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

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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 many-layered 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.

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... Blumenbachia (Figs. 2E,4B) and these two in turn are sister to the South Andean sect. Angulatae (Fig. 2C) as previously argued based on inflorescence, fruit and seed morphology (Weigend, 1997;Weigend & al., 2005;Henning & al., 2015). ...
... The serrate petal margins (Figs. 2K, L, 3A; lost in the most-derived clades of Caiophora: Fig. 3B, C), elaborate floral scales, finely dissected leaves (Fig. 4F-I), and deeply pitted seeds with fenestrate anticlinal walls (Weigend & al., 2005) support the molecular data from a morphological perspective. Scyphanthus and Caiophora share fruits opening with longitudinal dehiscence, whereas Loasa ser. ...
... Deserticolae) form a moderately-supported clade according to the plastid data. They have been consistently considered as closely related in the past (Urban & Gilg, 1900) and have similar flowers and floral scales (Weigend & al., 2004b) (Fig. 3F-H), as well as seeds unique in Loasoideae for their subterminal hilum or hilar scar (Weigend & al. 2005). This clade is resolved into two sister clades, which are clearly consistent with vegetative, flower, fruit, and seed morphology (Urban & Gilg, 1900;Weigend & al., 2004bWeigend & al., , 2005: Large-seeded and large-leaved ser. ...
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Loasaceae, a mostly American group, is one of the largest families of Cornales. In spite of considerable progress over the last 20 years, the relationships of some clades remain controversial, especially in the "South Andean Loasas" (SAL—Blumenbachia, Caiophora, Loasa, Scyphanthus). The present study addresses the phylogenetic relationships in SAL employing four plastid markers (rps16, trnL-trnF, trnS-trnG, matK) and ITS and aims at resolving the systematics and evolution of the group. Sequences obtained from a total of 59 SAL species (ca. 70% of the total, representing all lineages in the group) and 25 outgroup taxa were analysed using maximum likelihood and Bayesian inference approaches. ML best and BI strict consensus trees showed no significant differences in their topologies. Our results confirm that two species of Loasa ser. Malesherbioideae are not part of the SAL clade, but should be included in Presliophytum, a result which is here formalized. Blumenbachia (including sect. Angulatae and sect. Gripidea) is confirmed as a monophylum with high support. Loasa has to be redefined and restricted to a clade including only ser. Deserticolae, ser. Floribundae, ser. Loasa and ser. Macrospermae. Scyphanthus and Caiophora both are each monophyla and sister groups, but with two clades of Loasa as successive sister groups: (((Caiophora + Scyphanthus) + Loasa ser. Pinnatae) + (L. ser. Volubiles + L. ser. Acaules)) in a very well-supported clade. Accordingly, Caiophora, Loasa ser. Pinnatae, L. ser. Volubiles, L. ser. Acaules and Scyphanthus could be included into a single genus, with Scyphanthus taking priority over Caiophora, creating a fairly heterogenous genus of ca. 52 species and requiring 50 new names. Alternatively, the clades Loasa ser. Pinnatae and L. ser. Volubiles + ser. Acaules can be removed into new segregate genera, which is here argued for and which requires the creation of only 16 new names. The new genus names and some of the new combinations are here formalized.
... Large dust particles from these events were detected in traps in the north of the province of Buenos Aires, and according to the modelling of particle deposition and concentration, these dust storms may have also transported large dust particles from the Puna-Altiplano Plateau to the Uruguayan and Brazilian grasslands (Gaiero et al. 2013). Interestingly, the seeds of most of the BSouth Andean Loasas^, which include Loasa, Caiophora, Scyphanthus and Blumenbachia, are dispersed by wind (Weigend et al. 2005). Seeds present specialized testa structures that constitute an adaptation for aeolic transportation (Weigend et al. 2005). ...
... Interestingly, the seeds of most of the BSouth Andean Loasas^, which include Loasa, Caiophora, Scyphanthus and Blumenbachia, are dispersed by wind (Weigend et al. 2005). Seeds present specialized testa structures that constitute an adaptation for aeolic transportation (Weigend et al. 2005). Based on the weight, the linear measurements and the winglike structures of C. arechavaletae seeds (Weigend et al. 2005), strong winds may have transported Caiophora seeds from the Puna-Altiplano Plateau to the Pampas grasslands, where these may have been deposited by rain (Gaiero, personal communication). ...
... Seeds present specialized testa structures that constitute an adaptation for aeolic transportation (Weigend et al. 2005). Based on the weight, the linear measurements and the winglike structures of C. arechavaletae seeds (Weigend et al. 2005), strong winds may have transported Caiophora seeds from the Puna-Altiplano Plateau to the Pampas grasslands, where these may have been deposited by rain (Gaiero, personal communication). ...
Andean orogeny and the ecological changes that followed promoted diversification in plant and animal lineages since the Early Miocene. The angiosperm genus Caiophora (Loasaceae, subfam. Loasoideae) comprises around 50 species that are endemic to South America. These are distributed from southern Ecuador to Central Chile and Argentina. Bee pollination and distribution at low-intermediate elevations probably represent the ancestral condition in the lineage that includes Caiophora and its allied genera. The majority of Caiophora species grow at high elevations in the Andes, where some depend on vertebrate pollination. Previous studies did not resolve phylogenetic relationships within Caiophora, which precluded the dating of the origin and divergence of this group. We used markers of one nuclear (ITS) and one plastid region (trnSGCU-trnGUUC) to solve phylogenetic relationships among 19 Caiophora species (including different accessions). We also included 10 species of the allied genera Blumenbachia and Loasa. Aosa rostrata and Xylopodia klaprothioides were used as outgroups. Phylogenetic reconstruction strongly supports the monophyly of Caiophora, and although several clades within this genus are poorly supported, our study yielded a better infra-generic resolution than previous studies. The origin of Caiophora is dated to the Early-Middle Miocene and can be related to the uplift of the Cordilleras Frontal and Principal and to Pacific marine transgressions. According to our estimations, Caiophora began to diversify during the Middle-Late Miocene and this unfolding proceeded eastwards during the Pliocene and the Pleistocene, in parallel to the uplift of different Andean mountain ranges.
... In spite of its relatively moderate number of species, the family is morphologically highly diversified (Fig. 1). Numerous studies have revealed a high level of diversity for growth-and life-forms 57 , leaf morphology and wood anatomy [58][59][60][61] , pollen-and seed morphology 62,63 , indumentum [64][65][66] and especially floral morphology [67][68][69][70] . Most of the floral diversification is found in subfam. ...
... It has been argued that "…much plant taxonomy relies on flower structure in which plasticity is minimized" Trewavas (p. 15 111 ) It is undoubtedly true in general terms that the basic architecture of Loasoideae-flowers is remarkably conserved 112 . This argument could be contrasted with the notable behavioural diversity documented here for the first time, but this would underestimate the extreme diversification in the details of flower morphology (Fig. 1), in regards to aspects of function and signalling 60,62,63,70 . Similarly, the primary floral reward in Loasoideae is highly diverse and the broad range of nectar amounts and concentrations has been shown to correlate with pollination syndromes 78 . ...
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Obvious movements of plant organs have fascinated scientists for a long time. They have been studied extensively, but few behavioural studies to date have dealt with them, and hardly anything is known about their evolution. Here, we present a large experimental dataset on the stamen movement patterns found in the Loasaceae subfam. Loasoideae (Cornales). An evolutionary transition from autonomous-only to a combination of autonomous and thigmonastic stamen movement with increased complexity was experimentally demonstrated. We compare the stamen movement patterns with extensive pollinator observations and discuss it in the context of male mating behavior. Thigmonastic pollen presentation via stamen movements appears to be a crucial component of floral adaptation to pollinator behaviour, evolving in concert with complex adjustments of flower signal, reward and morphology. We hypothesize that rapid adjustments of pollen presentation timing may play a significant role in the diversificationof this plant group, representing a striking example for the evolutionary significance of plant behaviour.
... Fig. 3A-B; compareWeigend et al. 2005), or respectively with a LEO 1450 SEM(Fig. 3C-F) at a voltage of 10 or 15 kV. ...
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.
... Taxa are primarily kept apart by ecogeographical isolation. Secondary contact from human impact leads to hybrid formation , while under natural conditions the effi ciently wind-dispersed seed of Caiophora ( Weigend et al., 2004a Weigend et al., , 2005 ) may ensure the occasional contact between different taxa. Problems with taxon delimitation in Caiophora are likely at least partly due to the presence of specimens with " mixed " or " intermediate " morphological characters as a result of hybridization and possibly introgression. ...
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Interspecific hybridization is considered a possible mechanism of plant diversification. 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 interspecific 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 floral biology. Interspecific hybrids reported from the wild appear to be the result of secondary contact due to human impact.
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Acuña, R. & M. Weigend. 2018. The generic affinity of Caiophora pulchella (Loasaceae, Loasoideae). Darwiniana, nueva serie 6(1): 94-98. The history of the nomenclatural changes and past hypotheses on the relationships of Caiophora pulchella are presented. The identity of voucher specimens, identified as C. nivalis in previous phylogenetic studies, is reassessed based on fruit morphology. The validity of C. pulchella is reconsidered and can be confirmed as a member of the genus Caiophora based on both morphological and molecular data. Alternative generic placements are rejected.
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Members of Loasaceae subfamily Gronovioideae have distinctive flowers compared to those of other members of the family. In contrast to all other Loasaceae, they have haplostemonous androecia and uniovulate gynoecia. The haplostemonous androecia are paedomorphic. The two major clades of gronovioids-(1) Petalonyx and (2) Cevallia, Fuertesia, and Gronovia-differ in their avenues of diversification. The dade consisting of Cevallia, Fuertesia, and Gronovia has floral variation that is manifested most prominently in stamen, stigma, and nectary forms, although petal shapes and the positions of anthers relative to petals and stigmas also vary. The Petalonyx clade exhibits variation that is manifested in morphological differentiation among stamens (Petalonyx crenatus has staminodes) and in postgenital fusion in the corolla (lacking only in Petalonyx linearis and P. crenatus). Developmental studies have also shown that Gronovioideae share with their sister group Mentzelia the loss of a petal-stamen synorganization that is present in other investigated members of Loasaceae.
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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 Killip, = Caiophora pavonii Urb. & Gilg). Sobre la base de esta aclaracion de los nombres importantes se propone una division infragenerica provisional dei genero Caiophora en un total de 10 grupos, 8 de los cuales estan presentes en el Peru, los grupos C. pterosperma, C. chuqllitensis. C. contorta, C. coranata, C. cirsiifolia, C. lateritia, C. cardllifolia. Se presenta una tabla con los caracteres diagnosticos de los grupos y un listado de las especies pertenecientes a cada uno.
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The pharmaceutically interesting Δ6-FA 18∶3Δ6c, 9c, 12c (γ-linolenic acid) and 18∶4Δ6c,9c,12c,15c (stearidonic acid) appear to have evolved independently several times during plant phylogenetic evolution. They typically occur in “clusters” of a few closely related species or genera in about a dozen different plant families throughout the plant kingdom. A hither-unknown “cluster of occurrence” has now been discovered in the New World plant family Loasaceae. γ-Linolenic and stearidonic acids occur exclusively in representatives of the newly described genus Nasa at significance levels of between 3 and 10% each. Nasa had recently been separated from the older, more broadly circumscribed genus Loasa. The two Δ6-FA were not found in the closely related genus Loasa sensu stricto, nor in a number of other representatives of Loasaceae.
The seed functions as the reproductive unit of the Spermatophyta (seed plants), and links the successive generations. Other important seed functions concern dispersal and survival under cold, dry or other inclement conditions. There is an immense diversity in the internal and external structure of seeds. These differences are, to an appreciable extent, related to dispersal and germination strategies, and may involve the size and position of endosperm and embryo, structure, texture, and color of seed-coat, and the shape and dimensions of the seed as a whole. The minute seeds of orchids may weigh as little as 0,000002 g each. The seeds of the legume Mora oleifera are possibly the heaviest, and the weight of each exceeds 1,000 g, whereas a one-seeded fruit of the palm Lodoicea maldivica weighs more than 20,000 g. The seed consists of three components: embryo, endosperm (sometimes perisperm), and seed-coat. Both endosperm and embryo are the products of double fertilization, whereas the seed-coat develops from the maternal, ovular tissues. The seed habit is a significant advancement in the evolution of higher plants. Seed plants show several evolutionary advantages over spore-producing plants. Fertilization takes place within the protective tissues of the mother plant, and during its development the embryo is nourished by the mother plant. Besides, the embryo is covered by the protective seed-coat and often provided with storage material. These factors made the seed plants so successful that they became the dominant component in the terrestrial environments of the earth.
Seed morphology of Hydrangeaceae was explored to describe the parameters of diversity, define characters and character states, and hypothesize monophyletic groups in the family on the basis of derived features. Eleven seed characters were defined that delimit four basic aspects of morphology, including (1) shape, (2) primary sculpture, (3) secondary sculpture, and (4) appendages. Character states could not be delimited readily in some aspects of diversity, including transectional shape and length of seeds. Despite continuous variation in seed length, when mapped on previous cladograms these data supported the hypothesis that seeds less than 1.0 mm long were plesiomorphic and that seed size had increased separately in different subclades. Seed size increased in association with wing loss in Fendlera, Fendlerella, and Whipplea. A shift from apical, interstylar dehiscence of fruits that produce a very limited gap for seed emergence in most members of the hydrangea clade to fruit wall fragmentation may have p...
Morphological features of seeds of Eucnide and species of subfamily Loasoideae, including the genera Blumenbachia, Cajophora, Loasa, Sclerothrix, and Scyphanthus, were examined to better understand relationships among Loasaceae. Seeds of Eucnide are bipolar with the hilum at one pole. They are essentially cylindrical with a radial transectional symmetry. Longitudinal ridges extend between the poles of each seed. The seed surface is also characterized by irregular granulations that may be large and rough, small and fine, or even largely indistinct, depending upon the species. As has been reported for other loasaceous genera, morphological features of seeds of Eucnide are generally conservative at the sectional level. Differences at the sectional level include size, shape, and secondary surface sculpture. Examination of herbarium collections has shown that E. hirta is more morphologically diverse than other species of Eucnide. Secondary surface sculpture of seeds is used to support the proposition that E. pringlei, recently considered conspecific with E. hirta, is a distinct species. Seeds of a disjunct population of E. hypomalaca have been valuable for showing potential evolutionary relationships between this species and E. pringlei and E. cordata. Seeds of Eucnide sect. Sympetaleia are, perhaps, paedomorphic derivatives of an ancestor with a seed morphology similar to that currently presently in E. bartonioides. Among examined species of subfamily Loasoideae, seeds appear primarily to have reticulate surface sculpturing. This reticulate pattern may be a plesiomorphic condition of the family shared by certain sections of various genera of subfamily Loasoideae and Mentzelia (subfamily Mentzelioideae). Seeds of Loasa sect. Huidobria have a surface sculpture composed of longitudinal ridges or folds that superficially cause seeds of these species to resemble those of Eucnide. Certain realignments of species of Cajophora with other genera in subfamily Loasoideae that were suggested by Poston do not seem necessarily appropriate based on data of seed structures reported in her study and this one.
Klaprothieae includes two monotypic genera from Central and South America, Klaprothia and Sclerothrix. These are the only two neotropical Loasaceae with 4-merous flowers, reduced floral scales, staminodial structures, and chromosome numbers that suggest a polyploid relationship. Merger of the two genera is proposed and Sclerothrix fasciculata becomes Klaprothia fasciculata, comb. nov.
Six hundred seeds belonging to the genusMentzelia (Loasaceae) were examined using the Scanning Electron Microscope. The seeds represented all six sections of the genus and approximately 76% of the total number of species from northern Mexico and western United States. Emphasis was placed on seed material from Wyoming and adjoining states. Ovule serial sectioning was done to determine testa ontogeny. In all cases seed surface structures could be related to excrescences developing from the integument cells. Six basic seed coat relief features could be distinguished, corresponding to the six sections in the genus. Minor variations in the basic types are constant and characteristic for species or, in very few instances, for species groups. A dendrogram was constructed from the seed testa data which supports the phylogeny established from chromosomal and morphological data. Practical application of seed coat microcharacter specificity to identification ofMentzelia seeds from prehistoric sites is suggested.
The phylogeny of Loasaceae subfam. Loasoideae is investigated with sequences of the chloroplast trnL(UAA) intron, all genera and infrageneric entities are included in the analysis. Loasaceae subfam. Loasoideae is monophyletic, and the two most speciose, andmonoph yletic, clades (which account for approximately 90% of the species total) are Nasa andthe so-called Southern Andean Loasas (Blumenbachia, Caiophora, Loasa s.str., Scyphanthus), but the phylogeny of the remainder is not completely resolved. The data underscore a basal position for Chichicaste, Huidobria, Kissenia, andKlapr othieae (Xylopodia, Klaprothia, Plakothira). High bootstrap support values confirm the monophyly both of Klaprothieae and Presliophytum (when expanded to include Loasa ser. Malesherbioideae). Aosa and Blumenbachia are not resolvedas monophyletic, but have clear morphological apomorphies. Within Nasa, ‘‘N. ser. Saccatae’’ is paraphyletic, and‘‘ N. ser. Carunculatae’’ is polyphyletic. However, the N. triphylla group in ‘‘N. ser. Saccatae’’ is a well-supportedmonop hyletic group, as is N. ser. Grandiflorae. ‘‘Loasa’’ in its traditional circumscription is paraphyletic, but Loasa s.str. (L. ser. Macrospermae, L. ser. Deserticolae, L. ser. Floribundae) is monophyletic. The remainder of ‘‘Loasa’’ (L. ser. Pinnatae, L. ser. Acaules, L. ser. Volubiles) is probably closely alliedto the essentially Patagonian-High Andean group comprising also Scyphanthus and Caiophora. These findings are congruent with morphology andphy togeography. Nasa seems to have undergone its primary radiation at moderate elevations (1500–2500 m) in the Andes of northern Peru (Amotape-Huancabamba Zone) and subsequently diversified into high elevations (above 4000 m) of the tropical Central Andes. South Andean Loasas appear to have undergone their primary diversification in the southern temperate and mediterranean regions of Chile and Argentina, with a subsequent northwards expansion of Caiophora into the high elevations of the tropical Andes. Hummingbird pollination has evolved independently from melittophily in High Andean clades of Nasa and Caiophora.
The presence of transfer cells (TCs) in the seeds of Boraginales (Boraginaceae s.s., Hydrophyllaceae, Heliotropiaceae, Ehretiaceae, Cordiaceae and Lennoaceae) has been reported but has not hitherto been studied systematically. This study, surveying the seed anatomy of 50 species of Boraginales, demonstrates that in Heliotropiaceae, Cordiaceae, Ehretiaceae and Lennoaceae, TCs are found in an uninterrupted course from the placenta via the funicle to the seed coat. These families are characterized by indehiscent fruits with a protective endocarp. The TCs may act as a sponge, thus promoting rapid germination when sufficient water is available. In Hydrophyllaceae, which have capsular fruits, TCs are often found but have a different structure and are restricted to the seed coat. Boraginaceae s.s. on the other hand are characterized by a complete absence of TCs in their seeds. The presence of TCs in seeds is considered as a synapomorphic trait and appears to be phylogenetically informative. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 140, 155−164.
Seed release mechanisms, seed production, seed weights, and fatty acid composition are investigated for Loasaceae subfam. Loasoideae. A total of ca. 60 species are studied. Nearly all species have erect, xerochasious capsules which restrict the release of the (balistochorous and/or anemochorous) seeds to dry, windy conditions. Very few taxa have tardily dehiscent capsules that are dispersed as a unit (anemochoria, epizoochoria, Blumenbachia sect. Blumenbachia, Klaprothia mentzelioides) or one-seeded cypselas (anemochoria, Kissenia). Dehiscent, but non-xerochasious capsules are only found in Blumenbachia, and Blumenbachia sylvestis (B. sect. Angulatae) has seeds with good floating ability and may be occasionally hydrochorous. The seeds of all other Loasoideae taxa tested rapidly sink when placed in water. Seed weights range from less than 0.02 g per 1000 seeds (Huidobria, Presliophytum) to over 23 g per 1000 seeds (Loasa ser. Macrospermae), thus spanning 3 orders of magnitude. Seed weight seems to correlate to some degree with substrate and seeds are heavier in species from “deep substrates” such as gravel and leaf litter. Seed numbers per fruit range from 1 (Kissenia) to over 3000 (Presliophytum) and annual seed production of individual mature plants ranges from less than 100 (Kissenia) to over 4 ×106 (Presliophytum). Fatty acid composition of Loasoideae seeds is mostly of common fatty acids, but exact composition varies considerably between groups. Most taxa have high levels of poly-unsaturated fatty acids (over 40% in Caiophora and Nasa), very low levels (<2%) are only found in Presliophytum. Nasa is the only genus of Loasaceae which has γ-linolenic acid and stearidonic acid in its seeds (at levels of 3.5–10% and 2–8.5% respectively). There is no obvious connection between the degree of unsaturation of the seed oils and habitat.
Phylogenetic analyses of Loasaceae that apply DNA sequence data from the plastid trnL-trnF region and matK gene in both maximum-parsimony and maximum-likelihood searches are presented. The results place subfamily Loasoideae as the sister of a subfamily Gronovioideae-Mentzelia clade. Schismocarpus is the sister of the Loasoideae-Gronovioideae-Mentzelia clade. The Schismocarpus-Loasoideae-Gronovioideae-Mentzelia clade is the sister of Eucnide. Several clades in Loasoideae receive strong support, providing insights on generic circumscription problems. Within Mentzelia, several major clades receive strong support, which clarifies relationships among previously circumscribed sections. Prior taxonomic and phylogenetic hypotheses are modeled using topology constraints in parsimony and likelihood analyses; tree lengths and likelihoods, respectively, are compared from constrained and unconstrained analyses to evaluate the relative support for various hypotheses. We use the Shimodaira-Hasegawa (SH) test to establish the significance of the differences between constrained and unconstrained topologies. The SH test rejects topologies based on hypotheses for (1) the placement of gronovioids as the sister of the rest of Loasaceae, (2) the monophyly of subfamily Mentzelioideae as well as Gronovioideae and Loasoideae, (3) the monophyly of Loasa sensu lato as circumscribed by Urban and Gilg, and (4) the monophyly of Mentzelia torreyi and Mentzelia sect. Bartonia.
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901), mature seed (TEM). (6) Endosperm and testa. (7) Detail of testa with anticlinal and periclinal walls. (8) L. gayana (LoVo, Weigend et al. 7057), transverse section of mature seed with 3
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L. nitida (LoMc, Weigend et al. 2000/901), mature seed (TEM). (6) Endosperm and testa. (7) Detail of testa with anticlinal and periclinal walls. (8) L. gayana (LoVo, Weigend et al. 7057), transverse section of mature seed with 3-layered testa (LM).
On Gripidea, a new genus in Loasaceae
  • Miers
Miers, J., 1866. On Gripidea, a new genus in Loasaceae. Trans. Linn. Soc. Bot. 25, 227-238 table 28.
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Loasa volubilis Domb. – LiPi – (sem) Gay, s.n. anno, 1839 (P);
Seed oil FA of Loasaceae – a new source of g-linolenic and stearidonic acid
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  • A Bru¨
  • L Weigend
References Aitzetmuï, A., Bru¨, L., Weigend, M., 2004. Seed oil FA of Loasaceae – a new source of g-linolenic and stearidonic acid. JAOCS 81, 259–263.
Nasa and the conquest of South America
  • M Weigend
Weigend, M., 1997. Nasa and the conquest of South America. Doktorarbeit. Eigenverlag, Mu¨nchen.
& Gilg – CaCh – (sem) Kraus, s.n. (M)
  • Caiophora
  • Urb
Caiophora andina Urb. & Gilg – CaCh – (sem) Kraus, s.n. (M);
Embryology of Angiosperms The Seeds of Dicotyledons Transfer cells in the seeds of Boraginales
  • Spring-Er
  • Berlin
  • E J H Corner
In: Johri, B.M. (Ed.), Embryology of Angiosperms. Spring-er, Berlin. Corner, E.J.H., 1976. The Seeds of Dicotyledons, vol. 2. Cambridge University Press, Cambridge, UK. Diane, N., Hilger, H.H., Gottschling, M., 2002. Transfer cells in the seeds of Boraginales. Bot. J. Linn. Soc. 140, 155–164.
6924), longitudinal section of young seed (LM)
  • L Lolo
  • Weigend
L. acanthifolia (LoLo, Weigend et al. 6924), longitudinal section of young seed (LM).
transverse section of mature seed, 3-layered testa, massively thickened anticlinal wall (SEM). (10–12) L. asterias (LoPi, Weigend 6991), young seeds. (10) Two young seeds (LM). (11) Detail of 3–4-layered testa with secondary sculpturing in young seed (LM). (12) Details of secondary sculpturing (SEM)
  • L Martinii
L. martinii (LoVo, Montero 7905), transverse section of mature seed, 3-layered testa, massively thickened anticlinal wall (SEM). (10–12) L. asterias (LoPi, Weigend 6991), young seeds. (10). Two young seeds (LM). (11) Detail of 3–4-layered testa with secondary sculpturing in young seed (LM). (12) Details of secondary sculpturing (SEM). (aw ¼ anticlinal wall, es ¼ endosperm, em ¼ embryo, em ¼ embryo removed, it ¼ inner testa layers, opw ¼ outer periclinal wall, te ¼ testa epidermis).
Los nombres antiguos en el ge´ Caiophora (Loasa´ subfam. Loasoı´ )
  • M Weigend
  • M Ackermann
  • M Una Article In Press
  • Weigend
Weigend, M., Ackermann, M., 2003. Los nombres antiguos en el ge´ Caiophora (Loasa´ subfam. Loasoı´ ) y una ARTICLE IN PRESS M. Weigend et al. / Flora 200 (2005) 569–591 590
USM); (sem, tem)); (sem)
  • Weigend
Caiophora cirsiifolia K.Presl – CaCi – (sem) Weigend & Foerther, 97/797 (M, USM); (sem, tem) Weigend et al. et al., 97/465 (M, USM); (sem) Weigend et al., 97/ 314 (M, USM); (sem) Weigend et al., 97/464 (M, USM); (lm) C. cirsiifolia – CaCi – Dostert, 98/156 (M, USM);
Ack-erm. – CaCo – (sem) Weigend & Dostert
  • ( G Caiophora
  • Don
Caiophora grandiflora (G.Don) Weigend & M.Ack-erm. – CaCo – (sem) Weigend & Dostert 97/31 (M, USM);
– BlBl – (sem) Schlind-wein, 495 (TUEB); (sem) Reineck & Czermack 9 (F)
  • Blumenbachia
  • Camb
Blumenbachia latifolia Camb. – BlBl – (sem) Schlind-wein, 495 (TUEB); (sem) Reineck & Czermack 9 (F);
Don – (sem) Grandjot, s.n
  • Scyphanthus
Scyphanthus elegans D.Don – (sem) Grandjot, s.n. March, 1933 (M);
– LoVo – (sem) Po¨ , s.n. (M)
  • Loasa
  • Poepp
Loasa micrantha Poepp. – LoVo – (sem) Po¨, s.n. (M);
& Gilg -CaLa -(sem) Kraus, s
  • Urb Caiophora Buraeavii
Caiophora buraeavii Urb. & Gilg -CaLa -(sem) Kraus, s.n. (M);
Bariloche, Argentina) for help in the field We would like to thank Mr
  • G Vobis
  • C Ezcurra
G. Vobis and C. Ezcurra (Bariloche, Argentina) for help in the field. We would like to thank Mr. H. Lu¨nserLu¨nser (Berlin) for arranging the photographic plates, Dr. E. Facher and Dr. R. Melzer (Munich), and Mrs. Ch.
– LoVo – (sem) Po¨ppigPo¨ppig, s.n
  • Poepp Loasa
Loasa micrantha Poepp. – LoVo – (sem) Po¨ppigPo¨ppig, s.n. (M);
LoDe -(sem) King, s.n. (SGO)
  • Phil Loasa Urmenetae
Loasa urmenetae Phil. -LoDe -(sem) King, s.n. (SGO);
Presl – CaCa – (sem)
  • K Caiophora Carduifolia
  • Weigend
Caiophora carduifolia K.Presl – CaCa – (sem) Weigend et al., 2000/326; (lm) Weigend et al., 5415 (BSB, HUT, M, USM);
  • Deutsche Forschungsgemeinschaft
  • Grant No
Deutsche Forschungsgemeinschaft (Grant No. WE 2330/1, 2001–2003), Lewis B. and Dorothy Cullman Laboratory for Molecular Systematics Studies at the New York Botanical Garden (1999–2000), botconsult GmbH (1999–present).
CaCh -Caiophora chuquitensis group, CaCi -Caiophora cirsiifolia group, CaLa -Caiophora lateritia group, CaCl -Caiophora clavata group, CaCa -Caiophora carduifolia group, CaNi -Caiophora nivalis group, CaCr -Caiophora coronata group
  • Caar -Caiophora Arechavaletae Pinnatae
  • Group
Pinnatae, CaAr -Caiophora arechavaletae group, CaCh -Caiophora chuquitensis group, CaCi -Caiophora cirsiifolia group, CaLa -Caiophora lateritia group, CaCl -Caiophora clavata group, CaCa -Caiophora carduifolia group, CaNi -Caiophora nivalis group, CaCr -Caiophora coronata group, CaCo -Caiophora contorta group).
BlGr -(sem) Schlindwein
  • Urb Blumenbachia Eichleri
Blumenbachia eichleri Urb. -BlGr -(sem) Schlindwein, 610 (TUEB);
BlBl -(lm, sem) Weigend, s.n. (KW_124
  • Urb Blumenbachia Hieronymii
Blumenbachia hieronymii Urb. -BlBl -(lm, sem) Weigend, s.n. (KW_124, Bot. Garden Mu¨nchen, M);
BlBl -(sem) Schlindwein, 495 (TUEB); (sem) Reineck & Czermack 9
  • Camb Blumenbachia Latifolia
Blumenbachia latifolia Camb. -BlBl -(sem) Schlindwein, 495 (TUEB); (sem) Reineck & Czermack 9 (F);
Don) Urb. & Gilg -CaPt -(sem)
  • Weigend
Caiophora cf. pterosperma (Ruiz & Pav. ex G.Don) Urb. & Gilg -CaPt -(sem) Weigend et al., 5118 (BSB, HUT, M, USM); (lm) Weigend et al., 5484 (BSB, HUT, M, USM);
97/465 (M, USM); (sem) Weigend et al., 97/ 314 (M, USM); (sem) Weigend et al., 97/464 (M, USM); (lm) C. cirsiifolia -CaCi -Dostert
  • Weigend
Caiophora cirsiifolia K.Presl -CaCi -(sem) Weigend & Foerther, 97/797 (M, USM); (sem, tem) Weigend et al. et al., 97/465 (M, USM); (sem) Weigend et al., 97/ 314 (M, USM); (sem) Weigend et al., 97/464 (M, USM); (lm) C. cirsiifolia -CaCi -Dostert, 98/156 (M, USM);
Caiophora contorta (Desr.) K.Presl -CaCo -(sem)
  • Weigend
Caiophora contorta (Desr.) K.Presl -CaCo -(sem) Weigend et al., 97/323 (M, USM);
Loasa asterias Dusen -LoPi -(sem, lm) Weigend
  • Bsb Brco
Loasa asterias Dusen -LoPi -(sem, lm) Weigend, 6991 (BRCO, BSB, M);
LoVo -(sem) Po¨ppig, s.n
  • Poepp Loasa Micrantha
Loasa micrantha Poepp. -LoVo -(sem) Po¨ppig, s.n. (M);
LiPi -(sem) Gay, s.n. anno
  • Domb Loasa Volubilis
Loasa volubilis Domb. -LiPi -(sem) Gay, s.n. anno, 1839 (P);