THE GENUS PELECYPHORA (CACTACEAE): RESOLUTION OF A CONTROVERSY

The four species considered in this article fall into two natural groups. The evidence presented for this conclusion involves not only external features such as floral, fruit, and seed morphology, but also that obtained from internal structure, including tubercle and areole development, the anatomy of surface layers of mature tubercles, the structure of crystals in the ground tissues, and features of tracheary elements. When all evidence is considered, it becomes apparent that Pelecyphora aselliformis and Encephalocarpus strobiliformis are cogeneric as are P. valdeziana and P. pseudopectinata. Yet, the two cogeneric pairs are not closely related to each other. We do not propose to place them in any phylogenetic scheme as yet because general knowledge of the family Cactaceae is still too fragmentary. The first pair is assigned to Pelecyphora, of which a formal taxonomic treatment is presented, and the second pair is tentatively assigned to Thelocaclus.     

AREOLE DIMORPHISM IN CORYPHANTHA

Boke , Norman H. (U. Oklahoma, Norman.) Areole dimorphism in Coryphantha. Amer. Jour. Bot. 48(7): 593–603. Illus. 1961.—Coryphantha clava and C. erecta have both grooved and grooveless tubercles. The latter occur on young plants and emergent branches and may occur mixed with grooved tubercles on adult shoots, especially in C. erecta. Grooved tubercles have monomorphic areoles and are the only ones which bear flowers. The dimorphic areoles on grooveless tubercles develop exactly as in the mammillarias. In both species capitate glands occur in the axils of most tubercles and may occur at any position along an areole groove. The glands are initiated by the areole meristem in the same manner as spines, with which they are homologous. Close relationship between the 2 species is substantiated by both ectomorphic and endomorphic characters. The possession of both monomorphic and dimorphic areoles may mean that these cacti are evolving toward the complete areole dimorphism manifested by the mammillarias.     

ANATOMY AND DEVELOPMENT IN SOLISIA

Boke , Norman H. (U. Oklahoma, Norman.) Anatomy and development in Solisia. Amer. Jour. Bot. 47(1): 59—65. Illus. 1960.–The genus Solisia contains a single species of small cacti which resemble certain mammillarias in having: pink, lateral flowers; milky juice; and dimorphic areoles. Adult specimens have elongate spiniferous areoles with an unusual sequence of spine initiation. The first four or five primordia appear at the posterior end of the areole meristem; the next are initiated near its center, after which initiation proceeds both acropetally and basipetally until the single, elliptical series of primordia is complete. A similar pattern of spine initiation occurs in Pelecyphora aselliformis, but this species differs markedly from S. pectinata in other respects. In seedlings of S. pectinata the areoles are broadly elliptical and spine initiation is strictly acropetal, a situation found in certain species of Mammillaria. Seeds of S. pectinata are black with a large hilum and a small perisperm. Since the perisperm has apparently been overlooked, it appears that only the black seed coat and relatively large hilum keep the species out of Mammillaria. If Buxbaum's postulates concerning the value of seed structure in tracing phylogeny in the Cactaceae are valid, S. pectinata must have diverged from the main line of evolution somewhere below Neobesseya. In that event, the species probably merits generic rank; otherwise, it seems preferable to return it to Mammillaria.     

STRUCTURE AND DEVELOPMENT OF THE SHOOT IN DOLICOTHELE

Boke , Norman H. (U. Oklahoma, Norman.) Structure and development of the shoot in Dolicothele. Amer. Jour. Bot. 48(4): 316–321. Illus. 1961.—A study of 2 species of Dolicothele reveals that although they have dimorphic areoles and a pattern of spine development similar to those of certain mammillarias, they share a significant number of ectomorphic and endomorphic characters with coryphanthas of the “vivipara group.” These include a tendency toward a cespitose habit; relatively large flowers; green fruits; pitted seeds; medullary vascular systems; forking of the main tubercle traces in the bases of the tubercles; lack of mucilage cells; thin-walled epidermis and hypodermis, both devoid of crystals; and large, druse-like crystalline aggregates in older parts of the pith and cortex. The evidence suggests that Coryphantha vivipara and closely allied species are the nearest extant relatives of Dolicothele. It would, therefore, seem inconsistent to return Dolicothele to Mammillaria unless an author's viewpoint were so conservative that he was willing also to return most, if not all, coryphanthas and escobarias to that genus.     

How and why does the areole meristem move in Echinocereus (Cactaceae)?

Background and Aims In Cactaceae, the areole is the organ that forms the leaves, spines and buds. Apparently, the genus Echinocereus develops enclosed buds that break through the epidermis of the stem adjacent to the areole; this trait most likely represents a synapomorphy of Echinocereus. The development of the areole is investigated here in order to understand the anatomical modifications that lead to internal bud development and to supplement anatomical knowledge of plants that do not behave according to classical shoot theory.Methods The external morphology of the areole was documented and the anatomy was studied using tissue clearing, scanning electron microscopy and light microscopy for 50 species that represent the recognized clades and sections of the traditional classification of the genus, including Morangaya pensilis (Echinocereus pensilis).Key Results In Echinocereus, the areole is sealed by the periderm, and the areole meristem is moved and enclosed by the differential growth of the epidermis and surrounding cortex. The enclosed areole meristem is differentiated in a vegetative or floral bud, which develops internally and breaks through the epidermis of the stem. In Morangaya pensilis, the areole is not sealed by the periderm and the areole meristem is not enclosed.Conclusions The enclosed areole meristem and internal bud development are understood to be an adaptation to protect the meristem and the bud from low temperatures. The anatomical evidence supports the hypothesis that the enclosed bud represents one synapomorphy for Echinocereus and also supports the exclusion of Morangaya from Echinocereus.     

A STUDY OF THE PROPOSED GENUS OBREGONIA (CACTACEAE)

The monotypic genus Obregonia Frič was compared with Ariocarpus, Lophophora, Strombocactus, and certain other cactus genera. Obregonia and Ariocarpus are similar in characters of seeds, seedlings, habitat, fruits, and general habit. They differ in time of flowering, point of flower origin, areole structure, presence or absence of druses and a mucilage system, and presence or absence of spines. Obregonia and Lophophora are similar in characters of seeds, habitat, basic areole structure, and fruits. They differ in seedling form, habit of adult plants, presence or absence of spines, and chemical analysis. Strombocactus resembles Obregonia in few ways except in basic areole structure and some aspects of anatomy. The author concludes that Obregonia is an intermediate form between Ariocarpus and Lophophora and deserves generic rank. A formal taxonomic treatment of the genus follows the conclusions.     

Cotyledon areoles in Bobgunnia (Fabaceae: Swartzieae)

Cotyledon areoles, a seed character unique to papilionoid legumes, are reported for the first time in the legume tribe Swartzieae, an anomalous group sometimes considered to be intermediate between subfamilies Caesalpinioideae and Papilionoideae. Bobgunnia madagascariensis has a linear, slightly branched cotyledon areole on the abaxial surface of each cotyledon. Previous reports of other papilionoid seed features in this species are confirmed and extended. These highly distinctive papilionoid seed structures, and recent reports from floral ontogeny and molecular systematics which show Bobgunnia to be closely related to Swartzia and several other genera of Swartzieae, add further evidence to assertions that Swartzia seed structures are derived by simplification from a papilionoid seed, and not by retention of less specialized caesalpinioid seed features. The function and evolutionary advantage of cotyledon areoles are unknown and need investigation. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 287–291.     

A REVISION OF ARIOCARPUS (CACTACEAE). I. THE STATUS OF THE PROPOSED GENUS ROSEOCACTUS

Anderson, Edward F. (Claremont Graduate School, Claremont, California.) A revision of Ariocarpus (Cactaceae). I. The status of the proposed genus Roseocactus. Amer. Jour. Bot. 47(7) : 582–589. Illus. 1960.—The proposed genus Roseocactus Berger is found to be a subgenus of Ariocarpus. The subgenera Ariocarpus and Roseocactus are basically similar in habitat, seedlings, presence of mucilage systems, tubercle structure, seed structure, fruiting habit, flower origin and structure, alkaloidal properties, trichomes, pollen grains, and chromosome number. They differ in flower color, presence or absence of an areolar groove, tubercle divergence, and trichome texture. A hypothesis is proposed to explain variations in the areoles. The differences in structure of mature tubercles are thought to be due to variability in the location of growth and elongation with respect to the floral and spinous areolar areas.     

Morphology and anatomy of the Cephalocereus columna-trajani cephalium: why tilting?

The morphology and anatomy of the Cephalocereus columna-trajani flowering region was described and compared with data on other species. The vegetative and reproductive regions were described in detail. The results showed that after the flowering region is differentiated, morphological changes take place which are correlated with anatomical changes. The flowering region in this species is termed a lateral cephalium because of its reduced interareolar space, increased areole size and abundant long bristles and trichomes in the areoles. Periderm development near the apical meristem, lack of chlorenchyma and a delay in xylem fiber differentiation are also traits characteristic of a lateral cephalium. The lateral cephalium of C. columna-trajani shared the same combination of morpho-anatomical characters with its sister taxon, C. senilis, except for the number of ribs in the cephalium. Both species survive in high temperature environments and their cephalium faces north; however, only C. columna-trajani tilts, thus we hypothesize that incorporation of fewer ribs associated with periderm development in the cephalium contributes to stem tilting.     

HORMONAL CONTROL OF ORGANOGENESIS IN OPUNTIA POLYACANTHA (CACTACEAE)

An axillary bud (areole) of Opuntia polyacantha is composed of an extremely short axis which bears highly modified leaves (spines). After producing the spines, the bud apical meristem becomes quiescent. When the axillary bud is excised and cultured with benzylaminopurine (BAP), the apical meristem increases in complexity and produces leaves on an elongated axis. Gibberellic acid (GA) induces spine production with no elongation of the axis. Naphthaleneacetic acid (NAA) causes no structural change in the meristem but induces root formation in adjacent tissue. The initiation of roots, spines, or leaves is visible within three days. Explants on medium lacking hormones show no structural change, but by the tenth day are insensitive to BAP or NAA. These aged explants can be made to respond to BAP or NAA by merely rewounding them at the time of hormone application. With concentrations previously found to be optimal for single hormone responses, NAA in combination with BAP prevents shoot formation, or in combination with GA prevents spine production. When all three hormones are combined, spines are produced, indicating that GA is active morphologically while BAP and NAA counteract each other. If the level of NAA is decreased, BAP counteracts the NAA and also overrides the GA and leafy shoots are produced. By varying the proportions of BAP and GA, in the absence of NAA, four types of lateral appendage can be produced: leaves, mildly altered leaves, leaf-spine transition forms, and spines.     

Surface topography of the tegument of adult Schistosoma nasale Rao, 1933 from Sri Lanka

Scanning electron microscopical studies of adult male and female Schistosoma nasale are reported. The tubercles on the dorsal and dorso-lateral surfaces of unpaired male S. nasale are devoid of spines. In paired male worms the tubercles on the dorsal surface are large and also are devoid of spines, but some tubercles on the dorso-lateral surface possess spines. Pit-like openings are visible on the surface of the smooth tubercles. The oral and ventral suckers on the male worm are well developed and are invested with spines, as are the gynaecophoric canal and flap. Ciliated sensory receptors are distributed over the surface of the male worm. The oral and ventral suckers of the female worm are much smaller than those of the male: spines occur on both suckers. The surface of the female is non-tuberculate and is thrown into transverse folds. Pit-like openings are visible at higher magnifications. The anterior end of the female is heavily invested in ciliated receptors, whereas the posterior end is heavily spined. The surface topography of S. nasale is discussed in relation to other species of Schistosoma.     

Quantitative survey of sieve tube distribution in foliar terminal veins of ten dicot species

Quantitative data on sieve tubes in foliar terminal veins (vein endings) were added to the meager published information from only five dicot species. Correlations with other minor vein configurations were also explored. Leaf samples from ten species of dicots (Oxalis nelsonii, O. pes-capri, O. rubra, O. stricta [Oxalidaceae], Caesalpinia pulcherrima. Glycine max, Trifolium repens [Leguminosae], Ampelamus albidus [Asclepidaceae], Eupatorium rugosum [Asteraceae], and Polygonum convolvulus [Polygonaceae]) were selected for two quantitative procedures: 1) a survey of the arrangement of terminal veins and distribution of sieve tubes in terminal veins in 100 areoles per species using stained leaf clearings; and 2) a search for correlations of sieve tube distribution with number and branching patterns of terminal veins, and with sizes of areoles using image analysis. Two Oxalis species (O. pes-capri and O. stricta) had the smallest areoles and virtually no sieve tubes in any terminal vein. Polygonum convolvulus, at the other extreme, had sieve tubes extending to the tips of most terminal veins. The other species had various intermediate sieve tube configurations. The data indicate that species with few or no sieve tubes associated with their terminal veins, regardless of the number of terminal veins per areole, have smaller areoles. These results may have implications regarding the entry of leaf photosynthates into the vascular system.     

SIEVE TUBES IN FOLIAR VEIN ENDINGS: REVIEW AND QUANTITATIVE SURVEY OF RUDBECKIA LACINIATA (ASTERACEAE)

Sieve tube distribution in foliar vein endings of angiosperms is mentioned in 33 publications. Their collective judgment is that sieve tubes may be lacking, may end partway along, or may extend to the vein tip. Most reports conclude that all vein endings are the same within a species, but a few studies mention variation. Only three studies include quantitative data, all based on reconstruction from microtome sections. The present quantitative study, the most extensive one to date, surveyed sieve tubes in vein endings from cleared and stained leaf pieces of Rudbeckia laciniata (Asteraceae, tribe Heliantheae), a North American perennial herb. Of 203 areoles sampled, 32 (15.8%) lacked vein endings whereas 171 (84.2%) had one to six endings per areole. There were 385 individual vein endings in the 171 areoles: 115 lacked sieve tubes, 38 had them to the vein tip, but in most (232) they ended at some intermediate point. Serial cross sections of the latter showed phloem parenchyma continuing beyond the sieve tubes for some distance, beyond which occurred a distal zone consisting solely of tracheary elements. R. laciniata vein endings exhibit the same range of variation reported for dicots in general. More studies combining clearings with selected sections are needed to establish the range of vein ending patterns among angiosperms.     

FLORAL STRUCTURE AND EVOLUTION IN LOPEZIEAE (ONAGRACEAE)

The Lopezieae present an interesting mixture of ancestral and derived characters: some members of the tribe retain the basic onagraceous chromosome number (n = 11), but the flowers are advanced in that they are mostly zygomorphic and always have a two-merous androecium. Species differ in the position of the nectaries, also in the way in which floral parts are united above the inferior ovary. These differences, when analyzed with information from a new monograph of the Lopezieae, provide the basis for a phylogenetic tree. It is inferred that ancestral Lopezieae were bird-pollinated woody perennials with regular flowers, two fertile stamens, and no floral tube distal to the ovary. Evolutionary events accompanying the emergence of modern taxa included abortion of the abaxial stamen (all surviving Lopezieae except Lopezia lopezioides), development of an epigynous floral tube (L. riesenbachia, L. semeiandra), decrease in floral symmetry without conversion to insect pollination (in two independent lines), and decrease in floral symmetry with conversion to insect pollination (in at least two independent lines). The prominent tubercles on upper petals of certain insect-pollinated species apparently evolved from the less prominent swollen areas still present in some of the bird-pollinated species. The tubercles and an associated snapping mechanism arose in response to increasing selection for fly pollination. Densely staining areas in some specimens may be osmophores; if so, scent plays a supplementary role in the orienting of insects to the upper petals. Interstaminal nectaries and the absence of a floral tube link the Lopezieae to Ludwigia; the relationship of these two taxa to Epilobium is presently unclear. Fossil records indicate that the Onagraceae had evolved by the beginning of the Tertiary Period and that the Ludwigia line is very old. The family's ancestral features are retained to a greater degree in Fuchsia, however, than in Ludwigia.     

SUPERMAN regulates floral whorl boundaries through control of auxin biosynthesis

Proper floral patterning, including the number and position of floral organs in most plant species, is tightly controlled by the precise regulation of the persistence and size of floral meristems (FMs). In Arabidopsis, two known feedback pathways, one composed of WUSCHEL (WUS) and CLAVATA3 (CLV3) and the other composed of AGAMOUS (AG) and WUS, spatially and temporally control floral stem cells, respectively. However, mounting evidence suggests that other factors, including phytohormones, are also involved in floral meristem regulation. Here, we show that the boundary gene SUPERMAN (SUP) bridges floral organogenesis and floral meristem determinacy in another pathway that involves auxin signaling. SUP interacts with components of polycomb repressive complex 2 (PRC2) and fine‐tunes local auxin signaling by negatively regulating the expression of the auxin biosynthesis genes YUCCA1/4 (YUC1/4). In sup mutants, derepressed local YUC1/4 activity elevates auxin levels at the boundary between whorls 3 and 4, which leads to an increase in the number and the prolonged maintenance of floral stem cells, and consequently an increase in the number of reproductive organs. Our work presents a new floral meristem regulatory mechanism, in which SUP, a boundary gene, coordinates floral organogenesis and floral meristem size through fine‐tuning auxin biosynthesis.     

INFLORESCENCE AND FLOWER DEVELOPMENT IN THE PIPERACEAE. I. PEPEROMIA

Flowers of Peperomia species are the simplest structurally of any of the members of the Piperaceae. The spicate inflorescences form terminally and in axillary position; in each, the apex first is zonate in configuration with a two-layered tunica while 3-4 leaves are initiated. Later, when the inflorescence apical meristem begins bract initiation, the biseriate tunica persists, but zonal distinctions diminish and the apex can be described in terms of a simple tunicacorpus configuration. The inflorescence apex aborts after producing 30-40 bracts in acropetal succession an abscission layer forms across the base of the apex, and the meristem dries and drops off. Bracts are produced by periclinal divisions in T₂ (and occasionally also in the third layer as well); the later-formed floral apices arise by periclinal divisions in T₂ and the third layer. Each floral apex is at first a long transverse ridge in the axil, perpendicular to the long axis of the inflorescence. This establishes bilateral symmetry in the flower, which persists throughout subsequent growth. The floral meristem becomes saddle-shaped, and two stamen primordia are delimited, one at either end and lower than the central floral apex. A solitary carpel is initiated abaxially, and soon forms a circular rim which heightens as a tube with an apical pore. Within the open carpel, a solitary ovule is initiated from the entire remains of the floral apical meristem; it, hence, is terminal in the flower, and its placentation is basal. Carpellary closure in P. metallica results from accelerated growth of the abaxial lip, and the two margins become appressed. Species differ greatly as to whether the abaxial or the adaxial lobe predominates in late stages of carpel development. In P. metallica, the receptive portion of the stigma forms from the shorter lobe which is overtopped. Stigmatoid tissue forms internal to the receptive stigma. The prevailing bilateral floral symmetry, absence of a perianth, and the spicate inflorescence are features which distinguish Peperomia (and Piperaceae) from the magnolialian line of angiosperms.     

Separation of AG function in floral meristem determinacy from that in reproductive organ identity by expressing antisense AG RNA

The Arabidopsis floral homeotic gene AGAMOUS (AG) is a regulator of early flower development. The ag mutant phenotypes suggest that AG has two functions in flower development: (1) specifying the identity of stamens and carpels, and (2) controlling floral meristem determinacy. To dissect these two AG functions, we have generated transgenic Arabidopsis plants carrying an antisense AG construct. We found that all of the transgenic plants produced abnormal flowers, which can be classified into three types. Type I transgenic flowers are phenocopies of the ag-1 mutant flowers, with both floral meristem indeterminacy and floral organ conversion; type II flowers are indeterminate and have partial conversion of the reproductive organs; and type III flowers have normal stamens and carpels, but still have an indeterminate floral meristem inside the fourth whorl of fused carpels. The existence of type III flowers indicates that AG function can be perturbed to affect only floral meristem determinacy, but not floral organ identity. Furthermore, the fact that floral meristem determinacy is affected in all transformants, but floral organ identity only in a subset of them, suggests that the former may required a higher level of AG activity than the latter. This hypothesis is supported by the levels of AG'mRNA detected in different transformants with different frequencies of distinct types of abnormal antisense AG transgenic flowers. Finally, since AG inhibits the expression of another floral regulatory gene AP1, we examined AP1 expression in antisense AG flowers, and found that AP1 is expressed at a relatively high level in the center of type II flowers, but very little or below detectable levels in the inner whorls of type III flowers. These results provide further insights into the interaction of AG and AP1 and how such an interaction may control both organ identity and floral meristem determinacy.