Markus Ackermann | Universität Koblenz (original) (raw)

Papers by Markus Ackermann

Research paper thumbnail of A question of data quality-Testing pollination syndromes in Balsaminaceae

PloS one, 2017

Pollination syndromes and their predictive power regarding actual plant-animal interactions have ... more Pollination syndromes and their predictive power regarding actual plant-animal interactions have been controversially discussed in the past. We investigate pollination syndromes in Balsaminaceae, utilizing quantitative respectively categorical data sets of flower morphometry, signal and reward traits for 86 species to test for the effect of different types of data on the test patterns retrieved. Cluster Analyses of the floral traits are used in combination with independent pollinator observations. Based on quantitative data we retrieve seven clusters, six of them corresponding to plausible pollination syndromes and one additional, well-supported cluster comprising highly divergent floral architectures. This latter cluster represents a non-syndrome of flowers not segregated by the specific data set here used. Conversely, using categorical data we obtained only a rudimentary resolution of pollination syndromes, in line with several earlier studies. The results underscore that the use ...

Research paper thumbnail of Erratum to: Diversification of Caiophora (Loasaceae subfam. Loasoideae) during the uplift of the Central Andes

Organisms Diversity & Evolution

Research paper thumbnail of Phylogenetic relationships and generic re-arrangements in "South Andean Loasas" (Loasaceae)

Taxon

Ver impacto de la revista Resumen Loasaceae, a mostly American group, is one of the largest famil... more Ver impacto de la revista Resumen Loasaceae, a mostly American group, is one of the largest families of Cornales. In spite of

Research paper thumbnail of Flower reshaping in the transition to hummingbird pollination in Loasaceae subfam. Loasoideae despite absence of corolla tubes or spurs

Evolutionary Ecology, 2016

Many angiosperm lineages present transitions from bee to hummingbird pollination. The flower desi... more Many angiosperm lineages present transitions from bee to hummingbird pollination. The flower design in most of these lineages includes either corolla tubes or nectar spurs, structures that commonly experienced an elongation with the acquisition of hummingbird pollination. It is proposed that this increases the fit between the bird head and flower structures, and isolates or partially blocks bees from the interaction. But can this transition occur if the ancestral flower design lacks tubes or spurs? Here we focus on the transition from bee to hummingbird pollination in the Loasaceae subfamily Loasoideae. Loasoideae flowers have radial corollas with separated petals; therefore, they do not display corolla tubes nor nectar spurs. These flowers also present a whorl of nectar scales and staminodes, unique to the subfamily, which is involved in flower–pollinator fit and in nectar harvesting. To explore flower shape adaptation to hummingbird pollination, we tested for correspondence between pollinators and flower shape in Loasoideae. In order to achieve this, we first compared the evolutionary history of flower phenotype and pollination mode, and then used stochastic character mapping and geometric-morphometric variables in a comparison of alternative evolutionary models. The results of our study suggest that the transition from bee to bird pollination was accompanied by changes in the shape of the staminodial complex, along with the evolution of relatively closed corollas. Moreover, while bird pollination seems to be the end point in the evolution of pollination syndromes in many angiosperm lineages, rodent pollinated flowers probably evolved from ancestral bird pollinated flowers in Loasoideae. Our findings suggest that the evolution of bird pollinated flowers from ancestral bee pollinated flowers does not require the presence of corolla tubes or spurs, and can take place as long as the flower design includes structures participating in flower–pollinator fit.

Research paper thumbnail of Notas sobre el Género Caiophora (Loasoideae, Loasaceae) en Chile y países limítrofes

Darwiniana

Esta es la primera revisión del género Caiophora (Loasaceae) en Chile. El género comprende alrede... more Esta es la primera revisión del género Caiophora (Loasaceae) en Chile. El género comprende alrededor de 60 especies, y está ampliamente distribuido en los Andes, desde Argentina/Chile en el Sur, hasta el centro de Ecuador en el Norte. Se conocen solamente cinco especies de Chile: Caiophora chuquitensis, C. cirsiifolia, C. coronata, C. deserticola sp. nov. y C. rosulata. El material de Caiophora rosulata se divide entre dos subespecies, C. rosulata subsp. rosulata comb. nov., de los Andes Occidentales (presente en el norte de Chile y el sur de Perú, y C. rosulata subsp. taraxacoides stat. y comb. nov., de los Andes Orientales. Los nombres C. superba syn. nov. y C. macrocarpa syn. nov. son sinonimizados bajo C. chuquitensis, y C. rahmeri syn. nov., sinonimizado bajo C. rosulata subsp. rosulata. Estas cinco especies comprenden el rango completo de hábitos conocido para el género: sufrútices, hierbas perennes en forma de cojines, hierbas rosuladas acaules, y hierbas trepadoras. Proporcionamos una clave, descripciones morfológicas y la sinonimia completa para todos los taxones, incluyendo ilustraciones, notas sobre la distribución, el hábitat, la biología floral y números cromosómicos.

Research paper thumbnail of Los nombres antiguos en el género Caiophora (Loasaceas subfam. Loasoideae) y una clasificación infragenérica preliminar

EI presente trabajo se enfoca en establecer la identidad sistemälica de los nombres antiguos en e... more EI presente trabajo se enfoca en establecer la identidad sistemälica de los nombres antiguos en el genero Caiophora sobre la base de estudios de campo y herbario. incluyendo varios e1ementos tipicos originales. Se propone aceptar los siguientes nombres para los taxones centrales de la flora peruana : Caiophora pedullcu/aris (K.Presl) Weigend & M.Ack., combo llOV. ( - Loasa pedullCli/aris K.Presl, = Caiophora pallciseta Killip), Caioplwra cirsiifolia K.Presl. (= Caiophora sepiaria (Ruiz & Pav. ex G.Oon) J.F.Macbr., = Caiophora preslii Urb. & Gilg), Caioplwra carduifolia K.Presl (= B/limenbachia punicea Ruiz & Pav. ex G.Don, = Caiophora cymbifera Urb. & Gilg), Caioplrora gralldiflora (G.Oon) Weigend & M.Ack. , combo nov. (=Bllimenbachia grandijlora G.Oon =Loasaphysopetala Ruiz & Pav.) y Caioplwra pterosperma (Ruiz & Pav. ex. G.Oon) Urb. & Gilg (",Bllimenbachia pterasperma Ruiz & Pav. ex G.Don =Loasa pterasperma Ruiz & Pav. = Caiophora serropetala Macbr., = Caiophora smithii Killi...

Research paper thumbnail of Notes on the genus Caiophora (Loasoideae, Loasaceae) in Chile and neighbouring countries

This is the first revision for the representatives of the genus Caiophora (Loasaceae) in Chile. T... more This is the first revision for the representatives of the genus Caiophora (Loasaceae) in Chile. The genus is widely distributed in the Andes from Argentina/Chile in the South to Central Ecuador in the North, and comprises approximately 60 species. In Chile only five species are present, Caiophora chuquitensis, C. cirsiifolia, C. coronata, C. deserticola sp. nov. and C. rosulata. Caiophora rosulata is here subdivided into two subspecies: western Andean C. rosulata subsp. rosulata (present in northern Chile and southern Peru), and eastern Andean C. rosulata subsp. taraxacoides, stat. and comb. nov. Furthermore C. superba syn. nov. and C. macrocarpa syn. nov. are placed into synonymy under C. chuquitensis, and C. rahmeri syn. nov. is synonymized to C. rosulata subsp. rosulata. These five species comprise the complete range of growth forms known for the genus, i.e., subshrubs, cushionforming herbs, acaulescent, rosulate herbs and vines. For all taxa a key and full morphological descript...

Research paper thumbnail of Nectar, floral morphology and pollination syndrome in loasaceae subfam. Loasoideae (Cornales)

Annals of Botany, 2006

Background and Aims Loasaceae subfam. Loasoideae are mostly distributed in South America (sea lev... more Background and Aims Loasaceae subfam. Loasoideae are mostly distributed in South America (sea level to over 4500 m) with a wide range of animals documented as pollinators. The aim was to investigate correlations between nectar parameters, flower morphology, pollination syndrome and phylogeny. Methods Nectar was collected from 29 species from seven genera in the subfamily. Concentration and volumes were measured and the amount of sugar calculated. Correlations of nectar data were plotted on a ternary graph and nectar characteristics compared with flower visitors, floral morphology and phylogenetic data. Key Results Sugar concentrations are generally higher than reported for most plant families in the literature. The species investigated can be roughly grouped as follows. Group I: plants with approx. 1Á5(-3Á5) mL nectar with (40-)60-80 % sugar and 0Á19-2 mg sugar flower À1 ; with small, white, star-shaped corollas, pollinated by shorttongued bees. Groups II, III and IV: plants with mostly orange, balloon-, saucer-, bowl-or bell-shaped corollas. Group II: plants with approx. 9-14 mL nectar with 40-60 % sugar and 4-10 mg sugar flower À1 ; mostly visited by long-tongued bees and/or hummingbirds. Group III: plants with 40-100 mL nectar with 30-40 % sugar and 14-36 mg sugar flower-1 , mostly visited by hummingbirds. Group IV: geoflorous plants with 80-90 mL with 10-15 % sugar and 8Á5-12 mg sugar flower-1 , presumably visited by small mammals. Groups II and III include species visited by bees and/or hummingbirds. Conclusions Pollinator switches from short-tongued bees via long-tongued bees to hummingbirds appear to have taken place repeatedly in the genera Nasa, Loasa and Caiophora. Changes in nectar amount and concentration appear to evolve rapidly with little phylogenetic constraint.

Research paper thumbnail of A revision of loasoid Caiophora (Caiophora pterosperma-group, Loasoideae, Loasaceae) from Peru

ABSTRACT Caiophora is a taxonomically difficult, nearly exclusively Andean genus of the largely S... more ABSTRACT Caiophora is a taxonomically difficult, nearly exclusively Andean genus of the largely South American family Loasaceae subfam. Loasoideae. Elevational distribution and flower morphology argue for a relatively basal position of loasoid Caiophora in the genus. Caiophora has not been revised since 1900, and details of their morphology, distribution and species delimitation are incompletely understood. The Caiophora pterosperma-group clearly belongs to Caiophora based on habit, fruit morphology, karyology and molecular data, but is florally similar to members of the closely allied genera Loasa and Scyphanthus. The Peruvian members of the Caiophora pterosperma-group are here revised. Three species are recognized: C. pterosperma, endemic to the departments of Junín and Pasco (including C. smithii, C. serropetala and C. pavonii), C. stenocarpa from the departments Cuzco and Huancavelica, and the new species Caiophora dederichiorum, endemic to the department of Ancash. All three species occur at low elevations for the genus (down to 2200 m a.s.l.) and in seasonally dry habitats, unlike most representatives of the genus (usually found in mesic habitats at higher elevations). A key to the Peruvian members of the group, diagnoses, drawings and photographs are provided for all species recognized.

Research paper thumbnail of A preliminary phylogeny of Loasaceae subfam. Loasoideae (Angiospermae: Cornales) based on L sequence data, with consequences for systematics and historical biogeography

Organisms Diversity & Evolution, 2004

... 7080). (C) Serrate petal margin (spm) in the bee-pollinated flower of Caiophora pterosperma (... more ... 7080). (C) Serrate petal margin (spm) in the bee-pollinated flower of Caiophora pterosperma (CaPt, Weigend & Dostert, 97/27). ... absent. (D) Pendulous flower ofbee-pollinated Nasa carunculata (NaCa, Weigend et al., 5035). ...

Research paper thumbnail of The seeds of Loasaceae subfam. Loasoideae (Cornales) II: Seed morphology of “South Andean Loasas” (Loasa, Caiophora, Scyphanthus and Blumenbachia)

Flora - Morphology, Distribution, Functional Ecology of Plants, 2005

South Andean Loasas (Blumenbachia, Caiophora, Loasa, Scyphanthus) are a monophyletic group of tax... more South Andean Loasas (Blumenbachia, Caiophora, Loasa, Scyphanthus) are a monophyletic group of taxa within Loasaceae subfam. Loasoideae, comprising some 100 species, 49 of which are investigated here. They retain a manylayered testa in the mature seeds and usually have passive transfer testas with complex, spongiose wall outgrowths. Additional modifications concern the undulations of the testa epidermis, presence or absence of the outer periclinal wall, secondary sculpturing, the presence or absence of spines, warts and finally spongiose structures on the anticlinal walls of the testa epidermis and the inner periclinal wall. Seeds of the widespread ''deeply pitted'' type are plesiomorphic, while various subclades of South Andean Loasas have derivations underscoring their relationships and confirming the relationships found with molecular markers and other morphological characters. The genus Blumenbachia has either seeds with a many-layered testa forming longitudinal lamellae (sect. Angulatae), or balloon seeds with a loose outer testa layer and spongiose wall outgrowths on the inner periclinal walls (sect. Blumenbachia and sect. Gripidea) and is clearly monophyletic. Loasa s.str. (ser. Loasa, ser. Macrospermae, ser. Floribundae, ser. Deserticolae) is characterized by the presence of a subterminal hilum or hilar scar and one subgroup (ser. Loasa, ser. Macrospermae) by very large and heavy seeds with a collapsed testa. L. ser. Pinnatae, ser. Acaules, ser. Volubiles, Scyphanthus and Caiophora share more or less one seed types with minor modifications. Within Caiophora various derivations are observed, of which the gradual loss of the secondary sculpture of the inner periclinal wall is the most striking one. Anemochoria is the most widespread dispersal mechanism in South Andean Loasas and is achieved in at least five structurally different ways.

Research paper thumbnail of De Liliifloris Notulae 9. The only hitherto known Spiloxene species (Hypoxidaceae) from Namibia is a new species, Spiloxene etesionamibensis, and a new Spiloxene species from Namaqualand (Northern Cape), S. namaquana

Feddes Repertorium, 2011

The only Namibian species of Spiloxene is described and illustrated by drawings and photos as Spi... more The only Namibian species of Spiloxene is described and illustrated by drawings and photos as Spiloxene etesionamibensis U.MÜLL.-DOBLIES, MARK.ACKER-MANN, WEIGEND &. D.MÜLL.-DOBLIES. It was hitherto misinterpreted as Spiloxene scullyi (BAKER) GARSIDE in SÖLCH & ROESSLER (1969) in MERX-MÜLLER's Prodromus einer Flora von Südwestafrika, but its closest ally is S. flaccida (NEL) GARSIDE, found in the Western and the Eastern Cape. Spiloxene etesionamibensis, which was found independently by GIESS & MERXMÜLLER and by MÜLLER-DOBLIES, is only known from two or three independent collections from a single locality. S. scullyi is no longer known from Namibia. In addition to the eight Spiloxene species already known from the Northern Cape Spiloxene namaquana U.MÜLL.-DOBLIES, MARK.ACKERMANN, WEIGEND & D.MÜLL.-DOBLIES is described and illustrated by drawings and photos. Its closest ally is Spiloxene trifurcillata (NEL) FOURC. from the Eastern Cape. For Spiloxene namaquana the morphological analysis of the branching pattern of an annual shoot taken in 1988 in the field is evaluated by a longitudinal and a horizontal diagram with explanatory remarks. The unusual innovation system of extreme anisoiteration is discussed. On Map 1 the quarter degree squares of the type localities (± only known localities) of both species is given.

Research paper thumbnail of Reloading the revolver - male fitness as a simple explanation for complex reward partitioning in Nasa macrothyrsa (Loasaceae, Cornales)

Biological Journal of the Linnean Society, 2010

Reward partitioning and replenishment and specific mechanisms for pollen presentation are all gea... more Reward partitioning and replenishment and specific mechanisms for pollen presentation are all geared towards the maximization of the number of effective pollinator visits to individual flowers. An extreme case of an apparently highly specialized plant-pollinator interaction with thigmonastic pollen presentation has been described for the morphologically complex tilt-revolver flowers of Caiophora arechavaletae (Loasaceae) pollinated by oligolectic Bicolletes pampeana (Colletidae, Hymenoptera). We studied the floral biology of Nasa macrothyrsa (Loasaceae) in the field and in the glasshouse, which has very similar floral morphology, but is pollinated by polylectic Neoxylocopa bees (Apidae, Hymenoptera). We investigated the presence of thigmonastic anther presentation, visitor behaviour (pollinators and nectar robbers), coordination of pollinator visits with flower behaviour and the presence of nectar replenishment. The aim of this study was to understand whether complex flower morphology and behaviour can be explained by a specialized pollination syndrome, or whether alternative explanations can be offered. The results showed that Nasa macrothyrsa has thigmonastic pollen presentation, i.e. new pollen is rapidly (<< 10 min) presented after a pollinator visit. Nectar secretion is independent of removal and averages 7-14 mL h-1. The complex flowers, however, fail to exclude either native (hummingbirds) or introduced (honeybees) nectar robbers, nor does polylectic Neoxylocopa actively collect the pollen presented. The findings do not support a causal link between complex flower morphology and functionality in Loasaceae and a highly specialized pollination. Rapid pollen presentation is best explained by the pollen presentation theory: the large proportion of pollinators coming shortly after a previous visit find little nectar and are more likely to move on to a different plant. The rapid presentation of pollen ensures that all these valuable 'hungry pollinators' are dusted with small pollen loads, thus increasing the male fitness of the plant by increasing the likelihood of siring outcrossed offspring.

Research paper thumbnail of Hybridization and crossability in Caiophora (Loasaceae subfam. Loasoideae): Are interfertile species and inbred populations results of a recent radiation?

American Journal of Botany, 2008

Interspecifi c hybridization is considered a possible mechanism of plant diversifi cation. The An... more Interspecifi c hybridization is considered a possible mechanism of plant diversifi cation. The Andes are a hotspot of biodiversity, but hybridization in Andean taxa has so far not been investigated intensively. The current study investigates crossability in Caiophora (Loasaceae subfam. Loasoideae) by experimental interspecifi c hybridization of seven different species. Hand pollination was undertaken, developing fruits counted, thousand (seed) grain weights, and seed viability were examined. Cross pollination led to some fruit set in 36 of the 37 different combinations. Overall fruit set was virtually identical irrespective of the combination of parental plants. Mean germination rates were much higher in hybrid seeds, indicating a marked heterosis effect and the possible presence of an inbreeding depression in the source populations: In experimental hybridization the divergent taxa of Caiophora behave like isolated, inbred populations of a single species. Allopatry and different habitat preferences seem to be the key factors keeping the (interfertile) taxa of Caiophora apart in the apparent absence of both postmating isolating mechanisms and obvious isolating mechanisms in phenology and fl oral biology. Interspecifi c hybrids reported from the wild appear to be the result of secondary contact due to human impact.

Research paper thumbnail of Flower reshaping in the transition to hummingbird pollination in Loasaceae subfam. Loasoideae despite absence of corolla tubes or spurs

Many angiosperm lineages present transitions from bee to hummingbird pollination. The flower desi... more Many angiosperm lineages present transitions from bee to hummingbird pollination. The flower design in most of these lineages includes either corolla tubes or nectar spurs, structures that commonly experienced an elongation with the acquisition of hummingbird pollination. It is proposed that this increases the fit between the bird head and flower structures, and isolates or partially blocks bees from the interaction. But can this transition occur if the ancestral flower design lacks tubes or spurs? Here we focus on the transition from bee to hummingbird pollination in the Loasaceae subfamily Loasoideae. Loasoideae flowers have radial corollas with separated petals; therefore, they do not display corolla tubes nor nectar spurs. These flowers also present a whorl of nectar scales and staminodes, unique to the subfamily, which is involved in flower–pollinator fit and in nectar harvesting. To explore flower shape adaptation to hummingbird pollination, we tested for correspondence between pollinators and flower shape in Loasoideae. In order to achieve this, we first compared the evolutionary history of flower phenotype and pollination mode, and then used stochastic character mapping and geometric-morphometric variables in a comparison of alternative evolutionary models. The results of our study suggest that the transition from bee to bird pollination was accompanied by changes in the shape of the staminodial complex, along with the evolution of relatively closed corollas. Moreover, while bird pollination seems to be the end point in the evolution of pollination syndromes in many angiosperm lineages, rodent pollinated flowers probably evolved from ancestral bird pollinated flowers in Loasoideae. Our findings suggest that the evolution of bird pollinated flowers from ancestral bee pollinated flowers does not require the presence of corolla tubes or spurs, and can take place as long as the flower design includes structures participating in flower–pollinator fit.

Research paper thumbnail of A question of data quality-Testing pollination syndromes in Balsaminaceae

PloS one, 2017

Pollination syndromes and their predictive power regarding actual plant-animal interactions have ... more Pollination syndromes and their predictive power regarding actual plant-animal interactions have been controversially discussed in the past. We investigate pollination syndromes in Balsaminaceae, utilizing quantitative respectively categorical data sets of flower morphometry, signal and reward traits for 86 species to test for the effect of different types of data on the test patterns retrieved. Cluster Analyses of the floral traits are used in combination with independent pollinator observations. Based on quantitative data we retrieve seven clusters, six of them corresponding to plausible pollination syndromes and one additional, well-supported cluster comprising highly divergent floral architectures. This latter cluster represents a non-syndrome of flowers not segregated by the specific data set here used. Conversely, using categorical data we obtained only a rudimentary resolution of pollination syndromes, in line with several earlier studies. The results underscore that the use ...

Research paper thumbnail of Erratum to: Diversification of Caiophora (Loasaceae subfam. Loasoideae) during the uplift of the Central Andes

Organisms Diversity & Evolution

Research paper thumbnail of Phylogenetic relationships and generic re-arrangements in "South Andean Loasas" (Loasaceae)

Taxon

Ver impacto de la revista Resumen Loasaceae, a mostly American group, is one of the largest famil... more Ver impacto de la revista Resumen Loasaceae, a mostly American group, is one of the largest families of Cornales. In spite of

Research paper thumbnail of Flower reshaping in the transition to hummingbird pollination in Loasaceae subfam. Loasoideae despite absence of corolla tubes or spurs

Evolutionary Ecology, 2016

Many angiosperm lineages present transitions from bee to hummingbird pollination. The flower desi... more Many angiosperm lineages present transitions from bee to hummingbird pollination. The flower design in most of these lineages includes either corolla tubes or nectar spurs, structures that commonly experienced an elongation with the acquisition of hummingbird pollination. It is proposed that this increases the fit between the bird head and flower structures, and isolates or partially blocks bees from the interaction. But can this transition occur if the ancestral flower design lacks tubes or spurs? Here we focus on the transition from bee to hummingbird pollination in the Loasaceae subfamily Loasoideae. Loasoideae flowers have radial corollas with separated petals; therefore, they do not display corolla tubes nor nectar spurs. These flowers also present a whorl of nectar scales and staminodes, unique to the subfamily, which is involved in flower–pollinator fit and in nectar harvesting. To explore flower shape adaptation to hummingbird pollination, we tested for correspondence between pollinators and flower shape in Loasoideae. In order to achieve this, we first compared the evolutionary history of flower phenotype and pollination mode, and then used stochastic character mapping and geometric-morphometric variables in a comparison of alternative evolutionary models. The results of our study suggest that the transition from bee to bird pollination was accompanied by changes in the shape of the staminodial complex, along with the evolution of relatively closed corollas. Moreover, while bird pollination seems to be the end point in the evolution of pollination syndromes in many angiosperm lineages, rodent pollinated flowers probably evolved from ancestral bird pollinated flowers in Loasoideae. Our findings suggest that the evolution of bird pollinated flowers from ancestral bee pollinated flowers does not require the presence of corolla tubes or spurs, and can take place as long as the flower design includes structures participating in flower–pollinator fit.

Research paper thumbnail of Notas sobre el Género Caiophora (Loasoideae, Loasaceae) en Chile y países limítrofes

Darwiniana

Esta es la primera revisión del género Caiophora (Loasaceae) en Chile. El género comprende alrede... more Esta es la primera revisión del género Caiophora (Loasaceae) en Chile. El género comprende alrededor de 60 especies, y está ampliamente distribuido en los Andes, desde Argentina/Chile en el Sur, hasta el centro de Ecuador en el Norte. Se conocen solamente cinco especies de Chile: Caiophora chuquitensis, C. cirsiifolia, C. coronata, C. deserticola sp. nov. y C. rosulata. El material de Caiophora rosulata se divide entre dos subespecies, C. rosulata subsp. rosulata comb. nov., de los Andes Occidentales (presente en el norte de Chile y el sur de Perú, y C. rosulata subsp. taraxacoides stat. y comb. nov., de los Andes Orientales. Los nombres C. superba syn. nov. y C. macrocarpa syn. nov. son sinonimizados bajo C. chuquitensis, y C. rahmeri syn. nov., sinonimizado bajo C. rosulata subsp. rosulata. Estas cinco especies comprenden el rango completo de hábitos conocido para el género: sufrútices, hierbas perennes en forma de cojines, hierbas rosuladas acaules, y hierbas trepadoras. Proporcionamos una clave, descripciones morfológicas y la sinonimia completa para todos los taxones, incluyendo ilustraciones, notas sobre la distribución, el hábitat, la biología floral y números cromosómicos.

Research paper thumbnail of Los nombres antiguos en el género Caiophora (Loasaceas subfam. Loasoideae) y una clasificación infragenérica preliminar

EI presente trabajo se enfoca en establecer la identidad sistemälica de los nombres antiguos en e... more EI presente trabajo se enfoca en establecer la identidad sistemälica de los nombres antiguos en el genero Caiophora sobre la base de estudios de campo y herbario. incluyendo varios e1ementos tipicos originales. Se propone aceptar los siguientes nombres para los taxones centrales de la flora peruana : Caiophora pedullcu/aris (K.Presl) Weigend & M.Ack., combo llOV. ( - Loasa pedullCli/aris K.Presl, = Caiophora pallciseta Killip), Caioplwra cirsiifolia K.Presl. (= Caiophora sepiaria (Ruiz & Pav. ex G.Oon) J.F.Macbr., = Caiophora preslii Urb. & Gilg), Caioplwra carduifolia K.Presl (= B/limenbachia punicea Ruiz & Pav. ex G.Don, = Caiophora cymbifera Urb. & Gilg), Caioplrora gralldiflora (G.Oon) Weigend & M.Ack. , combo nov. (=Bllimenbachia grandijlora G.Oon =Loasaphysopetala Ruiz & Pav.) y Caioplwra pterosperma (Ruiz & Pav. ex. G.Oon) Urb. & Gilg (",Bllimenbachia pterasperma Ruiz & Pav. ex G.Don =Loasa pterasperma Ruiz & Pav. = Caiophora serropetala Macbr., = Caiophora smithii Killi...

Research paper thumbnail of Notes on the genus Caiophora (Loasoideae, Loasaceae) in Chile and neighbouring countries

This is the first revision for the representatives of the genus Caiophora (Loasaceae) in Chile. T... more This is the first revision for the representatives of the genus Caiophora (Loasaceae) in Chile. The genus is widely distributed in the Andes from Argentina/Chile in the South to Central Ecuador in the North, and comprises approximately 60 species. In Chile only five species are present, Caiophora chuquitensis, C. cirsiifolia, C. coronata, C. deserticola sp. nov. and C. rosulata. Caiophora rosulata is here subdivided into two subspecies: western Andean C. rosulata subsp. rosulata (present in northern Chile and southern Peru), and eastern Andean C. rosulata subsp. taraxacoides, stat. and comb. nov. Furthermore C. superba syn. nov. and C. macrocarpa syn. nov. are placed into synonymy under C. chuquitensis, and C. rahmeri syn. nov. is synonymized to C. rosulata subsp. rosulata. These five species comprise the complete range of growth forms known for the genus, i.e., subshrubs, cushionforming herbs, acaulescent, rosulate herbs and vines. For all taxa a key and full morphological descript...

Research paper thumbnail of Nectar, floral morphology and pollination syndrome in loasaceae subfam. Loasoideae (Cornales)

Annals of Botany, 2006

Background and Aims Loasaceae subfam. Loasoideae are mostly distributed in South America (sea lev... more Background and Aims Loasaceae subfam. Loasoideae are mostly distributed in South America (sea level to over 4500 m) with a wide range of animals documented as pollinators. The aim was to investigate correlations between nectar parameters, flower morphology, pollination syndrome and phylogeny. Methods Nectar was collected from 29 species from seven genera in the subfamily. Concentration and volumes were measured and the amount of sugar calculated. Correlations of nectar data were plotted on a ternary graph and nectar characteristics compared with flower visitors, floral morphology and phylogenetic data. Key Results Sugar concentrations are generally higher than reported for most plant families in the literature. The species investigated can be roughly grouped as follows. Group I: plants with approx. 1Á5(-3Á5) mL nectar with (40-)60-80 % sugar and 0Á19-2 mg sugar flower À1 ; with small, white, star-shaped corollas, pollinated by shorttongued bees. Groups II, III and IV: plants with mostly orange, balloon-, saucer-, bowl-or bell-shaped corollas. Group II: plants with approx. 9-14 mL nectar with 40-60 % sugar and 4-10 mg sugar flower À1 ; mostly visited by long-tongued bees and/or hummingbirds. Group III: plants with 40-100 mL nectar with 30-40 % sugar and 14-36 mg sugar flower-1 , mostly visited by hummingbirds. Group IV: geoflorous plants with 80-90 mL with 10-15 % sugar and 8Á5-12 mg sugar flower-1 , presumably visited by small mammals. Groups II and III include species visited by bees and/or hummingbirds. Conclusions Pollinator switches from short-tongued bees via long-tongued bees to hummingbirds appear to have taken place repeatedly in the genera Nasa, Loasa and Caiophora. Changes in nectar amount and concentration appear to evolve rapidly with little phylogenetic constraint.

Research paper thumbnail of A revision of loasoid Caiophora (Caiophora pterosperma-group, Loasoideae, Loasaceae) from Peru

ABSTRACT Caiophora is a taxonomically difficult, nearly exclusively Andean genus of the largely S... more ABSTRACT Caiophora is a taxonomically difficult, nearly exclusively Andean genus of the largely South American family Loasaceae subfam. Loasoideae. Elevational distribution and flower morphology argue for a relatively basal position of loasoid Caiophora in the genus. Caiophora has not been revised since 1900, and details of their morphology, distribution and species delimitation are incompletely understood. The Caiophora pterosperma-group clearly belongs to Caiophora based on habit, fruit morphology, karyology and molecular data, but is florally similar to members of the closely allied genera Loasa and Scyphanthus. The Peruvian members of the Caiophora pterosperma-group are here revised. Three species are recognized: C. pterosperma, endemic to the departments of Junín and Pasco (including C. smithii, C. serropetala and C. pavonii), C. stenocarpa from the departments Cuzco and Huancavelica, and the new species Caiophora dederichiorum, endemic to the department of Ancash. All three species occur at low elevations for the genus (down to 2200 m a.s.l.) and in seasonally dry habitats, unlike most representatives of the genus (usually found in mesic habitats at higher elevations). A key to the Peruvian members of the group, diagnoses, drawings and photographs are provided for all species recognized.

Research paper thumbnail of A preliminary phylogeny of Loasaceae subfam. Loasoideae (Angiospermae: Cornales) based on L sequence data, with consequences for systematics and historical biogeography

Organisms Diversity & Evolution, 2004

... 7080). (C) Serrate petal margin (spm) in the bee-pollinated flower of Caiophora pterosperma (... more ... 7080). (C) Serrate petal margin (spm) in the bee-pollinated flower of Caiophora pterosperma (CaPt, Weigend &amp;amp; Dostert, 97/27). ... absent. (D) Pendulous flower ofbee-pollinated Nasa carunculata (NaCa, Weigend et al., 5035). ...

Research paper thumbnail of The seeds of Loasaceae subfam. Loasoideae (Cornales) II: Seed morphology of “South Andean Loasas” (Loasa, Caiophora, Scyphanthus and Blumenbachia)

Flora - Morphology, Distribution, Functional Ecology of Plants, 2005

South Andean Loasas (Blumenbachia, Caiophora, Loasa, Scyphanthus) are a monophyletic group of tax... more South Andean Loasas (Blumenbachia, Caiophora, Loasa, Scyphanthus) are a monophyletic group of taxa within Loasaceae subfam. Loasoideae, comprising some 100 species, 49 of which are investigated here. They retain a manylayered testa in the mature seeds and usually have passive transfer testas with complex, spongiose wall outgrowths. Additional modifications concern the undulations of the testa epidermis, presence or absence of the outer periclinal wall, secondary sculpturing, the presence or absence of spines, warts and finally spongiose structures on the anticlinal walls of the testa epidermis and the inner periclinal wall. Seeds of the widespread ''deeply pitted'' type are plesiomorphic, while various subclades of South Andean Loasas have derivations underscoring their relationships and confirming the relationships found with molecular markers and other morphological characters. The genus Blumenbachia has either seeds with a many-layered testa forming longitudinal lamellae (sect. Angulatae), or balloon seeds with a loose outer testa layer and spongiose wall outgrowths on the inner periclinal walls (sect. Blumenbachia and sect. Gripidea) and is clearly monophyletic. Loasa s.str. (ser. Loasa, ser. Macrospermae, ser. Floribundae, ser. Deserticolae) is characterized by the presence of a subterminal hilum or hilar scar and one subgroup (ser. Loasa, ser. Macrospermae) by very large and heavy seeds with a collapsed testa. L. ser. Pinnatae, ser. Acaules, ser. Volubiles, Scyphanthus and Caiophora share more or less one seed types with minor modifications. Within Caiophora various derivations are observed, of which the gradual loss of the secondary sculpture of the inner periclinal wall is the most striking one. Anemochoria is the most widespread dispersal mechanism in South Andean Loasas and is achieved in at least five structurally different ways.

Research paper thumbnail of De Liliifloris Notulae 9. The only hitherto known Spiloxene species (Hypoxidaceae) from Namibia is a new species, Spiloxene etesionamibensis, and a new Spiloxene species from Namaqualand (Northern Cape), S. namaquana

Feddes Repertorium, 2011

The only Namibian species of Spiloxene is described and illustrated by drawings and photos as Spi... more The only Namibian species of Spiloxene is described and illustrated by drawings and photos as Spiloxene etesionamibensis U.MÜLL.-DOBLIES, MARK.ACKER-MANN, WEIGEND &. D.MÜLL.-DOBLIES. It was hitherto misinterpreted as Spiloxene scullyi (BAKER) GARSIDE in SÖLCH & ROESSLER (1969) in MERX-MÜLLER's Prodromus einer Flora von Südwestafrika, but its closest ally is S. flaccida (NEL) GARSIDE, found in the Western and the Eastern Cape. Spiloxene etesionamibensis, which was found independently by GIESS & MERXMÜLLER and by MÜLLER-DOBLIES, is only known from two or three independent collections from a single locality. S. scullyi is no longer known from Namibia. In addition to the eight Spiloxene species already known from the Northern Cape Spiloxene namaquana U.MÜLL.-DOBLIES, MARK.ACKERMANN, WEIGEND & D.MÜLL.-DOBLIES is described and illustrated by drawings and photos. Its closest ally is Spiloxene trifurcillata (NEL) FOURC. from the Eastern Cape. For Spiloxene namaquana the morphological analysis of the branching pattern of an annual shoot taken in 1988 in the field is evaluated by a longitudinal and a horizontal diagram with explanatory remarks. The unusual innovation system of extreme anisoiteration is discussed. On Map 1 the quarter degree squares of the type localities (± only known localities) of both species is given.

Research paper thumbnail of Reloading the revolver - male fitness as a simple explanation for complex reward partitioning in Nasa macrothyrsa (Loasaceae, Cornales)

Biological Journal of the Linnean Society, 2010

Reward partitioning and replenishment and specific mechanisms for pollen presentation are all gea... more Reward partitioning and replenishment and specific mechanisms for pollen presentation are all geared towards the maximization of the number of effective pollinator visits to individual flowers. An extreme case of an apparently highly specialized plant-pollinator interaction with thigmonastic pollen presentation has been described for the morphologically complex tilt-revolver flowers of Caiophora arechavaletae (Loasaceae) pollinated by oligolectic Bicolletes pampeana (Colletidae, Hymenoptera). We studied the floral biology of Nasa macrothyrsa (Loasaceae) in the field and in the glasshouse, which has very similar floral morphology, but is pollinated by polylectic Neoxylocopa bees (Apidae, Hymenoptera). We investigated the presence of thigmonastic anther presentation, visitor behaviour (pollinators and nectar robbers), coordination of pollinator visits with flower behaviour and the presence of nectar replenishment. The aim of this study was to understand whether complex flower morphology and behaviour can be explained by a specialized pollination syndrome, or whether alternative explanations can be offered. The results showed that Nasa macrothyrsa has thigmonastic pollen presentation, i.e. new pollen is rapidly (<< 10 min) presented after a pollinator visit. Nectar secretion is independent of removal and averages 7-14 mL h-1. The complex flowers, however, fail to exclude either native (hummingbirds) or introduced (honeybees) nectar robbers, nor does polylectic Neoxylocopa actively collect the pollen presented. The findings do not support a causal link between complex flower morphology and functionality in Loasaceae and a highly specialized pollination. Rapid pollen presentation is best explained by the pollen presentation theory: the large proportion of pollinators coming shortly after a previous visit find little nectar and are more likely to move on to a different plant. The rapid presentation of pollen ensures that all these valuable 'hungry pollinators' are dusted with small pollen loads, thus increasing the male fitness of the plant by increasing the likelihood of siring outcrossed offspring.

Research paper thumbnail of Hybridization and crossability in Caiophora (Loasaceae subfam. Loasoideae): Are interfertile species and inbred populations results of a recent radiation?

American Journal of Botany, 2008

Interspecifi c hybridization is considered a possible mechanism of plant diversifi cation. The An... more Interspecifi c hybridization is considered a possible mechanism of plant diversifi cation. The Andes are a hotspot of biodiversity, but hybridization in Andean taxa has so far not been investigated intensively. The current study investigates crossability in Caiophora (Loasaceae subfam. Loasoideae) by experimental interspecifi c hybridization of seven different species. Hand pollination was undertaken, developing fruits counted, thousand (seed) grain weights, and seed viability were examined. Cross pollination led to some fruit set in 36 of the 37 different combinations. Overall fruit set was virtually identical irrespective of the combination of parental plants. Mean germination rates were much higher in hybrid seeds, indicating a marked heterosis effect and the possible presence of an inbreeding depression in the source populations: In experimental hybridization the divergent taxa of Caiophora behave like isolated, inbred populations of a single species. Allopatry and different habitat preferences seem to be the key factors keeping the (interfertile) taxa of Caiophora apart in the apparent absence of both postmating isolating mechanisms and obvious isolating mechanisms in phenology and fl oral biology. Interspecifi c hybrids reported from the wild appear to be the result of secondary contact due to human impact.

Research paper thumbnail of Flower reshaping in the transition to hummingbird pollination in Loasaceae subfam. Loasoideae despite absence of corolla tubes or spurs

Many angiosperm lineages present transitions from bee to hummingbird pollination. The flower desi... more Many angiosperm lineages present transitions from bee to hummingbird pollination. The flower design in most of these lineages includes either corolla tubes or nectar spurs, structures that commonly experienced an elongation with the acquisition of hummingbird pollination. It is proposed that this increases the fit between the bird head and flower structures, and isolates or partially blocks bees from the interaction. But can this transition occur if the ancestral flower design lacks tubes or spurs? Here we focus on the transition from bee to hummingbird pollination in the Loasaceae subfamily Loasoideae. Loasoideae flowers have radial corollas with separated petals; therefore, they do not display corolla tubes nor nectar spurs. These flowers also present a whorl of nectar scales and staminodes, unique to the subfamily, which is involved in flower–pollinator fit and in nectar harvesting. To explore flower shape adaptation to hummingbird pollination, we tested for correspondence between pollinators and flower shape in Loasoideae. In order to achieve this, we first compared the evolutionary history of flower phenotype and pollination mode, and then used stochastic character mapping and geometric-morphometric variables in a comparison of alternative evolutionary models. The results of our study suggest that the transition from bee to bird pollination was accompanied by changes in the shape of the staminodial complex, along with the evolution of relatively closed corollas. Moreover, while bird pollination seems to be the end point in the evolution of pollination syndromes in many angiosperm lineages, rodent pollinated flowers probably evolved from ancestral bird pollinated flowers in Loasoideae. Our findings suggest that the evolution of bird pollinated flowers from ancestral bee pollinated flowers does not require the presence of corolla tubes or spurs, and can take place as long as the flower design includes structures participating in flower–pollinator fit.