; transverse commissures always connected to longitudinal veins, widely separated from each other, less abundant than in Calamus. Plication with proximal marginal folds reduplicate; distal inner marginal folds induplicate; outer marginal folds reduplicate. Epidermal cells rectangular; adaxial and abaxial anticlinal walls sinuous. Hairs absent. Stomata superficial, abundant on abaxial surface, arranged in regular lines at intercostal regions; elongate terminal cells do not overarch guard cells. Hypodermis an indistinct colorless layer with fibrous bundles at regular intervals; lumen widely open. Chlorenchyma differentiated; palisade layer distinct; spongy mesophyll with less than five layers lacking fibrous bundles. Expansion cells double-layered. Major veins associated with ridges, attached to adaxial and abaxial hypodermal layers; OS distinct. Median veins free, oriented toward abaxial sides. Minor veins not buttressed, oriented toward abaxial side; OS around adaxial end, forming a cap-shaped layer; an extra layer of radially arranged OS is present. Midrib abaxially prominent, vascularized by several simple vascular bundles. Marginal rib with fibrous layers. Petiole transverse section deeply concave. Phloem strands two. Metaxylem vessels single, widely open. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular and fibrous bundles; tannins scattered, appearing to be more concentrated on palisade parenchyma.

Germination of Pigafetta has been described by Davis and Kuswara (1987).

Plectocomia sp.

Seed remaining above plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root persistent for a long period before being replaced by shoot-borne roots; distinct disk collar; secondary roots branched; shoot-borne roots abundant, arising above first node; collar roots and root hairs absent. Hyperphyll inconspicuous. Cotyledonary sheath not evident. Coleoptile distinct apical opening; splitting side opposite to seed. Cataphyll single. Eophyll entire, lanceolate; apex acute. Venation pattern pinnate; leaf axis distinct; midvein not distinct from other longitudinal veins; veins convergent at apex; transverse commissures always connected to longitudinal veins, widely separated from each other. Plication with proximal marginal folds induplicate; distal marginal folds reduplicate and induplicate. Epidermal cells rectangular; dentate walls. Hairs absent. Stomata superficial, arranged in regular rows at intercostal regions; elongate terminal cells overarching guard cells. Hypodermal layer replaced by fibrous bundles at irregular intervals. Chlorenchyma undifferentiated; spongy mesophyll with less than five layers. Expansion cells single- or double-layered. Major veins associated with ridges, prominent at adaxial side, attached to adaxial and abaxial hypodermal layers; OS at lateral sides; cells smaller than surrounding mesophyll cells. Median veins free, oriented toward abaxial side; multilayered sclerotic IS. Minor veins not buttressed, oriented toward abaxial side, sometimes attached to epidermal layer; OS cap-shaped; extra layer of radially arranged OS present. Midrib abaxially prominent, vascularized by several simple vascular bundles (Fig. 5F). Marginal rib with fibrous layers. Petiole transverse section hemi-ellipse. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular and fibrous bundles; tannins abundant, some in scattered full sacs.

SUMMARY OF CALAMEAE

Plumular-radicular axis angular; primary root either straight or oblique, persistent or ephemeral; collar roots developed or not developed; hyperphyll inconspicuous; cotyledonary sheath apical opening; coleoptile present; cataphylls 1-2; acute and hard apex in Plectocomia; eophyll bifid, pinnate, acute, praemorse or lobed at apex; venation pinnate.

Dransfield (1979) has discussed the morphology of Calameae eophylls.

2. Lepidocaryeae

Mauritia flexuosa L. f.

Seeds remaining above plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root persistent; collar disk distinct and swollen; shoot-borne roots abundant, emerging above and below collar node; secondary roots branched; collar roots and root hairs absent. Hyperphyll not distinct. Cotyledonary sheath absent. Coleoptile distinct, shorter than cataphylls; ventral and lateral splitting; conspicuous tongue-like projections at lateral sides. Cataphylls two, opening lengthwise; second cataphyll elongate with a closed sheath; apex acute. Eophyll palmate; segments linear-lanceolate; apex acute; margins spiny; splitting at grooves along abaxial side. Venation pattern palmate; leaf axis inconspicuous; veins radiate from distal end of petiole; midvein distinct in each segment; veins convergent at apex; transverse veins very conspicuous, connected to longitudinal major and minor veins, abundant, closely arranged. Plication with proximal marginal fold induplicate; distal fold reduplicate. Epidermal cells rectangular; dentate lateral walls (Fig. 7B). Hairs present; few basal cells. Stomata superficial, arranged in regular rows at intercostal regions, scarce in adaxial surface; elongate terminal cells overarching guard cells. Hypodermal colorless parenchyma layer replaced by fibrous bundles arranged at irregular intervals. Chlorenchyma differentiated; palisade layer distinct; spongy mesophyll with less than five layers; fibrous bundles oriented toward abaxial side; fiber lumen small. Expansion cells single- or double-layered; scattered fibrous bundles. Major veins associated with ridges, attached to hypodermal layer; IS partially sclerotic; OS distinct, formed by small regular cells. Median veins free, oriented toward abaxial side; multilayered sclerotic IS. Minor veins not buttressed, oriented toward or attached to abaxial side; OS surrounding vascular bundle or restricted to adaxial end; radial OS present. Midrib abaxially prominent, triangular, vascularized by simple vascular bundles. Marginal rib with fibrous layers. Petiole transverse section terete; vascular bundles uniformly arranged in a V-fashion. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular and fibrous bundles; tannins abundant some in full sacs.

SUMMARY FOR LEPIDOCARYEAE

Plumular-radicular axis angular; primary root persistent; hyperphyll not distinct; cotyledonary sheath absent; coleoptile distinct, shorter than cataphylls; cataphylls two; eophyll palmate; plication with proximal marginal fold induplicate, distal fold reduplicate.

III. NYPOIDEAE

Nypa fruticans Wurmb

Seed remaining horizontal to plumular extension. Plumular-radicnlar axis not identifiable; plumular axis emerges horizontally, later diverts with a negative geotropism. Primary root never develops into an external structure; secondary roots simple; shoot-borne roots present; collar roots and root hairs absent. Because of its unique morphology, structures such as hyperphyll and cotyledonary sheath are not easily identified. Coleoptile opening lateral; apex with hook-like extensions. Cataphylls four or more, coriaceous; apex with a hook-like projection; distichous arrangement. Eophyll bifid or pinnate; segments linear-lanceolate; apex acute; splitting along grooved side. Venation pattern pinnate; leaf axis conspicuous; midvein not distinct from other longitudinal veins; veins convergent at apex; transverse commissures always connected to longitudinal veins, widely separated from each other. Plication data unavailable for proximal section; pinnae distal marginal folds induplicate. Epidermal cells rectangular; adaxial anticlinal walls sinuous; thick cuticular layer. Hairs present; basal cells few. Stomata sunken; short terminal cells overarching guard cells. Hypodermal layer present at adaxial and abaxial sides; cells ellipsoid; parallel orientation, occasionally interrupted by sunken stomata cells. Chlorenchyma distinct; palisade layers two, distinct; spongy mesophyll with more than five layers; fibers arranged in solid circular or ellipsoid bundles, made up of more than 15 strands, distributed at adaxial and abaxial sides; lumen wide. Expansion cells double-layered. Major veins not associated with ridges; OS lateral. Median veins free, oriented toward abaxial side. Minor veins equidistant or toward abaxial side; OS surrounding vascular bundle. Midrib prominent at adaxial and abaxial sides with a single multivascular bundle (Fig. 5G). Marginal rib with fibrous bundles. Petiole transverse section ellipsoid; hypodermal layers two; distinct circular intercellular spaces. Phloem strands single. Metaxylem vessels two. Cell inclusions: silica bodies hat-shaped, margins spinulose, distributed around vascular and fibrous bundles; tannins abundant, scattered, full sacs, traces are found within vascular bundles and hypodermal cells.

Germination of Nypa has been described by Tomlinson (1971), Fong (1986), and Bacon (2001).

SUMMARY OF NYPOIDEAE

Plumular axis emerges horizontally; primary root does not emerge from seed; secondary roots simple; shoot-borne roots present; hyperphyll and cotyledonary sheath are not easily identified; coleoptile opening lateral, apex with hook-like extensions; cataphylls four or more, coriaceous, apex with a hook-like projection, distichous arrangement; eophyll bifid or pinnate; segments linear-lanceolate, apex acute; splitting along grooved side.

IV. CEROXYLOIDEAE

1. Cyclospatheae

Pseudophoenix sargentii H. Wendl.

Seed remaining above plumular-radicular node. Plumular-radicular axis symmetric, straight. Primary root persistent; secondary roots simple; pneumatophores at base; root hairs abundant. Hyperphyll moderate; single groove at adaxial side; connection to seed flat. Cotyledonary sheath grooved; splitting lengthwise opposite to hyperphyll. Coleoptile absent. Cataphylls two, plicate; apical opening; apex acute. Eophyll entire, lanceolate; apex acute. Venation pattern pinnate; leaf axis distinct; veins convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal and distal marginal folds reduplicate (Fig. 8C). Epidermal cells rectangular; adaxial and abaxial anticlinal walls linear; regular rounded cells in transverse section; walls thickened; cuticle thick. Hairs absent. Stomata sunken; epidermal layer around stomata complex sunken; short terminal cells overarching guard cells; two sets of lateral subsidiary cells at each side of guard cells. Hypoderreal colorless layer absent, replaced by a continuous fibrous layer; double or triple layers at ridges; lumen wide. Chlorenchyma differentiated; palisade layer distinct; spongy mesophyll with more than five layers. Expansion cells single-layered, elongate, ellipsoid; adjacent epidermal cells papillose. Major veins prominent, associated with ridges; fibrous buttresses at abaxial side, attached to hypodermis; IS multilayered, sclerotic; OS distinct. Median veins buttressed to both abaxial and adaxial sides. Minor veins small, equidistant; OS surrounding vascular bundle. Midrib adaxially prominent; single vascular bundle (Fig. 5H). Marginal rib with minor vein; IS fibrous, solid, multilayered. Petiole transverse section heart-shaped. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles; raphids abundant, equidistant.

Read (1968) has described germination in Pseudophoenix.

SUMMARY FOR CYCLOSPATHEAE

Plumular-radicular axis straight; primary root persistent; secondary roots simple; pneumatophores present; root hairs abundant; hyperphyll moderate size; cotyledonary sheath grooved; coleoptile absent; cataphylls two, plicate, apex acute; eophyll entire, lanceolate, apex acute; venation pattern pinnate; leaf axis distinct; plication with proximal and distal marginal folds reduplicate.

2. Ceroxyleae

Ceroxylon sp.

Seed horizontal, neither above nor below plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root persistent; collar disk distinct; secondary roots simple; shoot-borne roots present; collar roots present. Hyperphyll inconspicuous, about 5-6 mm long; connection to seed flat; connected to base of coleoptile. Cotyledonary sheath absent. Coleoptile short; splitting at ventral and lateral sides; corrugated. Cataphylls two, elongate; apical opening; apex acute. Eophyll entire, broadly lanceolate; apex acute; several leaves similar to eophyll appearing before first split leaf; seventh leaf splitting along grooved side (Fig. 6D). Venation pattern pinnate; leaf axis distinct; midvein not distinct; veins converge gradually to marginal vein; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal marginal folds induplicate, distal marginal folds induplicate and margin reduplicate. Epidermal cells rectangular or rhombohedral; adaxial and abaxial anticlinal walls sinuous; cuticle think; abaxial epidermis sunken around hair base. Hairs multicellular, wart-like, abundant at abaxial surface; base multicellular, associated with ribs (Fig. 4C). Stomata superficial; short terminal cells overarching guard cells. Hypodermal colorless cells absent, replaced by a continuous fibrous layer; lumen small. Chlorenchyma undifferentiated; spongy mesophyll with fewer than five layers. Expansion cells single-layered, rectangular, short, perpendicular; adjacent epidermis papillose. Major veins associated with ridges, attached to adaxial and abaxial epidermal layers; OS distinct. Median veins buttressed to adaxial side. Minor veins oriented toward abaxial side; OS cap-shaped; radially arranged OS present. Midrib abaxially prominent; transverse section ax-shaped; single large and several small vascular bundles grouped. Marginal rib with fibrous layers. Petiole transverse section heart-shaped. Phloem strands single. Metaxylem vessels single or double. Cell inclusions: silica bodies irregular, margins spinulose, distributed around vascular bundles; raphids scarce, equidistant; tannins abundant in full sacs

Karsten (1847) has described and illustrated germination of Ceroxylon.

Oraniopsis appendiculata (F. Bailey) Dransf., Uhl & Irvine

Seed horizontal, neither above nor below plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root persistent; disk collar present, resembling Caryoteae collar disk; secondary roots simple; shoot-borne roots present. Hyperphyll very short, around 4-5 mm long, smooth; connection to seed flat, connecting at base of coleoptile. Cotyledonary sheath inconspicuous. Coleoptile short, leathery, splitting opposite to hyperphyll. Cataphylls two; first cataphyll apical opening; apex acute; second cataphyll elongate, opening laterally; both covered by brownish integument. Eophyll entire, linear-lanceolate; apex acute. Venation pattern pinnate; leaf axis distinct; midvein not distinct from other longitudinal veins; veins convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal marginal folds induplicate; distal marginal folds, one induplicate, the other reduplicate. Epidermal cells rectangular or rhombohedral; adaxial and abaxial anticlinal walls linear; cuticle thick. Hairs multicellular, abundant at abaxial surface, associated with ribs; multicellular base associated with ribs. Stomata superficial, scattered on intercostal regions; short terminal cells overarching guard cells. Hypodermis single-layered, at adaxial and abaxial sides; rounded or ellipsoid cells in transverse section; interrupted by small fibrous bundles at irregular intervals, more concentrated at ridges and grooves. Chlorenchyma undifferentiated; spongy mesophyll up to five layers; fibrous bundles at adaxial and abaxial sides; lumen wide. Expansion cells single-layered, large, rectangular, perpendicular; adjacent epidermal cells papillose. Major veins associated with ridges, buttressed to adaxial hypodermis; IS multilayered, sclerotic; OS distinct at lateral sides of vascular bundle. Median veins buttressed to adaxial side. Minor veins oriented toward abaxial side; OS cap-shaped; radially arranged OS present. Midrib abaxially prominent, triangular, with simple and multivascular vascular bundles grouped. Marginal rib with fibrous layers. Petiole transverse section rounded in shape. Phloem strands one. Metaxylem vessels single or double. Cell inclusions: silica bodies irregular, margins spinulose, distributed around fibrous bundle or vascular bundle.

Ravenea rivularis Jum. & H. Perrier

Seed above plumular-radicular node. Plumular-radicular axis symmetric, straight. Primary root persistent; distinct swollen disk collar; secondary roots branched; shoot-borne roots present; root hairs abundant. Hyperphyll short, smooth, connection to seed flat. Cotyledonary sheath inconspicuous. Coleoptile short, rugulose, splitting at opposite sides. Cataphylls two; apical opening. Eophyll bifid; apex acute; splitting side abaxial. Venation pattern pinnate; leaf axis distinct; midvein present; veins convergent at apex; transverse commissures widely separated from each other, connecting to some longitudinal veins and ending at intercostal areas. Plication with proximal and distal marginal fold reduplicate on one side, induplicate on the other. Epidermal cells rectangular or rhombohedral; adaxial and abaxial anticlinal walls linear; uniform rounded cells in transverse section. Hairs present; multicellular base. Stomata superficial; short terminal cells overarching guard cells. Hypodermis single-layered, present at adaxial and abaxial sides, replaced by small fibrous bundles at regular intervals; lumen small. Chlorenchyma undifferentiated; spongy mesophyll with fewer than five layers. Expansion cells single-layered, rectangular, perpendicular; adjacent epidermal cells papillose. Major veins associated with ridges, buttressed to adaxial hypodermis by multilayered fibrous IS; abaxial side attached to expansion layer; OS distinct. Median veins free, equidistant. Minor veins equidistant; OS surrounding vascular bundle. Midrib abaxially prominent; simple or multivascular bundles grouped. Marginal rib with fibrous layers. Petiole transverse section semi-ellipsoid; adaxial side ridges. Phloem strands one. Metaxylem vessels single or double. Cell inclusions: silica bodies shape irregular, margins spinulose, distributed around vascular bundles; raphids present.

SUMMARY FOR CEROXYLEAE

Plumular-radicular axis straight or angular; primary root straight and persistent; collar roots developed or not developed; hyperphyll elongate or contracted; cotyledonary sheath opening laterally or apically; coleoptile present; cataphylls two; eophyll entire, bifid or pinnate; apex acute; venation palmate or pinnate; proximal marginal folding induplicate; distal folding induplicate.

3. Hyophorbeae

Chamaedorea microspadix Burret

Seed attachment displaced below plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root persistent; distinct disk collar; secondary roots simple; shoot-borne roots present; root hairs abundant. Hyperphyll undeveloped. Cotyledonary sheath absent. Coleoptile short, splitting lengthwise opposite to hyperphyll. Cataphylls two, elongate; apical opening; apex acute. Eophyll bifid; segments sigmoid; apex acute; splitting side abaxial. Venation pattern pinnate; leaf axis distinct; midvein not distinct from other longitudinal veins; veins converging gradually toward apex; transverse commissures widely separated from each other, connecting some longitudinal veins and ending at intercostal areas. Plication with proximal marginal folding induplicate; distal folds varying on each side, one margin reduplicate and the other induplicate. Epidermal cells rhombohedral, papillose; adaxial and abaxial anticlinal walls linear. Stomata superficial, distribution irregular at intercostal regions; short or elongate terminal cells overarching guard cells. Hypodermis not distinct. Chlorenchyma undifferentiated; spongy mesophyll with fewer than five layers; fewer layers at folding regions. Expansion cells not distinct. Major veins multivascular; distribution associated with ridges; attached to adaxial and abaxial hypodermis; OS restricted to lateral sides. Median veins free, equidistant. Minor veins equidistant; OS restricted to lateral sides; radial OS present. Midrib abaxially prominent, spatula-shaped in transverse section; simple and multivascular bundles grouped. Margins lacking veins or fibrous bundles. Petiole heartshaped; abaxial side deeply concave. Phloem strands one. Metaxylem vessels two. Cell inclusions: silica bodies hat-shaped, smaller than silica bodies in the Caryoteae, margins spinulose, distributed around vascular and fibrous bundles, more concentrated at adaxial and abaxial sides of bundle; tannins scattered; raphids abundant.

Gaussia maya (O. F. Cook) Quero & Read

Seed attachment displaced below plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root persistent; distinct disk collar present; secondary roots simple; shoot-borne roots present; collar roots present; root hairs present, abundant. Hyperphyll absent. Cotyledonary sheath absent. Coleoptile short; apical opening; small slit opposite to seed. Cataphylls two; apical opening. Eophyll bifid; segments lanceolate; apex acute. Venation pattern pinnate; leaf axis distinct; midvein not distinct from other longitudinal veins; veins converging toward inner veins; transverse commissures widely separated from each other, connecting some longitudinal veins and ending at intercostal areas. Plication with proximal marginal folds induplicate; distal marginal outer folding induplicate; inner folding reduplicate. Epidermal cells rectangular or rhombohedral, papillose; adaxial and abaxial anticlinal walls linear. Stomata superficial; short or elongate terminal cells overarching guard cells. Hypodermis not distinct. Chlorenchyma undifferentiated; spongy mesophyll with fewer than five layers. Expansion cells single-layered, short, not too distinct from surrounding cells; adjacent epidermal cells papillose. Major veins multivascular, associated with ridges, attached to adaxial and abaxial epidermis; OS restricted to lateral sides. Median veins free, equidistant. Minor veins equidistant; OS restricted to lateral sides. Midrib abaxially prominent; simple vascular bundles grouped. Margins lacking vascular bundles or fibrous bundles. Petiole transverse section half-ellipsoid. Phloem strands one. Metaxylem vessels two. Cell inclusions: silica bodies hat-shaped, margins spinulose, distributed around vascular bundles; raphids equidistant; tannins scattered.

Synecanthusfibrosus (H. Wendl.) H. Wendl.

Seed attachment displaced below plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root persistent; disk collar distinct and swollen; shoot-borne roots present; collar roots present; root hairs abundant. Hyperphyll absent. Cotyledonary sheath absent. Coleoptile short, rugulose; apical opening opposite to seed. Cataphylls two, elongate; apical opening; apex acute or split and leathery. Eophyll bifid; segments lanceolate; apex acute; splitting side abaxial; short fibers visible in leaf clearings. Venation pattern pinnate; leaf axis distinct; midvein not distinct from other longitudinal veins; veins convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal marginal folds induplicate; distal outer marginal folding induplicate; inner marginal folding reduplicate. Epidermal cells rhombohedral; adaxial and abaxial anticlinal walls linear; papillose in transverse section. Hairs absent. Stomata superficial; short or elongate terminal cells overarching guard cells. Hypodermis not distinct. Chlorenchyma undifferentiated; spongy mesophyll with fewer than five layers. Expansion cells single-layered, conspicuous near major veins and midrib. Major veins multivascular, associated with ridges, attached to adaxial and abaxial epidermis; OS not distinct. Median veins free, equidistant. Minor veins equidistant; OS surrounding vascular bundle; radial OS present. Midrib abaxiaily prominent, rounded, sinuous; simple bundles grouped. Margins lacking vascular bundles or fibrous bundles. Petiole semi-ellipsoid; adaxial side flat; abaxial side ellipsoid. Phloem strands single. Metaxylem vessels two. Cell inclusions: silica bodies hat-shaped, margins spinulose, distributed around vascular and fibrous bundles; raphids equidistant; tannins abundant.

SUMMARY FOR HYOPHORBEAE

Plumular-radicular axis angular; primary root either straight or oblique, persistent or ephemeral; collar roots develop or do not develop; hyperphyll absent; cotyledonary sheath apical opening, rugulose, resembling Plectocomia (Calameae); coleoptile present; two cataphylls; eophyll bifid or pinnate and acute at apex; proximal marginal folds induplicate; distal margins induplicate.

V. ARECOIDEAE

1. Caryoteae

Arenga hookeriana (Becc.) Whitmore

Seed remaining above plumular-radicular node. Plumular-radicular axis symmetric, straight. Primary root persistent; secondary roots simple; collar roots present; root hairs absent. Hyperphyll with an adaxial single groove; region connected to seed, forming a swollen disk. Cotyledonary sheath elongate, splitting lengthwise. Coleoptile absent. Cataphyll single, tubular; apical opening; apex acute. Eophyll entire, broadly lanceolate; apex praemorse (Fig. 9A). Venation pattern palmate; leaf axis reduced; midvein not distinct from other longitudinal veins; veins not convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal and distal marginal folds induplicate (Fig. 8D). Epidermal cells fusiform; adaxial and abaxial anticlinal walls linear; papillose in transverse section; cuticle thick. Hairs abundant; perpendicular and elongate cells; basal cells sunken. Stomata superficial; inner wall of guard cells striate; short terminal cells overarching guard cells. Hypodermis single-layered, large, rounded cells; present at adaxial and abaxial sides. Chlorenchyma differentiated; palisade layer present; spongy mesophyll with fewer than five layers; fibers in equidistant bundles, few; lumen wide. Expansion cells single-layered, short, rectangular. Major veins not associated with ridges, attached to adaxial and abaxial hypodermis; OS distinct. Median veins free, equidistant. Minor veins equidistant; OS lateral; radial OS present. Midrib abaxially prominent; single vascular bundle. Marginal rib with major vascular bundle; OS distinct, surrounding bundle; parallel sclerotic wall dividing vascular bundle. Petiole transverse section pentagonal; large vascular bundles in each corner. Phloem strands one. Metaxylem vessels single. Cell inclusions: silica bodies large, hat-shaped, linear margins, distributed around vascular and fibrous bundles (Fig. 4D); raphids abundant, large, equidistant.

Caryota mitis Lour.

Seed remaining above plumular-radicular node. Plumular-radicular axis symmetric, straight. Primary root persistent; secondary roots simple; collar roots present. Hyperphyll short, smooth, connecting to seed by a distinct swollen disk. Cotyledonary sheath split lengthwise; splitting opposite to hyperphyll. Coleoptile absent. Cataphyll single, splitting lengthwise to plumular-radicular node; apex concave. Eophyll bifid; praemorse apex; splitting at adaxial side (Fig. 9B). Venation pattern palmate; leaf axis reduced; midvein not distinct from other longitudinal veins; veins not convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal and distal marginal folds induplicate. Epidermal cells fusiform; adaxial and abaxial anticlinal walls sinuous; cells ellipsoid; orientation parallel (Fig. 7C). Hairs present; basal cells sunken. Stomata slightly sunken; inner wall of guard cells striate; short terminal cells overarching guard cells. Hypodermis single-layered, ellipsoid cells; orientation parallel; present at adaxial and abaxial sides. Chlorenchyma undifferentiated; spongy mesophyll with fewer than five layers. Expansion cells absent. Major veins not associated with ridges; large, multivascular, prominent at abaxial side; OS distinct. Median veins free, equidistant, attached to hypodermis. Minor veins equidistant; OS lateral; radially arranged OS layer present. Midrib not distinct; major veins more prominent. Marginal rib absent. Petiole transverse section pentagonal; adaxial side grooved. Phloem strands one. Metaxylem vessels single. Cell inclusions: silica bodies hat-shaped, linear margins, distributed around vascular and fibrous bundles; raphids abundant (Fig. 4E).

Germination of Caryota has been described by Gatin (1906a), Mahabale and Shirke (1967), and Sento (1971).

Wallichia densiflora Mart.

Seed remaining above plumular-radicular node. Plumular-radicular axis symmetric, straight. Primary root persistent; secondary roots simple; collar roots present; root hairs present. Hyperphyll short, smooth, connected to seed by a distinct swollen disk. Cotyledonary sheath with short apical opening; covered by short hairs. Coleoptile absent. Cataphyll single, tubular, covered by short hairs; apex acute. Eophyll entire, ellipsoid; praemorse apex. Venation pattern palmate; leaf axis reduced; midvein not distinct from other longitudinal veins; veins not convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal and distal marginal folds induplicate. Epidermal cells fusiform, papillose; adaxial and abaxial anticlinal walls linear; cuticle thick. Hairs perpendicular, columnar; sunken basal cells. Stomata slightly sunken; inner wall of guard cells striate; short and elongate terminal cells overarching guard cell. Hypodermis singlelayered, rounded or ellipsoid cells; parallel orientation; present at adaxial and abaxial sides. Chlorenchyma differentiated; palisade layer distinct; spongy mesophyll with fewer than five layers; fibers in equidistant bundles; lumen wide. Expansion cells short, square, single-layered. Major veins not associated with ridges; large multivascular vascular bundles; OS distinct. Median veins flee, equidistant. Minor veins equidistant; OS lateral. Midrib abaxially prominent; single multivascular vascular bundle. Marginal rib with major vein; parallel sclerotic wall dividing vein. Petiole transverse section pentagonal. Phloem strands one. Metaxylem vessels single. Cell inclusions: silica bodies hat-shaped, linear margins, distributed around vascular and fibrous bundles; raphids abundant, equidistant.

SUMMARY FOR CARYOTEAE

Plumular-radicular axis; primary root straight and persistent; collar roots well developed; hyperphyll elongate, swollen at proximal end; cotyledonary sheath opening laterally; coleopfile absent; one cataphyll; eophyll entire or bifid, apex praemorse; palmate; induplicate marginal folding.

2. Iriarteeae

Iriartea deltoidea R. & P.

Seed horizontal, neither above nor below plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root ephemeral; collar disk distinct; wide flat surface; secondary roots branched; pneumatophores present; shoot-borne roots thicker than primary root. Hyperphyll undeveloped. Cotyledonary sheath absent. Coleoptile short, splitting opposite to seed. Cataphylls two, leathery; apical opening; apex acute. Eophyll entire, ellipsoid; apex praemorse. Venation pattern pinnate; leaf axis distinct; midvein present; veins not convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal marginal folds induplicate; distal margins reduplicate. Epidermal cells shape rhombohedral; adaxial and abaxial anticlinal walls linear; papillose in transverse section. Hairs abundant, conical, conspicuous in surface view; few basal cells (Fig. 7D). Stomata superficial, distribution irregular; short or elongate terminal cells overarching guard cells. Hypodermis single-layered, squared cells, present at adaxial and abaxial sides. Chlorenchyma undifferentiated; spongy mesophyll with fewer than five layers; fibers forming solid equidistant bundles; lumen small. Expansion cells single-layered; double-layered when flanking midrib. Major veins multivascular, prominent at abaxial surface, not associated with ridges, attached to hypodermis on both sides; distinct OS. Median veins free, equidistant. Minor veins equidistant; OS surrounding vascular bundle. Midrib abaxially prominent, square-shaped; simple and multivascular bundles grouped. Marginal rib with compact fibrous bundle. Petiole transverse section semicircular. Phloem strands one. Metaxylem vessels single or double. Cell inclusions: silica bodies hat-shaped, linear margins, distributed around vascular and fibrous bundles; raphids equidistant.

Iriartella setigera (Mart.) H. Wendl.

Seed attachment horizontal, neither above nor below plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root ephemeral; secondary roots, shoot-borne roots, collar roots, and root hairs absent. Hyperphyll undeveloped. Cotyledonary sheath absent. Coleoptile present. Eophyll entire; praemorse apex. Venation pinnate; leaf axis distinct; midvein present; veins not convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal and distal marginal folds reduplicate. Epidermal cells rectangular or rhombohedral; adaxial and abaxial anticlinal walls linear; cuticle thick. Hairs conical; few basal cells. Stomata superficial; short or elongate terminal cells overarching guard cells. Hypodermis single-layered; large ellipsoid cells arranged parallel; present at adaxial and abaxial sides. Chlorenchyma undifferentiated; spongy mesophyll with fewer than five layers; fibrous bundles abundant, small, equidistant; lumen small. Expansion cells single-layered; scattered fibrous bundles. Major veins prominent at abaxial side, not associated with ridges, attached to both hypodermal layers; OS distinct. Median veins free, oriented toward abaxial sides. Minor veins oriented toward abaxial side; OS cap-shaped; radial OS present. Midrib abaxially prominent; single multivascular bundle. Marginal rib with nonvascular bundle. Petiole semicircular. Phloem strands one. Metaxylem vessels single. Cell inclusions: silica bodies hat-shaped, margins smooth, distributed around fibrous bundles; tannins abundant in full sacs.

Socratea exorrhiza (Mart.) H. Wendl.

Seed attachment horizontal, neither above nor below plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root ephemeral; disk collar large, flat with callous contour; secondary roots simple; pneumatophores present; shoot-borne roots thicker than primary root. Hyperphyll inconspicuous. Cotyledonary sheath absent. Coleoptile distinct, splitting apically. Cataphylls four; fourth cataphyll elongate; apex bifid. Eophyll bifid; segments broad; margins and apex crenulated; splitting along grooved side (Fig. 9C). Venation pattern pinnate; leaf axis distinct; midvein not distinct from other longitudinal veins; veins not convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal marginal folds induplicate, distal marginal folds reduplicate. Epidermal cells rhombohedral; adaxial and abaxial anticlinal walls linear; abaxial cells papillose in transverse section. Hairs conical, abundant in both surfaces; few basal cells. Stomata superficial; short terminal cells overarching guard cells. Hypodermis single-layered, squared cells, at adaxial and abaxial sides. Chlorenchyma undifferentiated; spongy mesophyll with fewer than five layers; fibers in equidistant bundles; lumen small. Expansion cells single-layered, mainly associated to midvein or major veins; adjacent epidermal cells papillose. Major veins muitivascular, not associated with ridges, attached to adaxial hypodermis and abaxial epidermis, prominent at abaxial surface; IS multilayered and sclerotic; OS not distinct. Median veins free, oriented toward abaxial side. Minor veins equidistant; OS surrounding vascular bundle. Midrib abaxially prominent, rounded; simple and multivascular bundles grouped. Marginal rib with fibrous bundle. Petiole transverse section heart-shaped. Phloem strands one. Metaxylem vessels single or double. Cell inclusions: silica bodies hat-shaped, linear margins, distributed around vascular and fibrous bundles; tannins abundant, some in full sacs.

SUMMARY FOR IRIARTEEAE

Plumular-radicular axis angular; primary root oblique and ephemeral; collar roots develop or do not develop; cotyledonary sheath apical opening; a coleoptile present; cataphylls 2-4; eophyll bifid in Socratea and simple in Iriartea and Iriartella; apex praemorse; venation pinnate; reduplicate.

3. Podococceae

Podococcus barteri Mann & H. Wendl.

Seed horizontal, neither above nor below plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root ephemeral. Hyperphyll undeveloped. Cotyledonary sheath absent. Coleoptile present. Cataphylls were not easy to evaluate from herbarium samples. Eophyll entire, rhombohedral; crenulated margin. Venation pattern pinnate; leaf axis distinct; midvein; veins not convergent at apex. Plieation with proximal marginal reduplicate; distal marginal folds reduplicate. Epidermal cells shape rhombohedral; adaxial and abaxial anticlinal walls linear; irregular and papillose in transverse section. Hairs conical; basal cells superficial. Stomata superficial; elongate terminal cells overarching guard cells. Hypodermis not distinct. Chlorenchyma undifferentiated; spongy mesophyll with fewer than five layers; fibers abundant, compact, small; equidistant bundles; lumen small. Expansion cells absent. Major veins not associated with ridges, attached to both epidermal layers; sclerotic IS; distinct OS. Median veins free, equidistant; OS lateral and large single cells. Minor veins equidistant; OS restricted to lateral sides. Midrib prominent at both adaxial and abaxial sides; single multivascular bundle. Margins lacking ribs. Petiole transverse section heart-shaped. Phloem strands one. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, around vascular and fibrous bundles; tannins abundant, some in full sacs.

SUMMARY FOR PODOCOCCEAE

Plumular-radicular axis angular; primary root ephemeral; hyperphyll undeveloped. cotyledonary sheath absent; coleoptile present; eophyll entire, rhombohedral, crenulated margin; venation pinnate; reduplicate.

4. Areceae

Archontophoenix alexandrae (F. Muell.) H. Wendl. & Drude

Seed attachment displaced below plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root ephemeral; secondary roots branched; shoot-borne roots protruding through coleoptile base. Hyperphyll absent. Cotyledonary sheath not distinct. Coleoptile lateral split; apex acute; texture granular. Cataphylls two; apex splitting into several sections. Eophyll bifid; segments lanceolate; apex acute; splitting side abaxial. Venation pattern pinnate; leaf axis distinct; midvein present; veins not convergent at apex; transverse commissures widely separated from each other; some veins connect longitudinal veins; some end at intercostal areas. Plication with proximal marginal folds induplicate; distal marginal outer folding induplicate and inner folding reduplicate. Epidermal cells rhombohedral; adaxial and abaxial anticlinal walls linear. Hairs present, polyhedral in transverse section; few basal cells. Stomata superficial, scattered; short terminal cells overarching guard cells. Hypodermis single-layered, present at adaxial and abaxial sides. Chlorenchyma undifferentiated; spongy mesophyll with fewer than five layers; cells oriented parallel; fibers as equidistant bundles; lumen small. Expansion cells single-layered. Major veins prominent at both surfaces, associated with ridges, attached to adaxial and abaxial epidermis; OS distinct. Median veins free, equidistant; OS distinct; radial OS present. Minor veins distribution abaxial side; OS surrounding vascular bundle; radially arranged OS present. Midrib abaxially prominent, irregularly ax-shaped in transverse section, vascularized by grouped simple vascular bundles. Marginal rib with minor veins. Petiole transverse section semi-ellipsoid. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular and fibrous bundles; tannins abundant; raphids equidistant.

Gatin (1906a) and Ginieis (1953a, 1953b) have studied germination of Archontophoenix.

Dictyosperma album (Bory) H. Wendl. & Drude

Seed attachment displaced below plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root ephemeral; distinct swollen collar; secondary roots branched; pneumatophores present; shoot-borne roots emerging throughout coleoptile base, thick as the primary root. Hyperphyll absent. Cotyledonary sheath absent. Coleoptile short; apex acute; splitting laterally opposite to seed. CataphyUs two; apex acute and sharp. Eophyll bifid; segments linear-lanceolate; apex acute; splitting side along grooved side. Venation pattern pinnate; axis distinct; midvein present; veins convergent at apex; transverse commissures widely separated from each other, connecting some longitudinal veins and ending at intercostal areas. Plication with proximal marginal folds induplicate, distal marginal folds reduplicate. Epidermal cells rhombohedral; adaxial and abaxial anticlinal walls linear; polyhedral cells in transverse section (Fig. 7E). Hairs present; multicellular base. Stomata superficial, abundant in abaxial surface, arrangement scattered; short terminal cells overarching guard cells. Hypodermis not distinct; fibrous bundles present as a discontinuous layer; lumen wide. Chlorenchyma undifferentiated; spongy mesophyll with more than five layers. Expansion cells single-layered. Major veins prominent at both surfaces, associated with ridges, attached to adaxial and abaxial epidermis; distinct OS. Median veins free; equidistant. Minor veins distribution abaxial side; OS surrounding vascular bundle; radially arranged OS present. Midrib abaxially prominent, simple; multivascular bundles grouped. Marginal rib with fibrous layers. Petiole transverse section heart-shaped. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles; raphids equidistant; tannins abundant.

Germination of Dictyosperma has been described by Gatin (1906a).

Dypsis lutescens (H. Wendl.) Beentje & J. Dransf.

Seed attachment displaced below plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root ephemeral; disk collar not swollen; secondary roots simple; pneumatophores present; shoot-borne roots distinct. Hyperphyll absent. Cotyledonary sheath absent. Coleoptile short, splitting opposite to hyperphyll. Cataphylls two; apical opening; apex acute. Eophyll bifid; linear segments lanceolate; apex acute; splitting side abaxial. Venation pattern pinnate; leaf axis distinct; midvein present; veins convergent at apex; transverse commissures widely separated from each other, connecting some longitudinal veins and ending at intercostal areas. Piication with proximal marginal folds induplicate; distal marginal folds reduplicate. Epidermal cells rhombohedral, rounded in transverse section; adaxiai and abaxial anticlinal walls linear. Hairs present; multicellular base. Stomata superficial; short terminal cells overarching guard cells; scattered arrangement, more concentrated at abaxial surface. Hypodermis not distinct. Chlorenchyma undifferentiated; spongy mesophyll with more than five layers; fibers in equidistant bundles; lumen wide. Expansion cells double-layered; rectangular; perpendicular; adjacent epidermal cells papillose. Major veins associated with ridges; OS distinct. Median veins free, equidistant. Minor veins equidistant; OS surrounding vascular bundle; radially arranged OS present. Midrib abaxially prominent, rounded, vascularized by grouped simple vascular bundles. Marginal rib with minor vein. Petiole transverse section heart-shaped. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles; raphids equidistant; tannins abundant in full sacs, some in OS.

Euterpe precatoria Mart.

Seed attachment displaced below plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root ephemeral; secondary roots branched; pneumatophores scattered; shoot-borne roots present; collar present; root hairs present. Hyperphyll absent. Cotyledonary sheath absent. Coleoptile short, splitting opposite to seed. Cataphylls two; apical opening; apex acute; plicate. Eophyll pinnate; linear-segments lanceolate; apex acute; split leaf first; split along grooved side. Venation pattern pinnate; leaf axis distinct; midvein present; veins convergent at apex; transverse commissures widely separated from each other, connecting some longitudinal veins and ending at intercostal areas. Plication with proximal marginal folds induplicate; distal outer margin indupticate; inner margin reduplicate. Epidermal cells rectangular or fusiform; adaxial anticlinal walls linear; abaxial anticlinal walls sinuous. Hairs present; muiticellular base. Stomata superficial, scattered; elongate terminal cells overarching guard cells. Hypodermis not distinct; fibrous strands present as a discontinuous layer; lumen small. Chlorenchyma undifferentiated; spongy mesophyll with fewer than five layers, rectangular, with a horizontal orientation. Expansion cells single-layered, elongate, perpendicular; few cells. Major veins prominent at adaxial and abaxial surfaces, associated with ridges; IS multilayered and sclerotic; OS distinct. Median veins buttressed, attached to both epidermal layers; OS large lateral cells. Minor veins equidistant; OS lateral. Midrib adaxially prominent, vascularized by grouped simple vascular bundles. Marginal rib with major vein; IS multilayered at marginal end. Petiole transverse section rounded; adaxial side slightly concave. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles.

Germination of Euterpe has been described by Belin-Depoux and de Queiroz (1971).

Hyospathe elegans Mart.

Seed displaced below plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root ephemeral; collar disk distinct; secondary roots branched; shoot-borne roots thicker than primary root. Hyperphyll absent. Cotyledonary sheath absent. Coleoptile short; apex acute; splitting opposite to seed. Cataphylls two; splitting lengthwise; apex acute. Eophyll bifid; segments lanceolate; splitting side abaxial. Venation pattern pinnate; leaf axis distinct; midvein present; veins not convergent at apex; transverse commissures widely separated from each other, connecting some longitudinal veins and ending at intercostal areas. Plication with proximal marginal folds induplicate; distal margin reduplicate. Epidermal cells rectangular or rhombohedral, polyhedral in transverse section; adaxial and abaxial anticlinal walls linear (Fig. 7F). Hairs present; multicellular base. Stomata slightly sunken; short or elongate terminal cells not overarching guard cells; scattered distribution, more concentrated at abaxial surface. Hypodermis not distinct. Chlorenchyma undifferentiated; spongy mesophyll with fewer than five layers, rectangular, with horizontal orientation. Expansion cells single-layered, distributed at lateral sides of major veins. Major veins abaxially prominent, associated with ridges; distinct OS. Median veins free, equidistant. Minor veins at abaxial side; OS surrounding vascular bundle; radially arranged OS present. Midrib adaxially prominent; single vascular bundle. Marginal rib with minor veins present. Petiole transverse section heartshaped. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles; raphids equidistant; tannins abundant, scattered in every cell.

[FIGURE 7 OMITTED]

Neonicholsonia watsonii Dammer

Seed horizontal, neither above nor below the plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root ephemeral; swollen collar present; secondary roots branched; pneumatophores present. Hyperphyll absent. Cotyledonary sheath absent. Coleoptile short; splitting ventral. Cataphylls two; apex acute. Eophyll pinnate; segments lanceolate; apex acute; splitting side abaxial. Venation pattern pinnate; leaf axis distinct; midvein present; veins convergent at apex; transverse commissures widely separated from each other, connecting longitudinal veins and ending at intercostal areas. Plication with proximal marginal folds induplicate; distal margins, one reduplicate the other induplicate. Epidermal cells rectangular or fusiform; adaxial and abaxial anticlinal walls linear or sinuous (Fig. 7G). Hairs absent. Stomata superficial, scattered; elongate terminal cells overarching guard cells. Hypodermis not distinct. Chlorenchyma undifferentiated; spongy mesophyll with fewer than five layers; fibers in abaxial and adaxial bundles; lumen wide. Expansion cells single-layered, flanking major veins. Major veins prominent adaxially, associated with ridges; OS distinct. Median veins free, equidistant. Minor veins equidistant; OS surrounding vascular bundle. Midrib adaxially prominent; single vascular bundle. Marginal rib with major vein; IS multilayered and fibrous at marginal end. Petiole transverse section heart-shaped. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular and fibrous bundles; tannins scattered, some in full sacs.

Nephosperma vanhoutteanum (H. Wendl.) Balfour

Seed displaced below plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root ephemeral; swollen disk collar; secondary roots simple; shootborne roots thicker than primary root, borne on flat collar surface Hyperphyll absent. Cotyledonary sheath absent. Coleoptile short; ventral splitting. Cataphylls three, longer than coleoptile; apex acute. Eophyll pinnate; segments lanceolate; apex acute; splitting at abaxial side. Venation pattern pinnate; leaf axis distinct; midvein present; veins convergent at apex; transverse commissures widely separated from each other, connecting some longitudinal veins and ending at intercostal areas. Plication with proximal marginal folds induplicate; distal margins, one reduplicate, the other induplicate. Epidermal cells rhombohedral; adaxial and abaxial anticlinal walls linear. Hairs present; few basal cells. Stomata superficial; short terminal cells overarching guard cells. Hypodermis not distinct; fibrous bundles and layers present; lumen small. Chlorenchyma undifferentiated; spongy mesophyll with more than five layers. Expansion calls double-layered, elongate, ellipsoid. Major veins associated with ridges, attached to adaxial epidermal layer and abaxial expansion cells; OS distinct. Median veins free, equidistant. Minor veins oriented toward abaxial side; OS cap-shaped; radially oriented OS present. Midrib abaxially prominent, rounded, vascularized by grouped simple vascular bundles. Marginal rib with fibrous layers. Petiole transverse section heart-shaped. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles; tannins abundant.

Orania regalis Zipp.

Seed displaced above plumular-radicular node. Plumular-radicular axis slightly asymmetric, angular. Primary root persistent; collar not distinct; secondary roots branched; pneumatophores present; shoot-borne roots abundant; collar roots present. Hyperphyll elongate; single apical groove; hyperphyll smooth in contrast to sheath; connection to seed swollen. Cotyledonary sheath thick, rugulose, and wrinkled; opening laterally opposite to hyperphyll. Coleoptile absent. Cataphylls two, thick, covered by dense tegument. Eophyll bifid; segments broadly lanceolate; splitting side abaxial. Venation pattern pinnate; leaf axis distinct; midvein present; veins not convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal marginal folds induplicate; distal margins reduplicate. Epidermal cells polyhedral; abaxial epidermis papillose in transverse section. Hairs large, bicellular; few basal cells. Stomata superficial. Hypodermis not distinct. Chlorenchyma differentiated; palisade with distinct two layers; spongy mesophyll with more than five layers; fibers in equidistant bundles. Expansion cells double-layered. Major veins multivascular, associated with ridges; distinct OS. Median veins free, toward abaxial side. Minor veins oriented toward abaxial side, some associated with grooves; OS surrounding vascular bundle. Midrib abaxial squared protuberance; simple and multivascular bundles grouped. Marginal rib with minor vein. Petiole transverse section heart-shaped. Phloem strands three. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular bundles; tannins scattered.

Phoenicophorium borsigianum (K. Koch) Stuntz

Seed displaced below plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root ephemeral; distinct flat disk collar present; secondary roots branched; pneumatophores present; shoot-borne roots few, thicker than primary root; collar roots; root hairs. Hyperphyll very short. Cotyledonary sheath absent. Coleoptile short; splitting opposite to seed. Cataphylls two; opening lengthwise; apex with acute projection; covered by large trichoma. Eophyll entire, lanceolate; apex acuminate; split leaf third; splitting side abaxial. Venation pattern pinnate; leaf axis distinct; midvein not distinct; veins convergent at apex; transverse commissures widely separated from each other, connecting some longitudinal veins and ending at intercostal areas. Plication with proximal marginal folds induplicate; distal margins reduplicate. Epidermal cells rectangular or rhombohedral; adaxial and abaxial anticlinal walls sinuous. Hairs present; few basal cells. Stomata superficial, scattered; short or elongate terminal cells overarching guard cells. Hypodermis not distinct; fibrous bundles present as a discontinuous layer; lumen small. Chlorenchyma undifferentiated; spongy mesophyll with more than five layers. Expansion cells single-layered; scattered fibers. Major veins associated with ridges; adaxial buttresses attached to epidermis and abaxial expansion cells; multilayered IS; distinct OS. Median veins free; toward abaxial side. Minor veins toward abaxial side; OS surrounding vascular bundle; radially arranged OS present. Midrib abaxially prominent; single bundle. Marginal rib with fibrous layers. Petiole transverse section heart-shaped. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular and fibrous bundles; raphids equidistant; tannins scattered.

Roystonea borinquena O. F. Cook

Seed horizontal, neither above nor below plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root ephemeral; disk collar distinct and swollen; secondary roots simple; shoot-borne roots absent; collar roots present; root hairs present. Hyperphyll short, appearing continuous to primary root with plumule erupting at adaxial side. Cotyledonary sheath absent. Coleoptile short; slight slit opposite to seed. Cataphylls two, elongate; first apical opening; second split lengthwise; extremely acute apex. Eophyll entire, broadly lanceolate; apex acute. Venation pattern pinnate; leaf axis distinct; midvein distinct; veins convergent at apex; transverse commissures widely separated from each other, connecting some longitudinal veins and ending at intercostal areas. Plication with proximal marginal folds induplicate; distal margins reduplicate (Fig. 8E). Epidermal cells rectangular or fusiform, papillose in transverse section; adaxial and abaxial anticlinal walls linear. Hairs present; few basal cells. Stomata superficial scattered distribution, more concentrated at abaxial surface; elongate terminal cells overarching guard cells. Hypodermis single-layered, large rounded cells, at adaxial and abaxial sides. Chlorenchyma undifferentiated; spongy mesophyll with fewer than five layers; fibrous bundles solid with around 10 strands, equidistant; lumen wide. Expansion cells single-layered; scattered fibrous bundles. Major veins not associated with ridges, attached to adaxial and abaxial hypodermis; distinct OS. Median veins free, equidistant. Minor veins equidistant; OS surrounding vascular bundle. Midrib adaxially prominent; single, large vascular bundle. Marginal rib with minor veins present. Petiole transverse section heart-shaped. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies hat-shaped, margins spinulose, distributed around vascular and fibrous bundles; raphids equidistant; tannins scattered.

[FIGURE 8 OMITTED]

Germination of Roystonea has been described by Gatin (1906a).

Veitchia montgomeryana H. E. Moore

Seed horizontal, neither above nor below plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root ephemeral; collar disk distinct; secondary roots branched; scattered pneumatophores; shoot-borne roots abundant, arising around primary root and through base of coleoptile; collar roots and root hairs absent. Hyperphyll absent. Cotyledonary sheath absent. Coleoptile short, splitting laterally. Cataphylls three, displaying diverse splitting forms; apex acute, split. Eophyll bifid; segments broadly lanceolate; apex praemorse; abundant rounded spots present on lamina; splitting along grooved side. Venation pattern pinnate; leaf axis distinct; midvein not distinct; veins not convergent at apex; transverse commissures widely separated from each other, connecting some longitudinal veins and ending at intercostal areas. Plication with proximal marginal folds reduplicate; distal outer margin reduplicate; inner margin induplicate. Epidermal cells rectangular or rhombohedral; adaxial and abaxial anticlinal walls linear or sinuous. Hairs present; multicellular base. Stomata slightly sunken, scattered; short and elongate terminal cells overarching guard cells. Hypodermis single-layered, rounded cells, present at adaxial and abaxial sides. Chlorenchyma undifferentiated; spongy mesophyll with fewer than five layers; fibrous bundles at adaxial and abaxial sides; lumen small. Expansion cells double-layered, ellipsoid or rectangular. Major veins associated with ridges, attached to adaxial hypodermis and abaxial epidermis; OS distinct; radial OS present. Median veins free, equidistant. Minor veins equidistant; OS surrounding vascular bundle; radial OS present. Midrib abaxially prominent, ax-shaped, vascularized by grouped simple vascular bundles. Marginal rib with minor veins present. Petiole transverse section heart-shaped. Phloem strands two. Metaxylem vessels one or two. Cell inclusions: silica bodies spherical or ellipsoid, margins spinulose, distributed around vascular and fibrous bundles; raphids adaxial; tannins abundant, some in full sacs.

SUMMARY FOR ARECEAE

Plumular-radicular axis, angular; primary root either straight or oblique, persistent or ephemeral; collar roots develop or do not develop; cotyledonary sheath apical opening; coleoptile present; cataphylls 2-3; eophyll bifid, rarely simple or pinnate, and usually acute at apex (sometimes praemorse); reduplicate inner margins and induplicate outer margins.

5. Cocoeae

Allagoptera leucocalyx (Mart.) Kuntze

Seed remaining above plumular-radicular node. Plumular-radicular axis symmetric, straight. Primary root stout and persistent; collar disk not distinct; secondary roots simple, short; pneumatophores present; collar roots present. Hyperphyll elongate, grooved all around; connection to seed flat. Cotyledonary sheath grooved; splitting lengthwise, opposite to hyperphyll. Coleoptile absent. Cataphylls two; apex acute and curved. Eophyll entire, linear-lanceolate; apex acute. Venation pattern pinnate; leaf axis distinct; midvein distinct; veins convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal and distal marginal folds varying on each side, one margin reduplicate, the other induplicate. Epidermal cells rectangular or rhombohedral, regular and uniform in transverse section; adaxial and abaxial anticlinal walls linear. Hairs present; few basal cells. Stomata slightly sunken, arranged in rows at intercostal regions, more abundant on abaxial surface; short and elongate terminal cells overarching guard cells. Hypodermis single-layered, present at adaxial and abaxial sides. Fibrous bundles present at irregular intervals; lumen small. Chlorenchyma undifferentiated; spongy mesophyll with more than five layers; fibers as subepidermal bundles present at ridges. Expansion cells single-layered. Major veins associated with ridges; OS distinct. Median veins buttressed to abaxial and adaxial sides. Minor veins buttressed to abaxial side; OS cap-shaped; some vascular bundles associated with abaxial grooves. Midrib abaxially prominent, with simple and multivascular vascular bundles grouped. Marginal rib with minor veins. Petiole transverse section crescent-shaped. Phloem strands two to four. Metaxylem vessels single. Cell inclusions: silica bodies irregular, margins spinulose, distributed around vascular bundles.

Astrocaryum alatum Loomis

Seed displaced below plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root ephemeral; shoot-borne roots abundant; secondary roots stout, branched; pneumatophores present; shoot-borne roots present. Hyperphyll inconspicuous. Cotyledonary sheath absent. Coleoptile short; splitting side opposite to seed; spiny. Cataphylls two; splitting lengthwise. Eophyll bifid; splitting side abaxial; margins spiny (Fig. 9D). Venation pattern pinnate; leaf axis distinct; midvein distinct; veins convergent near apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal marginal folds induplicate; distal outer marginal folds induplicate; inner fold reduplicate. Epidermal cells fusiform, papillose in transverse section; adaxial and abaxial walls linear; wax layer thick. Hairs present; few basal cells. Stomata sunken, occluded by wax layer; short terminal cells overarching guard cells. Hypodermis single-layered; cells ellipsoid; orientation parallel; present at adaxial and abaxial sides. Chlorenchyma with differentiated layers; palisade layer present; spongy mesophyll with fewer than five layers; fibers as compact bundles, abundant, scattered; small lumen. Expansion cells double-layered; fibrous bundles scattered. Major veins abaxially prominent, associated with ridges, attached to adaxial and abaxial epidermis, usually flanked by expansion cells; OS not distinct. Median veins free, equidistant. Minor veins at abaxial side, not associated to grooves; OS surrounding vascular bundle. Midrib abaxially prominent, rounded; simple and multi-vascular vascular bundles grouped. Marginal rib with minor vein present. Petiole transverse section heart-shaped. Phloem strands four. Metaxylem vessels single. Cell inclusions: silica bodies hat-shaped, margins spinulose, distributed around nonvascular bundles; tannins scattered.

[FIGURE 9 OMITTED]

Bactris killippii Burret

Seed displaced below plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root ephemeral; secondary roots simple; shoot-borne roots stout, thicker than primary root. Hyperphyll absent. Cotyledonary sheath inconspicuous. Coleoptile short, splitting opposite to seed. Cataphylls two; opening apical; apex acute. Eophyll bifid; linear segments lanceolate; splitting side abaxial. Venation pattern pinnate; leaf axis distinct; veins gradually convergent toward the apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal marginal folds induplicate; distal outer fold induplicate; inner fold reduplicate (Fig. 8F). Epidermal cells fusiform, papillose in transverse section; adaxial and abaxial anticlinal walls linear; cuticle thick. Hairs present; few basal cells. Stomata superficial; short or elongate terminal cells not overarching guard cells; guard cells with large ledges. Hypodermis single-layered, present at adaxial and abaxial sides, orientation parallel. Chlorenchyma undifferentiated; spongy mesophyll with fewer than five layers; fibrous bundles compact, equidistant, surrounded by distinct tannin-filled OS; lumen small. Expansion cells single-layered. Major veins adaxially prominent, associated with ridges, attached to adaxial hypodermis and abaxial expansion cells; IS multilayered, sclerotic; OS distinct. Median veins free, equidistant. Minor veins equidistant; OS surrounding vascular bundle. Midrib abaxially prominent, rounded; simple and multivascular vascular bundles grouped. Marginal rib with fibrous bundle. Petiole transverse section heart-shaped. Phloem strands four. Metaxylem vessels single. Cell inclusions: silica bodies hat-shaped, spinulose margins, distributed around fibrous bundles; tannins scattered in mesophyll, abundant in OS.

Elaeis guineensis Jacq.

Seed above plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root persistent; secondary roots branched; pneumatophores present; shoot-borne roots present; collar disk swollen; collar roots present. Hyperphyll short, almost inconspicuous. Cotyledonary sheath absent. Coleoptile short, not split. Cataphylls two, elongate; apex acute; opening lengthwise. Eophyll entire, broadly lanceolate; apex acute; five leaves similar to eophyll before first split leaf appearing; splitting side abaxial. Venation pattern pinnate; leaf axis distinct; midvein distinct; veins convergent gradually to midvein; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal marginal folds induplicate; distal folds induplicate. Epidermal cells rectangular or rhombohedral, polyhedral in transverse section; adaxial and abaxial anticlinal walls linear. Hairs present; few basal cells. Stomata slightly sunken, scattered at intercostal regions; short terminal cells overarching guard cells. Hypodermis single-layered, present at adaxial and abaxial sides. Chlorenchyma undifferentiated; spongy mesophyll with fewer than five layers; fibers in equidistant bundles; lumen wide. Expansion cells single-layered. Major veins adaxially prominent, displaced from ridge, attached to epidermal layers; OS not distinct. Median veins free, equidistant. Minor veins equidistant; OS surrounding vascular bundle; radial OS present. Midrib abaxially prominent; squared protuberance; simple and multivascular vascular bundles grouped. Marginal rib with fibrous layers. Petiole transverse section crescent-shaped. Phloem strands four. Metaxylem vessels single. Cell inclusions: silica bodies irregular, margins smooth, distribution around fibrous bundle or vascular bundle; raphids equidistant; tannins, abundant full sacs.

Yampolsky (1922) has described germination of Elaeis guineensis Jacq.

Jubaea chilensis (Molina) Baillon

Seed remaining above plumular-radicular node. Plumular-radicular axis symmetric, straight. Primary root stout and persistent; disk collar slightly swollen; secondary roots simple, very short; collar roots present. Hyperphyll short, smooth; connection to seed flat. Cotyledonary sheath short, splitting apically. Coleoptile apical opening; splitting opposite to seed. Cataphylls two, opening laterally; apical extensions bifid with distinct protuberant transverse lines. Eophyll entire, lanceolate; apex acute. Venation pattern pinnate; leaf axis distinct; midvein distinct; veins convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal marginal and distal folds varying on each side; one marginal fold reduplicate and the other induplicate. Epidermal cells rectangular or rhombohedral; adaxial and abaxial anticlinal walls linear; cuticle thick. Hairs present; few basal cells. Stomata slightly sunken, scattered; terminal cells short and elongate overarching guard cells. Hypodermis single-layered, rounded cells, present at adaxial and abaxial sides; compact fibrous bundles at irregular intervals. Chlorenchyma undifferentiated; spongy mesophyll with more than five layers; fibers in bundles at abaxial side; distinct bundles at ridges; lumen wide. Expansion cells double-layered; adjacent epidermis papillose. Major veins not associated with ridges, buttressed to adaxial and abaxial epidermis; IS multilayered, sclerotic; OS not distinct. Median veins buttressed to abaxial and adaxial sides; solid buttresses. Minor veins abaxially buttressed; OS not distinct. Midrib abaxially prominent, squared; simple and multivascular vascular bundles grouped. Marginal rib with minor vein. Petiole transverse section crescent-shaped. Phloem strands four. Metaxylem vessels single. Cell inclusions: spherical or ellipsoid, margins spinulose, around vascular bundles; tannins abundant.

Gatin (1906a) has described germination of Jubaea.

Syagrus coronata (Mart.) Becc.

Seed above plumular-radicular node. Plumular-radicular axis symmetric, straight. Primary root stout and persistent; disk collar swollen; secondary roots simple and short; pneumatophores present; collar roots present. Hyperphyll elongate, grooved all around; connection to seed flat. Cotyledonary sheath splitting lengthwise, deeply grooved. Coleoptile absent. Cataphylls two, elongate; apex acute. Eophyll entire, linear-lanceolate; apex acute. Venation pattern pinnate; leaf axis distinct; midvein distinct; veins convergent at apex; transverse commissures widely separated from each other, connecting longitudinal veins sporadically. Plication with proximal and distal marginal folds induplicate and reduplicate. Epidermal cells rectangular; adaxial and abaxial anticlinal walls linear (Fig. 7H). Hairs basal cells few. Stomata sunken, scattered, abundant in both surfaces; short and elongate terminal cells overarching guard cells. Hypodermis single-layered, present at adaxial and abaxial sides; fibrous bundles or layers at regular intervals. Chlorenchyma undifferentiated; spongy mesophyll with more than five layers; fibers in bundles at ridges; lumen small. Expansion cells doublelayered, elongate, ellipsoid; adjacent epidermis papillose. Major veins not associated with ridges, attached to both epidermal layers; OS not distinct. Median veins buttressed to both adaxial and abaxial sides. Minor veins buttressed to abaxial side; OS capshaped; some vascular bundles associated with grooves. Midrib abaxially prominent, triangular; single, large multivascular bundle. Marginal rib with minor vein. Petiole transverse section crescent-shaped. Phloem strands two and four. Metaxylem vessels single. Cell inclusions: silica bodies irregular, margins spinulose, around vascular bundles; raphids equidistant; tannins scattered.

[FIGURE 7 OMITTED]

Gatin (1906a) has described germination of Syagrus australis (as Cocos australis) and S. campestris (as Cocos campestris).

Voanioala gerardii J. Dransf.

Seed characters not recorded. Plumular-radicular axis symmetric. Primary root persistent; collar disk distinct; secondary roots simple; shoot-borne roots present. Hyperphyll elongate, connection to seed flat. Cotyledonary sheath rugulose, opening lengthwise. Coleoptile not seen. Cataphylls two. Eophyll entire, lanceolate; apex acute with long linear extension; third leaf splitting at abaxial side. Venation pattern pinnate; leaf axis distinct; midvein distinct; veins convergent at apex; transverse commissures abundant, very close to each other, connecting longitudinal veins or intercostal regions sporadically. Plication with proximal and distal marginal folds reduplicate and induplicate (Fig. 8G). Epidermal cells rectangular or rhombohedral, small and polyhedral in transverse section; adaxial and abaxial anticlinal walls linear; cuticle thick. Hairs present, with few cells; sunken base (Fig. 4F). Stomata sunken, scattered at hypodermal layer; short terminal cells overarching guard cells. Hypodermis single-layered, at adaxial and abaxial sides. Chlorenchyma differentiated; palisade layer distinct; spongy mesophyll up to five layers; fibers in bundles at adaxial side; lumen wide. Expansion cells double-layered, elongate, rectangular; adjacent epidermis papillose. Major veins associated with ridges, attached to adaxial hypodermis and abaxial expansion layer; OS not distinct. Median veins free, equidistant. Minor veins oriented toward abaxial side; OS cap-shaped. Midrib abaxially prominent, squared; simple and multivascular vascular bundles grouped. Marginal rib not vascularized. Petiole transverse section terete. Phloem strands two to four. Metaxylem vessels single. Cell inclusions: silica bodies irregular, margins spinulose, distributed around vascular bundles (only one sample in an advanced stage of development was available for study).

[FIGURES 4-8 OMITTED]

SUMMARY FOR COCOEAE

Plumular-radicular axis may be straight or angular; primary root straight and persistent or oblique and ephemeral; collar roots developed or not developed; hyperphyll short or elongated; cotyledonary sheath opening apically or laterally; coleoptile present or absent; cataphylls two; eophyll entire or bifid; apex acute; bifid eophyll with pinnate venation; margins either reduplicate on both sides or induplicate on one and reduplicate on the other.

6. Geonomeae

Geonoma interrupta (R. & P.) Mart.

Seed above plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root persistent; disk collar distinct; secondary roots branched; shoot-borne roots outgrowing primary root. Hyperphyll absent. Cotyledonary sheath absent. Coleoptile short, not split. Cataphylls two, elongate, opening lengthwise; apex acute, split. Eophyll bifid; segments sigmoid; apex acute; splitting side abaxial. Venation pattern pinnate; leaf axis distinct; midvein distinct from other longitudinal veins; veins convergent at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal marginal folds induplicate; distal outer margin induplicate; inner margin reduplicate. Epidermal cells rhombohedral, large and regular in transverse section; adaxial and abaxial anticlinal walls linear; costal cells shorter than intercostal cells. Hairs unicellular; globose basal cells few. Stomata superficial; short terminal cells overarching guard cells. Hypodermis not distinct. Chlorenchyma undifferentiated; spongy mesophyll with fewer than five layers; fibrous bundles equidistant. Expansion cells single-layered; large cuneate cells adjacent to midrib. Major veins prominent at abaxial and abaxial sides, associated with ridges, attached to epidermal layers; IS multicellular, sclerotic; OS distinct, at lateral sides of vascular bundle, filled with tannins. Median veins free, equidistant. Minor veins equidistant; OS surrounding vascular bundle, filled with tannins. Midrib abaxially prominent; simple and multivascular bundles grouped. Margins lacking ribs. Petiole crescentshaped. Phloem strands two. Metaxylem vessels single. Cell inclusions: silica bodies hat-shaped, margins spinulose, around vascular bundle; raphids equidistant; tannins abundant, large, in full sacs.

Welfia regia H. Wendl.

Seed above plumular-radicular node. Plumular-radicular axis asymmetric, angular. Primary root persistent; secondary roots branched; shoot-borne roots abundant; collar roots and root hairs absent. Hyperphyll extremely short. Cotyledonary sheath absent. Coleoptile very short; apical opening; not split. Cataphylls two; opening lengthwise, with apex split. Eophyll bifid; broad-sigmoid segments; apex acute; splitting side abaxial. Venation pattern pinnate; leaf axis distinct; midvein distinct from other longitudinal veins; veins gradually converging at apex; transverse commissures widely separated from each other, connected to longitudinal veins. Plication with proximal marginal folds induplicate; distal outer margin induplicate; inner margin reduplicate. Epidermal cells rhombohedral, papillose in transverse section; adaxial and abaxial anticlinal walls linear; cuticle present. Hairs unicellular, globular; few basal cells. Stomata superficial; short terminal cells overarching guard cells. Hypodermis not distinct. Chlorenchyma undifferentiated; spongy mesophyll with fewer than five layers; fibrous bundles equidistant. Expansion cells single-layered. Major veins associated with ridges; distinct OS. Median veins free, equidistant. Minor veins equidistant; OS surrounding vascular bundle. Midrib abaxially prominent, rounded; simple and multivascular bundles grouped. Margins lacking ribs. Petiole transverse section heart-shaped. Phloem strands one. Metaxylem vessels single. Cell inclusions: silica bodies irregular, margins spinulose, around vascular bundle; raphids scarce; tannins abundant, full sacks, present in vascular bundle and mesophyll cells.

SUMMARY FOR GEONOMEAE

Piumular-radicular axis curved; primary root either straight or oblique, persistent or ephemeral; collar roots develop or do not develop; cotyledonary sheath apical opening; coleoptile present; cataphylls two; eophyll bifid, apex acute; venation pinnate; margins reduplicate.
Table I
List of material examined

 Voucher
Taxa specimen *

I. CORYPHOIDEAE
1. Corypheae
Acoelorraphe wrightii (Griseb. & H. Wendl.) Chavez 909
 H. Wendl. ex Becc.
Chamaerops humilis L. Chavez 910
Chuniophoenix hainanensis Burret Chavez 964
Colpothrinax cookii Read Chavez 965
Copernicia baileyana Leon. Chavez 918
Corypha sp. Chavez 911
Cryosophila grayumii R. Evans Chavez 920
Itaya amicorum H. E. Moore Chavez 955
Livistona chinensis R. Br. Chavez 966
Nannorrhops ritchiana (Griff.) Aitchson Chavez 915
Pritchardia remote (Kuntze) Becc. Chavez 917
Rhapidophyllum hystrix (Pursh) H. Wendl. & Drude Chavez 963
Sabal minor (Jacq.) Persoon Chavez 912
Serenoa repens (Bartram) Small Chavez 959
Thrinax excelsa Lodd. ex Griseb. Chavez 903
Trachycarpus sp. Chavez 902
Trithrinax brasiliensis (Mart.) Chavez 967
Washingtonia filifera (Linden) H. Wendl. Chavez 930
2. Phoeniceae
Phoenix roebelinii O'Brien Chavez 904
3. Borasseae
Borassus sp. Chavez 968
Hyphaene coriacea Gaertn. Chavez 969
Latania loddegesii Mart. Chavez 957

II. CALAMOIDEAE
1. Calameae
Calamus flagellum Griff. Chavez 945
Pigafetta filaris (Gis.) Becc. Chavez 944
Plectocomia sp. Chavez 946
2. Lepidocaryeae
Mauritia flexuosa L. f. Chavez 948

III. NYPOIDEAE
Nypa fruticans Wurmb Chavez 949

IV. CEROXYLOIDEAE
1. Cyclospatheae
Pseudophoenix sargentii H. Wendl. Chavez 971
2. Ceroxyleae
Ceroxylon sp. Henderson 3019
Oraniopsis appendiculata (F. Bailey)
Dransf., Uhl & Irvine Henderson 3070
Ravenea rivularis Jum. & H. Perrier Chavez 972
3. Hyophorbeae
Chamaedorea microspadixBurret Chavez 937
Gaussia maya (O. F. Cook) Quero & Read Chavez 978
Synecanthus fibrosu. s (H. Wendl.) H. Wendl. Chavez 938

V. ARECOIDEAE
1. Caryoteae
Arenga hookeriana (Becc.) Whitmore Chavez 907
Caryota mitis Lour. Chavez 916
Wallichia densiflora Mart. Chavez 905
2. Iriarteeae
Iriartea deltoidea R. & P. Henderson 3015
Iriartella setigera (Mart.) H. Wendl. Henderson 647
Socratea exorrhiza (Mart.) H. Wendl. Chavez 935
3. Podococceae
Podococcus barteri Mann & H. Wendl. Reitsma 2840
4. Areceae
Archontophoenix alevandrae (F. Muell.) Chavez 932
 H. Wendl. & Drude
Dictyosperma album (Bory) H. Wendl. & Drude Chavez 934
Dypsis lutescens (H. Wendl.) Beentje & J. Dransf. Chavez 931
Euterpe precatoria Mart. Balslev 4813
Hyospathe elegans Mart. Chavez 929
Neonicholsonia watsonii Dammer Chavez 928
Nephosperma vanhoutteanum (H. Wendl.) Balfour Chavez 939
Orania regalis Zipp. Chavez 985
Phoenicophorium borsigianum (K. Koch) Stuntz Henderson 2063
Roystonea borinquena O. F. Cook Chavez 927
Veitchia montgomeryana H. E. Moore Chavez 977
5. Cocoeae
Allagoptera leucocalyx (Mart.) Kuntze Chavez 941
Astrocaryum alatum Loomis Stevenson 1200
Bactris killippii Burret Henderson 2015
Elaeis guineensis Jacq. Chavez 942
Jubaea chilensis (Molina) Baillon Chavez 975
Syagrus coronata (Mart.) Becc. Chavez 947
Voanioala gerardii J. Dransf. Chavez 976
6. Geonomeae
Geonoma interrupta (R. & P.) Mart. Henderson 30
Welfia regia H. Wendl. Henderson 301

VI. PHYTELEPHANTOIDEAE
Phytelephas seemanii O. F. Cook Chavez 950
Phytelephas tenuicaulis (Barfod) Henderson Chavez 951

* All vouchers are deposited at The New York Botanical Garden.
COPYRIGHT 2006 New York Botanical Garden
No portion of this article can be reproduced without the express written permission from the copyright holder.
Copyright 2006 Gale, Cengage Learning. All rights reserved.

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Author:Henderson, Flor M.
Publication:The Botanical Review
Geographic Code:1USA
Date:Oct 1, 2006
Words:18215
Previous Article:Xeric limestone prairies of eastern United States: review and synthesis.
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