Petal Appendages Research Articles

A new species of Tillandsia (Tillandsioidae, Bromeliaceae) from the high Andean forests near Bogotá (Colombia)

A new species of Tillandsia from high elevation forests of the Eastern Cordillera of Colombia is described here: Tillandsia chingacensis. This species is unique because of its floral parts densely covered with a resin-like substance, and exceptionally long fruits. Although this species has distichous flowers and bears petal appendages, it was not placed in Vriesea because it shows morphological affinities to species of Tillandsia subg. Allardtia, suggesting a better placement of this taxon in Tillandsia. The discovery of this species is the result of recent explorations to a former war zone in the proximity of Bogotá (Colombia). This novelty demonstrates that further scientific expeditions to these areas recently opened for exploration will bring numerous taxonomic discoveries for Colombia. Finally, the description of this new species is complemented with notes on its geographical distribution, habitat, conservation status and a discussion of its taxonomic affinities.

Can flowers help to disentangle the Vriesea oligantha complex (Bromeliaceae: Tillandsioideae)? Taxonomic implications and contributions to the floral anatomy and biology of Stigmatodon

Abstract The Vriesea oligantha complex comprises Vriesea lancifolia, Vriesea oligantha, and Vriesea pseudoligantha, which were recently transferred to Stigmatodon. Such species exhibit strong morphological convergence to live in rocky fields, which makes difficult their identification in the wild or in herbarium specimens. In a previous work, leaf characters helped in the species delimitation and evidenced a new taxon. Considering that morphology, anatomy, and aspects of floral biology can contribute to better define species limits and to indicate potential pollinators, individuals from nine populations of V. oligantha complex were comparatively studied. The typical V. oligantha is mainly distinguished by tubular flowers, with simple petal appendages and diurnal anthesis. The remaining individuals studied from different populations exhibit campanulate corollas, with denticulate petal appendages and nocturnal anthesis, not allowing for the separation of V. lancifolia, V. pseudoligantha, and a fourth taxon. Floral anatomy is less variable than the external morphology, with little contribution to the species delimitation; however, it is helpful in the circumscription of Stigmatodon. The infralocular septal nectaries and the floral features provide evidence that the main pollinators of campanulate and tubular flowers are bats and hummingbirds, respectively. Specific pollinator guilds in combination with fragmented populations may explain the limited gene flow amongst populations.

Erythroxylumaustroguangdongense (Erythroxylaceae), a new species from Guangdong, China.

Erythroxylumaustroguangdongense (Erythroxylaceae), a new species from Guangdong Province, China, is described and illustrated. This new species is morphologically most similar to E.calyptratum, but is distinguished by the leathery leaf blade with fewer pairs of secondary veins and flowers borne on leafless nodes of the basal part of the current branch with much longer pedicels and sub-rectangular petal appendages. This is the second native species of Erythroxylum recorded from China.

<p><strong>Re-evaluation of the Amazonian <em>Hylaeaicum</em> (Bromeliaceae: Bromelioideae) based on neglected morphological traits and molecular evidence</strong></p>

Generic status for the Amazonian Hylaeaicum is proposed within the Aechmea alliance, excluding it from the “Nidularioid complex” in general and from Neoregelia in particular. The monophyly of this new genus is supported by molecular phylogenetic analyses. The taxonomic circumscription of Hylaeaicum is based on the combination of geographical range and morphological characters, such as clonal growth, inflorescence structure, petal and corolla conformation, petal appendages, ovary, ovule, stigma, pollen, fruit, and seed, as well as seed anatomy, thoroughly documented and illustrated from field-collected specimens that flowered in cultivation in the Rio de Janeiro Botanical Garden, the Marie Selby Botanical Gardens, and in Refúgio dos Gravatás. The presence of seeds with long bicaudate appendages on both chalazal and micropylar ends is reported for the first time in Bromelioideae and considered an important character to distinguish Hylaeaicum from the related genera in the Aechmea alliance. In order to support the morphological distinctness of Hylaeaicum, the most varied and complete documentation of stigmata, fruits, and seeds of Bromelioideae is also presented for the first time, covering 24 genera and 17 subgenera. Fourteen new combinations, including 12 species and two varieties, are proposed.

Floral development and anatomy of two species ofAechmea(Bromeliaceae, Bromelioideae)

AbstractAechmea (Bromeliaceae) is a large genus with controversial systematics and distinct flower shapes and pollinators. We explored floral anatomy and development in two Aechmea spp. belonging to different subgenera to contribute useful information on reproductive biology and taxonomy. We examined floral buds using scanning electron and light microscopy to characterize the development of septal nectaries, petal appendages, ovules, stamens and carpels. In A. gamosepala, we confirmed that the petal appendages develop late, whereas in A. correia-araujoi they develop earlier during floral development. Petal appendage formation included positional changes, possibly affecting floral attributes and visitation by insects, rather than vertebrates. Nectar is released through three basal orifices distally on the ovary, and here we document the link between the nectary region, through discrete canals, upward to the conduplicate lobes of the wet stigma. Improved understanding of the floral development and morphology of Aechmea may help to explain the existence of polymorphic flowers in this genus and may have implications for studies on interactions with pollinators and systematics.

SILENE NELSONII, A NEW LARGE-FLOWERED SPECIES FROM THE TRINITY RIVER AREA OF NORTHWESTERN CALIFORNIA, USA, AND A RE-EVALUATION OF S. BOLANDERI GRAY

Silene nelsonii M.R. Mesler, M.S. Mayer, and S.K. Carothers (Caryophyllaceae), a species with large, day-blooming flowers from the Trinity River Basin of northwestern California, is described and illustrated. In the past, plants assignable to this new species have been referred to incorrectly as S. bolanderi A.Gray or S. hookeri Nutt. subsp. bolanderi (Gray) Abrams, names that rightfully apply to another member of the Silene hookeri complex. Here we argue that both S. nelsonii and S. bolanderi are worthy of recognition as separate species based on morphological distinction and a molecular phylogenetic analysis. Silene nelsonii differs from S. bolanderi by its more deeply lobed, white petals; lack of well-developed coronal petal appendages; densely ciliate petal bases; and shorter, sometimes branched sepal hairs. We present a key to the five members of the S. hookeri complex, all of which are in some degree rare in California and worthy of conservation concern.

Erythroxylum calyptratum (Erythroxylaceae), a new species from Mt. Fansipan, northern Vietnam

Erythroxylum calyptratum, sp. nov. (Erythroxylaceae), from Mt. Fansipan, Hoang Lien National Park, Lao Cai Province, northern Vietnam, is newly described. The new species is characterized by thinly chartaceous leaves with 2–3 mm long petiole, 1–3 flowered inflorescences, downward facing white flowers, well-developed petal appendages (1/3–1/2 length of petals) and entirely free 3 styles. A description, illustrations, DNA barcodes of rbcL and matK and a key to the species of Erythroxylum in Indochina are provided.

New circumscription of Cryptanthus and new Cryptanthoid genera and subgenera (Bromeliaceae: Bromelioideae) based on neglected morphological traits and molecular phylogeny

The authors provide a new circumscription for genera of the “Cryptanthoid complex” of Bromeliaceae subfam. Bromelioideae, originally composed of Cryptanthus, Lapanthus, Orthophytum, and Sincoraea, on the basis of new or re-evaluated ecological, geographical and morphological evidence, as well as molecular phylogenies. A new generic status is proposed for Cryptanthus subg. Hoplocryptanthus and two new genera, Forzzaea, and Rokautskyia, as well as four new subgenera in Orthophytum (Capixabanthus, Clavanthus, Krenakanthus, and Orthocryptanthus) are described to render taxonomic units monophyletic. The recognized taxa are well circumscribed by the combination of geographical range, ecology and morphological characters (sex distribution, leaf succulence, sepal and petal size and connation, petal appendages, pollen and stigma morphology, fruit size, calyx persistency, seed size and number per fruit). Field collected living specimens of 78 of the 81 species of Cryptanthus s.l., all species of Lapanthus, 58 of the 59 species of Orthophytum, and all species of Sincoraea were analysed in habitat and/or in cultivation, allowing the documentation and illustration of new and underutilized characters. The molecular analysis incorporated 91 accessions representing 33 species of Cryptanthus, all species (3) of Lapanthus, 42 species of Orthophytum, and 9 species of Sincoraea, including the type species for the first three genera and four outgroup taxa. The results suggest, that some morphological characters generally considered homoplasious for Bromelioideae, for the “Cryptanthoid complex”, are not homoplasious at least within the obtained, biogeographycally well delimited clades and their taxonomical utility is redeemed.

Anatomia floral de Aechmea distichantha Lem. e Canistropsis billbergioides (Schult. & Schult.f) Leme (Bromeliaceae)

RESUMO Aechmea Ruiz & Pav. e Canistropsis (Mez) Leme pertencem à subfamília Bromelioideae, detentora da maior diversidade morfológica em Bromeliaceae. Botões florais de Aechmea distichantha Lem. e Canistropsis billbergioides (Schult. & Schult. f.) Leme foram coletados, fixados e processados conforme técnicas usuais em anatomia vegetal. As espécies compartilham características, tais como presença de cristais de sílica esféricos nas células epidérmicas do perianto; idioblastos contendo ráfides; endotécio com espessamento do tipo anelar e ovário ínfero com placentação axilar. Apêndices petaloides não vascularizados foram observados apenas em flores de A. distichantha, dispostos aos pares em cada pétala. Em ambas as espécies ocorre um nectário septal cuja secreção é rica em proteínas e carboidratos. Tecido obturador placentário está presente, e testes histoquímicos revelaram que a secreção produzida por este tecido contém proteínas e carboidratos, provavelmente, relacionados com o direcionamento do tubo polínico aos óvulos.

And then came molecular phylogenetics—Reactions to a monographic study of Hypericum (Hypericaceae)

Completion of a detailed monograph of Hypericum using traditional methods has stimulated the publication of phylogenetic treatments of the genus using molecular methods. The relationships thereby revealed differ from those of the traditional account in several ways that are discussed here. A key point of conflict concerns taxa that share a specific set of morphological characters, i.e. the ‘Elodes’ syndrome (pseudotubular corolla, sterile stamen fascicles, stamen filaments ± united in the fascicle and sometimes petal appendages), which was found to be paraphyletic in each molecular study. The various groups with this syndrome were shown to have arisen from various parts of Hypericum, apparently suddenly, probably as the result of a genetic switch, the action of which can also be reversed. The formerly segregated genera with this floral syndrome should therefore all be re-incorporated in Hypericum. This necessitates the re-establishment of one section, namely Hypericum sect. Elodea, and recognition of two new sections within Hypericum, namely Hypericum sect. Lianthus and Hypericum sect. Thornea, which are proposed here. The remaining species have resolved in various places in recent molecular phylogenetic trees that differ from each other and from the classification adopted in the monograph. In particular, in molecular results the herbaceous species with ‘3’ (i.e. 2+2+1) stamen fascicles all form a single clade; whereas, in the traditional treatment adopted in the monograph, they form three distinct, distantly related groups. In light of recent molecular studies, Hypericum is now interpreted to comprise two distinct groups (respectively mainly Old World and mainly New World) that differ in the presence or absence of dark (hypericin-containing) glands and in the arrangement of the stamen fascicles. These two clades are described as subgenera, with the New World clade named Hypericum subgenus Brathys. By drawing attention to this grouping, the molecular work has improved on the monograph.

Three New Additions to the Guayana Shield-endemic <I>Steyerbromelia</I> (Bromeliaceae) for Colombia with Comments on the Problematic Generic Delimitation within the Subfamily Navioideae

Abstract Two new species and a new combination for Steyerbromelia (Bromeliaceae: Navioideae) are presented. These novelties represent the first records of Steyerbromelia for Colombia. The new species, Steyerbromelia naquenensis and Steyerbromelia nukakii, were placed in Steyerbromelia based on the combination of their 3- or 4-divided inflorescences, presence of smooth posterior sepals, and ovate, shortappendaged ovules. Both species resemble each other and S. thomasii, which also occurs in the basin of the Upper Rio Negro. Steyerbromelia garcia-barrigae is transferred from Navia, despite the lack of petal appendages, because its morphological characteristics fit better the limits of Steyerbromelia. An updated description of this species is presented. We now consider Steyerbromelia to be a genus of nine species. Additionally, notes on the geographic distribution, vernacular names, conservation status, and taxonomic comments of the three species are provided, as well as a key to all of the species. Finally,...

Vriesea(Bromeliaceae, Tillandsioideae): taxonomic history, and morphology of the Brazilian lineage1

The genus Vriesea Lindl. comprises about 290 species distributed into two sections: V. sect. Vriesea and V. sect. Xiphion. The genus is recognized as polyphyletic and is historically associated with Tillandsia L., differing from it mainly due to the presence of petal appendages. Here we focus our efforts on the Vriesea taxa in its main center of diversity in southeastern Brazil that are considered to constitute a monophyletic group based on molecular analyses as well as a total evidence approach. We present a morphological description of the species from the Brazilian lineage, and the taxonomic history of the genus. We based our results on extensive bibliographic research and morphological analyses of herbarium material and living specimens, as well as illustrations of the main taxonomic characteristics of the Brazilian lineage of the genus.

Alatococcus, a new genus of Sapindaceae from Espirito Santo, Brazil

The new genus Alatococcus (Sapindaceae) is described from Espirito Santo, Brazil. Alatococcus seems to be closely related to Scyphonychium of northeastern Brazil, as they both share similar vegetative morphology, flowers with zygomorphic calyx, fruits with indehiscent mericarps, and isopolar, tricolporate pollen grains. They however differ by the shape of the nectary disc, petal appendages, mericarps, and pollen shape and ornamentation. The new species Alatococcus siqueirae is described and illustrated.

Crassula fragarioides, a new sandstone outcrop species from the Bokkeveld Escarpment, Northern Cape, South Africa

Crassula fragarioides, a new dwarf succulent, is described from sandstone outcrops on the Bokkeveld Escarpment, south of Nieu-woudtville in the Northern Cape. It is currently known from only three subpopulations. The white flowers, with dorsal petal appendages and globular, translucent trichomes on the reddishpurple leaves distinguish it from C. clavata, C. namaquensis subsp. comptonii and C. elegans.

Nocturne for an unknown pollinator: first description of a night-flowering orchid (Bulbophyllum nocturnum)

Bulbophyllum nocturnum, a species of section Epicrianthes from New Britain, is described and illustrated. It is the first known example of an orchid species in which the flowers open after dark and close in the morning. The poorly understood pollination biology of section Epicrianthes, a clade with highly unusual flowers, is discussed. Attention is drawn to the close resemblance between the petal appendages of some species and the fruiting bodies of certain Myxogastria. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 167, 344–350.

<I>Lapanthus</I> (Bromeliaceae, Bromelioideae): A New Genus from the Southern Espinhaço Range, Brazil

In this paper we describe a new genus of Bromeliaceae, Lapanthus, restricted to the southern portion of the Espinhaco Range, Minas Gerais State, in southeastern Brazil. Two new combinations to accommodate species previously described in the genera Orthophytum and Cryptanthus and one new synonym are proposed. Lapanthus has morphological affinities with both Cryptanthus and Orthophytum, but nevertheless differs by the combination of margins of the petals ciliate, presence of lanceolate petal appendages and free stamens, and also by molecular data. Cryptanthus and Orthophytum have petals entire along the margins, and the filaments of the most internal whorl are adnate to the petals. Lapanthus stands out by having a pair of lanceolate petal appendages, which are almost completely adnate to the petals. In Orthophytum, however, appendages are cupuliform or sacciform and they are totally absent in the genus Cryptanthus. Lapanthus and Orthophytum present meiotic and mitotic chromosome numbers equal to n=25 and 2n=50, 100 and 150 respectively, while Cryptanthus presents meiotic and mitotic chromosome numbers n=17 and 2n=34, 36, 54 respectively, and this difference is considered to be an autapomorphic feature of Cryptanthus. Descriptions of the genus and species, identification keys, illustrations, photographs of living specimens, and taxonomic comments are provided.

Male and female booths with separate entrances in the tiny flowers of Guazuma ulmifolia (Malvaceae–Byttnerioideae). I. Structural integration

In spite of their tiny dimensions (3×5 mm 2), the strongly honey-scented flowers of Guazuma ulmifolia (Malvaceae–Byttnerioideae) provide six sexual chambers. A central female unit is surrounded by five interconnected male compartments. At the onset of anthesis the strap-shaped odoriferous petal appendages form an open star with the female chamber in the centre. Hereafter, they wilt, intermingle and finally lock access to the stigma. Meanwhile, wilting sepals liberate new entrances from the back between the petal claws. They give way to nectar-offering male chambers. In each of them a well-lit (but closed) window lures the visitor into a standard position for passive pollen uptake. Finally, the insects have to back out of one of the rear entrances. It remains unknown how the minute visitors (maximum height ca. 0.5 mm) – if ever – overcome the distance to another Guazuma tree.

Floral anatomy of Bromeliaceae, with particular reference to the evolution of epigyny and septal nectaries in commelinid monocots

Floral anatomy is described in ten genera of Bromeliaceae, including three members of subfamily Bromelioideae, three Tillandsioideae, and four genera of the polyphyletic subfamily Pitcairnioideae (including Brocchinia, the putatively basal genus of Bromeliaceae). Bromeliaceae are probably unique in the order Poales in possessing septal nectaries and epigynous or semi-epigynous flowers. Evidence presented here from floral ontogeny, vasculature, and the relative positions of nectary and ovules indicates that there could have been one or more reversals to apparent hypogyny in Bromeliaceae, although this hypothesis requires a better-resolved phylogeny. Such evolutionary reversals probably evolved in response to specialist pollinators, and in conjunction with other aspects of floral morphology of Bromeliaceae, such as the petal appendages of some species. The ovary is initiated in an inferior position even in semi-epigynous or hypogynous species. The ovary of all so-called hypogynous Bromeliaceae is actually semi-inferior, because the septal nectary is infralocular; in these species the nectaries have a labyrinthine surface and many vascular bundles. Brocchinia differs from most other fully epigynous species in that each carpel is secretory at the apex and reproductive, rather than secretory, at the base.

Floral Development in Mangrove Rhizophoraceae

The flowers of mangrove Rhizophoraceae (tribe Rhizophoreae) are adapted to three different pollination mechanisms. Floral development of representative species of all four genera suggests that the ancestral flower of the tribe was unspecialized, with successively initiated whorls of separate sepals, petals, antisepalous stamens, and antipetalous stamens; at its inception, the gynoecium had a united, half-inferior ovary and separate stigmatic lobes. This developmental pattern is found in Rhizophora mangle (wind-pollinated) and Ceriops decandra (insect-pollinated). In Kandelia, all floral organs distal to the sepals are initiated simultaneously, and there has apparently been an evolutionary amplification in the number of stamens to about six times the number of petals. Explosive pollen release evolved independently in C. tagal and in Bruguiera. In the former, all stamens belong to one whorl and arise simultaneously upon a very weakly differentiated androecial ring primordium. In Bruguiera, the androecial ring is pronounced, and two whorls of stamens arise upon it; the primordia of the antisepalous whorl arise first but are closer to the center of the apex than the antipetalous stamen primordia. The antisepalous stamens bend toward and are enclosed by the petals early in development. In all genera, the inferior ovary develops by zonal growth of receptacular tissue; additional intercalary growth above the placenta occurs in Bruguiera. In general, floral specialization is accompanied by an increase in the width of the floral apex compared to the size of the primordia, increasing fusion of the stylar primordia, and decreasing prominence of the superior portion of the ovary. Apparent specializations of petal appendages for water storage, including the presence of sub-terminal hydathodes (previously unreported in any angiosperm), were found in two species in which flowers remain open during the day but were absent from two species normally pollinated at night or at dawn. Distinctive tribal characteristics that may aid in phylogenetic analysis include the mode of development of the inferior ovary; the aristate, bifid, usually fringed petals that individually enclose one or more stamens; the intrastaminal floral disc; and the initially subepidermal laticiferous cell layer in the sepals and ovary.

FLORAL DEVELOPMENT IN MANGROVE RHIZOPHORACEAE

The flowers of mangrove Rhizophoraceae (tribe Rhizophoreae) are adapted to three different pollination mechanisms. Floral development of representative species of all four genera suggests that the ancestral flower of the tribe was unspecialized, with successively initiated whorls of separate sepals, petals, antisepalous stamens, and antipetalous stamens; at its inception, the gynoecium had a united, half‐inferior ovary and separate stigmatic lobes. This developmental pattern is found in Rhizophora mangle (wind‐pollinated) and Ceriops decandra (insect‐pollinated). In Kandelia, all floral organs distal to the sepals are initiated simultaneously, and there has apparently been an evolutionary amplification in the number of stamens to about six times the number of petals. Explosive pollen release evolved independently in C. tagal and in Bruguiera. In the former, all stamens belong to one whorl and arise simultaneously upon a very weakly differentiated androecial ring primordium. In Bruguiera, the androecial ring is pronounced, and two whorls of stamens arise upon it; the primordia of the antisepalous whorl arise first but are closer to the center of the apex than the antipetalous stamen primordia. The antisepalous stamens bend toward and are enclosed by the petals early in development. In all genera, the inferior ovary develops by zonal growth of receptacular tissue; additional intercalary growth above the placenta occurs in Bruguiera. In general, floral specialization is accompanied by an increase in the width of the floral apex compared to the size of the primordia, increasing fusion of the stylar primordia, and decreasing prominence of the superior portion of the ovary. Apparent specializations of petal appendages for water storage, including the presence of sub‐terminal hydathodes (previously unreported in any angiosperm), were found in two species in which flowers remain open during the day but were absent from two species normally pollinated at night or at dawn. Distinctive tribal characteristics that may aid in phylogenetic analysis include the mode of development of the inferior ovary; the aristate, bifid, usually fringed petals that individually enclose one or more stamens; the intrastaminal floral disc; and the initially subepidermal laticiferous cell layer in the sepals and ovary.