EARLY LEAF ONTOGENY AND THE SHOOT APEX OF COLOPHOSPERMUM MOPANE (LEGUMINOSAE)

Shoot tips of Colophospermum mopane (Kirk ex Benth.) Kirk ex Léonard produce leaves which at maturity are bifoliate and devoid of stipules. Investigation of their early ontogeny, however, shows that these leaves begin as trifoliate structures partially enclosed by their stipules. The latter are fused along their mid regions, forming a tongue-like “connector.” The lower chamber of this stipule pair harbors the apical meristem while the upper compartment enfolds the two lateral leaflets. The terminal leaflet, histologically resembling the stipules, also fulfills a similar function by covering the top portion of its sister leaflets. Anatomically, the shoot apex displays a pendulum symmetry, with rather steep elevation of that internode portion which subtends the newly formed primordium. Some comparisons with the shoot apex of Hymenaea are drawn.     

Hedyotis austrosinica sp. nov. (Rubiaceae) from southern China: A new heterostylous species similar to H. pulcherrima

Hedyotis austrosinica L. Wu & L.H. Yang is described as a new species from southern China. It is similar to H. pulcherrima, but can be separated by its herbaceous weak to ascending habit with dense pubescence on the surface of most organs, 4‐angled to subterete stems, longer petioles, and longer stipules with 5–9 apical linear lobes. The new species is distributed in Guangdong and Guangxi in southern China.     

Peltate trichomes in the dormant shoot apex of Metrodorea nigra,a Rutaceae species with rhythmic growth

• In Metrodorea nigra, a Rutaceae species with rhythmic growth, the shoot apex in the dormant stage is enclosed by modified stipules. The young organs are fully covered with peltate secretory trichomes, and these structures remain immersed in a hyaline exudate within a hood-shaped structure. Our study focused on the morpho-functional characterization of the peltate trichomes and cytological events associated with secretion.
• Shoot apices were collected during both dormant and active stages and processed for anatomical, cytochemical and ultrastructural studies.
• Trichomes initiate secretion early on, remain active throughout leaf development, but collapse as the leaves expand; at which time secretory cavities start differentiation in the mesophyll and secretion increases as the leaf reaches full expansion. The subcellular apparatus of the trichome head cells is consistent with hydrophilic and lipophilic secretion. Secretion involves two vesicle types: the smaller vesicles are PATAg-positive (periodic acid/thiocarbohydrazide/silver proteinate) for carbohydrates and the larger ones are PATAg-negative. In the first phase of secretory activity, the vesicles containing polysaccharides discharge their contents through exocytosis with the secretion accumulating beneath the cuticle, which detaches from the cell wall. Later, a massive discharge of lipophilic substances (lipids and terpenes/phenols) results in their accumulation between the wall and cuticle. Release of the secretions occurs throughout the cuticular microchannels.
• Continued protection of the leaves throughout shoot development is ensured by replacement of the collapsed secretory trichomes by oil-secreting cavities. Our findings provide new perspectives for understanding secretion regulation in shoot apices of woody species with rhythmic growth.

     

Conflict of use for multi-purpose tree species in the state of Pará, eastern Amazonia, Brazil

Although diversified forest management is promoted as a strategy aimed at slowing tropical deforestation, little is known about the viability of integrating timber and non-timber forest products in the same forest management plans. In this study we offer an initial characterization of multi-purpose tree species in the State of Pará, the principal Amazonian logging region. We identify the species used for both timber and non-timber extraction, and classify these according to their commercial value. We relate multi-purpose species to their ecological traits, the type of non-timber forest use and the fraction of the tree harvested. Although a high number of species present a potential conflict of use, this conflict is only relevant in four of them: D. odorata, T. serratifolia, T. impetiginosa and H. courbaril. Nevertheless, the nature and relevance of this conflict will ultimately depend on the importance that the non-timber use has for the livelihoods of forest-dependant people, the commercial value and the ecological resilience of these species.     

EARLY LEAF ONTOGENY AND SHOOT DEVELOPMENT IN BROWNEA ARIZA (LEGUMINOSAE)

Brownea ariza Benth. (Leguminosae: Caesalpinioideae) shows early shoot tip abortion and subsequent renewal growth from the pseudoterminal bud. This species is unusual in that the entire shoot system is formed before flushing from the bud occurs, shoot tip abortion occurs during flushing, and the aborting portion contains three to six leaves as well as primordial structures varying from hood to peg shape. This study focused on the morphological changes from initiation of scale and foliage leaf primordia in the “resting” renewal bud through bud elongation to flushing and bud abortion. Scanning electron microscopy revealed that embryonic scale leaves are hood-shaped while foliage leaf primordia show early segmentation into leaflets and stipules. No transitional stages were observed. Bud scales and foliage leaves show opposite developmental trends. In bud scales, length at maturity increases from first to last formed, while length decreases in sequentially formed foliage leaves. Early in leaf development the stipules keep pace with the elongation of the rachis. When the bud reaches about one half of its final length the leaf rachis begins to exceed the lengths of its stipules. This young rachis terminates in a distinct mucro that persists until maturity at which time it abscises. Growth patterns indicate that mucro and rachis are a single developmental unit. The early abortion of a shoot tip containing several leaves cannot be easily rationalized. Previous suggestions have involved maintenance of form and ecological adaptation. We add the possibility of elimination of cell progeny encumbered by mutations. From this and other studies of this group, it is clear that at maturity leaves of different species may look alike, e.g., Hymenaea and Colophospermum are bifoliolate; Brownea, Saraca, and others are multifoliolate. However, early stages of leaf ontogeny are quite diverse and may be of systematic value, since these early differences are lost or masked by later development.     

Dendrobium tamenglongense sp. nov. (Orchidaceae) from Manipur,India

Dendrobium tamenglongense R. Kishor, Y. N. Devi, H. B. Sharma, J. Tongbram & S. P. Vij sp. nov. from Manipur, India is described and illustrated as a new species. The new species differs from D. jaintianum in having larger and glabrous leaves with acute apices, green sepal apices, a 1–3‐flowered inflorescence, smaller and non‐fragrant flowers and a 3‐lobed labellum with fimbriate midlobe.     

The morphology of stipules and inflorescences in <Emphasis Type="Italic">Geissois sensu stricto</Emphasis> (<Emphasis Type="Italic">Cunoniaceae</Emphasis>)

Summary   Geissois sensu stricto (i.e. excluding the Australian species of this genus) has 19 species restricted to islands in the south-west Pacific and is the sole group in Cunoniaceae in which the stipules are intrapetiolar (axillary) in the adult foliage. In apical buds, paired opposite stipules are coherent round their margins and are either clearly intrapetiolar in relation to the most distal pair of developed leaves (Type I, most species), or they appear interpetiolar due to the abortion of the pair of leaves associated with them and the failure of the internode proximal to them to elongate (Type II, a few species). Stipules are often accrescent and the largest in the family occur in this group, reaching 10 cm long in some species. The ornithophilous racemose inflorescences of members of this group normally consist of either axillary triads (G. hirsuta only) or apparently simple, unbranched racemes which are ramiflorous or occasionally axillary. However, bract scars on the axes proximal to the flowers show that both types of inflorescence module usually consist of two metamers. The structure of the inflorescences and stipules in Geissois sensu stricto supports the distinction between this group and the Australian species traditionally included in Geissois sensu lato.     

Die Stipeln derIrvingioideae undRecchioideae und ihre systematische Wertung nebst Bemerkungen zur Holzanatomie und Palynologie

TheSimaroubaceae generally have no true stipules. The stipule-like appendages of some genera proved to be pseudo- or metastipules (Weberling & Leenhouts 1965). There seem to be some exceptions, however: the generaCadellia (incl.Guilfoylia) andRecchia on the one hand, and theIrvingioideae on the other. As these taxa, with exception ofRecchia, have simple leaves, there are no indications that their stipule-like appendages might be pseudo- or metastipules. In regard to their position and ontogeny these appendages behave completely like true stipules. Assuming the view ofForman, one could conceive a morphological line from the long, broadly inserted axillary stipules of mostIrvingioideae to the small scaly triangular stipules ofIxonanthoideae. The similarities between the stipules ofIrvingioideae andErythroxylaceae (already emphasized byHallier and others), become even more evident when their ontogeny is investigated. TheIrvingioideae, therefore, might be regarded as a separate family (perhaps with some relation to theErythroxylaceae,Hallier) or as a subfamily ofIxonanthaceae (Forman).—In addition to data on stipules some results on the palynology and shoot anatomy of the generaCadellia (incl.Guilfoylia) andRecchia are reported. Their relationship with theSimaroubaceae also appears doubtful. If they are to be included, they represent a somewhat isolated group near the base of the family which otherwise has lost its stipules.

Herrn Univ.-Prof. Dr.Walter Leinfellner zum 70. Geburtstag gewidmet.     

Tree seedling development in tropical dry abandoned pasture and secondary forest in Costa Rica

Development of planted seedlings of four canopy tree species in recently abandoned pastures (mown and unmown) and in ca. 40-yr old secondary dry forest in Guana-caste National Park, Costa Rica, was studied from July 1989–June 1992. The species were the light-demanding Cedrela odorata and Swietenia macrophylla, and the shade-tolerant Hymenaea courbaril and Manilkara chicle. Seedling mortality was high and primarily correlated with dry season drought. After the first dry season, M. chicle and C. odorata showed < 5 % survival in the pasture and 20 % in the forest, but after 3 yr survival had decreased to 3% in all plots. Survival of S. macrophylla was highest in the mown pasture (45 % after 3 yr) and lowest in the forest (10 %). For H. courbaril, survival was 40–55 % under all three conditions. Growth rates were low, with a height increment of < 15 cm/yr, possibly due to low precipitation in 1990 and 1991. All species grew taller in the pasture than in the forest. Differences in soil depth, texture and drainage appeared to contribute to variation in the results both within and between plots. Herbivory contributed to the lower survival of C. odorata and S. macrophylla in the forest.     

Comparative analysis of qualitative anatomical characters of <Emphasis Type="Italic">Trifolium</Emphasis> L. (Fabaceae) and their taxonomic implications: preliminary results

A comparative anatomical study of 20 Trifolium species was carried out using light and scanning electron microscopy in order to investigate the structure of their vegetative and reproductive organs and to evaluate the taxonomic significance of qualitative characters for distinguishing between the taxa. Characters were coded and analysed by multiple correspondence analysis. The combination of selected qualitative anatomical characters of the petiole, stipules, stem, peduncle and calyx (not previously studied in detail) were significant in delimiting sections of the genus Trifolium, but not for identification of species. The characters that contributed most to the separation of sections were peduncle cross-section shape, the presence of trichomes on the calyx tubes and the presence of trichomes and crystals in the stipules.     

STIPULES IN SOME MEMBERS OF THE VITACEAE: RELATING PROCESSES OF DEVELOPMENT TO THE MATURE STRUCTURE

The development of stipules especially their spatial and temporal pattern of initiation in relation to the leaf was investigated in Vitis riparia Michx., cv. Concord, Parthenocissus tricuspidata (Sieb. & Zucc.) Planch., Cissus oblonga (Benth.) Planch., Cissus hypoglauca (F.v.M.) A. Gray, and Cissus rhombifolia Vahl. Early initiation is characterized by the occurrence of a single primordium with a wide insertion on the flank of the shoot apex. Distinguishing between stipule primordia and the leaf primordium is impossible at this early stage. Distinct primordia can only be seen in later stages of development. At maturity, the stipules occupy free lateral positions. Developmental processes such as timing of initiation and zonal growth seem to play an important role in early development. In five of the six taxa examined in this study, the early initiation of stipules, their close association with the leaf and also their faster relative rate of growth during early development appear to give them a characteristic protective function. In contrast, C. rhombifolia stipules are initiated later than the leaf and seem to develop at a slower rate than the leaf proper. Consequently, they never enclose their associated leaf but instead cover the next youngest leaf. Many different criteria are used to distinguish the broad category of stipules, and therefore many interpretations have been made depending on the type of approach that is used. This study attempts to look at stipules in terms of developmental processes and demonstrates a more accommodating leaf/stipule concept which provides a clearer comprehension of the nature of the stipule.     

Leaf defoliation of tropical dry forest tree seedlings – implications for survival and growth

 The effect of herbivory on survival and growth of seedlings of four species, Cedrela odorata, Hymenaea courbaril, Manilkara chicle and Swietenia macrophylla, was studied in secondary dry forests in Guanacaste, Costa Rica. Potted seedlings were planted at two sites in a 2×2 factorial design, combining thinning to increase light levels at the forest floor, and trenching to reduce root competition around the planted seedlings. C. odorata and S. macrophylla were repeatedly severely defoliated by insects, while H. courbaril became less affected. M. chicle did not show any significant signs of defoliation and was not further analysed. Defoliation levels were generally higher in both thinned and trenched treatments, and also positively correlated with larger initial seedlings sizes. Decreased growth rates caused by defoliation were seen in S. macrophylla and C. odorata in the growing season. Defoliation of more than 50% in combination with abiotic factors, particularly drought, were sufficient to contribute to retarded seedling development and increased seedling mortality of C. odorata and S. macrophylla. Received: 10 April 1997 / Accepted: 26 June 1998     

EFFECTS OF GIBBERELLIC ACID ON STEM APICES OF VERNALIZABLE GRASSES

Koller , D. (Hebrew U., Jerusalem, Israel), H. R. Highkin , and O. H. Caso . Effects of gibberellic acid on stem apices of vernalizable grasses. Amer. Jour. Bot. 47(6) : 518–524. Illus. 1960.—Gibberellic acid was found to have distinct morphogenic effects on the stem apices of 3 vernalizable but unvernalized grasses: Hordeum vulgare var. ‘Kentucky,’ Hordeum bulbosum and Secale cereale var. ‘Winter Petkus.’ In these 3 species, GA caused the activation of lateral meristems on the embryonic nodes of the stem apices, under both short- and long-day conditions. These lateral meristems differentiated into either flower primordia or vegetative shoots. The flower primordia developed almost invariably in Hordeum, while in Secale they seldom did, the apical meristem usually resuming normal vegetative growth after treatment. Despite the occurrence of flowering, it is concluded that the role of GA in this phenomenon is restricted to the activation of lateral meristems in the apex.     

Reinvestigation of Coprosma for ability to fix atmospheric nitrogen

Summary Coprosma robusta Raoul plants grown in plus- and minus-nitrogen sand culture showed no significant response to removal of stipules. Acetylene reduction assays for nitrogenase on leaves and stipules gave negative results for six Coprosma species. Two unwashed root samples showed slight acetylene reduction and this was ascribed to casual aerobic rhizosphere organisms.     

<Emphasis Type="Italic">Psychotria kupensis</Emphasis> (<Emphasis Type="Italic">Rubiaceae</Emphasis>) a new dwarf,litter-gathering species from western Cameroon

Summary   Psychotria kupensis Cheek is described as a new species in Psychotria sect. Confertiflorae Hiern. Attention is drawn to its monopodial, dwarf, litter-gathering and adventitious root-forming habit and to its anomalous stipules. The species appears to be endemic to the submontane evergeen forest of Mt Kupe in Cameroon. Its conservation status is assessed.     

Taxonomic revision of the genus Heliotropium (Boraginaceae s.l.) in south Yemen

The flora and fauna of Arabia, particularly southern Yemen, has recently attracted the interest of many authors. In this study, the genus Heliotropium L. (Boraginaceae) is taxonomically revised in southern Yemen. Ten species are recognized. Nomenclature, typification, representative specimens and a diagnostic key to all species are provided, along with their distribution in southern Yemen. The taxonomically most valuable characters in the genus are those of flowers (calyx, corolla, androecium and gynoecium) and nutlets, and those characters are thoroughly investigated and discussed. Both light and scanning microscopy are used in the investigation. Papillate anther apices (brush‐like) were recognized in some species. Connate anthers and deep or shallow ventral circular depressions on the nutlets were found to be useful in distinguishing members of H. sect. Orthostachys (viz. H. strigosum, H. rariflorum and H. ovalifolium), while winged nutlets were found only in H. pterocarpum. The length and hairiness of the style and stigma also proved to be taxonomically useful.     

Development of leaves and shoot apex protection in Metrodorea and related species (Rutaceae)

Most members of Sapindales are characterized by compound leaves, but several genera also (or only) produce simple or unifoliolate leaves. A few genera may bear stipules or pseudostipules. Little is known about the morphological structure and morphogenesis of these types of leaves in Sapindales, but this information is required for comparative and evolutionary studies. Metrodorea is a Neotropical genus of Rutaceae, comprising species presenting compound and unifoliolate leaves, plus heterophylly, together with an intriguing bud‐protecting structure at the leaf base. The aims of the present study are: (1) to examine leaf morphogenesis in Metrodorea and in closely related species (four Esenbeckia spp., Helietta apiculata and Raulinoa echinata); and (2) to improve our understanding of the morphological evolution of leaves in Metrodorea and Rutaceae. Our data show that the hood‐shaped structure at the base of the leaf in Metrodorea, usually interpreted as a sheath, is, in fact, a pair of united stipules, a synapomorphy of the genus. In the species studied, it is possible to recognize two main types of unifoliolate leaf: early unifoliolate leaves and late unifoliolate leaves. We also found that the number of leaflets in the studied species is dependent on the late or early determination of the leaf primordium, and that loss of leaflets may have been favoured by the restriction of space available for development within the cavity formed by the pair of united stipules. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 178 , 267–282.     

A morphological revision of the genus Herbertus S.Gray (Herbertaceae,Marchantiophyta) in Africa,including the East African islands

Abstract

The genus Herbertus S.Gray in Africa and the East African islands is revised. The division of the genus into two broad species complexes centred on H. dicranus (Tayl.ex Gottsche et al.) Trev. and H. juniperoideus (Sw.) Grolle is confirmed. H. capensis (Steph.) Sim, H. capillaris (Steph.) H. doggeltianus (Steph.) Demaret, H. lobatus (Steph.) Demaret, H. mascarenicus (Steph.) S.Arn., H. mollis (Steph.) Dusén, H. stuhlmannii (Steph.) Demaret and Schisma kilimandjarense Steph. are synonymous with H. dicranus. H. grossevittatus (Steph.) S.Arn. ex Grolle is synonymous with H. juniperoideus. Three new species are described: H. spicatus N.G.Hodgetts sp. nov., which is related to H. juniperoideus and appears close to the neotropical H. pensilis, with long, narrow leaf lobe apices and a sheathing leaf base; H. mauritianus N.G.Hodgetts sp. nov., also related to H. juniperoideus, has fewer basal slime papillae and the vitta bifurcating lower down; and H. pocsii N.G.Hodgetts sp. nov., which is related to H. dicranus but has setaceous leaf lobe apices composed of long, narrow cells. The relationships of the taxa and patterns of distribution, taking account of recent molecular work, are discussed.     

Two new endemic species of Ophiorrhiza L. (Rubiaceae: Ophiorrhizeae) from Davao Oriental,Philippines

Two new species of Ophiorrhiza collected from Mt Hamiguitan Range Wildlife Sanctuary in Davao Oriental, Philippines, are herein described and illustrated. Ophiorrhiza erythropilosa is the third Philippine species possessing involucral bracts and is further characterized by its predominantly villose stems that appear red due to strikingly red-violet trichomes, subpersistent bifid stipules, linear involucral bracts without prominent midrib, and 4 mm long urceolate corolla that is villose outside and lightly puberulous inside. Ophiorrhiza hamiguitanensis is characterized by its coriaceous, lanceolate leaves with attenuate apex and base, brochidodromous venation, subpersistent bilobed stipules with slightly recurved acute tips, presence of 1.5–2.5 mm long linear ensiform bracts, heterostylous flowers, clavate calyces and broadly obcordate capsules.