DEVELOPMENT OF FOLIAR DIAPHRAGMS IN SPARGANIUM EURYCARPUM

Three types of diaphragms are produced in regular sequence by the basal intercalary meristem in the leaf of Sparganium eurycarpum Engelm. (Sparganiaceae). They bridge compartments formed by the collapse and disintegration of rib meristem derivatives. The adaptive nature of diaphragms, intercalary meristems, and linear photosynthetic organs is considered for emergent aquatic plants.     

DIAPHRAGMS AND AERENCHYMA IN SCIRPUS VALIDUS

After the short-lived apical meristem ceases activity, a basal intercalary meristem produces all new tissues in the aerial internode of Scirpus validus Vahl. These include extensions of the original vascular system and of the original partitioning walls as well as new vascular bundles and new walls which are produced in a predictable pattern. Diaphragms begin to differentiate well within the intercalary meristem. At first their cells are indistinguishable from those that will become the aerenchyma, but they undergo segmentation and form packets of daughter cells. Such continued mitotic activity allows the diaphragms to expand with the increasing girth of the stem above the intercalary meristem. Aerenchyma cells between the diaphragms become stellate by being stretched as the cells of the vertical partitions divide and enlarge in and above the intercalary meristem.     

COMPARATIVE FOLIAR HISTOGENESIS IN ACORUS CALAMUS AND ITS BEARING ON THE PHYLLODE THEORY OF MONOCOTYLEDONOUS LEAVES

A comparative histogenetic investigation of the unifacial foliage leaves of Acorus calamus L. (Araceae; Pothoideae) was initiated for the purposes of: (1) re-evaluating the previous sympodial interpretation of unifacial leaf development; (2) comparing the mode of histogenesis with that of the phyllode of Acacia in a re-examination of the phyllode theory of monocotyledonous leaves; and (3) specifying the histogenetic mechanisms responsible for morphological divergence of the leaf of Acorus from dorsiventral leaves of other Araceae. Leaves in Acorus are initiated in an orthodistichous phyllotaxis from alternate positions on the bilaterally symmetrical apical meristem. During each plastochron the shoot apex proceeds through a regular rhythm of expansion and reduction related to leaf and axillary meristem initiation and regeneration. The shoot apex has a three- to four-layered tunica and subjacent corpus with a distinctive cytohistological zonation evident to varying degrees during all phases of the plastochron. Leaf initiation is by periclinal division in the second through fourth layers of the meristem. Following inception early growth of the leaf primordium is erect, involving apical and intercalary growth in length as well as marginal growth in circumference in the sheathing leaf base. Early maturation of the leaf apex into an attenuated tip marks the end of apical growth, and subsequent growth in length is largely basal and intercalary. Marked radial growth is evident early in development and initially is mediated by a very active adaxial meristem; the median flattening of this leaf is related to accentuated activity of this meristematic zone. Differentiation of the secondary midrib begins along the center of the leaf axis and proceeds in an acropetal direction. Correlated with this centralized zone of tissue specialization is the first appearance of procambium in the center of the leaf axis. Subsequent radial expansion of the flattened upper leaf zone is bidirectional, proceeding by intercalary meristematic activity at both sides of the central midrib. Procambial differentiation is continuous and acropetal, and provascular strands are initiated in pairs in both sides of the primordium from derivatives of intercalary meristems in the abaxial and adaxial wings of the leaf. Comparative investigation of foliar histogenesis in different populations of Acorus from Wisconsin and Iowa reveals different degrees of apical and adaxial meristematic activity in primordia of these two collections: leaves with marked adaxial growth exhibit delayed and reduced expression of apical growth, whereas primordia with marked apical growth show, correspondingly, reduced adaxial meristematic activity at equivalent stages of development. Such variations in leaf histogenesis are correlated with marked differences in adult leaf anatomy in the respective populations and explain the reasons for the sympodial interpretation of leaf morphogenesis in Acorus and unifacial organs of other genera by previous investigators. It is concluded that leaf development in Acorus resembles that of the Acacia phyllode, thereby confirming from a developmental viewpoint the homology of these organs. Comparison of development with leaves of other Araceae indicates that the modified form of the leaf of Acorus originates through the accentuation of adaxial and abaxial meristematic activity which is expressed only slightly in the more conventional dorsiventral leaf types in the family.     

A CALAMITEAN SHOOT APEX FROM THE PENNSYLVANIAN OF IOWA

Melchior , Robert C., and John W. Hall . (U. Minnesota, Minneapolis.) A calamitean shoot apex from the Pennsylvanian of Iowa. Amer. Jour. Bot. 48(9): 811–815. Illus. 1961.—A shoot apex of a calamitean stem is described from the Des Moines Series, Middle Pennsylvanian. Internodal elongation of the 7 preserved internodes follows a sigmoid curve. A large apical cell has produced derivatives in a fashion apparently comparable to those in Equisetum arvense, except for the number of cells in the first leaf primordium ring and, possibly, the intercalary meristem. Pith meristem developed close to the apical cell. Data from internodal cell elongation of hypodermal cells of the cortex are presented which demonstrate intercalary internodal growth; no intercalary meristems are preserved and the existence of intercalary meristems which might have produced a jointed stem like that of Equisetum is only inferred.     

Development of the intercalary meristem in Chorda filum (Laminariales, Phaeophyceae) and other primitive Laminariales

Development of the intercalary meristem in the terete laminarialean species Chorda filum (L.) Stackhouse was studied in culture using light and transmission electron microscopy as well as by tracing elongation and cell divisions in various parts of the sporophyte. Growth of C. filum sporophytes could be classified into three developmental stages: (i) diffuse growth; (ii) basal meristematic growth; and (iii) intercalary meristematic growth. In the diffuse growth stage, elongation and cell division frequency were almost the same in each cell. In the basal meristematic growth stage, elongation and division of cells became localized in the tissues derived from the meristematic initial cell. Cells of the basal meristematic region contained smaller chloroplasts and many small opaque vesicles. In the intercalary meristematic growth stage, there was further elongation and differentiation of cells originating from the meristematic region, and this became more active in adjacent regions below the meristem than in regions above the meristem, causing the relative position of the intercalary meristem to shift towards the tip of the sporophyte. Meristematic cells of C. filum contained well-developed Golgi vesicles around the nucleus (perinuclear Golgi), many secretion vesicles and many small disk-shaped chloroplasts whose thylakoids were not well developed. Sporophytes of three other terete members of Laminariales, Chorda tomentosa Lyngbye, Pseudochorda nagaii (Tokida) Kawai et Kurogi, and Pseudochorda gracilis Kawai et Nabata, show diffuse growth and basal meristematic growth, but no intercalary meristematic growth. This suggests that the common ancestor of the Pseudochordaceae and Chordaceae had basal meristematic growth, and intercalary meristematic growth evolved more recently in C. filum.     

Carbohydrate metabolism in leaf meristems of tall fescue : I. Relationship to genetically altered leaf elongation rates

The physiological bases for genetic differences in leaf growth rates were examined in two genotypes of tall fescue (Festuca arundinacea Schreb.) selected for a 50% difference in leaf elongation rate. Genotypes had similar dark respiration rates and concentrations of carbohydrate fractions in the leaf meristem and in each daily growth segment above the meristem. Dark respiration rates and concentrations of nonreducing sugars, fructans, and takadiastase-soluble carbohydrates were highest in leaf intercalary meristems and declined acropetally with tissue age. Concentrations of reducing sugars were 1.0% of dry weight in leaf meristems, 3.7% of dry weight in tissue adjacent to the meristem, then decreased progressively with distance from the meristem. Glucose, fructose, and myo-inositol comprised over 90% of the monosaccharides present in leaf meristems. Soluble protein concentration was 9.7 milligrams per gram fresh weight in leaf meristems, 5.5 milligrams per gram in tissues immediately above the meristem and, thereafter, increased linearly with distance from the meristem.

Leaf meristems of the genotype exhibiting rapid leaf elongation contained 30% more soluble protein than those of the genotype selected for slow leaf elongation. The 4-fold difference in size of the leaf meristem appeared to be more important in influencing leaf elongation than were other characteristics examined.

     

COMPARATIVE MORPHOGENESIS OF THE DIMORPHIC LEAVES OF CYAMOPSIS TETRAGONOLOBA

Cyamopsis tetragonoloba, guar, produces two leaf forms: a simple leaf and a trifoliate leaf. The steps in the development of each of these forms have been investigated in an attempt to determine the precise point at which the leaf primordium becomes destined to produce one or the other characteristic leaf shapes. Up to 140 μ in length the leaf primordia are morphologically indistinguishable. If a simple leaf is to be formed the marginal meristem remains continuous and initiates only lamina. If a trifoliate leaf is to be formed the continuity of the marginal meristem is interrupted by a group of “pocketal” cells dividing it into an upper lamina meristem and a basal leaflet buttress.     

PHYLLODE THEORY IN RELATION TO LEAF ONTOGENY IN SANSEVIERIA TRIFASCIATA

Shoot apices of Sansevieria trifasciata have a three-layered mantle, a zone of subapical initials, a central meristem, and a peripheral meristem. Leaf initiation begins with periclinal divisions in L-3 and is followed by periclinal divisions in L-2 and anticlinal divisions in L-l. At first, the primordium is a mound of tissue at one point on the flank, but it soon takes the form of a low ridge encircling the apex. An ephemeral adaxial meristem differentiates in L-2 of the primordium when it is about 50 μ high and is active until the primordium is about 450 μ high. Then it ceases basipetally and is not observable after the primordium is about 600μ high. As the adaxial meristem ceases at the base of the radial tip, its two lateral regions become the submarginal meristems of the expanded portion. Marginal meristems differentiate from the protoderm, and oblique-anticlinal divisions of the marginal initials result in the formation of an abaxial and adaxial epidermis. These derivatives undergo a few anticlinal divisions, increasing marginal width, and then they divide periclinally, increasing marginal thickness. After the primordium is about 600-700 μ high it continues to grow in length by a diffuse basal intercalary meristem. When the leaf is 3 dm long, an adaxial rounding meristem differentiates in the region just above the sheath. Leaf vasculature consists of parallel bundles which anastomose acropetally. Vascular bundles are arranged in a semicircle in the expanded portion and in a circle in the radial tip. There is one centrally located bundle at the apex as a result of lateral anastomoses. Present evidence from leaf ontogeny and mature vasculature in S. trifasciata is interpreted as supporting the concept that the liliaceous leaf is homologous with the phyllodes of A corns and Acacia.     

Developmental study onHypolepis punctata (Thunb.) Mett.

The third petiolar bud ofHypolepis punctata appears on the basiscopic lateral side of the petiole above the fairly developed first petiolar bud. This investigation clarified the fact that the third bud is formed neither by the activity of the meristem of the first bud nor by the meristem directly detached from the shoot apical meristem, but is initiated in the cells involved in the abaxial basal part of the elevated portion of the leaf primordium. Thus the third bud is of phyllogenous origin. This investigation further revealed that the cells to initiate the third bud are originally located in the abaxial side of the leaf apical cell complex like the cells to initiate the first bud, but are not incorporated into the meristem of the first. After the first, second and third petiolar buds have been initiated, they are carried up into fairly high regions on the petiolar base by the intercalary growth which occurs in the leaf base below the insertion level of the first and the second buds.     

ADAPTIVE LEAF ARCHITECTURE IN EMERGENT AND FLOATING SPARGANIUM

The robust emergent leaves of Sparganium eurycarpum and S. americanum are supported by corner fiber masses and large bundle sheaths, but the thin floating leaves of S. fluctuans and S. minimum have only moderate bundle sheaths. In emergent types heavily photosynthetic diaphragms bearing vascular bundles are separated from each other in the leaf compartments by three lightly photosynthetic diaphragms without bundles, but in floating types only every other heavily photosynthetic diaphragm has a bundle. Palisade chlorenchyma occurs only at aerial surfaces—abaxial and adaxial in emergent leaves, but only adaxial in floating leaves. Extra photosynthetic areas are provided in emergent leaves by concentrations of chlorenchyma in limited areas on interior partitioning walls, while the remainder of the walls is translucent. Since only 25 % of the diaphragms are heavily photosynthetic, and the others essentially transparent because of their diffusely distributed chloroplasts and large intercellular spaces, a sieve effect exists which allows even the interior parts of thick emergent leaves to photosynthesize.     

Stem structure and its bearing on the systematics of Pleurothallidinae (Orchidaceae)

STERN, W. L., PRIDGEON, A. M. & LUER, C. A., 1985. Stem structure and its bearing on the systematics of Pleurothallidinae (Orchidaceae) . Pleurothallids comprise a subtribe of numerous orchids consisting of mostly diminutive New World epiphytes. The cauline system involves a branching rhizome bearing unifoliate ramicauls. The inflorescence usually originates from an adaxial invagination toward the apex of the ramicaul. Two or more cycles of vascular bundles traverse the ramicaul; typically, there is an outer, branching series of smaller strands and an inner, non-branching series of larger strands. In the ultimate internode subtending the leaf the smaller, outer bundles migrate between the larger, inner bundles and become aggregated in the medullary region. They bend adaxially to vascularize the inflorescence. The leaf is articulated at an abscission layer to the prolongated apex of the ramicaul more or less distal to the cauline invagination. In some pleurothallids, a cauline annulus appears at the insertion of the inflorescence and cither below or at the attachment of the leaf to the apex of the ramicaul. Anatomical studies show that the annulus is the external manifestation of an intercalary meristem. Except for Pleurothallis , genera are consistent in having or lacking an annulus. Genera with two pollinia either possess or lack the annulus; those with four to eight pollinia always lack the annulus. It is suggested that the intercalary meristem is a derived characteristic providing survival value and a selective advantage to those pleurothallids in which it occurs. Presence of the intercalary meristem is correlated with other structural features considered to represent specializations.     

DEVELOPMENT OF THE EPIPHYLLOUS INFLORESCENCE OF HELWINGIA JAPONICA (HELWINGIACEAE)

The inflorescence of Helwingia japonica (Thunb.) Dietr. is initiated adjacent to the leaf axil on the adaxial side of the base of a leaf primordium during its second plastochron. The inflorescence which develops from the resulting primordium comes to be situated on the midrib of the mature fertile leaf, through the action of a basal, intercalary meristem. In fertile leaves this meristem develops beneath, as well as above, the insertion of the inflorescence primordium on the leaf primordium. The same meristem is present in sterile leaves as well. A separate, adaxial vascular bundle departs from the leaf trace in the base of the petiole and leads to the inflorescence, in the mature fertile leaf. This adaxial vascular bundle is absent in sterile leaves. It is argued that the vascular anatomy does not conclusively confirm the hypothesis that the epiphyllous inflorescence is the congenital fusion product of a leaf and an axillary inflorescence. Instead, it is suggested that the interplay of changes in the position of primordium initiation, and intercalary growth, offers an ontogenetic explanation of the situation, which in turn may be related to the phylogeny of the species in question. It appears to be misguided and futile to look for homologies (i.e., 1:1 correspondences) between fertile and sterile leaves, since 1:1 correspondences do not exist in this case.     

LEAF DEVELOPMENT IN SENECIO ROWLEYANUS (COMPOSITAE)

Prolonged apical growth of the leaf primordium and the presence of distinct marginal meristems do not occur in Senecio rowleyanus. Intercalary cell divisions accompanied by radial expansion of derivatives from an adaxial meristem account for the spherical shape of the leaf. The “window” in the lamina marks the position of the adaxial meristem and precludes interpretation of the leaf as being unifacial. Stomata are mesoperigenous and anomocytic in type. Schizogenous secretory canals occur in both the leaf and the stem, and their association with vascular bundles is discussed. The anatomy of the leaf is interpreted in terms of xeromorphy.     

Comparative development of heavily asymmetric-cordate gametophytes of <Emphasis Type="Italic">Anemia phyllitidis</Emphasis> (Anemiaceae) focusing on meristem behavior

Development of heavily asymmetric cordate gametophytes of Anemia phyllitidis (Anemiaceae), one of the schizaeoid ferns, was examined using a sequential observation technique; epi-illuminated light micrographs of the same growing gametophytes were taken approximately every 24 h. The apical cell-like wedge-shaped cell was produced once from the terminal cell of a germ filament, but it stopped dividing soon after production of one or two derivative cells. Without a functional apical cell, the gametophyte developed by intercalary growth until the early stage of wing formation, and then the multicellular (pluricellular) meristem arose from the lower lateral side of the gametophyte. This was in sharp contrast to the observation that the multicellular meristem forms in place of the apical cell in typical cordate gametophytes. Loss of the functional apical cell probably caused a site-shift in the multicellular meristem of the Anemia phyllitidis gametophyte during evolution from apical to lateral. The results suggest that apical cell-based and multicellular meristems are primarily independent of each other. The multicellular meristem produced cells equally in the distal and proximal directions to form wings in both directions but proximally produced cells divided much less frequently. As a result, a heavily asymmetric gametophyte was formed.     

LEAF DIMORPHISM IN THE AQUATIC ANGIOSPERM CALLITRICHE HETEROPHYLLA

Depending on the position of the shoot tip relative to the water surface, the aquatic angiosperm Callitriche heterophylla produces either ovate land-form or linear water-form leaves. This paper is concerned with the developmental basis for the leaf dimorphism of this species. Little significant difference is observed between the apical meristems of submerged vs. emergent shoots; moreover, land-form and water-form primordia undergo similar, if not identical, patterns of initial development until they attain a length of 350 to 400 μm. These findings are interpreted to mean that the divergent leaf forms result from the marked sensitivity of the primordia to their respective environments rather than from the mode of their inception. Subsequent growth of the young water-form leaf emphasizes longitudinal extension, while the immature land-form leaf continues balanced expansion in both longitudinal and lateral directions. The lateral growth of the land-form primordium is accomplished in part by a more persistent marginal meristem, but the morphological difference between the two leaf forms is mostly attributable to the difference in the predominant direction of intercalary expansion. In addition, certain anatomical features, such as vasculature, stomates, and cuticle, are much more prominent in mature land-form leaves than in water-form leaves. These anatomical differences seem to represent structural adaptations of each leaf form to the specific physiological requirements of its environment.     

The floral anatomy of Victoria Schomb. (Nymphaeaceae)*

The vasculature and development of the flower of Victoria Schomb. are described. The vasculature is basically similar to that found in other genera of the Nymphaeaceae sensu stricto (e.g. Nymphaea L. and Nuphar Sm.). The early development of the flower is similar to that of a hypogynous flower, but meristematic activity shifts from the apex to the periphery in the form of an intercalary ring meristem. The innermost appendicular organs, including the gynoecium, arise by differentiation of tissues formed by this intercalary ring meristem. Evidence is assembled from the mature vasculature and developmental studies: (a) to refute Troll's interpretation that receptacular strips of tissue occur between the carpels and that the outer ovary wall is totally receptacular; (b) to propose that the occurrence of epeltate carpels in Victoria, as correctly described by Troll, has been phylogenetically ‘read’ in the wrong direction; (c) to propose that the flower of Victoria has evolved by (1) the adnation and connation of the proximal portions of the appendicular organs which now envelop the syncarpous gynoecium and (2) the concomitant condensation from a primitive ranalian floral apex.     

ENDOMORPHIC AND ECTOMORPHIC CHARACTERS IN PELECYPHORA AND ENCEPHALOCARPUS

Boke , Norman H. (U. Oklahoma, Norman.) Endomorphic and ectomorphic characters in Pelecyphora and Encephalocarpus. Amer. Jour. Bot. 46(3) : 197-209. Illus. 1959.—Outstanding ectomorphic characters of Pelecyphora valdeziana include its small size; pectinate, hairy spines; broad, truncate, floral buds; dehiscent, berry-like fruits; and black, tuberculate seeds. The leaves are vestigial, and although the areole meristem originates on the adaxial face of the tubercle primordium, it is soon elevated to the summit by intercalary growth. The first primordium of the single, elliptical series of spines is initiated immediately in front of the rudimentary leaf. Others form in acropetal sequence on either side of the areole meristem. The last ones form across the areole, leaving a meristem, which may be floral or vegetative, on the anterior side. Whether areoles of P. valdeziana can be considered dimorphic is doubtful. However, they approach the type of dimorphism found in Epithelantha. Pelecyphora aselliformis has acuminate floral buds; dry, papery fruits; and brown, curved, reticulate seeds. The leaves are reduced almost to extinction. The areole meristem becomes separated into spiniferous and axial portions early in ontogeny, but the 2 parts remain connected by a band of trichomes, which probably represents a vestigial groove. The axial meristem may be reproductive or vegetative. The sequence of spine initiation in P. aselliformis is unusual in that it begins at the anterior side of the spiniferous meristem and proceeds toward the posterior side. Areoles in this species are clearly dimorphic, much as in the mammillarias, but the vestigial groove is reminiscent of Coryphantha and related genera. Although adult specimens of Encephalocarpus strobiliformis bear scale-like tubercles, which are very different from the laterally compressed tubercles of P. aselliformis, their flowers, fruits, and seeds are almost identical. The two species share the same type of areole dimorphism, including the vestigial groove. Tubercles on seedlings and young branches of E. strobiliformis are prismatic rather than scale-like. Since they tend to be laterally compressed at the summit and bear elliptical areoles with many more spines than the adult, they resemble seedling tubercles of P. aselliformis. Tubercles on adult specimens likewise resemble each other in the structure of the epidermis and hypodermis. It does not seem possible that P. valdeziana can be retained in the genus Pelecyphora. If seed structure has any systematic value, the species belongs in or near the genus Thelocactus, to which it was assigned by Bravo. Pelecyphora aselliformis and Encephalocarpus strobiliformis, on the other hand, share so many important characters that they could well be considered cogeneric. Both seed structure and the rudimentary grooves on the tubercles suggest that their affinities may lie with certain coryphanthas or mammillarias rather than with Ariocarpus and Epithelantha.     

Carbohydrate Metabolism in Leaf Meristems of Tall Fescue : II. Relationship to Leaf Elongation Rates Modified by Nitrogen Fertilization

Our objective was to examine alterations in carbohydrate status of leaf meristems that are associated with nitrogen-induced changes in leaf elongation rates of tall fescue (Festuca arundinacea Schreb.). Dark respiration rates, concentrations of nonstructural carbohydrates, and soluble proteins were measured in leaf intercalary meristems and adjacent segments of elongating leaves. The two genotypes used differed by 43% in leaf elongation rate. Application of high nitrogen (336 kilograms per hectare) resulted in 140% higher leaf elongation rate when compared to plants receiving low nitrogen (22 kilograms per hectare). Leaf meristems of plants receiving high and low nitrogen had dark respiration rates of 5.4 and 2.9 microliters O₂ consumed per milligram structural dry weight per hour, respectively. Concentrations of soluble proteins were lower while concentrations of fructan tended to be slightly higher in leaf meristems of low-nitrogen plants when compared to high-nitrogen plants. Concentrations of reducing sugars, nonreducing sugars, and takadiastase-soluble carbohydrate of leaf meristems were not affected by nitrogen treatment. Total nonstructural carbohydrates of leaf meristems averaged 44 and 39% of dry weight for low- and high-nitrogen plants, respectively. Within the leaf meristem, approximately 74 and 34% of the pool of total nonstructural carbohydrate could be consumed per day in high- and low-nitrogen plants, respectively, assuming no carbohydrate import to the meristem occurred. Plants were able to maintain high concentrations of nonstructural carbohydrates in leaf meristems despite a 3-fold range in leaf elongation rates, suggesting that carbohydrate synthesis and transport to leaf intercalary meristems may not limit leaf growth of these genotypes.     

EARLY HISTOGENESIS OF THE ADULT LEAVES OF DARLINGTONIA CALIFORNICA (SARRACENIACEAE) AND ITS BEARING ON THE NATURE OF EPIASCIDIATE FOLIAR APPENDAGES

In order to assess the validity of various interpretations of tubular leaves of angiosperms, a histogenetic study of the ontogeny of adult leaves of Darlingtonia californica was undertaken. The adult leaf of Darlingtonia is characterized by a sheathing leaf base, an elongate ascidium, an overarching hood, and two “fishtail” appendages which arise near the leaf apex. A keellike growth, with two rows of alternate vascular bundles, traverses the tube from base to mouth. Ontogenetic studies show that the primordium arises by a monopodial rather than a sympodial mode of growth as previously reported. After the formation of a small, erect primordium, a restricted adaxial meristem is initiated that expands both adaxially and upwards. This “querzone” serves, in effect, to congenitally combine the two primordial margins during its extension. Growth and maturation of the subjacent portions cause tubular elongation in the leaf. Primordial apical divisions are later replaced by more general intercalary growth with acropetal and centrifugal maturation. The hood and fishtails are established early in ontogeny by adaxial growth of the primordial apex and subsequent activation of juxtaposed localized meristems. Comparative morphology has established that the epiascidiate leaf is a foliar appendage that undergoes certain specific morphogenetic modifications. It has a structural relationship to ensiform appendages of Acacia and Acorus as well as to peltate foliar organs. The early ontogeny of Darlingtonia leaves is considered to be homologous with other epiascidiate foliar organs, including some supposedly primitive carpels.