taxonID	type	description	language	source
181BB66C65150D569677FA9700A8FEF3.taxon	description	— The neotropical Luxemburgieae consists of two genera, Luxemburgia and Philacra Dwyer. The latter was included in the original sampling for the present study but did not pass the final filtering. However, there is strong support for this clade from morphology (Amaral, 1991; Amaral & Bittrich, 2014) and molecular data (Schneider & al., 2014). Both genera are characterized by flowers that are obliquely zygomorphic already in bud. Their stamens surround the ovary only adaxially, and the filaments are basally or completely fused; staminodes are absent (Amaral & Bittrich, 2014). Luxemburgia is a Brazilian endemic and contains 20 species (Feres, 2001, 2010). The only comprehensive molecular phylogenetic study to date (Feres, 2001) was based on the internal transcribed spacer region only and did not provide any phylogenetic resolution. In the same study, a morphological cladistic analysis retrieved two main clades, separating species based on the presence or absence of a petiole, a character that was already used by Beauverd (1915) for his infrageneric classification. However, the poor phylogenetic resolution in Luxemburgia in the present study precludes the evaluation of this phylogenetic hypothesis. The poor resolution, even with the increased molecular data of this study, is most likely the result of a recent and rapid diversification, a scenario that is supported by divergence time estimation (Schneider & al., 2017).	en	Schneider, Julio V., Jungcurt, Tanja, Cardoso, Domingos, Amorim, André Márcio, Töpel, Mats, Andermann, Tobias, Poncy, Odile, Berberich, Thomas, Zizka, Georg (2021): Phylogenomics of the tropical plant family Ochnaceae using targeted enrichment of nuclear genes and 250 + taxa. TAXON 70 (1): 48-71, DOI: 10.1002/tax.12421, URL: http://dx.doi.org/10.1002/tax.12421
181BB66C65150D5595C1F8B6064DFAF3.taxon	description	— Since APG III (2009), Ochnaceae comprises Ochnaceae s. str., the monotypic Medusagynaceae, endemic to the Seychelles archipelago in the Indian Ocean, and the neotropical Quiinaceae, now recognized at the rank of subfamily (Schneider & al., 2014). The three subfamilies diverged around the CretaceousPalaeogene boundary (Schneider & Zizka, 2017; Schneider & al., 2017). The relationships among them have remained unclear, with different inferences across earlier studies (Fay & al., 1997; Savolainen & al., 2000; Davis & Chase, 2004). Recent more data-rich studies (Xi & al., 2012; Schneider & al., 2014) retrieved Medusagynoideae as sister to Quiinoideae, both sister to Ochnoideae, but with weak to moderate support. This pattern is also observed in the present study, and some uncertainty remains due to the moderate support in the CC analysis. Such clades recalcitrant to phylogenetic resolution may be explained by model misspecification (Eytan & al., 2015), but are more likely due to rapid divergence combined with long branches (Rokas & Carroll, 2006; King & Rokas, 2017) as observed in the split of the subfamilies of Ochnaceae. Quiinoideae were recovered with strong support, with Froesia Pires being sister to a clade of Lacunaria Ducke and Touroulia Aubl. The only missing genus of that subfamily, Quiina Aubl., was originally included but was removed during the filtering process because of too many missing data. Nevertheless, the relationships among the Quiinoideae genera were already well resolved in previous studies, with Quiina being sister to the clade of Lacunaria and Touroulia (Schneider & al., 2014; Schneider & Zizka, 2017). The position of the African Testulea – the only member of tribe Testuleeae – as sister to the rest of Ochnoideae was already recovered with maximum support in Schneider & al. (2014). The morphology-based hypothesis that Testulea is sister to a clade of Cespedesia, Godoya, Krukoviella and Rhytidanthera (Amaral, 1991) is thus rejected. Testulea strongly differs in morphology from the rest of Ochnoideae by having tetramerous flowers with a single bracteole, a single fertile stamen and staminodes fused into a column for two-thirds of their length (Schneider & al., 2014).	en	Schneider, Julio V., Jungcurt, Tanja, Cardoso, Domingos, Amorim, André Márcio, Töpel, Mats, Andermann, Tobias, Poncy, Odile, Berberich, Thomas, Zizka, Georg (2021): Phylogenomics of the tropical plant family Ochnaceae using targeted enrichment of nuclear genes and 250 + taxa. TAXON 70 (1): 48-71, DOI: 10.1002/tax.12421, URL: http://dx.doi.org/10.1002/tax.12421
181BB66C65160D579591FE9703A2FC13.taxon	description	— Sauvagesieae is a blend of neotropical and palaeotropical taxa and was recovered as sister to Ochneae. This sister-group relationship received maximum support in the FAM analysis but much weaker support in the SLT analysis, which may be due to the relatively high amount of missing data of the two representative species of Ochneae in the latter (see Methods). In Schneider & al. (2014), the neotropical Blastemanthus came out as sister to the remaining Sauvagesieae. A moderately supported clade united the neotropical genera Godoya, Krukoviella, Cespedesia and Rhytidanthera, and these were sister to a clade of the African Fleurydora and a clade of the remaining genera of that tribe. In the present study, Blastemanthus is sister to and forms a strongly supported clade together with Godoya, Krukoviella, Cespedesia, Rhytidanthera and Fleurydora. The African Fleurydora is sister to the remainder, which are all neotropical, with Rhytidanthera retrieved as sister to a poorly resolved clade of Godoya, Krukoviella and Cespedesia. This confirms the findings of a recent study based on five DNA regions (Reinales & Parra-O., 2020). There is also support from morphology for this clade: Godoya, Krukoviella, Cespedesia and Rhytidanthera are all united, among others, by the same carpel and ovule numbers, the coriaceous capsules with alate seeds and a peculiar leaf venation with scalariform net-like third-order veins (Amaral, 1991; Schneider & al., 2014; Schneider & al., 2017). In addition to its position as sister to the remaining three genera, Rhytidanthera also exhibits an odd morphology. It is the only genus with pinnately compound leaves in the subfamily. All four genera also share the presence of colleters (i. e., glandular hairs) on stipules, bracts and sepals (lost on sepals in Cespedesia and Krukoviella; Reinales & Parra-O., 2020). Flower zygomorphy only at anthesis (in contrast to its presence already in bud as in Luxemburgieae) unites this clade with Blastemanthus and Fleurydora, but this character was identified as plesiomorphic (Schneider & al., 2014). The sister clade of these Sauvagesieae genera comprises a clade of the neotropical Poecilandra and Wallacea, which in turn is sister to the remaining Sauvagesieae, congruent with the findings of Schneider & al. (2014) and the morphology-based cladistic analysis of Amaral (1991). Both genera share retuse or emarginate leaves, very dense and closely parallel secondary veins, many leaf traces, and anthers covered by wax crystals (Amaral, 1991; Schneider & al., 2017). The remaining Sauvagesieae fell into two main clades, an Asian clade and a neotropical clade. The Asian clade comprises Schuurmansia and Schuurmansiella as sister to Neckia, Euthemis, and the two genera Indovethia and Indosinia, which were included for the first time in a molecular phylogenetic study. This relationship is in contrast to Schneider & al. (2014), in which Neckia was sister to the neotropical and Asian clades, albeit support was weak to moderate in the Asian clades in that study. The clade of Schuurmansia and Schuurmansiella was found in an earlier morphology-based study, supported by the shared presence of a papillose seed surface (Amaral, 1991). The position of Indosinia in a clade together with Euthemis and Schuurmansia is also supported by morphology because all three share a similar exine structure (Amaral, 1991). Hence, the clade of Indosinia and Adenarake retrieved in a morphology-based cladistic analysis and predicated on the shared presence of an androgynophore and valvate sepals was obviously the result of homoplastic characters as already suspected by Amaral (1991). The neotropical clade includes Tyleria and Sauvagesia with Adenarake nested within the latter. Tyleria includes the formerly separate genus Adenanthe Maguire & al. (A. bicarpellata Maguire & al.; ≡ T. bicarpellata (Maguire & al.) M. C. E. Amaral) in agreement with previous findings from morphological and molecular studies (Amaral, 1991; Schneider & al., 2014). All three genera share a reticulate-striate exine sculpture (Amaral, 1991). In contrast to Schneider & al. (2014), the support for all the relationships along the backbone of this part of Sauvagesieae is consistently high, thus providing strong arguments in favour of the here presented phylogenetic hypothesis. Sauvagesia and allies. — The circumscription of Sauvagesia has changed considerably over time. Early allies were the genera Lavradia Vell. and Leitgebia Eichl. that differed from Sauvagesia in the structure of the external whorl of free staminodes and the internal whorl of fused or free and overlapping petaloid staminodes that form a corona enveloping the reproductive organs (Saint-Hilaire, 1824; Eichler, 1871). Later, Gleason (1931, 1933) added the genera Pentaspatella Gleason and Roraimanthus Gleason (the latter a segregate of Leitgebia) that also differed in minor characters from core Sauvagesia. In subsequent treatments based on evidence from numerical taxonomy, Sastre (1970, 1971, 1973) expanded the concept of Sauvagesia by putting all the allied genera mentioned above in its synonymy. Moreover, he expanded the circumscription by also including the SE Asian genus Neckia, thereby making this genus fully pantropical. This concept of an expanded Sauvagesia was supported by Amaral’ s (1991) morphology-based cladistic analysis, which also added a second SE Asian genus, Indovethia, to the list of synonyms. The most recent expansion of Sauvagesia was the inclusion of the Chinese endemic Sinia because of similarities in flower and seed morphology (Amaral, 2006). This concept was challenged first by the molecular phylogenetic analysis of Schneider & al. (2014), in which Sauvagesia serrata (= Neckia) appeared independent from Sauvagesia. In the present study, we also included Indovethia (i. e., Sauvagesia calophylla (Boerl.) M. C. E. Amaral), which also came out as independent from Sauvagesia. Indovethia is sister to Euthemis, and Neckia is sister to Indosinia, although both relationships received weak to moderate support only. Thus, at least some of the morphological traits that were thought to unite all the above genera under an expanded concept of Sauvagesia are most likely homoplastic, as observed for some characters of diagnostic value in the ancestral state reconstructions in Schneider & al. (2014). As a consequence of the present study, Indovethia and Neckia have to be kept as own genera. These are important findings because it does not only require a change in the circumscription of Sauvagesia, but it also has immediate consequences for the interpretation of the historical biogeography of Ochnaceae. The position of the third Asian representative of Sauvagesia (i. e., Sinia) could not be clarified here because of the lack of material for our study. For the first time, two distinct clades were retrieved for core Sauvagesia here. One comprises the type, S. erecta L., and allies, including representatives of the former separate genera Pentaspatella, Roraimanthus and Leitgebia (clade B). This clade is sister to Adenarake. Clade A comprises some species of the formerly independent Lavradia (S. capillaris (A. St. - Hil.) Sastre, S. glandulosa (A. St. - Hil.) Sastre). Sastre’ s (1978, 1981) infrageneric classification of Sauvagesia might provide a solution to the issue of the two clades with a nested Adenarake. He subdivided Sauvagesia into S. sect. Sauvagesia and sect. Imthurnianae Dwyer ex Sastre, and the first into two subsections with the newly erected S. subsect. Vellozianae Sastre containing species formerly assigned to Lavradia. Thus, Adenarake might be kept as a distinct genus if clade A is established as a different genus (e. g., as Lavradia). However, without a comprehensive taxonomic treatment of the genus at hand, it is beyond the scope of the present study to judge if this is a sound solution to the problem or whether a broad concept is to be preferred, including both clades plus Adenarake in Sauvagesia.	en	Schneider, Julio V., Jungcurt, Tanja, Cardoso, Domingos, Amorim, André Márcio, Töpel, Mats, Andermann, Tobias, Poncy, Odile, Berberich, Thomas, Zizka, Georg (2021): Phylogenomics of the tropical plant family Ochnaceae using targeted enrichment of nuclear genes and 250 + taxa. TAXON 70 (1): 48-71, DOI: 10.1002/tax.12421, URL: http://dx.doi.org/10.1002/tax.12421
181BB66C65170D4995C1FC77004FFC93.taxon	description	— Ochneae as defined in Schneider & al. (2014) contains the African Lophirinae (with Lophira as the sole genus) as sister to a clade of the neotropical Elvasiinae (with Elvasia and Perissocarpa) and the pantropical Ochninae (Brackenridgea, Campylospermum, Idertia, Ochna, Ouratea, Rhabdophyllum). This classification was supported in the present study, too, with all clades receiving strong support, including the sister relationship of Elvasia and Perissocarpa, which was analyzed here for the first time using molecular data. Ochninae is the most species-rich clade of Ochnaceae, containing approximately two-thirds of the family’ s species. The associated radiations most likely benefitted from the emergence of the savanna biome in the Old and New World as inferred from the time frame of the divergence events and the actual species distributions. The split into the six currently accepted genera occurred over a very short period of about a maximum of 5 – 10 million years during the Miocene (Schneider & al., 2017), which is certainly also one reason for the hitherto unclear relationships among them. In the present study, all backbone nodes of Ochninae were recovered with strong support, thus resolving its relationships for the first time. The first split divides Ochninae into the neotropical Ouratea and the remaining genera, which are all palaeotropical. Perhaps most remarkable within the palaeotropical clade is the polyphyly of Campylospermum, which was already revealed in Bissiengou (2014), but which is here recovered with strong support. Clade A of Campylospermum contains all Central and West African species of this genus (except C. elongatum (Oliv.) Tiegh.) and forms a clade with the African Rhabdophyllum. Clade B of Campylospermum contains the Malagasy (here, only C. obtusifolium Tiegh. included) and East African species (plus C. elongatum) and is sister to Brackenridgea, both being sister to the West to Central African Idertia. It is important to mention that both clades share their own set of characters: Clade A usually has terminal inflorescences, and the embryos are either accumbent or incumbent and similar or dissimilar in size, whereas clade B shares usually axillary inflorescences, accumbent embryos that are similar in size, and a distinct reticulate tertiary leaf venation (Bissiengou, 2014). Sosef (2008) also noted that the leaf venation of Campylospermum lecomtei (Tiegh.) Farron and C. paucinervatum Sosef, both belonging to our clade A, resembles that of Rhabdophyllum, thus providing further support from morphology for the here inferred sister-group relationship between both groups. However, we are awaiting a modern revision of the Malagasy species (Bissiengou, 2014 only treated the continental species) before making the necessary nomenclatural changes. Brackenridgea originally comprised only species from SE Asia, Australia and Oceania (Van Tieghem, 1902). The morphologically similar genus Pleuroridgea Tiegh. was erected by Van Tieghem (1902) to unite species that share embryos with laterally disposed cotyledons and lateral, deeply divided caducous stipules, initially all from continental Africa. This concept was followed by Perrier de la Bathie (1941) but with an expanded Pleuroridgea to include new Malagasy species. Subsequent authors (Robson, 1963; Du Toit & Obermeyer, 1976; Verdcourt, 2005; Callmander & al., 2010), however, merged both genera under a broad concept of Brackenridgea, judging the morphological differences too small for maintaining them separate. However, it is remarkable that in our phylogeny the former Brackenridgea s. str. and Pleuroridgea form well-supported geographically distinct clades within a monophyletic Brackenridgea. One clade unites the Asian-OceanianAustralian species B. palustris Bartell., B. foxworthii Furtado (= B. palustris Bartell.), B. nitida A. Gray (= B. nitida subsp. australiana (F. Muell.) P. O. Karis) and B. forbesii Tiegh., the second one unites the African-Malagasy species B. madecassa (H. Perrier) Callm., B. arenaria (De Wild. & T. Durand) N. Robson and B. zanguebarica Oliv., the latter itself divided into a Malagasy and a continental (African) clade. Thus, the disjunct African-Asian distribution, which is shared with Ochna and Campylospermum, most likely resulted from a single crosscontinental dispersal event, perhaps involving Madagascar as an intermediate step. Here, we present the by far most comprehensive phylogenetic framework for Ochna, comprising about 50 % of its species. While a modern taxonomic revision of this genus is still lacking, this framework serves as important baseline data for future infrageneric classification. Earlier attempts for an infrageneric classification included the one by Van Tieghem (1902). He segregated 15 genera using characters such as anther dehiscence (poricidal versus longicidal), carpel number or embryo morphology (iso- versus heterocotyledonous). More recent regional treatments on African (Robson, 1963; Du Toit & Obermeyer, 1976; Verdcourt, 2005; Callmander & Phillipson, 2012) or Asian (Kanis, 1968) species of Ochna did not follow Van Tieghem’ s narrow concept. In the present study, there is a remarkable congruence of our phylogeny with the infrageneric classification of Robson (1963), who divided the genus into three sections based on carpel shape and anther dehiscence (see also Verdcourt, 2005). Our clade A contains species belonging to Robson’ s Ochna sect. Ochna, which is characterized by bi-porous anthers. Clade B contains species of his O. sect. Schizanthera in which anthers open by longitudinal slits. In contrast, his species-poor O. sect. Renicarpus, which differs by having reniform drupelets that are attached near the centre of the long side of the drupelet, appears to be polyphyletic according to the placement of Ochna arborea Burch. ex DC. and O. pulchra Hook. However, a more comprehensive taxon sampling of this section is required for a sound evaluation of its status. Besides the two large clades, there are two smaller clades. One contains the morphologically close O. polycarpa Baker and O. louvelii (H. Perrier) Callm. & Phillipson, which are from Madagascar (Callmander & Phillipson, 2012). The other one is sister to the rest of Ochna and comprises O. andravinensis Baill., O. pulchra Hook. and O. latisepala (Tiegh.) Bamps. How these smaller clades are integrated into an infrageneric classification and how species of O. sect. Reniformis are accommodated, is, however, beyond the scope of the present study and needs to be assessed in the framework of a modern taxonomic revision of this genus. From an ecological perspective remarkable is the observation that several of the species for which a fire-adapted geoxylic growth form has been reported (which are elements of the so-called underground forests; see White, 1976) were positioned in a subclade of clade C. The geoxylic species include, for example, O. leptoclada Oliv., O. katangensis De Wild., O. confusa Burtt Davy & Greenway and O. pygmaea Hiern. From a biogeographical perspective noteworthy are our findings that the Asian species (O. integerrima (Lour.) Merr., O. obtusata DC.) included in this study form a clade that is nested in clade D and that the Malagasy species (e. g., O. brachypoda Baill., O. macrantha Baker, O. polycarpa, O. andravinensis) are spread across three major clades of Ochna. The largest genus of Ochnaceae is Ouratea, which is among the larger woody genera of the New World with about 200 (Amaral & Bittrich, 2014) to 310 accepted species (Schneider, unpub. data), comparable in size to other well-known radiations in tropical plant families such as Inga Mill. (ca. 300 spp., Fabaceae; Richardson & al., 2001), Ocotea Aubl. (ca. 300 spp., Lauraceae; Madriñán, 2004), Clusia L. (ca. 300 spp., Clusiaceae; Gustafsson & al., 2007), or Guatteria Ruiz & Pav. (ca. 265 spp., Annonaceae; Erkens & al., 2007). The lack of a comprehensive taxonomic revision of the genus hampers the interpretation of the here presented phylogeny. However, different infrageneric classification systems are available. First attempts for an infrageneric classification of this genus were largely based on unreliable characters such as leaf indumentum and texture (Erhard, 1849). Later, Engler (1876) erected the two series, Ouratea ser. Cardiocarpae and ser. Oocarpae, using fruit characters for the then known 76 species. The first differs by its obcordate drupelets from the second, which is characterized by mostly ovoid to obovoid drupelets. With Van Tieghem’ s (1902) classification, the number of subdivisions increased drastically. He distinguished 22 genera for his subtribe Orthospermeae (which corresponds to modern Ouratea; see Sastre & Offroy, 2016) using differences in inflorescence type and insertion, indumentum, number of styles, embryo shape or the persistence of stipules, among others. Subsequent authors disregarded Van Tieghem’ s system (e. g., Gilg, 1925). Perhaps the more satisfying classification of Ouratea was introduced by Sastre (1988). He subdivided Ouratea into the six sections O. sect. Cardiocarpae (Engl.) Sastre, sect. Kaieteuria (Dwyer) Sastre, sect. Polyouratea (Tiegh.) Sastre, sect. Ouratea, sect. Ouratella (Tiegh.) Sastre and sect. Persistens Sastre. This classification was superseded by his more recent one (Sastre, 1995), in which O. sect. Caducae Sastre was newly erected, whereas sect. Persistens was placed in the synonymy of sect. Ouratea. Characters of diagnostic value for the different sections are, for example, the possession of 2 – 4 fused sepals in O. sect. Kaieteuria, fruits with the drupelets borne horizontally in sect. Cardiocarpae. Other important characters are the number of carpels (e. g., 6 – 10 in sect. Polyouratea), fruits with persistent sepals (sect. Ouratea) and the position of the inflorescence (terminal versus axillary, the first characteristic of sect. Caducae). Here, based on our phylogenetic analysis, including roughly half the species reported for Ouratea, we tentatively defined five major clades for the ease of discussion, all except clade D with maximum support. Mapping Sastre’ s (1995) sections for species for which we took the relevant information from his publications (Sastre, 1988, 1995, 2001, 2007) revealed that all sections were polyphyletic (suppl. Fig. S 5). In particular, Ouratea sect. Kaieteuria and sect. Ouratella were spread across most of the clades. Topological artefacts as resulting from methodological issues and sometimes associated with either the concatenation or the coalescent approach can be ruled out as an explanation for the lack of congruence between our phylogeny and the sectional classification because all five major clades are recovered identically with both methods of phylogenetic inference. We also evaluated Van Tieghem’ s (1902) classification but did not find any congruent pattern. His genus Trichouratea Tiegh. is the only one which coincides in part with a subclade of our clade E. However, it is also polyphyletic and intermingled with species from two other genera he had defined. Therefore, many of the characters used for the infrageneric classification are supposedly homoplastic. This is most likely due to the explosive radiation during the early history of Ouratea as inferred from the short branches along its backbone, and which also might explain why previous morphology-based infrageneric classifications in such speciose angiosperm genera often fail in defining monophyletic groups (e. g., Simon & al., 2011; Goldenberg & al., 2018; Moonlight & al., 2018). Only at the level of smaller subclades, we discern some congruence with character combinations that defined Sastre’ s sections. For example, in clade D, we found a subclade uniting O. guildingii (Planch.) Urb., O. grosourdyi (Tiegh.) Steyerm., O. mexicana (Bonpl.) Engl. and O. pseudoguildingii Sastre, which all share the set of characters of Ouratea sect. Ouratella. Another subclade of clade D unites species belonging to O. sect. Kaieteuria (O. steyermarkii Sastre, O. thyrsoidea Engl., O. clarkia Sastre, O. arbobrevicalyx Sastre), and there is also a clade uniting the Caribbean species O. illicifolia (DC.) Baill., O. agrophylla (Tiegh.) Urb., O. elliptica (A. Rich.) M. Gómez, O. lenticellosa Urb., O. laurifolia (Sw.) Engl. and O. jamaicensis (Planch.) Urb., some of which have been included in Van Tieghem’ s (1902) Camptouratea. In clade E, we observe a subclade uniting species of Ouratea sect. Polyouratea (O. scottii Sastre, O. discophora Ducke, O. decagyna Maguire). Although we may find consistent character combinations for smaller species groups, it might be difficult to find such combinations for the major clades in view of the short branches during early divergence events creating a scenario of incomplete lineage sorting at deeper time (Xu & Yang, 2016) and many clades with not yet well-fixed traits. However, morphological characters of diagnostic value that describe the here retrieved major clades will have to be searched and evaluated in an urgently needed taxonomic revision of that genus.	en	Schneider, Julio V., Jungcurt, Tanja, Cardoso, Domingos, Amorim, André Márcio, Töpel, Mats, Andermann, Tobias, Poncy, Odile, Berberich, Thomas, Zizka, Georg (2021): Phylogenomics of the tropical plant family Ochnaceae using targeted enrichment of nuclear genes and 250 + taxa. TAXON 70 (1): 48-71, DOI: 10.1002/tax.12421, URL: http://dx.doi.org/10.1002/tax.12421
