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a-f. SEMs of Campanulaceae (a, b) and Goodeniaceae (c-f) pollen. Scale bars for whole pollen grains 5 10 mm; for fractured grains scale bars 5 1 mm. a, b. Campanula rotundiflora; c-f. Goodenia ovata.
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... from SEM and TEM (Patel, 1976;Skvarla et al., 1977;Polevova, 2006) were used to determine pollen surface and ultrastructural characteristics of Goodeniaceae pollen grains. Most gen- era within the family have spinulate surfaces, with two genera (Dampiera and Anthotium) being characterized by the presence of striae and rugulae. TEM micrographs document a relatively thin endoexine and a thick footwall with large, distally bifurcating columellae (see Good- enia, Fig. 5f). This bifurcating pattern can be simple (e.g., Dampiera) or of a higher order (e.g., Brunonia). In Scaevola, the higher order columellae are fused. More recent LM/SEM/TEM studies in the Good- eniaceae ( Gustafsson et al., 1997) sup- ported the three groups based on pollen morphology originally established by Duigan (1961) but with an expanded list of taxa. Their Scaevola-Goodenia group (5Brunonia and Goodenia groups of Duigan, 1961), with tricolporate, microspinu- lose grains having distally branched colu- mellae was indicated by Gustaffson et al. (1997) as showing similarity to Asteraceae and Calyceraceae. Finally, Carolin's review (2007) of Goodeniaceae further supports the pollen data of Duigan (1961) and Gustafsson ...
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... Molecular phylogenetic studies have suggested an origin of Asteraceae at c. 69.5 Mya (Panero and Crozier 2016). A similar time perspective was provided by Denham et al. (2016) who obtained a value of 76.5 Mya for the divergence between Asteraceae and Calyceraceae, the latter being the sister family based on pollen morphology (DeVore et al. 2007), morphological studies (Pozner et al. 2012(Pozner et al. , 2021, and molecular phylogenetic analyses (Gustafsson et al. 1996, Jansen and Kim 1996, Lundberg and Bremer 2003, Mandel et al. 2017. More recent phylotranscriptomic investigations by Zhang et al. (2021), using 1087 nuclear genes from transcriptomes and genomes of 243 species, have placed the divergence of Calyceraceae and Asteraceae at c. 83 Mya, and Barnadesioideae at c. 81 Mya. ...
Based on molecular phylogenetic studies, Barnadesioideae have been proposed to be the basal subfamily of Asteraceae. This is a complex of
10 genera and 87 species distributed primarily along the Andean mountains, Patagonia, and into southern Brazil and Uruguay. Phylogenetic
analyses have recovered all genera as monophyletic groups and have provided insights to their inter-relationships. Four generic clades have
been substantiated: (1) Chuquiraga, Doniophyton, and Duseniella; (2) Dasyphyllum; (3) Barnadesia and Huarpea; and (4) Archidasyphyllum,
Arnaldoa, and Fulcaldea. The remaining genus, the monospecific Schlechtendalia, has been an outlier in the subfamily, with some previous
analyses recovering it as basal for the entire subfamily, and others showing it as sister to Barnadesia and Huarpea (with weak support) as
well as to other genera. Recent massive sampling of loci has confirmed Schlechtendalia as the sister genus for the subfamily. Schlechtendalia
luzulifolia has morphology atypical for Asteraceae. The capitula are loose aggregations of florets, and the leaves are long and strap shaped,
more reminiscent of monocots. Morphological and anatomical investigations of the leaves reveal long, laminar blades with parallelodromous
vascularization. The vesture is often with ‘barnadesioid trichomes’, especially towards the base of the plant, plus additional uniseriate trichomes consisting of 3 to many cells, newly reported for the subfamily. Some glandular trichomes with 2-4 short cells also occur. The transverse
anatomy of the leaves reveals a single epidermal layer on both surfaces, which also contain the stomata (the leaf being amphistomatic). The
mesophyll is undifferentiated; the vascular traces are surrounded by sclerenchyma that not only encircles the traces but also extends towards
the epidermis and connects with it. The morphology and anatomy of the leaves of Schlechtendalia are divergent in comparison with other
genera of the subfamily. Chuquiraga, Doniophyton, and Huarpea have leaf adaptations for survival in xeric habitats, such as dense pubescence,
grey surfaces, and revolute margins. Schlechtendalia, in contrast, is adapted to a more mesic environment, especially near the Atlantic Ocean
and along the Uruguay and La Plata rivers. The leaves are oriented upright, which correlates with undifferentiated mesophyll and stomata
on both epidermal layers. The stem is an underground rhizome, an adaptation that permits survival during seasonal drought in the austral
summer in Uruguay and adjacent regions. It is hypothesized that Schlechtendalia may have become adapted to more mesic environments in
the Miocene prior to the rise of the Andes and development of the modern arid environments, into which many of the other genera of the
subfamily subsequently radiated.
... Pollen of Calyceraceae has similarities in polarity, symmetry and microspinulate spines, but different in the shape of pollen. Asteraceae has spheroidal-shaped and macro-echinate pollen, while Calyceraceae has triangular-type and micro-echinate pollen (De Vore et al. 2007). Goodeniaceae also have highly similar pollen characteristic to the Asteraceae with a slight difference in shape ratio (suboblate 6:8-7:8; spheroidal 7:8-8:7; prolate 8:6-8:4). ...
Salamah A, Luthfikasari R, Dwiranti A. 2019. Pollen morphology of eight tribes of Asteraceae from Universitas Indonesia Campus, Depok, Indonesia. Biodiversitas 20: 152-159. Asteraceae is one of the largest flowering plant families in the world consisting of a high diversity of species that possess many macro-morphological characters. Ecological pressures have resulted in this highly varied morphology. Our research aims to describe the pollen morphology of the Asteraceae for species occurring in Depok Campus of Universitas Indonesia. Pollens of 14 species from eight tribes were extracted using acetolysis method and observed under a light microscope. The result showed that pollen unity, shape and pollen surface appeared to be differentiating characters of the Asteraceae from other families, while polarity and symmetry characters can be distinguishing characters at higher levels of the family such as subdivisions. Aperture characters were useful to differentiate between tribes. The shape of pores and size of pollen were not useful in differentiating between tribes, but may differentiate between lower taxa. The results of this study could be used as supporting data for regrouping taxa within the Asteraceae using morphological features. © 2019, Society for Indonesian Biodiversity. All rights reserved.
... Reitz (1988) was one of the first authors to suggest synonymy of Boopis, Gamocarpha and Nastanthus due to poor morphological differentiation. The palynological results published by Hansen (1992) showed at least two main pollen morphologies in the family, which are not uniform within genera (DeVore & al., 2007). According to Carlquist & DeVore (1998), the diversity of wood anatomy in Calyceraceae (14 species of five genera sampled) suggests adaptation to particular ecological conditions, with little phylogenetic information. ...
Calyceraceae is a small family with six traditionally recognized genera and 47 species from southern South America. Most species grow along the Andes (of both Argentina and Chile) and in arid regions of the Patagonian steppe. This family belongs to the well-supported MGCA clade within Asterales, which includes Menyanthaceae + Goodeniaceae + Calyceraceae + Asteraceae. Calyceraceae is monophyletic and sister to Asteraceae, one of the five largest families of angiosperms. Although Calyceraceae is clearly distinct as a family, its genera are not, and taxonomic revisionary effort has confirmed the lack of sharp boundaries among genera. We performed a phylogenetic analysis of Calyceraceae with a broad taxon sampling (41 of 47 species), and with sequence data from multiple regions from the nuclear (ITS) and plastid genomes (ycg6-psbM, psbM-trnD, trnS-trnG, trnH-psbA, trnD-trnT) using maximum parsimony and Bayesian approaches. We aimed at identifying monophylectic groups, their putative morphological synapomorphies and their geographical distribution; we also estimated divergence times and examined chromosomes numbers in an evolutionary context. We obtained well-resolved and strongly supported phylogenies that show Calyceraceae to be divided into two major clades with geographically structured subclades within each. Our results indicate that an early split within Calyceraceae occurred about 27.4 Ma, probably related to differential changes in chromosome numbers, which allowed the two lineages to evolve in sympatry. We found that major natural subgroups diverged 15–12 Ma, following the Early-Miocene South Andes construction stage. Finally, the diversification of the extant species is probably associated to Andean orogeny and climate changes in the last 5–4 Myr. We recovered Acicarpha as monophyletic, while the remaining traditionally recognized genera of Calyceraceae are para- or polyphyletic. Most species of Moschopis are included in the Glutinose group, but M. monocephala is more closely related to some Calycera species. Calycera is divided into two clades: the Calycera group and the Pilose group. All species of Nastanthus are placed in a well-supported main group with species of Gamocarpha and Boopis. Gamocarpha could be monophyletic after exclusion of G. dentata and G. angustifolia, but is nested within Nastanthus and Boopis species. Boopis is clearly polyphyletic with its species distributed in all main groups.
... When considering Tubulifloridites lilliei, Zavada and de Villiers (2000) and Zavada and Lowrey (2010) find these tricolporate, psilate to microechinate pollen grains with lalongate endoapertures could represent the ancestral pollen type of Asteraceae but are difficult to separate from those of other families. This combination is found in Calyceraceae where it is hypothesized to be ancestral (Gamocarpha type, DeVore et al., 2007), and Goodeniaceae (Gustafsson et al., 1997) as well as in Asteraceae in Chuquiraga, Doniophyton and some species of Dasyphyllum of the Barnadesioideae. T. lilliei type A pollen grains are ascribed to extant Dasyphyllum based on the optimal tree resulting from maximum parsimony analysis of 26 pollen morphological characters, but trees one step longer place type A pollen in Barnadesioideae, Calyceraceae, Goodeniaceae or the Asteraceae-Calyceraceae stem, and two-step longer trees place type A grains in Goodeniaceae, Menyanthaceae, Alseuosmiaceae, Campanulaceae or stems leading to more inclusive clades . ...
... lilliei type A relatedness also (Panero, 2016). Type A fossils' exine width ranges from 1.2 to 2 lm , smaller than recorded Barnadesioideae (Urtubey and Tellería, 1988;Tellería et al., 2015) but typical of Calyceraceae (DeVore et al., 2007), whereas Dasyphyllum has exine layers that are mostly around 3 lm (Urtubey and Tellería, 1988) with D. velutinum as small as around 2.5 lm . The fossil would not share a synapomorphy with Barnadesioideae had the cut-off between larger categories been 2 lm. ...
Spatial and temporal differences in ecological opportunity can result in disparity of net species diversification rates and consequently uneven distribution of taxon richness across the tree of life. The largest eudicotyledonous plant family Asteraceae has a global distribution and at least 460 times more species than its South American endemic sister family Calyceraceae. In this study, diversification rate dynamics across Asteraceae are examined in light of the several hypothesized causes for the family’s evolutionary success that could be responsible for rate change. The innovations of racemose capitulum and pappus, and a whole genome duplication event occurred near the origin of the family, yet we found the basal lineages of Asteraceae that evolved in South America share background diversification rates with Calyceraceae and their Australasian sister Goodeniaceae. Instead we found diversification rates increased gradually from the origin of Asteraceae approximately 69.5 Ma in the late Cretaceous through the Early Eocene Climatic Optimum at least. In contrast to earlier studies, significant rate shifts were not strongly correlated with intercontinental dispersals or polyploidization. The difference is due primarily to sampling more backbone nodes, as well as calibrations placed internally in Asteraceae that resulted in earlier divergence times than those found in most previous relaxed clock studies. Two clades identified as having transformed rate processes are the Vernonioid Clade and a clade within the Heliantheae alliance characterized by phytomelanic fruit (PF Clade) that represents an American radiation. In Africa, subfamilies Carduoideae, Pertyoideae, Gymnarrhenoideae, Cichorioideae, Corymbioideae, and Asteroideae diverged in a relatively short span of only 6.5 million years during the Middle Eocene.
... Several angiosperm families produce triaperturate pollen grains with microechinate exine sculpture. Here we used the apomorphy-based method [in the sense of Sauquet et al. (10)] as a first attempt to estimate the closest living relatives of the fossil T. lilliei type A, conducting exhaustive comparisons with members of Lamiaceae, Ranunculaceae, Cleomaceae, Sol-anaceae, Hectorellaceae, Rhamnaceae, Euphorbiaceae, Rubiaceae, Caprifoliaceae, and families of Asterales (Stylidiaceae, Campanulaceae, Goodeniaceae, Calyceraceae and Asteraceae) by using information available in the literature (36,(50)(51)(52)(53)(54)(55)(56)(57)(58)(59)(60). Despite gross similarities, however, most of these families have significant differences in apertures, structure, or sculpture with T. lilliei type A: Lamiaceae (Clerodendrum type) are tricolpate, have short colpi, uniformly microechinate sculpture, and perforate tectum (53); Ranunculaceae (Clematis) are larger (approximately 40 μm), oblate to spheroidal, tricolpate, have a clearly columellate exine structure and variable sculpture (microechinatemicrogranulate), with minutely perforate tectum (52,54,55); Cleomaceae (Cleome) are tricolporoidate, with thin exine (< 1 μm) and conspicuous columellae (50); Solanaceae (Latua) have indistinct ora and very thin exine (< 1 μm) (50); Hectorellaceae (Hectorella) are tricolpate, larger (approximately 40 μm), clearly columellate and with perforate tectum (54); Rhamnaceae (Pomaderris) have circular ora and sparingly perforate tectum (54); Euphorbiaceae (Neoscortechinia) differ in having shorter colpi, clearly columellate exine, and stout microspines with acute ends (57); Rubiacae, (Bikkia) have shorter colpi with poorly defined margins (56); Caprifoliaceae (Symphoricarpos, Plectritis) have tricolporate apertures, but with short colpi, clearly columellate exine, and conical microspines with acute ends (51). ...
Significance
The flowering plant family Asteraceae (e.g. sunflowers, daisies, chrysanthemums), with about 23,000 species, is found almost everywhere in the world except in Antarctica. Asteraceae (or Compositae) are regarded as one of the most influential families in the diversification and evolution of a large number of animals that heavily depends on their inflorescences to survive (e.g. bees, hummingbirds, wasps). Here we report the discovery of pollen grains unambiguously assigned to Asteraceae that remained buried in Antarctic deposits for more than 65 million years along with other extinct groups (e.g. Dinosaurs, Ammonites). Our discovery drastically pushes back the assumed origin of Asteraceae, because these pollen grains are the oldest fossils ever found for the family.
... Pollen samples were taken from specimens deposited in the following herbaria: HAO, HUT, LP, MCNS, TRU (Holmgren et al., http://sciweb.nybg.org/science2/IndexHerbariorum.asp). Some exine data were taken from Skvarla et al. (1977), Gustafsson et al. (1997), Zao et al. (2000), Tellería et al. (2003), De Vore et al. (2007) (Table 1). ...
Barnadesioideae (94 species) is the sister subfamily to the rest of the Asteraceae (23,000 species). Pollen grains in this subfamily are structurally and sculpturally distinctive and diverse. Although pollen morphology has contributed to the taxonomy of the subfamily, there is a gap of knowledge concerning the evolution of the exine structure. This study aims at exploring the systematic and phylogenetic significance of optimizing selected pollen characters of Barnadesioideae on the latest molecular phylogenetic tree. Transmission electron microscope (TEM) observations on pollen of selected species, some of them never explored so far, show that the exine probably evolved from a thin pattern (ca. 1-3 μm), with a well-developed foot layer and solid and free columellae, present in sister family Calyceraceae, towards a thicker (> 6-11 μm) and a more complex columellate-granulate bilayered exine in Barnadesioideae (with very delicate columellae). The particular exine structure observed in the monotypic Schlechtendalia luzulaefolia, which combines compact and independent columellae (common in more derived Asteraceae) with a granular internal tectum as the inner ectexine layer (as in Barnadesioideae), reinforces its distant phylogenetic position within Barnadesoideae. More derived lineages within Asteraceae (e.g. Mutisioideae) retained some ancestral exine features although evolved an even thicker exine and a columellate trilayered exine (with robust columellae), rare in the Angiosperm pollen grains.
... Their shape in polar view is reminiscent of species of Acarpha Griseb. (Calyceraceae) (De Vore et al. 2007) or species of Lopezia Cav. (Onagraceae) (third author). ...
Pollen grains of Rhigiophyllum squarrosum Hochst., Siphocodon spartioides Turcz. and S. debilis Schltr., are flattened and triangular with pores at the angles. This morphology is radically different from known pollen of the Campanulaceae s.sfr:: the Campanulaceae are treated here as a family separate from the Lobeliaceae, Cyphiaceae, Nemacladaceae, Pentaphragmataceae and Sphenocleaceae (Lammers 1992). As traditionally conceived, the Campanulaceae is very heterogeneous and, in many classifications, these families were treated as subfamilies of a much-enlarged Campanulaceae. The consistently different floral morphology, biochemistry and pollen structure of the Lobeliaceae favours the recognition of this predominantly tropical group as a separate family.The pollen grains of these species are described in comparison with other members of the Campanulaceae. Based on surface characteristics of their pollen grains, we conclude that they represent an early offshoot o f the wahlenbergioid lineage in southern Africa. We suggest that this unique pollen may also be the result of a highly selective regime in the fynbos, associated with specialized pollinators, and base-poor soils, in addition to possible adaptations for ant dispersal and fire. Rhigiophyllum Hochst. and Siphocodon Turcz. are also unique in having free carpel-like structures within the ovary. These shrink to form seed pockets around the seeds and disperse as units when the capsule matures. Data from molecular studies support the contention that these taxa form a sister group to all other wahlenbergioids and that this should be formally recognized in a classification system. We treat Rhigiophyllum and Siphocodon within the Campanulaceae: Wahlenbergioideae, as a separate tribe, the Rhigiophylleae tribus nov., the species of which are distinguishable from other wahlenbergioids by unique angulaperturatc pollen, epipetalous stamens, free carpel-like structures and seed pockets.
... Their shape in polar view is reminiscent of species of Acarpha Griseb. (Calyceraceae) (De Vore et al. 2007) or species of Lopezia Cav. (Onagraceae) (third author). ...
Pollen grains of Rhigiophyllum squarrosum Hochst., Siphocodon spartioides Turcz. and S. debilis Schltr., are flattened and triangular with pores at the angles. The morphology is radically different from known pollen of the Campanulaceae s.str. The pollen grains of these species are described in comparison with other members of the Campanulaceae. Based on surface characteristics of their pollen grains, we conclude that they represent an early offshoot of the wahlenbergioid lineage in southern Africa. We suggest that this unique pollen may also be the result of a highly selective regime in the fynbos, associated with specialized pollinators, and base-poor soils, in addition to possible adaptations for ant dispersal and fire. Rhigiophyllum Hochst. and Siphocodon Turcz. are also unique in having free carpel-like structures within the ovary. These shrink to form seed pockets around the seeds and disperse as units when the capsule matures. Data from molecular studies support the contention that these taxa form a sister group to all other wahlenbergioids and that this should be formally recognised in a classification system. We treat Rhigiophyllum and Siphocodon within the Campanulaceae: Wahlenbergioideae as a separate tribe, the Rhigiophylleae tribus nov., the species of which are distinguishable from other wahlenbergioids by unique angulaperturate pollen, epipetalous stamens, free carpel-like structures and seed pockets.
... Specimen location of the slides is according to the England finder coordinates. Fossil pollen grains have been compared with extant specimens illustrated by Gustafsson et al. (1997) and DeVore et al. (2007). In some cases, recent pollen grains were re-analyzed to observe specific features of the nearest living forms. ...
... The pollen morphology of Calyceraceae was examined by Skvarla et al. (1977), Avetisjan (1980), Hansen (1992) and DeVore et al. (2007). The pollen grains are ecaveate, tectate, microperforate and microechinate. ...
... These two groups are mainly characterized by having (Calycera type) or lacking (Gamocarpha type) intercolpar concavities (depressions between colpi). The Calycera type is further distinguished by having angulaperturate apertures and colpar ledges (ridges of exine that line the inner margin of the colpi) according to DeVore et al. (2007). The exine has short, thin and unbranched columellae, similar to those found in some Barnadesioideae genera such as Dasyphyllum. ...
A new fossil pollen species (Psilatricolporites protrudens sp. nov) is described from Miocene sedimentary sections of the Chenque and Puerto Madryn formations (Chubut province, Argentina). The fossil pollen grains are characterized by being small, tricolporate, subspheroidal to suboblate in shape; rhombic outline in equatorial view and subtriangular in polar view. The exine is tectate and columellate; the nexine is thickened toward endoapertures resulting in a typical wall protrusion on the external surface. These morphological features point to a possible relationship with Gamocarpha type of the Calyceraceae. Most species of this type grow in high-altitude arid habitats or in coastal locations under extreme climatic condition. The gradual spread of the stress-adapted Calyceraceae as well as other phylogenetically related taxa (e.g. Barnadesioideae, Mutisioideae) during the Miocene in southern South America may have been triggered by the increasing aridity and seasonality caused by Andean uplift. This fossil record represents the first finding of Calyceraceae, the most closely related family of Asteraceae, and provides evidence for the timing of their geographic radiation.
Based on molecular phylogenetic studies, Barnadesioideae have been proposed to be the basal subfamily of Asteraceae. This is a complex of
10 genera and 87 species distributed primarily along the Andean mountains, Patagonia, and into southern Brazil and Uruguay. Phylogenetic
analyses have recovered all genera as monophyletic groups and have provided insights to their inter-relationships. Four generic clades have
been substantiated: (1) Chuquiraga, Doniophyton, and Duseniella; (2) Dasyphyllum; (3) Barnadesia and Huarpea; and (4) Archidasyphyllum,
Arnaldoa, and Fulcaldea. The remaining genus, the monospecific Schlechtendalia, has been an outlier in the subfamily, with some previous
analyses recovering it as basal for the entire subfamily, and others showing it as sister to Barnadesia and Huarpea (with weak support) as
well as to other genera. Recent massive sampling of loci has confirmed Schlechtendalia as the sister genus for the subfamily. Schlechtendalia
luzulifolia has morphology atypical for Asteraceae. The capitula are loose aggregations of florets, and the leaves are long and strap shaped,
more reminiscent of monocots. Morphological and anatomical investigations of the leaves reveal long, laminar blades with parallelodromous
vascularization. The vesture is often with ‘barnadesioid trichomes’, especially towards the base of the plant, plus additional uniseriate trichomes consisting of 3 to many cells, newly reported for the subfamily. Some glandular trichomes with 2-4 short cells also occur. The transverse
anatomy of the leaves reveals a single epidermal layer on both surfaces, which also contain the stomata (the leaf being amphistomatic). The
mesophyll is undifferentiated; the vascular traces are surrounded by sclerenchyma that not only encircles the traces but also extends towards
the epidermis and connects with it. The morphology and anatomy of the leaves of Schlechtendalia are divergent in comparison with other
genera of the subfamily. Chuquiraga, Doniophyton, and Huarpea have leaf adaptations for survival in xeric habitats, such as dense pubescence,
grey surfaces, and revolute margins. Schlechtendalia, in contrast, is adapted to a more mesic environment, especially near the Atlantic Ocean
and along the Uruguay and La Plata rivers. The leaves are oriented upright, which correlates with undifferentiated mesophyll and stomata
on both epidermal layers. The stem is an underground rhizome, an adaptation that permits survival during seasonal drought in the austral
summer in Uruguay and adjacent regions. It is hypothesized that Schlechtendalia may have become adapted to more mesic environments in
the Miocene prior to the rise of the Andes and development of the modern arid environments, into which many of the other genera of the
subfamily subsequently radiated.
