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Trichomes. A-G. Leaf hairs.-A. Oblique septate flagellate hair: Gochnatia tortuensis (Ekman H-3553, S). B, C. 2-armed hair.-B. Gochnatia polymorpha (Pedersen 8587, LP).-C. Cyclolepis genistoides (Correa 3172 & Nicora, LP).-D. 3-to 5-armed hair: Gochnatia barrosii (Hatschbach 16945, LP).-E. Multistoried T-shaped hair: Ianthopappus corymbosus (Palacios & Cuezzo 2304, LP). F, G. Biseriate glandular hairs.-F. Gochnatia discoidea (Blanchet 3345, LP).-G. Gochnatia glutinosa (Fabris 1343, LP). H-L. Achenial hairs. H-K. Duplex hairs: Wunderlichia azulensis (Harley et al. 25209, MO). H-I. Duplex hairs with one of the hair cells shorter.-J. Septate duplex hair.-K. Duplex hair with only one hair cell.-L. Capitate glandular biseriate hair: Gochnatia cowellii (Britton & Cowell 10183, NY).
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Gochnatia is one of the largest genera of the tribe Mutisieae (Asteraceae) and has been traditionally characterized by its homogamous capitula with isomorphic corollas. A morphological study of Gochnatia and associated genera, i.e., Actinoseris, Chucoa, Cnicothamnus, Cyclolepis, Hyalis, Ianthopappus, Nouelia, Pleiotaxis, and Wunderlichia, was carri...
Contexts in source publication
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... Oblique septate flagellate hairs: one or two foot cells, one-or more-celled stalks or stipes, and unicellular, very long, flagellate, tubular heads (Fig. 7A). This trichome type is present in most species of Gochnatia, Chucoa, Cnicothamnus, Pleiotaxis, and ...
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... Two-armed hairs (or T-shaped, malpighia- ceous, anvil, dolabriform hairs): one or two foot cells, uniseriate, generally two-to more-celled stalks, and unicellular heads. The apical cell, which constitutes the head, initially assumes the shape of a hammer and later becomes T-shaped by further outgrowth of the two ends (Fig. 7B, C). This trichome type is present in Gochnatia sect. Moqui- niastrum (Fig. 7B) and G. cordata, Cyclolepis ( 7C), and Hyalis argentea. In Nouelia one end of the apical cell is very ...
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... dolabriform hairs): one or two foot cells, uniseriate, generally two-to more-celled stalks, and unicellular heads. The apical cell, which constitutes the head, initially assumes the shape of a hammer and later becomes T-shaped by further outgrowth of the two ends (Fig. 7B, C). This trichome type is present in Gochnatia sect. Moqui- niastrum (Fig. 7B) and G. cordata, Cyclolepis ( 7C), and Hyalis argentea. In Nouelia one end of the apical cell is very ...
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... Three-to 5-armed hairs (or 3-to 5-branched, stellate hairs sensu Cabrera, 1971): similar to the 2-armed hairs, but the apical cell has 3 to 5 branches (Fig. 7D). The apical cell does not divide and thus the head remains one-celled. This type is found only in Gochnatia barrosii and G. rusbyana of section ...
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... Multistoried T-shaped hairs: similar to the 2-armed hairs, but further periclinal divisions take place in the apical cell. The head is thus comprised of 3 or 4 one-celled layers, all oriented transver- sally and parallel, but of different lengths (Fig. 7E). This type of trichome, not very common in Aster- aceae, has been reported in the tribe Senecioneae (Robinson, 1989). Ianthopappus is the only genus with this type of ...
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... Biseriate glandular hairs: comprised of 2 rows of cells in the body, with two to many cells in each row, enclosed by a persistent or collapsed cu- ticular vesicle (Fig. 7F, G). Glandular hairs are widespread in all the taxa studied, and especially in species of Gochnatia sect. Pentaphorus (i.e., Gochnatia foliolosa, G. glutinosa) where they cover almost the entire surface of the leaf, with the fla- gellate hairs restricted to the ...
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... Duplex or twin hairs: Twin hairs are the common type in Asteraceae, comprised of two tri- angular or rectangular basal cells, one sometimes reduced, and two cylindrical or elliptical hair cells, equal or subequal in length, generally in contact up to their tips (Hess, 1938;Freire & Katinas, 1995) (Fig. 7H, I). All the taxa studied, in- cluding most species of Gochnatia, have achenes with twin hairs, usually very long and filiform. In some cases twin hairs have one hair cell very short (e.g., Gochnatia purpusii, G. recurva, G. tortuensis, Cyclolepis, Wunderlichia azulensis, W. mirabilis) (Fig. 7I), they are septate (e.g., Gochnatia hatsch- ...
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... up to their tips (Hess, 1938;Freire & Katinas, 1995) (Fig. 7H, I). All the taxa studied, in- cluding most species of Gochnatia, have achenes with twin hairs, usually very long and filiform. In some cases twin hairs have one hair cell very short (e.g., Gochnatia purpusii, G. recurva, G. tortuensis, Cyclolepis, Wunderlichia azulensis, W. mirabilis) (Fig. 7I), they are septate (e.g., Gochnatia hatsch- bachii, G. oligocephala, Hyalis, Pleiotaxis eximia, Wunderlichia azulensis, W. mirabilis) (Fig. 7J), or have only one hair cell (Wunderlichia azulensis) (Fig. ...
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... twin hairs, usually very long and filiform. In some cases twin hairs have one hair cell very short (e.g., Gochnatia purpusii, G. recurva, G. tortuensis, Cyclolepis, Wunderlichia azulensis, W. mirabilis) (Fig. 7I), they are septate (e.g., Gochnatia hatsch- bachii, G. oligocephala, Hyalis, Pleiotaxis eximia, Wunderlichia azulensis, W. mirabilis) (Fig. 7J), or have only one hair cell (Wunderlichia azulensis) (Fig. ...
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... hairs have one hair cell very short (e.g., Gochnatia purpusii, G. recurva, G. tortuensis, Cyclolepis, Wunderlichia azulensis, W. mirabilis) (Fig. 7I), they are septate (e.g., Gochnatia hatsch- bachii, G. oligocephala, Hyalis, Pleiotaxis eximia, Wunderlichia azulensis, W. mirabilis) (Fig. 7J), or have only one hair cell (Wunderlichia azulensis) (Fig. ...
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... Achenial biseriate glandular hairs: These are similar to those on the leaves (Fig. 7F, G) and are very widespread in the group under study, occur- ring with the other types. A modification of the typ- ical biseriate glandular hair with a very enlarged head, i.e., capitate glandular hair (Metcalfe & Chalk, 1950) (Fig. 7L), was found in the Caribbean species of Gochnatia sect. ...
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... Achenial biseriate glandular hairs: These are similar to those on the leaves (Fig. 7F, G) and are very widespread in the group under study, occur- ring with the other types. A modification of the typ- ical biseriate glandular hair with a very enlarged head, i.e., capitate glandular hair (Metcalfe & Chalk, 1950) (Fig. 7L), was found in the Caribbean species of Gochnatia sect. ...
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... multistoried T-shaped hair (Fig. 7E), the 2-, and 3-to 5-armed hairs (Fig. 7B-D), the capitate glandular achenial hair (Fig. 7L), and the pappus types established here (Fig. 8) are revealed as new diagnostic characters. In fact: (1) the multistoried T-shaped hair is exclusive to Ianthopappus and be- comes another character to distinguish it; (2) the 2-armed hairs are ...
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... multistoried T-shaped hair (Fig. 7E), the 2-, and 3-to 5-armed hairs (Fig. 7B-D), the capitate glandular achenial hair (Fig. 7L), and the pappus types established here (Fig. 8) are revealed as new diagnostic characters. In fact: (1) the multistoried T-shaped hair is exclusive to Ianthopappus and be- comes another character to distinguish it; (2) the 2-armed hairs are present in Gochnatia cordata, G. sect. ...
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... multistoried T-shaped hair (Fig. 7E), the 2-, and 3-to 5-armed hairs (Fig. 7B-D), the capitate glandular achenial hair (Fig. 7L), and the pappus types established here (Fig. 8) are revealed as new diagnostic characters. In fact: (1) the multistoried T-shaped hair is exclusive to Ianthopappus and be- comes another character to distinguish it; (2) the 2-armed hairs are present in Gochnatia cordata, G. sect. Moquiniastrum, Cyclolepis, Hyalis, and Noue- lia; (3) the ...
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... cordata, placed by Cabrera in section Hedraiophyllum, has characters that link it to sec- tion Moquiniastrum such as the 2-armed foliar hairs (Fig. 7B), numerous capitula arranged in glomeru- lose pseudopanicles (Fig. 2H), and pappus type C (Fig. 8C) Cabrera (1971) established two groups in his key to section Gochnatia, the ''South American species'' and the ''Caribbean species.'' Species of this sec- tion have some characters in common, such as sol- itary or 2 to 3 capitula and ...
Citations
... Indeed, genera like Moquiniastrum (Cabrera) G. Sancho and Cnicothamnus Griseb. are distinguished from other closely related genera by this type of trichome (Sancho and Otegui 2000;Freire et al. 2002;Funk et al. 2014;Gostel et al. 2022). In Vernonieae, these trichomes are informative in various degrees by supporting DNA evidence and distinction at different taxonomic levels (Faust and Jones 1973;Keeley et al. 2007;Hayat et al. 2009;Robinson et al. 2009;Redonda-Martínez et al. 2012). ...
Premise of the Research. While carrying out a taxonomic revision of Kaunia, uncommon types of trichomes and arrangement of trichomes on the leaves of K. gynoxymorpha and K. pachanoi were observed. Moreover, the trichome arrangement is not usual for either Kaunia or most of Eupatorieae. In this study, we aim to describe the leaf blade trichomes and indumentum in these two species and discuss, together with other anatomical characteristics, their ecological role, and potential adaptation to arid environments.
Methodology. We used traditional light microscopy, scanning electron microscopy (SEM), and fluorescence microscopy to examine leaf blade anatomical characteristics. Besides, we carried out histochemical techniques on the leaves of both studied species to determine the presence of cutinized cell walls which may influence apoplastic water flow in trichomes.
Pivotal Results. Our results show two-armed trichomes in Kaunia gynoxymorpha that are unusual in the tribe Eupatorieae. The arrangement of glandular and non-glandular leaf trichomes of the two studied species in tufts of 3 to 10 trichomes, sunken in surface crypts, is also uncommon.
Conclusions. These tufts of trichomes and other anatomical and histochemical evidence suggest that non-glandular trichomes and their arrangement in tufts in Kaunia gynoxymorpha and K. pachanoi provide adaptive advantages by combining physical, mechanical, and biochemical functions, thus protecting plants against drought and high light intensity among other adverse conditions.
... Although support has been weak, it is notable that these recent, deeply sampled phylogenetic studies have demonstrated a close relationship between Wunderlicheae and Gochnatieae. In agreement to previous authors (Cabrera, 1977;Freire et al., 2002), this study indicates Cyclolepis does belong within the Gochnatieae and we recognize the tribe as such, comprising ten genera (with Anastraphia, Cnicothamnus, Cyclolepis, Gochnatia, Moquiniastrum, Nahuatlea, Pentaphorus, Richterago, Tehuasca, and Vickia). ...
... 3). In addition to molecular studies, key morphological characters (Sancho, 2000;Freire et al., 2002;Roque and Funk, 2013;Roque and Sancho, 2020) support the identity of Moquiniastrum and Pentaphorus as independent genera from Gochnatia as currently circumscribed.We believe that taxonomic decisions should be integrative and involve as much evidence as possible to build a robust classification. In light of all available evidence, we recognize Moquiniastrum and Pentaphorus as evolutionary lineages independent from Gochnatia as proposed by Sancho et al. (2013) and Funk et al. (2014). ...
... Some morphological characters in Moquiniastrum show transitional states toward floral and sexual differentiation. Considering the taxonomic studies carried out in Moquiniastrum (Cabrera, 1971;Sancho, 2000;Freire et al., 2002;Sancho et al., 2013;Roque et al., 2019;Freitas et al., 2020) and according to the classification by Radford et al. (1974), we can describe four distinct sexual systems in the genus (Table 1): gynodioecious (10 spp.), in which the functionally female and hermaphroditic flowers occur on different individuals; gynomonoecious (3 spp.), individuals with functionally female marginal and hermaphroditic central flowers arranged in the same head; polygamodioecious (7 spp.), some individuals with marginal functionally female and central hermaphroditic flowers are arranged in the same head (heterogamous heads); some individuals with functionally female flowers (homogamous heads); and some individuals with hermaphroditic flowers (homogamous heads); hermaphroditic (3 spp.), individuals with hermaphroditic flowers only (homogamous heads) (Fig. 7A, Table 1). ...
Understanding the evolution of the tribe Gochnatieae (Compositae) has been the subject of considerable effort in the past decade. This is due to the key position of this tribe in the phylogeny of the sunflower family and the corresponding implications for biogeographic and morphological evolution of Compositae. Previous studies have confirmed the monophyly of this tribe as well as most of the genera that belong to it. However, phylogenetic resolution of Gochnatieae at both the genus- and species-level has remained poor. A subset of new phylogenomic loci used in this study has proven effective and has improved phylogenetic resolution in this group. The results of this work demonstrate Gochnatieae is a well-supported clade comprised of nine genera (Anastraphia, Cnicothamnus, Cyclolepis, Gochnatia, Moquiniastrum, Nahuatlea, Pentaphorus, Richterago, Tehuasca). One recently described genus, Vickia, was not included in this study; but its placement in Gochnatieae as a tenth genus in the tribe is well-justified. The monospecific Cyclolepis, which had been circumscribed within the tribe since its inception but was subsequently removed and designated as incertae sedis since 2014, is also shown to belong to Gochnatieae. We confirmed the monophyletic Moquiniastrum with two well-supported subclades. Ancestral area reconstruction analyses show that Gochnatieae originated in Eastern South America about 53 my. Apparently, except for Cyclolepis and Richterago, the ancestors of the other genera of Gochnatieae originated about 44 my from an area that now corresponds to the central Andes. The presence of the genera in the Chaco phytogeographic province, central Chile, and Mexico-United States-Caribbean is a result of dispersal from the central Andes. The ancestral distribution of Moquiniastrum corresponds to a large area comprising Eastern South America and the current central Andes, about 32 my. Ancestral character state reconstruction that included four characters indicates several states associated with complex plant reproductive biology such as gynodioecy, gynomonoecy, and polygamodioecy are derived in Gochnatieae as are heterogamous capitula (in Moquiniastrum and Richterago), dimorphic and subdimorphic corollas (in Cnicothamnus, Moquiniastrum, and Richterago), and the presence of marginal female corollas (in Moquiniastrum and Richterago). Within Moquiniastrum, two subclades (Densicephalum and Polymorphum) exhibit divergent patterns of trait evolution associated with these reproductive characters which suggests this genus can serve as a model to understand the sexual system evolution in plants.
... The leaves were classified as microphylls 244 20.55 ± 2.55 mm length by 2.55 ± 0.75 mm latitude; with acute attenuated and slightly sheeted base,245 entire margins, and acute apex (Fig. 2A). The primary venation was characterized as paralelodromous 246 three-veined, instead of actinodromous as described byFreire et al. (2002) andRoque and Sancho 247 (2020). The three primary veins run in convergent arches toward the leaf apex and are originated beside 248 each other at the leaf base(Fig. ...
... among This preprint research paper has not been peer reviewed. Electronic copy available at: https://ssrn.Gochnatia complex and many other Asteraceae(Freire et al. 2002;Metcalfe and Chalk, 1950; 293 Sancho, 1999;Youssef et al., 2013), whereas multistoried T-shaped trichome are not very common in294 Asteraceae and only have been reported in the tribe Senecioneae(Robinson, 1989) and in Ianthopappus 295 genus (Freire et al., 2002). 296 G. glutinosa leaves in transversal section showed heterogenous isolateral mesophyll with dense 297 content, formed by three to four layers of short adaxial palisade chlorenchyma, three to four layers of 298 compact spongy tissues with rounded cells and one or two layers of palisade abaxial parenchyma (Fig. 299 4A). ...
Gochnatia glutinosa is a shrub that grown in the Argentinean semiarid region (Monte region) used in the ancestral medicine as an antiseptic and anti-inflammatory agent.This study was aimed to examine the morpho-anatomical characteristics of G. glutinosa aerial parts, identify the chemical composition of traditionally used preparations to assess its pharmacobotanical characterization and evaluate its activity as antiseptic and anti-inflammatory to give scientific support to its traditional uses. G. glutinosa morpho-anatomical description was performed following standard histological techniques. Tincture and infusion of its aerial parts were prepared and were subjected to phytochemical analysis. Xanthine oxidase (XOD) and lipoxygenase (LOX) inhibition experiments, as well as ABTS•+, superoxide radical, and hydrogen peroxide scavenging activity, were carried out. The growth inhibition of methicillin-resistant Staphylococcus aureus (MRSA) strains was also determined.The morpho-anatomical traits of G. glutinosa leaves and stems were reported for the first time. The medicinal preparations exhibited a large amount of phenolic chemicals mainly flavonoids such as rhamnetin, arcapillin, rhamnacin, hesperetin, isorhamnetin, centaureidin, europetin 7-O-mehylmyricetin, cirsiliol, sakuranetin, genkwanin and eupatorine and also phenolic acids and diterpenoid derivatives. Both preparations had free radical scavenging activity and were able to reduce both XOD and LOX activity, indicating their anti-inflammatory properties. Besides, tincture was effective against all MRSA strains (MIC values ranging from 60 to 240 g DW/mL).The results obtained in this work scientifically support the medicinal popular use of G. glutinosa as an antiseptic and anti-inflammatory. The identification of bioactive compounds and their morpho-anatomical description contribute to the quality control of this medicinal plant from Argentine Calchaquí Valley.
... Si bien el melendre ha sido descrito botánicamente por varios autores (Cabrera, 1950;Cabrera, 1971;Cabrera, 1977;Beltrán y Ferreyra, 2001;Freire et al., 2002;Sancho et al., 2005;Panero y Funk, 2008), en este trabajo se han realizado observaciones complementarias asociadas al valor forrajero de esta especie, en zonas semi-áridas. La raíz del melendre el pivotante, bastante ramificada y muy profunda. ...
Resumen. En varios municipios del Cono Sur de Cochabamba, la actividad pecuaria es muy importante para la seguridad alimentaria y la economía de los pobladores. El mane-jo del ganado es semi estabulado, por lo tanto, gran parte del año, los bovinos, caprinos y ovinos deben buscar su alimento en los bosques y praderas naturales de la zona, si-tuación que genera condiciones de sobrepastoreo que provoca una severa disminución de muchas especies forrajeras altamente valiosas para el medio. Es así que los bosques actuales están compuestos por especies espinosas y muy pocas forrajeras sin espinos. Una de estas especies forrajeras sin espinos que ha sobrevivido a este proceso, es el melendre (Gochnatia palosanto Cabrera). Es una especie que los criadores de ganado la citan en primer lugar cuando son cuestionados sobre las plantas nativas que consume el ganado. El presente estudio etnobotánico permitió corroborar el potencial forrajero del melendre. Palabras clave: Forrajera nativa; Restauración ecológica; Pastoreo en bosque Abstract: Ethnobotanical study of the melendre (Gochnatia palosanto Cabrera), a promising forage tree in the Southern Cone of Cochabamba. In several municipalities of the Southern Cone of Cochabamba, livestock activity is very important for food security and for the economy of their inhabitants. Livestock management is semi-stabled, therefore , much of the year, cattle, goats and sheep must seek their food in the forests and natural grasslands of the area, a situation that generates overgrazing conditions that causes a severe decrease in many highly valuable forage species for the environment. Thus, today's forests are made up of spiny species and very few thornless foragers. One of these thornless forage species that has survived this process is the melendre (Gochnatia palosanto Cabrera). It is a species that cattle breeders cite first when they are questioned about the native plants consumed by cattle. This ethnobotanical study allowed understanding the forage potential of melendre.
... from Brazil) and 26 species from the West Indies. This concept was maintained until Freire & al. (2002), based on the glabrous and three-veined leaves, white corollas, and anther appendage attenuate, considered G. rotundifolia distinct enough from the other sections of Gochnatia and liable to be placed in its own, monotypic section, Gochnatia sect. Rotundifolia S.E. ...
... Morphological study. -Data were derived from the study of herbarium specimens from ALCB, GH, LP, NY, S, and US (Appendix 1) and previous literature (Cabrera, 1971;Freire & al., 2002;Sancho & al., 2005;Tellería & al., 2013). For light microscopy examination, floral and vegetative parts were re-hydrated in water and stained in 2% safranin. ...
... Morphological study. -Morphological features of Gochnatia rotundifolia were described in Cabrera (1971), Freire & al. (2002), , and Tellería & al. (2013), and our results mostly agree with these authors. In this paper, we will focus on leaf venation and pappus characteristics that were not previously studied in detail. ...
The tribe Gochnatieae is restricted to America. A recent molecular study including over 60% of the species of the tribe was carried out to untangle evolutionary relationships among the taxa. Eight supported clades were recovered as monophyletic genera, while Gochnatia s.l. was revealed as paraphyletic and currently restricted to the Central Andes. Gochnatia rotundifolia was not included in this phylogeny since extracted DNA from few and old collections from Brazil was unusuable. Since Cabrera's treatment of Gochnatia s.l., this species has received special attention due to its distinct morphological characters when compared with the other monophyletic genera of Gochnatieae. Therefore, this paper aims to provide new evidence that supports Gochnatia rotundifolia to be described as a separate monospecific genus based on a morphological study and a maximum parsimony (MP) analysis. A total of 21 species were studied. The leaf venation was analysed using digital X-ray capture (VIVA-Varian Image viewing & Aquisition). Seventeen morphological characters obtained from vegetative and floral parts of the plants were included in the data matrix. Our results reinforce the placement of Gochnatia rotundifolia within the tribe Gochnatieae. Gochnatia rotundifolia is consistently recovered in a clade as sister to the Cnicothamnus-Richterago group. The species differs from the other genera of the tribe by an actinodro-mous basal leaf venation with three primary veins, and pappus elements composed by (50-)60-75 bristles, sometimes almost flat setae, arranged in 2 or 3 series, unequal in length. Our results support Gochnatia rotundifolia as the sole species Vickia rotundifolia comb. nov. in a new genus of Gochnatieae, Vickia gen. nov. The new genus is described, illustrated, and a generic identification key for the tribe Gochnatieae is provided.
... These two characters are also present in members of the tribe Hyalideae of the subfamily Stifftioideae (Hyalis, Ianthopappus, Leucomeris, Nouelia). On the other hand, the pappus type E, i.e. half of the bristles long, relatively wide and flat, somewhat paleaceous, and plumose at the apex, and half of the bristles short, thin, and all scabrid (Freire et al. 2002), relates the genus to the subfamily Wunderlichioideae. Another option would be to treat it as a separate tribe or subfamily. ...
A thorough morphological circumscription of the tribe Mutisieae (Asteraceae) was established in 1977 by Angel L. Cabrera (1908–1999), who made fundamental worldwide contributions to the taxonomy of the family Asteraceae. The advance of molecular phylogenetic studies resulted in portrayals of relationships that have impacted the classification of the whole family and particularly of the Mutisieae. The publication of the book Systematics, Evolution and Biogeography of the Compositae in 2009, with the collaboration of the main worldwide specialists in the family, represented another fundamental milestone in the classification of the tribe. Mutisieae, which was previously thought to be nested far up in the phylogenetic tree of the family, is now known to be a basal grade, becoming a focus of attention for synantherologists. Traditional Mutisieae, or Mutisieae sensu Cabrera, is now split into numerous new subfamilies, tribes and subtribes. As a result of these changes, genera within the tribe have been reduced from 89 to 48 through new synonymisations, transfers to other tribes, and partly offset by description of 23 new genera. The remaining genera of Mutisieae sensu Cabrera have been placed in tribes of different subfamilies. A synthesis of these changes is presented in tables, in an illustrated key, and in a synopsis of the subfamilies with keys to tribes within the subfamilies.
... Subfamily Gochnatioideae was described (Panero & Funk, 2002) to include four genera of Mutisieae sensu Cabrera (1977), Cnicothamnus, Cyclolepis, Gochnatia, and Richterago, that form a well-supported clade in phylogenetic analysis of chloroplast DNA sequence data (Panero & Funk, 2008). There are no morphological synapomorphies for the subfamily; characters that have been used to define a Gochnatia complex (Freire et al., 2002) are also found in closely related taxa outside Gochnatioideae since the two Southeast Asian species of Gochnatia section Leucomeris were found to be sister to the genus Nouelia tribe Hyalideae of southwestern China (Panero and Funk, 2008). Sancho and Freire (2009) point out that genera of Gochnatioideae can be recognized by a combination of three characters including dorsally glabrous style branches, apiculate apical anther appendages and deeply dissected 5-lobed corollas. ...
... Cabrera recognized six sections in the genus including Discoseris, Gochnatia, Hedraiophyllum, Leucomeris, Moquiniastrum, and Pentaphorus. Freire et al. (2002) synonymized section Moquiniastrum under sect Hedraiophyllum and recognized three additional sections, namely Anastraphioides, Glomerata and Rotundifolia. The process of defining a natural Gochnatia started with the removal of species of Gochnatia section Discoseris and placing them in Richterago (Roque & Pirani, 2001). ...
... Hind (2007) recognized the two species of section Pentaphorus at the genus level but Telleria et al. (2013) did not find pollen characters to differentiate it from other South American gochnatias. Anastraphia was recently resurrected (Ventosa-Rodriguez & Herrera-Olivier, 2011b) to accommodate all the Caribbean species of Gochnatia placed by Freire et al. (2002) in section Anastraphioides. A phylogenetic study using nrITS concatenated with several chloroplast markers (Funk et al., 2014) shows that Gochnatia s. l. is paraphyletic with the South American species placed in two clades, one with members of Gochnatia section Moquiniastrum sister to Richterago, and the other clade including taxa of Gochnatia sections Pentaphorus and Gochnatia and sister to all other species of the subfamily except Cyclolepis. ...
Subfamily Gochnatioideae was described (Panero & Funk, 2002) to include four genera of Mutisieae sensu Cabrera (1977), Cnicothamnus, Cyclolepis, Gochnatia, and Richterago, that form a well-supported clade in phylogenetic analysis of chloroplast DNA sequence data (Panero & Funk, 2008). There are no morphological synapomorphies for the subfamily; characters that have been used to define a Gochnatia complex (Freire et al., 2002) are also found in closely related taxa outside Gochnatioideae since the two Southeast Asian species of Gochnatia section Leucomeris were found to be sister to the genus Nouelia tribe Hyalideae of southwestern China (Panero and Funk, 2008). Sancho and Freire (2009) point out that genera of Gochnatioideae can be recognized by a combination of three characters including dorsally glabrous style branches, apiculate apical anther appendages and deeply dissected 5-lobed corollas. Except for Cnicothamnus and three species of Richterago (Roque & Pirani, 2014) members of the subfamily have discoid capitula. Gochnatioideae contains approximately 90 species distributed in South America, the Caribbean and Mexico.
A stable generic classification of the Gochnatioideae has not been reached mainly because of the difficulty in circumscribing the limits of the genus Gochnatia. Cladistic analyses of morphological characters show that Gochnatia as circumscribed by Cabrera (1971) is a polyphyletic genus (Bremer, 1994; Jiménez Rodríguez et al., 2004; Ventosa-Rodriguez and Herrera-Olivier, 2011a). Cabrera recognized six sections in the genus including Discoseris, Gochnatia, Hedraiophyllum, Leucomeris, Moquiniastrum, and Pentaphorus. Freire et al. (2002) synonymized section Moquiniastrum under sect Hedraiophyllum and recognized three additional sections, namely Anastraphioides, Glomerata and Rotundifolia. The process of defining a natural Gochnatia started with the removal of species of Gochnatia section Discoseris and placing them in Richterago (Roque & Pirani, 2001). Hind (2007) recognized the two species of section Pentaphorus at the genus level but Telleria et al. (2013) did not find pollen characters to differentiate it from other South American gochnatias. Anastraphia was recently resurrected (Ventosa-Rodriguez & Herrera-Olivier, 2011b) to accommodate all the Caribbean species of Gochnatia placed by Freire et al. (2002) in section Anastraphioides. A phylogenetic study using nrITS concatenated with several chloroplast markers (Funk et al., 2014) shows that Gochnatia s. l. is paraphyletic with the South American species placed in two clades, one with members of Gochnatia section Moquiniastrum sister to Richterago, and the other clade including taxa of Gochnatia sections Pentaphorus and Gochnatia and sister to all other species of the subfamily except Cyclolepis. The Mexican species of Gochnatia that were strongly supported as sister to Anastraphia have been recognized subsequently as a new genus, Nahuatlea (Funk et al., 2017). Despite a lack of significant statistical support for the relationships of the three clades of South American Gochnatia representing sections Gochnatia, Moquiniastrum and Pentaphorus, Funk et al. (2014) choose to recognize these three groups at the genus level.
The phylogenetic relationships of Cyclolepis to other Gochnatioideae and Wunderlichioideae in nuclear genomic studies of Asteraceae (Mandel et al., 2019) is equivocal, and conflicts with earlier Asteraceae phylogenies based on plastid data (Panero & Funk, 2008; Panero et al., 2014). In concatenated analysis of nuclear genomic data, a Cyclolepis –Wunderlichioideae clade is moderately supported as sister to Gochnatioideae. In coalescent-based species tree estimation, Cyclolepis is sister to Gochnatioideae but these are collectively sister to Wunderlichioideae rather than a Hecastocleidoideae - Asteroideae clade as reconstructed from chloroplast DNA.
To ascertain generic relationships in Gochnatioideae I undertook a sequence analysis of ITS and ETS regions of the nuclear ribosomal DNA. In particular, I aimed to (1) evaluate the monophyly of the South American Gochnatia and (2) determine the relationships of Nahuatlea to other genera of Gochnatioideae.
... In the species studied here, these trichomes were observed in members of different tribes (Eupatorieae, Vernonieae, Inuleae, Neurolaeneae, Millerieae, Gnaphalieae and Heliantheae), as well as reported in other studies involving species of different tribes of the family: Eupatorieae (Castro et al. 1997, Monteiro et al. 2001, Del-Vecho-Vieira et al. 2008, Trindade 2013, Trindade et al. 2014, Fernandes et al. 2016, Heliantheae (Empinotti 2005, Martins et al. 2006, Bombo et al. 2012, Amrehn et al. 2013, Oliveira et al. 2013, Vernonieae (Narayana 1979, Castro et al. 1997, Empinotti 2005, Empinotti & Duarte 2008, Santos 2013, Sosa et al. 2014, Neurolaeneae (Castro et al. 1997, Budel et al. 2006, Millerieae (Araújo et al. 2013), Mutisieae (Castro et al. 1997, Freire et al. 2002, Melo-de-Pinna 2004, Astereae (Cortadi et al. 1999, Budel & Duarte 2007 and Inuleae (Castro et al. 1997). Although the literature (cited above) indicates the occurrence of uniseriate to multiseriate vesicular-capitate trichomes in representatives of the tribes Mutisieae and Astereae, these were not observed among the species of these tribes studied here, possibly due to the analysis of only one species of each of these tribes. ...
... Trichomes like the oblique-flagellate type were observed by Cabrera & Ragonese (1978) in American species of the genus Pterocaulon (including Pterocaulon angustifolium and Pterocaulon alopecuroides, however, in our study this last species did not present this type of thichome), but these authors use the term "non-septate flagelliform" to identify this trichome. Freire et al. (2002) also indicated the occurrence of these trichomes in some species of the genus Gochnatia (Gochnatieae). ...
... Trichomes of this type are frequent in the species of the Vernonieae tribe and can be used in the taxonomy of the representatives of this tribe (Narayana 1979, Redonda-Martínez et al. 2012, Wagner et al. 2014. Similar trichomes were also observed in some species of the genus Gochnatia (Gochnatieae) (Freire et al. 2002). ...
Asteraceae is the largest family of angiosperms and occurs predominantly in grassland areas. This study aimed to identify and characterize the foliar trichomes of 34 Asteraceae species from Sand-fields of the Pampa biome, by means of epidermal analyzes (front and transverse view) under light and scanning electron microscopy. Eleven types of trichomes were identified and characterized: three glandular (recurved on the epidermis, erect-capitate and uniseriate to multiseriate vesicular-capitate) and eight non-glandular (simple conical, flagellate-filiform, aseptate-flagellate, whip-like, oblique-flagellate, branched with one arm, branched with two T-shaped arms, branched with three or more arms). The most representative glandular type was the uniseriate to multiseriate vesicular-capitate (58%) and the non-glandular type was the simple conical (35%). A large number of trichomes is an adaptive strategy to the adverse conditions of the Pampa biome and its morphological diversity can be useful in the family systematics.
... It is a perennial rhizomatous sub-shrub of ca. 1 m height used in environment restoration (Dalmasso, 2010). Hyalis argentea has capitula (inflorescences) arranged in terminal pseudocorymbes (Freire et al., 2002). Each capitulum has five to six violet florets: four or five bilabiated ray florets and one tubular pentasect disk floret (Cabrera, 1963;Fig. ...
... 1). The style is bilobed, without sweeping hairs and the achene has a pappus with heterogeneous bristles (Cabrera, 1963;Freire et al., 2002;Torres & Galetto, 2007;Roque & Funk, 2013). Hyalis argentea is a nondormant species that germinates during autumn and blooms from December to February (Camina et al., 2013;Forcone & Andrada, 2006), and its inflorescences are visited mainly by insects (Camina, 2011). ...
... The absence of the disk floret contrasts with the description made by Cabrera (1963) for this species. It is common that in Asteraceae the proportion of florets per capitulum usually vary (Freire et al., 2002), but the variability observed here had not been previously reported for H. argentea. Notably, the morphological traits studied here at the capitulum level have been little investigated in other Asteraceae species with homogamous capitula (but see Lane, 1996). ...
p> Introducción y objetivos: Una asignación diferencial de recursos entre la función de atracción, recompensa y sexual ha sido observada en capítulos de especies derivadas de Asteraceae. Los capítulos heterógamos están compuestos por flores pistiladas, estaminadas o estériles y perfectas, con lo cual las funciones sexual y de recompensa son realizadas por diferentes tipos de flores y en distintos momentos. Esta distribución espacial y temporal de las recompensas dentro del capítulo no es tan clara en especies con capítulos homógamos, donde todas las flores son perfectas y producen polen y néctar. Aquí evaluamos la distribución espacial y temporal de las recompensas florales en los capítulos homógamos de Hyalis argentea.
M&M: Comparamos la fenología floral, el número de granos de polen y la concentración y volumen de néctar entre las flores marginales y centrales, y registramos el comportamiento de forrajeo de los visitantes florales.
Resultados: Los capítulos tienen un patrón de floración centrípeto y también alterno y son visitados por abejas, hormigas, mariposas, polillas, escarabajos y trips, siendo Apis mellifera su principal polinizador. No encontramos un patrón temporal en la oferta de recompensas dentro de los capítulos, pero sí un patrón espacial en el volumen de néctar que aumenta desde las flores marginales hacia las del centro del capítulo.
Conclusiones: Dicha variabilidad espacial en la cantidad de néctar podría afectar el comportamiento de forrajeo de los polinizadores y así aumentar las probabilidades de polinización cruzada, mejorando la reproducción sexual de esta especie autoincompatible.</p
... It is a perennial rhizomatous sub-shrub of ca. 1 m height used in environment restoration (Dalmasso, 2010). Hyalis argentea has capitula (inflorescences) arranged in terminal pseudocorymbes (Freire et al., 2002). Each capitulum has five to six violet florets: four or five bilabiated ray florets and one tubular pentasect disk floret (Cabrera, 1963;Fig. ...
... 1). The style is bilobed, without sweeping hairs and the achene has a pappus with heterogeneous bristles (Cabrera, 1963;Freire et al., 2002;Torres & Galetto, 2007;Roque & Funk, 2013). Hyalis argentea is a nondormant species that germinates during autumn and blooms from December to February (Camina et al., 2013;Forcone & Andrada, 2006), and its inflorescences are visited mainly by insects (Camina, 2011). ...
... The absence of the disk floret contrasts with the description made by Cabrera (1963) for this species. It is common that in Asteraceae the proportion of florets per capitulum usually vary (Freire et al., 2002), but the variability observed here had not been previously reported for H. argentea. Notably, the morphological traits studied here at the capitulum level have been little investigated in other Asteraceae species with homogamous capitula (but see Lane, 1996). ...
Differential resource allocation to attraction, reward, and sexual functions has been observed at capitula level of derived species of Asteraceae. Heterogamous capitula contain a combination of pistillate, staminate or sterile and perfect florets, thus rewarding and sexual functions are performed by different kinds of florets and at different times. Such spatial and temporal distribution of rewards within the capitula is not as clearly established in species with homogamous capitula, where all florets are perfect and produce pollen and nectar. We evaluated the spatial and temporal distribution of floral rewards in homogamous capitula of Hyalis argentea. We compared the floral phenology, the number of pollen grains and nectar volume and concentration between ray and disk florets, and registered the foraging behaviour of floral visitors. Capitula have a centripetal and alternate flowering pattern and they were visited by bees, ants, butterflies, moths, beetles and thrips, but pollinated mainly by Apis mellifera. We did not find a temporal pattern in the offer of rewards within the capitula, but we do find a spatial pattern in nectar volume increasing from outer to inner florets. This spatial variability in nectar quantity could impact pollinator behavior and thus enhance outcrossing likelihoods improving sexual reproduction in this self-incompatible species.
