The study was based on plants observed, collected and photo-documented in the field during three collecting trips in Bolivia and Peru, as well as physical specimens loaned to B from AAU, F, G, K, LPB, MA, MO, NY and P (Thiers, continuously updated). Specimens that were physically examined are listed in Suppl. material 1. In addition, high resolution digital images of herbarium specimens, in particular types, were consulted online either accessed through JSTOR Global Plants (https://plants.jstor.org/), GBIF (https://www.gbif.org/) or directly through online databases of the individual herbaria.

The species inventory of the Gynoxyoids was built in a database using the EDIT Platform for Cybertaxonomy (Berendsohn 2010), based on imports of names and associated data (authors, protologue citations) from the International Plant Names Index (IPNI) (https://www.ipni.org/) supplemented by TROPICOS (https://tropicos.org/home), the Global Asteraceae Database (https://www.Asteraceae.org/aphia.php?p=stats) and the World Flora online (http://www.worldfloraonline.org/)

The first round of assessing the morphological variation in the Gynoxyoid group included all species of the genera Aequatorium, Nordenstamia, Paracalia and Paragynoxys and a representative selection in terms of morphological diversity of Gynoxys species, altogether 65 species. We examined the diagnostic characters stated in the protologues and in other studies of the five genera, but also compared specimens to detect morphological variance to develop a list of characters and their states. For this investigation, a character state was considered taxonomically relevant and selected for further processing if its expression marked morphological discontinuities at supra-specific level. For each such character, unordered categorical states were defined following the terminology by Roque et al. (2009) and Beentje (2010). In cases where a more detailed homology statement was needed due to conflicting or unclear use of character definitions or terms, a description and illustration were included. For later reconstruction of character evolution, a specimen-based matrix of characters and states suitable for reliable delimitation and characterisation of supra-specific entities was constructed using the specimens included in the plastid phylogenomic analysis of Escobari et al. (2021). For certain characters, for example, the plant habit, the respective states were recorded from literature if not given on the specimen label.

Only Bayesian trees obtained from complete plastome sequences with indels coded and alignments manually corrected as provided by Escobari et al. (2021) were used as the hypothesis of the phylogenetic relationships in the Gynoxyoids, because lack of resolution rendered the use of nrDNA marker trees impossible. The reconstruction of character states at ancestral nodes was performed with a Bayesian approach using BayesTraits version 2.0 (Pagel and Meade 2006), which uses a selection of post-burn-in trees obtained from the t.files of the Bayesian analysis. This random selection of 800 of the total of 1600 post-burn-in trees taken from Escobari et al. (2021) was obtained through Mesquite version 3.7 (Maddison and Maddison 2021). The file stating the relevant nodes of the tree to be addressed by the analyses of BayesTraits was generated with TreeGraph v.2.14beta (Stöver and Müller 2010). The inference of the ancestral character state reconstruction was performed using the reverse jump MCMC approach with 5,050,000 iterations, with a burn-in of 50000, a sample frequency of 1000 and, following the recommendation by Pagel and Meade (2006), a hyper-prior where the mean of the exponential is drawn from a uniform 0–100 distribution. TreeGraph v.2.14beta (Stöver and Müller 2010) was used to plot the results from the BayesTraits output log file with the function Import BayesTraits data on the Bayesian major consensus tree. We excluded the other genera of the Tussilagineae that were present in the plastid phylogenomic investigation, considering that the outgroup sampling in their dataset is incomplete with respect to the morphological diversity.

To achieve a better overview on species-level phylogenetic relationships within the Gynoxyoid clade and to test if groups of samples identified with the same species name appeared in terminal subclades, 171 samples belonging to 50 species (Suppl. material 1) were included into an extended molecular dataset. These samples were selected to cover morphological and geographical variation as much as possible and also included the samples that were already part of the plastid phylogenomic study. The nrITS and nrETS regions were used as they provided some variable and informative characters in a short marker that was possible to sequence with little effort per sample. Additionally, by representing the nuclear genome, the dataset could be used to test for incongruence between trees inferred from different genomic compartments. Plastid regions often applied to assess the tree space of speciose clades (Mansion et al. 2012) were not suitable in the Gynoxyoid clade due to extremely low genetic distances (Escobari et al. 2020). Genomic DNA was extracted using the CTAB method by Doyle and Doyle (1987), with three fractions for each sample as modified by Borsch et al. (2003). PCR amplification of ITS followed White et al. (1990), ETS was amplified with the primers AST-1 (f) and 18-S-ETS (r) (Markos and Baldwin 2001), following Pelser et al. (2010). PCR was performed in a peqSTAR Thermocycler 1107D (PeqLab, Erlangen, Germany). The PCR products were electrophoresed on 1.5% agarose, the bands were cut out and cleaned with the GenepHlow Gel/PCR kit (Geneaid, New Taipei, Taiwan). Samples were sequenced by Macrogen Europe (Amsterdam, The Netherlands). Sequence files were aligned using MAFFT v.7.394 (Katoh and Standley 2013) and manually edited using PhyDE version 0.9971 (Müller et al. 2010), following the rules of Löhne and Borsch (2005). Indels were coded as binary characters using the simple-indel-coding method (Simmons and Ochoterena 2000) in SeqState version 1.4.1 (Müller 2005). Altogether, 146 ETS and 166 ITS sequences were newly generated and the sequences were deposited in the European Nucleotide Archive (ENA) using the annonex2embl submission pipeline (Grünstäudl 2020) and can be retrieved from ENA under study number PRJEB53579 (https://www.ebi.ac.uk/ena/submit/webin/study/PRJEB53579).

Phylogenetic trees were inferred from the ITS, ETS and a concatenated matrix of both belonging to the corresponding samples in the plastid tree presented in Escobari et al. (2021). A Bayesian analysis was performed with MrBayes v.3.2.6 (Ronquist and Huelsenbeck 2003), using four parallel Markov Chain Monte Carlo (MCMC) runs for a total of 50 million generations under the GTR+G+I model. The convergence of the Markov chains was checked with Tracer v.1.7 (Rambaut et al. 2018). The initial 25% of all trees were discarded as burn‐in and the remaining trees were used to summarise the 50% majority consensus tree.

Despite the extended nuclear ribosomal sequence dataset generated in this investigation, not all species could be included into phylogenetic analysis. This was largely due to the unavailability of suitable material, for example, in species only known from type or historical specimens. We, therefore, used our set of eleven morphological characters with their states in conjunction with the results from ancestral state reconstruction, to assign all species to a genus and, in the case of Gynoxys species, also to informal infrageneric groups of morphologically similar species that can be used as a hypothesis on close relationships. The genera and informal infrageneric entities were described and a taxonomic key for their determination was created. At species level, all protologues were consulted to check for the correct typification of names. Type specimens of all names, with the exception of only a few unavailable ones (indicated in the taxonomic treatment part, below), were examined from high resolution digital images provided by JSTOR Global Plants, GBIF and the herbarium websites of individual herbaria. The digital images of type specimens were referenced in the checklist to the type citation. Where necessary, new combinations were made and names were lectotypified. As a general principle, a morpho-species concept, delimiting species purely based on morphological discontinuities, was applied. Type specimens and additional specimens (see Suppl. material 1) were examined to assess the qualitative differences and possible infraspecific variation with the aim to hypothesise a name as accepted or as a synonym. The citation of authors follows the international standards by Brummitt and Powell (1992); the citation of publications follows BPH (Bridson et al. 2004) and TL-2 (Stafleu and Cowan 1976–1986; Stafleu and Mennega 1992–2009); the latter was also consulted for actual publication dates. Accepted names were provided with full synonymies and type citations. Type specimens that were online include only the herbarium acronym. Specimens that were physically examined are marked with (!).

The evaluation of morphological characters with respect to discontinuities at supra-specific level resulted in a matrix of eleven characters. These characters and their states are defined in Table 1 and, where appropriate, illustrated in Fig. 1.

Figure 1. 

Characters and states from floral morphology and indumentum for the Gynoxyoid clade a capitula: radiate (Gynoxys calyculisolvens left), discoid (G. longifolia right) b involucral outer phyllaries: absent (Paracalia pentamera left), present (Paragynoxys martingrantii right) c ratio corolla lobe/tube length: deeply lobed (Paragynoxys venezuelae left) (ratio > 0.6), shortly lobed (G. asterotricha right) (ratio < 0.6) d anther-base shape: obtuse (Paracalia jungioides left), sagittate (G. ignaciana right) e lenght of style branches: large (P. jungioides left), short (G. ignaciana right) f style branches apex shape: rounded (G. ignaciana left), acute (G. baccharoides right) g trichome architecture: multicellular hairs (A. jamesonii left above, N. kingii, right above), unicellular hairs (G. violacea below line). Scale bars: 2 mm (a–c); 200 μm (d); 500 μm (e); 100 μm (f, g).

Our evaluation for consistent presence and absence of sets of diagnostic character states in Gynoxyoid species resulted in the recognition of four morphologically and phylogenetically defined genera. The morphological matrix with the diagnostic characters applied to the genera and species of the Gynoxyoids represented in the sampling for the plastome tree is given in Table 2.

The first of these four genera is Aequatorium with all species sharing the combination of multicellular trichomes, radiate capitula, white flowers, a low number of disc flowers (< 8) and an obtuse shape of the anther base. Diagnostic for this genus is the unique combination of white flowers and radiate capitula.

Further genera are Paracalia and Paragynoxys, the species of which are differentiated from the other Gynoxyoid genera by a deep-lobed corolla, white flowers and discoid capitula. Paracalia can be distinguished from Paragynoxys by a scandent habit, absence of outer phyllaries and a central Andean distribution. In contrast, Paragynoxys has a woody habit, an involucrum with outer phyllaries and a north-Andean distribution.

The genus Nordenstamia cannot be delimited morphologically. The presence of stellate hairs by which this genus was originally distinguished from Gynoxys (Lundin 2006) is not only highly variable amongst the Nordenstamia species, but also shared with many Gynoxys species. If Nordenstamia is included in Gynoxys, this genus can be differentiated from all the others by the combination of yellow flowers and a shallowly divided disc corolla.

Gynoxys is notably the most diverse taxon within the Gynoxyoid clade, displaying a wide range of morphological variation. Within the genus, three informal groups can be discerned, based on distinct characteristics, including phyllotaxis, the number of ray flowers and the type of trichomes. The first group encompasses species with discoid capitula. In contrast, the second group comprises species with multiseriate stellate hairs, primarily featuring alternate leaves. Finally, the third and largest group is characterised by opposite leaves, radiate capitula and simple hairs.

In addition to the trees of the Gynoxyoids, based on a representative set of complete plastid genome sequences (Escobari et al. 2021), this study attempted to provide further phylogenetic evidence from nrDNA, amongst many others also including the same set of samples present in the plastid tree. Three phylogenetic analyses were performed, based on the ribosomal nuclear markers ETS, ITS and a concatenation of both (Suppl. material 2). In contrast to the tree, based on the plastid genome (Suppl. material 2: appendix 2a), the Bayesian ETS and ITS trees are poorly resolved (Suppl. material 2: appendix 2d). In all trees, the members of the Gynoxys clade form a single polytomy. The sister group relationship between the two species of Paragynoxys is the only clear congruence between the two nuclear ribosomal trees (however, with low support in the ITS inference) and is, moreover, in conformity with the plastid genome tree. Only the ETS tree resolved the two species of Paracalia as a (moderately supported) clade (Suppl. material 2: appendix 2c), whereas they were resolved in separate clades in the ITS (Suppl. material 2: appendix 2b) and in the concatenated ETS+ITS tree (Suppl. material 2: appendix 2d).

Employing the eleven characters of Table 1, a species-based matrix was created for the 17 members of the Gynoxyoid clade represented in the phylogenetic tree by Escobari et al. (2021) (Table 2) and used for ancestral character reconstruction. Two character states for G. ignaciana (colour of flowers) and G. violacea (length ratio disc flower corolla lobe/tube) were coded as missing because they were not accessible in the material at hand. The accession Gynoxys sp. in Escobari et al. (2021) was identified as G. calyculisolvens during this study. The ancestral character reconstructions for the eleven characters in the Gynoxyoid clade are presented in Figs 2–4.

Figure 2. 

Bayesian inference of ancestral character state reconstruction of four morphological characters of the Gynoxyoid clade in the consensus plastome tree by Escobari et al. (2021). Each pie chart represents a single character and each colour represents a character state as described in the legend. The actual state of the characters is represented by boxes next to the species names. The pie charts at the stem of the tree show the character abbreviations as mentioned in Table 1. Missing data are represented as (-).

Figure 3. 

Bayesian inference of ancestral character state reconstruction of four morphological characters of the Gynoxyoid clade in the consensus plastome tree by Escobari et al. (2021). Each pie chart represents a single character and each colour represents a character state which is described in the legend. The actual state of the characters is represented by boxes next to the species names. The pie charts at the stem of the tree show the character abbreviations as mentioned in Table 1.

Figure 4. 

Bayesian inference of ancestral character state reconstruction of four morphological characters of the Gynoxyoid clade in consensus plastome tree by Escobari et al. (2021). Each pie chart represents a single character and each colour represents a character state which is described in the legend. The actual state of the characters is represented by boxes next to the species names. The pie charts at the stem of the tree show the character abbreviations as mentioned in Table 1. Missing data are represented as (-).

The Gynoxyoids exhibit various evolutionary changes in their characteristics. The shrubby habit was initially considered primitive, but two independent shifts to a scandent habit occurred in the two Paracalia species, while a shift from shrub to tree habit was observed in Paragynoxys and within the Gynoxys clade. Opposite phyllotaxis was revealed as the ancestral state, but shifts to alternate phyllotaxis occurred in Paracalia pentamera, the stem node of Paragynoxys, two (out of three) species of Nordenstamia and Paracalia jungioides. Unicellular trichomes were revealed as ancestral for all Gynoxyoids, but Paracalia pentamera became glabrous. Multicellular hairs emerged in the most recent common ancestor of Aequatorium and Paragynoxys, as well as in certain species within the Gynoxys clade. White flowers were revealed as the ancestral state, retained by the earliest diverging clades (Aequatorium, Paragynoxys and Paracalia pentamera), while yellow flowers appeared at the stem node of Gynoxys and Nordenstamia. A reversal to white flowers occurred in Paracalia jungioides, nested within the Gynoxys clade. A higher number of inner phyllaries was ancestral, but both Paracalia and Paragynoxys species showed a decrease in this number. Radiate flowers were ancestral, but discoid capitula emerged in all Paracalia and Paragynoxys species, with additional losses of ray flowers in some Gynoxys species. A high number of disc flowers was the ancestral state, but reductions occurred at the stem node of Aequatorium and Paragynoxys and in all Paracalia species, partially within the Gynoxys clade. A shallow division of the corolla into lobes was revealed as plesiomorphic and retained in Aequatorium and all Gynoxys, but changed in Paracalia and Paragynoxys to a deep division. The style branch apex was rounded ancestrally, retained in Aequatorium and Paragynoxys, but an acute apex appeared in the earliest diverging species, Paracalia pentamera, with further shifts and reversals in Gynoxys, Nordenstamia and Paracalia jungioides.

A summary of the BayesTraits analysis of all state shifts for each character in Figs 2–4 is given in Fig. 5. Characters are represented by numbers and states with the codes given in Table 1. A high number of shifts occur in the two species of Paracalia because the genus is retrieved as non-monophyletic in the plastid topology, although its species share most morphological character states. The clade represented by both Paragynoxys species shared all derived characters with Aequatorium jamesonii in addition to five derived characters that characterise the clade. A single character (corolla colour) was retrieved as synapomorphic for the clade containing Aequatorium, Gynoxys, Paragynoxys, Nordenstamia and Paracalia jungioides and even this character shows several reversals at the MCRA of Aequatorium and Paragynoxys and of both species of Paracalia. The analysis retrieved most of the morphological characters as highly homoplastic with the style branch apices being the most variable character throughout the tree at many nodes. A summary of the BayesTraits analysis with each character at each node is given in Suppl. material 3.

Figure 5. 

Summary tree based on the results of the BayesTraits analysis (Figs 2–4) of state shifts in morphological character. A threshold of 0.75 was used to define the character shifts between states. Characters with multiple state shifts (homoplasies) are shown with white boxes, reversals are indicated by * and unresolved shifts are indicated by an open circle. Numbers at the left of the branches represent the nodes in Suppl. material 3.

The initial revision of the different databases resulted in a variable number of species. The Compositae Global Database lists 270 names (May 2023), The World Flora online includes 257 (Dec 2022), IPNI registers 242 (May 2023) and Tropicos has 243. This study resulted in a checklist comprising a total of 209 names of which 158 are accepted. Additionally, eleven names were newly synonymised, two were lectotypified and eight were newly transferred to other genera.

In the present investigation, we attempted to compare, for the members of the Gynoxyoid clade, tree reconstructions, based on the plastid genome and on the ITS and ETS nuclear ribosomal regions. Variation amongst the plastid genomes was extremely low (Escobari et al. 2021) and even more so in the nuclear ribosomal DNA. The lack of resolution in the nuclear ribosomal trees severely limits the comparison of phylogenetic signal from the organellar and nuclear genomic compartments. Nevertheless, there are some noteworthy exceptions. One is the unequivocal support for the sister group relationship of the two Paragynoxys members. The second is the missing support for the monophyly of the three Nordenstamia members in all three trees. The most significant result is, however, the gene tree incongruence concerning the two Paracalia species. The plastid tree placed Paracalia jungioides within Gynoxys and far distant from the second species, P. pentamera (Escobari et al. 2021). In contrast, the two species of Paracalia are supported as monophyletic in the ETS tree in conformity with morphology, although not in the ITS nor in the concatenated ETS-ITS tree (Suppl. material 2: appendix 2d). This finding is surprising because Paracalia jungioides is morphologically very distinct from all members of Gynoxys. It is scandent (instead of a tree or shrub), has white (instead of yellow) flowers and an involucre without outer phyllaries (instead of present). Moreover, P. jungioides and P. pentamera are morphologically very similar and the plastome phylogeny would suggest that these species have accumulated a high number of independent parallel state shifts (i.e. scandent habit, alternate leaves, absent outer phyllaries, few inner phyllaries, discoid capitula, few number of disc flowers, deep-lobed corolla, obtuse anther base) (Figs 3, 5). We assume that the incongruence with respect to the position of the two Paracalia species between the morphological data and the ETS topology on the one hand and the plastome tree topology on the other hand, is best explained by a chloroplast capture event. Chloroplast capture occurs when two species hybridise and go through extensive backcrossing to one of the ancestors (Rieseberg and Soltis 1991). The hybridisation event followed by extensive backcrossing swamp out the nuclear signal, but the captured plastid remains (Kandziora et al. 2022). In our case, we assume that P. jungioides, after introgression with a Gynoxys species, has captured the plastome of a member of the latter genus.

Nuclear-cytoplasmic incongruences have been reported in several studies within the Asteraceae family at higher and specific levels (Kilian et al. 2017; Pascual-Díaz et al. 2021; Senderowicz et al. 2021), especially in the Senecioneae (Pelser et al. 2007, 2010). It has also been shown by Stull et al. (2020) for the asterids that conflicts between nuclear and plastome trees are a relevant issue even at higher evolutionary scales. Phylogenetic inferences on nuclear data recovered different placements for several asterid lineages compared to topologies on plastid data (Yin et al. 2021; Kandziora et al. 2022). This is of some significance when we consider that current backbones of angiosperm phylogeny are largely based on plastid phylogenies (APG IV 2016). Amongst the principal reasons for these incongruences, horizontal gene flow amongst lineages, introgression, hybridisation and incomplete lineage sorting were suggested (Rieseberg and Soltis 1991; Maddison 1997; Vargas et al. 2017). The inclusion of different markers of different origins in a phylogenetic analysis has the capacity to elucidate signals of such events. Pelser et al. (2010) analysed potential causes for tree incongruences in the tribe Senecioneae comparing two nuclear (ITS/ETS) and six plastid markers. They concluded that hybridisation is a much more likely explanation than ILS, long-branch attraction or sampling error. Lee-Yaw et al. (2018) focused their study on organelle discordances by sequencing whole plastomes and over 1000 nuclear single-nucleotide polymorphisms in Helianthus L. The authors showed that incongruences in this genus can be expected at species level and amongst individuals of the same species. The Gynoxyoid clade is a further example of short molecular distances on plastid and ribosomal markers amongst species. The lack of molecular variability hampers the reconstruction of well-supported clades on this type of data; nevertheless, the great morphological variation enables the definition of morpho-species in many cases. On the other hand, the phylogenetic reconstruction, although with moderate support, can give evidence to support the assignment of morphologically similar individuals to the same entities (i.e. hypothesised species).

Gene tree discordance is expected to be more likely in rapid radiating lineages that can be found in young biodiversity hotspots, such as the Andean Region (Madriñán et al. 2013; Kandziora et al. 2022). The fast succession and accumulation of descendant species are prone to inter-breeding before reproductive barriers develop, increasing the probability of incomplete lineage sorting (ILS) (Vargas et al. 2017). In addition, young radiating groups have shown whole genome duplication and hybridisation events in the tropical high-altitude areas of South America (Lachemilla: Morales-Briones et al. 2018; Lupinus: Nevado et al. 2018; Diplostephium: Vargas et al. 2017; Espeletiinae: Cortés et al. 2018). Hybridisation may be a result of sexual selection, ecological adaptation, pollinator changes (Moreira-Munoz 2020; Kandziora et al. 2022) or due to the dynamic changes in habitat connectivity in this ecosystem with multiple topography changes during the Pleistocene (Flantua et al. 2019) which facilitated the contact between geographically isolated species before exhibiting strong barriers to gene flow (Vargas et al. 2017; Kandziora et al. 2022). Vargas et al. (2017) revealed complex patterns of reticulate evolution at generic and species level of Diplostephium.

Previous generic classifications of the Gynoxyoid group were based on morphological similarities and discontinuities between species assemblages. In this study, we tested these hypotheses by optimising character states on the full plastome phylogeny (Escobari et al. 2021, see also Suppl. material 2: appendix 2a). Morphological differentiation amongst the Gynoxyoids is shallow and limited to comparatively few and often rather subtle characters. The most recent ancestor of the Gynoxyoid had a shrubby habit, opposite leaves and it was vested by unicellular simple hairs. The capitula was radiate, equipped with outer phyllaries and 6–8 inner phyllaries and had up to eight disc flowers. The corolla was whitish and the corolla lobes were remarkably shorter than the corolla tube. Most of these plesiomorphic states (except the whitish corolla) were retained by most of the species of the genus Gynoxys during its evolution. Shifts in the character states are evident in the rest of the Gynoxyoid members. Specially, the switch from whitish to yellowish corolla (which is apparently the only synapomorphy under the given tree inference) resulted as unresolved due to a small difference of the PP values (59% yellow vs. 41% white). All shifts reconstructed under the ancestral character reconstruction were retrieved as highly homoplasious and are, therefore, unsuitable for genera characterisation under the given plastid inference presented in Escobari et al. (2021).

Our taxonomic backbone provides the best estimate of species diversity in the Gynoxyoid clade. Type information has been synthesised here for the first time in a comprehensive way. Further taxonomic knowledge turnover is expected at species level once species limits are tested in an integrative approach in a separate paper. Specially the examination of a reduced group of Bolivian species depicted shallow morphological differences, making the taxonomy complicated and predicting further nomenclatural changes. Additionally, the low number of collections available hinders a full examination of the species limits.

1. Aequatorium B.Nord. in Opera Bot. 44: 59. 1978 (Fig. 1f)

Type: Aequatorium asterotrichum B.Nord.

12 species

Colombia, Ecuador

Erect shrubs or trees, sometimes with sub-scandent branches. Indumentum tomentose, of subsessile stellate trichomes (with 1–3-tiered, irregularly star-shaped, subtended by a narrow pluricellular uniseriate stalk), with age, glabrescent, but with persistent greyish-white tomentum on petioles, abaxial side of leaves and sometimes involucres. Leaves alternate or subopposite or rarely opposite, petiolate, rounded-elliptic to lanceolate; margin entire, sinuate-dentate or denticulate, with small callous-tipped teeth; base acute, cuneate to rounded-truncate, subcordate or oblique; apex acute or rotund; coriaceous; leaves indumentum rusty-brownish on the adaxial side with two layers of peltate-stellate hairs, internal layer with sessile hairs and outer layer with subsessile hairs in patches, becoming grey-tomentose with age. Synflorescence terminal, rarely subterminal, (thyrsoid-) paniculiform or corymbiform. Capitula heterogamous. Receptacle flat or slightly convex. Involucre campanular; outer phyllaries usually < 6; inner phyllaries 5–10, biseriate. Ray flowers usually < 5 (–8); ligule white or cream-coloured. Disc flowers 5–10; corolla campanulate or funnel-shaped, white or pale greenish-yellow, shortly lobed, ratio lobes/tube < 0.8; lobes narrowly ovate, triangular or oblong, recurved to the outside or straight. Anther base sagittate-auriculate; filament collar narrowly cylindrical, uniform, not thicker than the filament. Style branches half contorted, apically obtuse or truncate. Distribution: Colombia, Ecuador.

Notes: We exclude Aequatorium venezuelanum from this genus, based on its yellow flowers and distribution and transfer this species to Gynoxys.

Aequatorium albiflorum (Wedd.) Cuatrec. & S.Díaz, Revista Acad. Colomb. Ci. Exact. 17(67): 665. 1990 ≡ Gynoxys albiflora Wedd., Chlor. Andina 1(3): 78. 1856 [“1855”]. – Syntypes: Colombia. Mariquita, sur la lisière du volcan de Tolima, 3900 m, Jan 1843, J. Linden 907 (F: V0076792F V0076793F (photo & fragments), K: K000497659, NY 178788, P: P00711390 P00711391 P02273078).

Aequatorium asterotrichum B.Nord., Opera Bot. 44: 59. 1978. – Holotype: Ecuador. Pichincha, lago Papallacta, thicket, 3300 m, 31 Oct 1955, E. Asplund 18263 (S: S-R–8297; isotypes: k: K000497658, LD 1821970, MO: MO–3237504, NY, P: P00971087, R, S: S18–7665, UPS, US).

Aequatorium caucanum S.Díaz & Cuatrec., Revista Acab. Colomb. Ci. Exact. 73: 248. 1994. – Holotype: Colombia. Cauca, Macizo Central Colombiano, Páramo de las Papas, El Boquerón, 3200–3510 m, 7–27 Sep 1958, J. Idrobo et al. 3221 (COL: COL000004758).

= Aequatorium caucanum var. abbreviatum S.Díaz & Cuatrec., Revista Acab. Colomb. Ci. Exact. 73: 248, f. 2. 1994. – Holotype: Colombia. Cauca, Volcán Puracé, alrededores de la Laguna San Rafael, 3340 m, 6 Jan 1972, A. M. Cleef & A. Fernandez 526 (COL: COL000004759).

Aequatorium jamesonii (S.F.Blake) C.Jeffrey, Kew Bull. 47(1): 61. 1992 ≡ Gynoxys jamesonii S.F.Blake, Acad. Sci. 18: 34. 1928. – Holotype: Ecuador. Pichincha, west side of Mount Pichincha, 3050 m, 2 Aug 1926, Jameson 227 (K: K000497657; isotype: US 00122911 (fragments & photo)).

= Senecio simulans Benoist, Bull. Soc. Bot. France 83: 808. 1937, nom. illeg. [non Senecio simulans Chiov. 1935] ≡ Gynoxys simulans Cuatrec., Brittonia 8: 158. 1955. – Syntype: Ecuador. Pichincha, 12 Jul 1931, Benoist 4572 (P: P02273075).

Aequatorium latibracteolatum S.Díaz & Cuatrec., Revista Acad. Colomb. Ci. Exact. 17(67): 661, 663, f. 1. 1990. – Holotype: Colombia. Cauca, Municipio de Puracé, Parque Nacional Natural del Puracé, cercanías de la Laguna San Rafael, 3300 m, 6 Oct 1984, C. G. Lozano 4667 (COL: COL000004762; isotypes: COL: COL000004760 COL000004761).

Aequatorium lepidotum B.Nord., Compositae Newslett. 31: 6, f. 3. 6B. 1997. – Holotype: Ecuador. Carchi, El Mirador, 15 km S of San Francisco, 00°37‘N, 77°31‘W, 3300 m, 2 Aug 1990, W. Palacios & D. Rubio 5286 (MO: MO–037535; isotype: US 01919680).

Aequatorium palealbum S.Díaz & A.Correa, Revista Acad. Colomb. Ci. Exact. 26(100): 345–346, f. 3. 2002. – Holotype: Colombia. Nariño, Ospina, páramo de Paja Blanca, alrededores de la bocatoma del acueducto, 1°58‘N, 77°34‘W, 3200 m, 2 Dic 1995, B. Ramirez-P. et al. 8904 (PSO: PSO0000058; isotype: MO: s.n.).

Aequatorium polygonoides B.Nord., Opera Bot. 44: 63. 1978 ≡ Senecio polygonoides Cuatrec., Notas a la Flora de Colombia 6: 20, f. 14. 1944, nom. illeg. [non Senecio polygonoides Muschl. 1911]. – Holotype: Colombia. Caldas, Cordillera Central, vertiente occidental, vertiente SE del Nevado del Ruiz, Termales, 3400 m, 4 May 1940, J. Cuatrecasas 9243 (COL).

Aequatorium repandiforme B.Nord., Compositae Newslett. 31: 9, f. 4. 1997. – Holotype: Ecuador. Pichincha, over high pass en route to Quito, 92 km E of Quevedo, 3400 m, 19 Sept 1959, B. Maguire & C. Maguire 44246 (NY 3468431; isotypes: K: K000497656, US 01919679).

Aequatorium sinuatifolium S.Díaz & A.Correa, Revista Acad. Colomb. Ci. Exact. 19(73): 251–252, f. 3. 1994. – Holotype: Colombia. Quindío, Mun. de Salento, arriba de Guayaquil, 3680 m, 10 Jan 1994, W. G. Vargas 1335 (COL: COL000004764; isotype: COL: COL000004763).

Aequatorium tatamanum S.Díaz & A.Correa, Revista Acad. Colomb. Ci. Exact. 23(88): 332, f. 1. 1999. – Holotype: Colombia. Risaralda, Municipio de Santua Río, Macizo de Tatamá, 200 m arriba del campamento El Reposo, 3700 m, 8 Feb 1983, J. H. Torres et al. 1720 (COL: COL000004765).

Note: This species is probably conspecific with A. albiflorum.

Aequatorium verrucosum (Wedd.) S.Díaz & Cuatrec., Revista Acad. Colomb. Ci. Exact. 17(67): 659–666. 1990 ≡ Gynoxys verrucosa Wedd., Chlor. Andina 1(3): 77. 1856 [“1855”] [non Gynoxys verrucosa V.M.Badillo 1946]. – Lectotype (Diaz & Cuatrecasas 1990: 663): Colombia. Nueva Granada, Mariquita, Cordillere de Quindiu, a Los Volcancitos, 3200 m, Jan 1843, Linden 1050 (P: P02273077; isolectotypes: F: V0076796F V0076797F V0076798F, K: K000497655).

= Senecio verrucosus Klatt, Abh. Naturf. Ges. Halle 15(2): 332. 1881 [1882]. –Syntypes: Triana s.n. (P; photo: F).

2. Paracalia Cuatrec., Brittonia 12: 183. 1960 (Fig. 1b, e)

Type: Paracalia pentamera (Cuatrec.) Cuatrec.

2 species

Bolivia, Peru

Scandent shrublets. Indumentum glabrate to glabrescent, of simple hairs, glabrate with age. Leaves alternate, petiolate, ovate; margin entire or angulate, with or without small callous-tipped teeth; base rotund or cordate; apex acuminate or mucronate; coriaceous; leaves glabrous or pilose on the adaxial site, but glabrescent with age. Synflorescence terminal or axillar, paniculiform or corymbiform. Capitula homogamous. Receptacle flat. Involucre tubular; outer phyllaries absent; inner phyllaries 5, uniseriate. Ray flowers absent. Disc flowers 5; corolla campanulate, white or pale greenish, deeply lobed, ratio lobes/tube ≈ 1; lobes linear, helically twisted. Anther base auriculate or obtuse; filament collar cylindrical, thicker than the filament. Style branches half contorted, apically obtuse or subtruncate. Distribution: Peru, Bolivia.

Note: Although the phylogenetic inferences suggest this genus to be not monophyletic, we kept the circumscription of Paracalia including two species. We substantiate this decision based on shared morphological characters, such as deeply lobed and white-flowered corolla and the central Andean distribution beginning from lowlands (800 m). Paracalia jungioides which is nested in the Gynoxys clade strikingly differs morphologically from the true Gynoxys species and its inclusion in this genus would break the continuity of the morphological characters and altitudinal distribution in this group. A possible explanation for the contradiction between morphological/ecological and molecular data may be chloroplast capture and this needs to be further studied and better understood before further nomenclatural decisions are made. In this context, we think the best practice is to retain the current circumscription of Paracalia and avoid suggesting further possibly wrong hypotheses of relationships of these species.

Paracalia jungioides (Hook. & Arn.) Cuatrec., Brittonia 12: 183. 1960 ≡ Pentanthus jungioides Hook. & Arn., Companion Bot. Mag. 1: 33. 1835. – Holotype: Perú. Purruchuca, Jun 1833, Matthews 1016 (K: K000497546(!); isotypes: E: E00414051 E00414052, K: K000497547(!)).

= Cacalia mikaniifolia DC., Prodr. 6: 328. 1837 ≡ Senecio mikaniifolius (DC.) Sch.Bip., Flora 28: 498. 1845. – Syntypes: Peru. San Buenaventura, Nee & Thibaud s.n. (not traced in G-DC, F: s.n. (photo)).

Paracalia pentamera (Cuatrec.) Cuatrec., Brittonia 12: 183. 1960 ≡ Senecio pentamerus Cuatrec., Fieldiana, Bot. 27: 57. 1951. – Holotype: Bolivia. La Paz, Larecaja, Copacabana (ca. 10 km. south of Mapiri), 850–950 m, 08 Oct – 15 Nov 1939, B. A. Krukoff 11150 (NY 259336(!); isotypes: A: A00010877, F: V0077069F, K: K000497545(!), S: S-R–7986, U 0105750, US 00123446).

3. Paragynoxys (Cuatrec.) Cuatrec., Brittonia 8: 153. 1955. (Fig. 1b, c)

≡ Senecio sect. Paragynoxys Cuatrec., Fieldiana, Bot. 27(2): 72. 1951.

Type: Paragynoxys neodrendoides (Cuatrec.) Cuatrec.

13 species

Colombia, Venezuela

Erect shrubs or trees. Indumentum tomentose of (always?) stellate T-shaped trichomes, persistent in all age states, becoming greyish-white on petioles, abaxial side of leaves and involucres. Leaves alternate or rarely opposite, petiolate, oblong-elliptic, obovate-elliptic or ovate; margin entire or repand, with or without small callous-tipped teeth; base cordate, obtuse or rarely cuneate; apex obtuse, attenuate or rarely acute; coriaceous; leaf indumentum shaggy rusty-brownish in the adaxial site, persistent with age. Synflorescence terminal rarely subterminal, (thyrsoid-) paniculiform or corymbiform. Capitula homogamous. Receptacle flat. Involucre campanular; outer phyllaries < 6; inner phyllaries 5 or 8, uniseriate. Ray flowers absent. Disc flowers 5–11; corolla campanulate, white, deeply lobed, ratio lobes/tube => 1; lobes linear, helically twisted. Anther base auricular or obtuse; filament collar cylindrical, thicker than the filament. Style branches fully contorted (forming a complete loop or even two), apically obtuse to subacute. Distribution: Colombia, Venezuela.

Note: We support the view of Correa (2003) who transferred Paragynoxys regis back to Gynoxys (as it was originally described), based on its radiate capitula with yellow flowers and distribution.

Paragynoxys angosturae (Cuatrec.) Cuatrec., Brittonia 8: 154. 1955 ≡ Senecio angosturae Cuatrec., Feddes Repert. Spec. Nov. Regni Veg. 55: 132. 1953. – Holotype: Colombia. Antioquia, Angostura, just outside town, 2000 m, 11 Mar 1944, F. R. Fosberg 21603 (US 00123252; isotypes: US 00123253 00123254).

Paragynoxys corei (Cuatrec.) Cuatrec., Brittonia 8: 154. 1955 ≡ Senecio corei Cuatrec., Feddes Repert. Spec. Nov. Regni Veg. 55: 136. 1953. – Holotype: Colombia. Antioquia, Alto El Oso, n. of Yarumal, 2320 m, 4 Mar 1944, E. L. Core 624 (F: V0051295F; isotype: US 00123277).

Paragynoxys cuatrecasasii Ruiz-Teran & López-Fig., Revista Fac. Farm. Univ. Andes 14: 14, f. 3, 4. 1974. – Holotype: Venezuela. Merida, Rangel, norte de la población Las Piedras, Cuenca del río Aracay, afluente del Santo Domingo, 2550–2700 m, 16 Dic 1972, Ruiz-Terán et al. 8258 (MERF; isotype: US).

Paragynoxys magnifolia Cuatrec., Brittonia 8(2): 154. 1955. – Holotype: Venezuela. Merida, Culata, 7000 ft., May 1847, N. Funck & Schlimm 1522 (P: P00711443 P00711444; isotypes: G: G00301285, P: P00711445 P00711446, US 00811048, VEN: VEN118056 (fragments of holotype)).

Paragynoxys martingrantii (Cuatrec.) Cuatrec., Brittonia 8: 156. 1955 ≡ Senecio martingrantii Cuatrec., Feddes Repert. Spec. Nov. Regni Veg. 55: 139. 1953. – Holotype: Colombia. Magdalena, Sierra de Perijá, Casacará Valley 23 km. East of Codazzi, 2 km from the Venezuelan border, 2450, 15 Feb 1945, M. L. Grant 10949 (F: V0051336F V0051337F; isotypes: COL: COL000005419, HUA: HUA0000364, NY 259292 259293, US 00123324 00123323, VEN: VEN209193, WIS: WISv0256984WIS WISv0256985WIS).

Paragynoxys meridana (Cuatrec.) Cuatrec., Brittonia 8(2): 156. 1955 ≡ Gynoxys verrucosa V.M.Badillo, Bol. Soc. Venez. Ci. Nat. 10: 312. 1946, nom. illeg. [non Gynoxys verrucosa Wedd. 1855] ≡ Senecio steyermarkii Cuatrec., Fieldiana, Bot. 27: 32–33. Jun 1950, nom. illeg. [non Senecio steyermarkii Greenm. Apr 1950] ≡ Senecio meridanus Cuatrec., Fieldiana, Bot. 27(2): 38. 1951. – Syntypes: Venezuela. Merida, Paramo de Pozo Negro between San José and Beguilla, 2590–3220 m, 3 May 1944, A. Steyermark 56268 (NY 259418, US 00123361).

Paragynoxys neodrendoides (Cuatrec.) Cuatrec., Brittonia 8(2): 156, f. 13, 14. 1955 ≡ Senecio neodendroides Cuatrec., Notas Fl. Colombia 6: 19, f. 13. 14. 1944. – Holotype: Colombia. Santander, Cordillera Oriental, Páramo de la Rusia, vertiente noroeste, 3300–3500 m, 4 Ago 1940, J. Cuatrecasas 10435 (COL; isotypes: F: V0051343F V0051344F V0051345F, P: P01816686).

Paragynoxys pileolanata S.Díaz, Caldasia 12(59): 379–381, f. 1. 1979. – Holotype: Colombia. Santander, Municipio de Onzaga, vereda Chaguaz, finca de Oliverio Mesa, en robledal, 2820 m, 29 Mar 1976, J. H. Torres et al. 500 (COL: COL000005310).

Paragynoxys santurbanensis (Cuatrec.) Cuatrec., Brittonia 8(2): 156. 1955 ≡ Senecio santurbanensis Cuatrec., Feddes Repert. Spec. Nov. Regni Veg. 55: 145. 1953. – Holotype: Colombia. Santander, Páramo de Santurbán, vert. W, 3100 m, 27 Jul 1940, J. Cuatrecasas & H. García Barriga 10326 (F: V0051361F V0051362F; isotype: P: P01816508).

Paragynoxys steyermarkii Cuatrec., Phytologia 40(1): 34. 1978. – Holotype: Venezuela. Táchira, Between Las Copas de Alto de Fila de Tierra Negra at the ridge dividing headwaters of rivers Quinimarí, Riofrio, Uribante and Talco (Oirá), 2870–2880 m, 16 Jan 1968, J. A. Steyermark & E. Dunsterville 101014 (US 00115958; isotypes: MA 638740, US 00115959, VEN: VEN74042).

Paragynoxys undatifolia Cuatrec., Proc. Biol. Soc. Washington 74: 15. 1961. – Holotype: Colombia. Magdalena, Sierra Nevada de Santa Marta. Southeastern slope: Hoya del Río Donachuí, below Sabanita Diricune, 3200 m, 29 Sep 1959, J. Cuatrecasas & R. Romero-Castañeda 24485 (US: US00115961 US00115962 US00115963 US00115964 US00115965; isotypes: COL: COL000005311 COL000005312 COL000005313 COL000005314, P: P00711449 P00711450, US: US00115960 US00930968).

Paragynoxys uribei Cuatrec., Phytologia 40(1): 33. 1978. – Holotype: Colombia. Boyaca, Arcabuco, 4 km. NE of town, 08 Jun 1966, L. U. Uribe 5633 (US: US00115966 US00115967; isotypes: COL: COL000005315 COL000005316).

Paragynoxys venezuelae (V.M.Badillo) Cuatrec., Brittonia 8(2): 156. 1955 ≡ Cacalia venezuelae V.M.Badillo, Bol. Soc. Venez. Ci. Nat. 10: 319. 1947 ≡ Senecio venezuelae (V.M.Badillo) Cuatrec., Fieldiana, Bot. 27(1): 31. 1950. – Holotype: Venezuela. Merida, Below páramo above San Isidro Alto, 1820 m, 14 May 1944, J. Steyermark 56560 (VEN: VEN32772; isotypes: F: V0049135F, NY 162855).

4. Gynoxys Cass. in Cuvier, Dict. Sci. Nat., ed. 2, 48(2): 455. 1827. (Fig. 1a, c–f)

Lectotype (Flann et al. 2010: 1225): Gynoxys baccharoides (Kunth) Cass.

= Nordenstamia Lundin, Compositae Newslett. 44: 15–16, f. 1. 2006, syn. nov. Type: Nordenstamia repanda (Wedd.) Lundin [≡ Gynoxys repanda Wedd.]

130 species

Argentina, Bolivia, Colombia, Ecuador, Peru, Venezuela

Erect shrubs or trees. Indumentum absent or tomentose, of simple or multicellular simple trichomes, glabrescent with age, but with persistent greyish-white tomentum on abaxial side of all leaves and involucres. Leaves alternate or opposite, petiolate or subsessile, elliptic, lanceolate, ovate, obovate; margin entire, repand, sinuate, sparsely angular or denticulate, with small callous-tipped teeth; base acute, attenuate, cordate, cuneate, obtuse, rotund, truncate or oblique; apex acute, acuminate, obtuse or mucronate; coriaceous or papyraceous; leaf indumentum absent, white or rusty-brownish in the adaxial site, persistent with age. Synflorescence terminal or axillar, paniculiform or corymbiform. Capitula homogamous (Sec 1) or heterogamous. Receptacle flat or convex. Involucre campanular; outer phyllaries 1–8; inner phyllaries usually 5–8(–13), biseriate. Ray flowers usually < 8 (–10–13); ligule yellow. Disc flowers usually 5–32 (–36); corolla tubular, campanulate or funnel-shaped, yellow, usually shortly lobed, ratio lobes/tube usually < 0.5; lobes triangular, oblong or narrowly ovate, recurved to the outside or straight. Anther base sagittate, auriculate or rarely obtuse; filament collar narrowly cylindrical, usually thicker than the filament. Style branches straight or half contorted, apically obtuse, truncate or acute.

Note: Gynoxys alternifolia and G. mandonii were described in literature as scandent. This information is certainly erroneous; in the field, we had a chance to trace several individuals of G. mandonii as large trees with thick branches and the type specimen of G. alternifolia also shows thick and erect branches with no sign of a liana-like growth.

Gynoxys campii Cuatrec., Brittonia 8(1): 39. 1954. – Holotype: Ecuador. Cañar, Near El Tambo (ca. 69 km. by railroad south of Sibambe), 9500–10000 ft., 5 Jul 1945, W. H. Camp E–3970 (F: V0076704F; isotypes: G: G00223899, GH: GH00008575, K: K000497540, NY 178793, P: P00711407, US 00122893, VEN: VEN34418).

Gynoxys dielsiana Domke, Biblioth. Bot. 116: 169. 1937. – Syntype: Ecuador. Chimborazo, Tipococha, untere Rand des Paramo, ca. 3230 m, 20 Aug 1933, L. Diels 675 (B, destroyed).

Gynoxys hutchisonii H.Rob. & Cuatrec., Novon 2(4): 414. 1992. – Holotype: Perú. Piura, above Huancabamba, road to Piura, 3000 m, 10 Oct 1957, P. C. Hutchison 1609 (US 00409556; isotype: F: V0076714F).

Gynoxys induta Cuatrec., Fieldiana, Bot. 27: 9. 1950. – Syntypes: Colombia. Valle, Cordillera Central, Hoya del río Bugalagrande, Barragán, Páramo de Bavaya, corrales, 3550–3400 m, 9 Apr 1946, J. Cuatrecasas 20546 (COL: COL000005215 COL000005216, F: V0076715F V0076716F V0076717F, P: P00711414, US 00122909, WIS: WIS00001047MAD).

Gynoxys leiotheca S.F.Blake, J.Wash. Acad. Sci. 18: 35. 1928. – Holotype: Ecuador. Borma, Sep 1904, Rivet 671 (P: P00711416; isotypes: US 00122915 (photo & fragments)).

Gynoxys littlei Cuatrec., Revista Acad. Colomb. Ci. Exact. 9: 242. 1954. – Holotype: Colombia. Huila, on foot of Cordillera Oriental, 20 km. SE of gigant, 103000 ft., 15 Sep 1944, E. L. Little 8658 (F: V0076752F; isotypes: COL: COL000005220, US 00650427).

Note: This species is described as having “pale flowers” in the protologue; nonetheless, the label of the type specimen describes the flowers as “pale yellow”.

Gynoxys longifolia Wedd., Chlor. Andina 1(3): 79. 1855. – Syntypes: Perú. Cuzco, Andes de Cuzco, Oct 1839 – Feb 1940, Gay s.n. (F: V0076718F, P: P00711417 P00711418 P00711419, US 00122917 (fragments)).

Gynoxys lopezii M.O.Dillon & Sagást., Brittonia 40(2): 223, f. 2. 1988 ≡ Paragynoxys lopezii (M.O. Dillon & Sagást.) Cuatrec., Phytologia 69(5): 314. 1990 ≡ Paracalia lopezii (M.O. Dillon & Sagást.) A.Correa, Brittonia 55(2): 167. 2003. – Holotype: Perú. La Libertad, Patáz, Yaupa (Llaupa), entre Chagual-Retanas, carretera a Tayabamba, en borde carretera, pedregoso, 2300 m, 24 Jan 1974, A. López & A. Sagástegui 8160 (HUT; isotypes: F: V0076719F, MO: MO–176388 s.n.).

Note: The taxonomic assignment of this species was discussed by Cuatrecasas (1990) and Correa (2003). Its ratio of corolla lobe and tube length resembles the genus Paracalia, but we agree with Correa (2003) that this species belongs to Gynoxys because of its yellow flowers, shrubby (non-climbing) habit and central Andean distribution.

Gynoxys megacephala Rusby, Bull. New York Bot. Gard. 4: 398. 1907. – Syntypes: Bolivia. M. Bang 1959 (F: V0076754F (fragments), GH: GH00008597, K: K000497526, MICH: MICH1107432, MO: MO–1183133, NY 178867 178868, PH: PH00013514, US 00122920, WIS: WISv0256704WIS). Bolivia. Huaycani, 11000 ft., May 1866, Pearce s.n. (MO: s.n.).

= Diplostephium foliosum Rusby, Bull. New York Bot. Gard. 8(28): 128–129. 1912. – Syntypes: Bolivia. Cargadira, 8000 ft., 29 Jul 1902, W. Roberts 1529 (BM: BM001024073, F: V0076745F, K: K000497534, NY 168221).

Gynoxys moritziana Sch.Bip. ex Wedd., Chlor. Andina 1: 79. 1855. – Syntypes: Venezuela. Merida, Sierra Nevada, 1844, Moritz 1385 (GH: GH00008598 GH00008599 (drawing & fragment), K: K000497525, P: P00711421 P00711422 P00711423, US 00122924 (fragments)).

Gynoxys pendula Sch.Bip. ex Wedd., Chlor. Andina 1(3): 78. 1855. – Syntypes: Colombia. Nouvelle-Grenade, Mariquita, Boqueron, Tolima, Jan 1843, J. J. Linden 954 (F: V0076760F V0076761F, GH: GH00008601, K: K000497523, NY 178870, P: P00711425 P00711426, US 00122929 (fragments)).

= Gynoxys pendula var. sinuata Cuatrec., Trab. Mus. Nac. Ci. Nat., Ser. Bot. 29: 38. 1935. – Syntypes: Colombia. Tolima, Andes, Cordillera Central, vert. merid. monte Tolima, loc. dict. Las Mesetas, 3600 m, 13 May 1932, J. Cuatrecasas 2851 (MA: MA240997 MA240997–2 (fragments)).

Gynoxys regis H.Rob. & Cuatrec., Phytologia 56: 370(–371), f. 1984 ≡ Paragynoxys regis (H.Rob. & Cuatrec.) H.Rob. & Cuatrec., Novon 2(4): 415. 1992. – Holotype: Ecuador. Azuday, 30 km S of cumbé on the road to Saraguro at an elevation of 9800 ft., 26 Jan 1979, R. M. King & F. Almeda 7804 (US 00122934; isotypes: K: K000497542, QCA: QCA17841).

Gynoxys soukupii Cuatrec., Bull. Soc. Bot. France 101: 245. 1954. – Holotype: Perú. Amazonas, Chachapoyas, cerro Puma Urco, Jun 1952, Soukup 4072 (F: V0076774F; isotype: US 00122939).

Gynoxys subhirsuta Cuatrec., Notas Fl. Colombia 6: 35. 1944. – Holotype: Colombia. Santander, Cordillera Oriental, Páramo de Tamá, alrededores ed la Cueva, 3000–3200 m, 28 Oct 1941, J. Cuatrecasas et al. 12714 (COL: COL000005229; isotypes: BC: BC634998, F: V0076778F, GH: GH00008611, U: U0001284, US 00122942).

Gynoxys aquifolia Cuatrec ≡ Scrobicaria aquifolia (Cuatrec.) B.Nord.

Gynoxys auriculata Turcz. = Aetheolaena patens (Kunth) B.Nord.

Gynoxys berlandieri DC. = Pseudogynoxys chenopodioides (Kunth) Cabrera

Gynoxys cordifolia Cass. ≡ Pseudogynoxys cordifolia (Cass.) Cabrera

Gynoxys cummingii Benth. ≡ Pseudogynoxys chenopodioides var. cummingii (Benth.) B.L.Turner

Gynoxys discolor Rusby = Pentacalia marinii (Cabrera) Cuatrec.

Gynoxys fragrans Hook. ≡ Pseudogynoxys fragrans (Hook.) H.Rob. & Cuatrec.

Gynoxys haenkei DC. ≡ Pseudogynoxys haenkei (DC.) Cabrera

Gynoxys heterophylla Turcz. ≡ Aetheolaena heterophylla (Turcz.) B.Nord.

Gynoxys ilicifolia (L.f.) Wedd. = Scrobicaria ilicifolia (L.f.) B.Nord.

Gynoxys incana Less. = Jacmaia incana (Sw.) B.Nord.

Gynoxys laciniata Less. = Odontocline laciniata (Sw.) B.Nord.

Gynoxys lucida Less. = Dendrophorbium lucidum (Sw.) C.Jeffrey

Gynoxys oerstedii Benth. ≡ Pseudogynoxys oerstedii (Benth.) Cuatrec.

Gynoxys poeppigii DC. ≡ Pseudogynoxys poeppigii (DC.) H.Rob. & Cuatrec.

Gynoxys prenanthifolia Turcz. = Aetheolaena patens (Kunth) B.Nord.

Gynoxys scabra Benth. ≡ Pseudogynoxys scabra (Benth.) Cuatrec.

Gynoxys sinclairii Benth. = Pseudogynoxys sonchoides (Kunth) Cuatrec.

For the following names in the Global Compositae Database, no publication is given:

Aequatorium castillense B.Nord.

Aequatorium stellatopilosum Cuatrec.

Gynoxys auriculata Sch.Bip

Gynoxys cumingii Sch.Bip.

Gynoxys glabrata Less.

Gynoxys incana (Sw.) Griseb.

Gynoxys lanceolata Weddel

Gynoxys napoensis H.Rob.

Gynoxys perbracteata Cuatrec.

Gynoxys unduaviana Cuatrec.

Gynoxys verrucosa var. magna Cuatrec.

The authors have declared that no competing interests exist.

No ethical statement was reported.

Funding for fieldwork was provided to BE by the Julia Krieg Forschungsfonds of the BGBM in the context of the collaboration with the Herbario Nacional de Bolivia and to NK by the Friends of the Botanic Garden Berlin.

Conceptualization: TB, NK. Data curation: BE. Funding acquisition: NK, TB. Investigation: BE. Methodology: NK. Supervision: NK, TB. Writing - original draft: BE. Writing - review and editing: BE, TB, NK.

Belen Escobari https://orcid.org/0000-0002-4588-3180

Tomas Borsch https://orcid.org/0000-0002-5724-2786

Norbert Kilian https://orcid.org/0000-0003-0890-4373

All of the data that support the findings of this study are available in the main text or Supplementary Information.